The Liverworts and Hornworts of Colombia and Ecuador (Memoirs of The New York Botanical Garden, 121) 3030494497, 9783030494490

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Memoirs of the New York Botanical Garden 121

S. Robbert Gradstein

The Liverworts and Hornworts of Colombia and Ecuador

Memoirs of the New York Botanical Garden Volume 121 Series Editors William R. Buck Institute of Systematic Botany New York Botanical Garden Bronx, NY, USA Lawrence M. Kelly NYBG Press New York Botanical Garden Bronx, NY, USA

Launched in 1900 by The New York Botanical Garden (NYBG), and with 120 volumes published as of 2019, Memoirs has traditionally focused on taxonomy and floristics, with an emphasis on plants of the Western Hemisphere. Most titles comprised a single volume, but quite a few were multi-volume titles and constitute some of the most iconic and substantial monographic publications of NYBG. In recent years, the Memoirs series has expanded to encompass field guides and botanical history as well. All in-print volumes of Memoirs through volume 120 are available exclusively through NYBG Shop: https://nybgshop.org/nybg-press/. Volumes 121 and going forward are available exclusively through Springer. More information about this series at http://www.springer.com/series/16366

S. Robbert Gradstein

The Liverworts and Hornworts of Colombia and Ecuador Illustrations Anna Luiza Ilkiu-Borges Manon Delval Laura Campos-Salazar Contributions Barbara Crandall-Stotler Tamás Pócs Catherine Reeb Jaime Uribe-Meléndez

S. Robbert Gradstein Muséum National d’Histoire Naturelle Paris, France University of Göttingen Göttingen, Germany

ISSN 0077-8931     ISSN 2662-2858 (electronic) Memoirs of the New York Botanical Garden ISBN 978-3-030-49449-0    ISBN 978-3-030-49450-6 (eBook) https://doi.org/10.1007/978-3-030-49450-6 © Springer Nature Switzerland AG 2021 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Population of Riccardia verticillata. Photograph by Alvaro J. Pérez

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Contents

Author Biography�����������������������������������������������������������������������������������������    xi Preface�����������������������������������������������������������������������������������������������������������   xiii Acknowledgments������������������������������������������������������������������������������������������   xvii Summary�������������������������������������������������������������������������������������������������������   xix Resumen���������������������������������������������������������������������������������������������������������    xxi Abbreviations �����������������������������������������������������������������������������������������������  xxiii Introduction����������������������������������������������������������������������������������������������������    1 Brief History of Exploration����������������������������������������������������������������������������     2 Underexplored Areas���������������������������������������������������������������������������������������     5 Format of the Book������������������������������������������������������������������������������������������     6  iversity and Endemism ������������������������������������������������������������������������������    9 D Phytogeographic Elements������������������������������������������������������������������������������    11 Regional Diversity ������������������������������������������������������������������������������������������    16 Classification��������������������������������������������������������������������������������������������������   27 Marchantiophyta����������������������������������������������������������������������������������������������    27 Haplomitriopsida������������������������������������������������������������������������������������������   27 Marchantiopsida������������������������������������������������������������������������������������������    28 Jungermanniopsida��������������������������������������������������������������������������������������    28 Anthocerotophyta��������������������������������������������������������������������������������������������    33  eys and Descriptions ����������������������������������������������������������������������������������    35 K Introductory Key����������������������������������������������������������������������������������������������    35 Marchantiophyta (liverworts) ��������������������������������������������������������������������������    36 Haplomitriaceae ����������������������������������������������������������������������������������������������    37 Aytoniaceae������������������������������������������������������������������������������������������������������    41

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Cleveaceae ������������������������������������������������������������������������������������������������������    45 Corsiniaceae����������������������������������������������������������������������������������������������������    47 Cyathodiaceae��������������������������������������������������������������������������������������������������    47 Dumortieraceae������������������������������������������������������������������������������������������������    50 Lunulariaceae��������������������������������������������������������������������������������������������������    51 Marchantiaceae������������������������������������������������������������������������������������������������    52 Monocleaceae��������������������������������������������������������������������������������������������������    57 Ricciaceae��������������������������������������������������������������������������������������������������������    58 Targioniaceae ��������������������������������������������������������������������������������������������������    67 Fossombroniaceae��������������������������������������������������������������������������������������������    70 Pallaviciniaceae������������������������������������������������������������������������������������������������    77 Pelliaceae ��������������������������������������������������������������������������������������������������������    85 Aneuraceae������������������������������������������������������������������������������������������������������    87 Metzgeriaceae��������������������������������������������������������������������������������������������������   105 Pleuroziaceae ��������������������������������������������������������������������������������������������������   122   Order Jungermanniales��������������������������������������������������������������������������������   124 Acrobolbaceae��������������������������������������������������������������������������������������������������   130 Adelanthaceae��������������������������������������������������������������������������������������������������   136 Antheliaceae����������������������������������������������������������������������������������������������������   142 Arnelliaceae ����������������������������������������������������������������������������������������������������   143 Balantiopsidaceae��������������������������������������������������������������������������������������������   145 Calypogeiaceae������������������������������������������������������������������������������������������������   152 Cephaloziaceae������������������������������������������������������������������������������������������������   160 Cephaloziellaceae��������������������������������������������������������������������������������������������   172 Gymnomitriaceae��������������������������������������������������������������������������������������������   179 Herbertaceae����������������������������������������������������������������������������������������������������   183 Jamesoniellaceae����������������������������������������������������������������������������������������������   191 Jungermanniaceae��������������������������������������������������������������������������������������������   201 Lepicoleaceae��������������������������������������������������������������������������������������������������   203 Lepidoziaceae��������������������������������������������������������������������������������������������������   204 Lophocoleaceae������������������������������������������������������������������������������������������������   243 Plagiochilaceae������������������������������������������������������������������������������������������������   266 Pseudolepicoleaceae����������������������������������������������������������������������������������������   320 Scapaniaceae����������������������������������������������������������������������������������������������������   324 Solenostomataceae������������������������������������������������������������������������������������������   338 Stephaniellaceae����������������������������������������������������������������������������������������������   342 Trichocoleaceae ����������������������������������������������������������������������������������������������   346   Order Porellales ������������������������������������������������������������������������������������������   349 Frullaniaceae����������������������������������������������������������������������������������������������������   350 Jubulaceae��������������������������������������������������������������������������������������������������������   383 Lejeuneaceae����������������������������������������������������������������������������������������������������   384

Contents

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Porellaceae ������������������������������������������������������������������������������������������������������   614 Radulaceae ������������������������������������������������������������������������������������������������������   618 Anthocerotophyta (hornworts)������������������������������������������������������������������������   640 Glossary������������������������������������������������������������������������������������������������������������  657 References ��������������������������������������������������������������������������������������������������������  673 Index of Scientific Names��������������������������������������������������������������������������������  699

Author Biography

S. Robbert Gradstein  is a Dutch Botanist and a retired Professor of Botany at the University of Göttingen and a Research Associate of the Natural History Museum of Paris. His research interests include the systematics and ecology of bryophytes and epiphytes, especially those of tropical America and Asia. He is a leading authority on tropical liverworts and has published almost 500 research papers and 15 books. He is the author of the Guide to the Bryophytes of Tropical America (2001) together with Steven P. Churchill (Missouri Bot. Garden) and Noris Salazar Allen (Smithsonian Tropical Research Institute) and the Catalogue of the Plants and Lichens of Colombia (2016) together with Rodrigo Bernal and Marcela Celis (Universidad Nacional de Colombia). Robbert Gradstein is a member of the Akademie der Wissenschaften zu Göttingen and the Academia Colombiana de Ciencias Exactas, Físicas y Naturales, and a Founder and former President of the International Association of Bryologists. He has been Editor of TAXON, Regnum Vegetabile, Flora Neotropica, and Bryophytorum Bibliotheca and Deputy Director of the Organization for Flora Neotropica (UNESCO) for 20 years. He has been awarded the Jesse M. Greenman Award of the Missouri Botanical Garden for best dissertation in systematics, the Prix Augustin-Pyramus de Candolle of the Societé de Physique et d'Histoire Naturelle de Genève for monographic work on liverworts, the first Fundraising Prize of the University of Göttingen for his directorship of the Botanical Garden of Göttingen, and the Hedwig Medal of the International Association of Bryologists for lifelong achievement in bryology.

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Preface

Work toward this book spans almost half a century. I started work on liverworts of Colombia in 1970–71 during my Ph.D. study at the University of Cincinnati. My advisor, Margaret H. Fulford, encouraged me to do some general identification of neotropical hepatics to familiarize myself with the flora. Supported by my friend Antoine Cleef who was studying vegetation of the páramos of Colombia for his doctorate and had offered to collect liverworts, I decided to pay special attention to oil bodies which at the time were virtually unknown in neotropical liverworts. Although Dr. Fulford advised me to abandon this idea as she believed it would be impossible to obtain fresh material necessary for this study, the project worked out well thanks to the help of Antoine who regularly sent me fresh samples via airmail. Out of this came my first paper on liverworts of Colombia (1977), focusing on oil bodies and habitat preferences and co-authored by Margaret Fulford who helped with identifications and Antoine Cleef who provided ecological descriptions. My involvement in the study of Colombian hepatics intensified after my return to Utrecht. Antoine Cleef had brought back thousands of collections from the Colombian páramos, including many liverworts, and numerous collections were also made in the framework of ECOANDES, a large-scale ecological project in the Colombian Andes under the inspiring leadership of Thomas van der Hammen of the University of Amsterdam. Many students and specialists, among them Dana Griffin III from the University of Florida, participated in the herculean task of working up these collections, results of which appeared in the series Studies on Colombian Cryptogams initiated by my supervisor Peter A. Florschütz. Wim Hekking, a specialist of neotropical Violaceae, helped in compiling the first edition of the Annotated Catalogue of the Liverworts and Hornworts of Colombia (1979), and through this work I gained an impression of the country’s flora and how little was known about its hepatics. In Colombia, a research program dedicated to the cryptogams of the country was set up at the Instituto de Ciencias Naturales under the leadership of Jaime Aguirre, who had come to Utrecht to study lichens and liverworts with Harrie Sipman and me for his M.Sc. degree. Early on, I was also involved in the study of hepatics of Ecuador. Having read Voyage of the Beagle in my school years, I had become eager to visit the Galápagos xiii

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Islands in Darwin's footsteps. In 1973, during the first congress of the newly establish International Association of Bryologists, I had a chance to meet William A.  Weber from the University of Colorado who had published a bryophyte catalogue of the Galápagos archipelago and knew the islands well. Together we decided to visit the islands and undertake a Galápagos Bryophyte Flora. In the spring of 1976, after having obtained my doctorate, Bill and I travelled to Ecuador and collected bryophytes throughout the archipelago. Out of this came a series of papers on the taxonomy of selected liverwort genera and an analysis of the bryogeography of the islands. The projected Flora, however, remained unfinished due to the insufficient taxonomic knowledge of many of the groups, necessitating full-scale revisions. Now, 45 years later, this work has largely been done, at least for liverworts. All my observations on the hepatics of the Galápagos Islands have been incorporated in this book. After my move to the University of Göttingen in 1995, I did little fieldwork in Colombia as the country became increasingly insecure. My last major fieldtrip in the country, together with Jan-Peter Frahm and Jaime Aguirre, was in 1992 to the Chocó, an area phenomenally rich in hepatics and very little explored. The identification of the liverworts collected during this trip took many years and yielded several species new to science. My main occupation with Colombian botany after this trip focused on doing taxonomic revisions and cataloging the flora. This resulted in three monographs, two new editions of the hepatic catalogue prepared in collaboration with Jaime Uribe-M, who had taken charge of liverwort research at the Instituto de Ciencias Naturales, and the Catalogue of the Plants and Lichens of Colombia produced in collaboration with Rodrigo Bernal and Marcela Celis from the same institution. While in Göttingen, fieldwork took me back to Ecuador. The German Research Foundation had initiated a large-scale ecological research program in the south of the country, based at the San Francisco Biological Station, with focus on sustainable use of the tropical montane forest. During the years 2001–2009, I worked there with Ph.D. students on human impact on epiphyte diversity. Numerous liverwort collections were made in the framework of this program and a catalogue of the bryophytes of the reserve was prepared together with Harald Kürschner and Gerhard Parolly from the Free University of Berlin who had studied the bryophyte communities of the reserve. Susana León-Yánez at the Pontifícia Universidad Católica del Ecuador in Quito was my counterpart in this project and our fruitful cooperation resulted in the Catalogue of the Liverworts and Hornworts of Ecuador, the Red List of Endemic Bryophytes of Ecuador, the Guía de briófitas comunes de Los Andes de Quito, and several other publications. I also undertook fieldtrips elsewhere in the country, including exploration of the Río Topo and other tributaries of the Pastaza River together with orchid specialist and conservationist Lou Jost from Baños and in the footsteps of Richard Spruce, who had first explored the area 150 years earlier. This resulted in the rediscovery of the famous Myriocolea irrorata Spruce, considered lost for many years. Another highlight was the rediscovery, during a trip to the coastal lowlands, of Spruceanthus

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theobromae Spruce, a narrowly endemic species of Ecuador that is only known from cacao plantations. In 2009, after the publication of Guide to the Plants of Central French Guiana. Part 4. Liverworts and Hornworts, Bill Buck asked me to write this book. For various reasons it took me a few years to get started. Liverworts and Hornworts of Brazil, written 12 years earlier in cooperation with Denise Pinheiro da Costa, served as a blueprint for the new book project. In spite of all the preparatory work, major hurdles remained as large groups such as Bazzania, Riccardia and parts of Lejeunea and Plagiochila were without appropriate treatment and needed revision. It took me more than 6 years. The manuscript of the book was completed in September 2019. Paris, France Göttingen, Germany

S. Robbert Gradstein

Acknowledgments

Many people helped me in the preparation of this book. First of all I want to thank William R. Buck, Editor of the Memoirs of the New York Botanical Garden, who encouraged me to write this book and spent endless hours in meticulously checking and correcting the manuscript. I express my special gratitude to Anna Luiza Ilkiu-­ Borges (Museu Paraense Emílio Goeldi, Belém) and Manon Delval (Muséum National d'Histoire Naturelle, Paris, UMS 2700) who generously prepared most of the illustrations. I am also grateful to Didier Geffard-Kuriyama and Ludivine Longou from the Muséum National d'Histoire Naturelle (UMS 2700) for help with artwork and to Laura Campos-Salazar (Universidad de La Salle, Bogotá) for contributing some of the Lejeuneaceae plates. I cordially thank the contributors who helped in preparing some of the generic treatments: Barbara Crandall-Stotler (Fossombronia), Catherine Reeb (Riccardia), and Jaime Uribe-Meléndez (Marchesinia). I am very grateful to Anna Luiza Ilkiu-­ Borges and Tamás Pócs who critically read the entire manuscript and made numerous useful comments and corrections. Tamás Pócs also identified the rare Myriocoleopsis tixieri, helped describing new taxa in Drepanolejeunea and Zoopsidella, and contributed to the treatment of Lepidoziaceae. I furthermore thank Juan Carlos Villarreal for checking the hornwort chapter and providing photographs of spores; Tomoyuki Katagiri for checking the Trichocoleaceae text; Cid Bastos Elena Reiner-Drehwald, Alfons Schäfer-Verwimp, and Rui-Liang Zhu for data on selected species of Frullaniaceae and Lejeuneaceae; Juan Carlos Benavides for information on Colura stotleri; Catherine Reeb for help in the identification of Riccia huebeneriana; Frank Müller for the photograph of Phaeomegaceros squamuligerus spores; and Françoise Bouazzat and Didier Geffard-Kuriyama for preparing the hornwort photoplate. I have greatly profited from the rich holdings of the Paris herbarium, where I have a working desk since 2010, for the work on this book. I cordially thank the director of the herbarium, Serge Muller, as well as Sebastian Leblond and Lionel Kevran for supporting my work on the collections. Frequent visits were made to the Göttingen herbarium, where most of my Colombian and Ecuadorian collections are housed, and to the herbarium of Geneva, which holds the largest collection of type xvii

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specimens of tropical liverworts in the world. I am very grateful to Michelle Price in Geneva and Elvira Hörandl and Marc Appelhans in Göttingen for hosting my stays. I have also greatly profited from visits to the herbaria of the Pontifícia Universidad Católica del Ecuador in Quito and the Universidad Nacional de Colombia in Bogotá, and I express my warmest gratitude to Susana León-Yánez and Jaime Uribe-Meléndez for supporting my work in these institutions. I also thank Ángel Benitez-Chavez at the Universidad Técnica Particular de Loja for his collaboration in the study of the Ecuadorian liverwort and hornworts. Finally, I cordially thank the curators of the herbaria cited in the text for making type specimens and other materials available on loan.

Summary

S.  Robbert Gradstein, Muséum National d'Histoire Naturelle—Sorbonne Universités, Institut de Systématique, Évolution, Biodiversité (UMR 7205), 75005 Paris, France; Department of Systematics, Biodiversity and Evolution of Plants (with Herbarium), Albrecht von Haller Institute for Plant Sciences, University of Göttingen, 37073 Göttingen, Germany. The Liverworts and Hornworts of Colombia and Ecuador: Memoirs of the New York Botanical Garden 121. This book provides keys, descriptions, and illustrations for about 850 species of liverworts and hornworts, in 148 genera and 47 families, of Colombia and Ecuador. The largest genera are Lejeunea (66 spp.), Plagiochila (65), Frullania (54), Radula (33), Metzgeria (33), Cololejeunea (32), Cheilolejeunea (30), Bazzania (26), Drepanolejeunea (25), Ceratolejeunea (18), Diplasiolejeunea (18), and Syzygiella (18). Species descriptions include brief morphological characterization and discussion with emphasis on characters for identification, world range, as well as distribution and habitat in Colombia and Ecuador. Classes, orders, families, and genera are also described and the main features for recognition of the genera are briefly discussed. The introduction includes chapters on history of exploration, diversity and endemism, and classification. A glossary, bibliography, and index to scientific names are also provided. Colombia has about 720 species of liverworts and hornworts, and endemism is 3.5%. Ecuador with a surface area four times smaller than that of Colombia has also 720 species while endemism is higher, 6%. Highest endemism is seen on the Galápagos Islands (10%). Almost 95% of the species of Colombia and Ecuador are found in the Andean region and about one hundred species, including three genera (Chaetocolea, Platycaulis, Reinerantha), are endemic to the northern Andes. More than one hundred new synonyms and numerous new lectotypifications are proposed in this book; new names are Acrobolbus setaceus (Steph.) Gradst. comb. nov., Blepharolejeunea saccata subsp. harlingii (S.W.Arnell) Gradst. comb. nov., Drepanolejeunea orthophylla subsp. saxicola Gradst. & Pócs subsp. nov., Frullania

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Summary

intumescens var. sabanetica (Gottsche) Gradst. comb. nov., Frullania rio-­janeirensis var. megalostipa (Spruce) Gradst. comb. nov., Lejeunea killipii (Herzog) Gradst. comb. nov., Lejeunea mojandae Gradst. sp. nov., Lejeunea schultesii Gradst. sp. nov., Lejeunea yasuniensis Gradst. & C.Bastos sp. nov., Marchesinia brachiata var. robusta (Mitt.) Gradst. & Uribe comb. nov., Myriocoleopsis tixieri Pócs & Gradst. nom. nov., Radula bischlerae Gradst. & Ilkiu-Borges sp. nov., and Xylolejeunea pellucidissima (Spruce) Gradst. comb. nov.

Resumen

S. Robbert Gradstein, Muséum National d'Histoire Naturelle – Sorbonne Universités, Institut de Systématique, Évolution, Biodiversité (UMR 7205), 75005 Paris, France; Department of Systematics, Biodiversity and Evolution of Plants (with Herbarium), Albrecht von Haller Institute for Plant Sciences, University of Göttingen, 37073 Göttingen, Germany. The Liverworts and Hornworts of Colombia and Ecuador: Memoirs of the New York Botanical Garden 121. Este libro provee claves, descripciones e ilustraciones para unas 850 especies de hepáticas y antocerotes, en 148 géneros y 47 familias, de Colombia y Ecuador. Los géneros más diversos son Lejeunea (66 spp.), Plagiochila (65), Frullania (54), Radula (33), Metzgeria (33), Cololejeunea (32), Cheilolejeunea (30), Bazzania (26), Drepanolejeunea (25), Ceratolejeunea (18), Diplasiolejeunea (18) and Syzygiella (18). Las descripciones incluyen caracteres morfológicos importantes, rasgos salientes utilizados en la identificación, distribución mundial así como distribución y habitat de las especies en Colombia y Ecuador. También se describen clases, órdenes, familias y géneros. La introducción provee capítulos sobre la historia de exploración, diversidad, endemismo y clasificación. Se incluyen también un glosario de términos técnicos, una bibliografía y un índice de nombres científicos. Colombia tiene unas 720 especies de hepáticas y antocerotes; el endemismo es 3.5%. Ecuador con una superficie casi cuatro veces menor tiene tambien 720 especies mientras el endemismo es 6%. Las Islas Galápagos tienen el endemismo más alto (10%). Casi 95% de las especies de Colombia y Ecuador ocurren en la región Andina y unas cien especies y tres géneros (Chaetocolea, Platycaulis, Reinerantha) son endémicas en los Andes del Norte. Se proponen unos cien nuevos sinónimos; los siguentes nombres son nuevos: Acrobolbus setaceus (Steph.) Gradst. comb. nov., Blepharolejeunea saccata subsp. harlingii (S.W.Arnell) Gradst. comb. nov., Drepanolejeunea orthophylla subsp.

xxi

xxii

Resumen

saxicola Gradst. & Pócs subsp. nov., Frullania intumescens var. sabanetica (Gottsche) Gradst. comb. nov., Frullania rio-janeirensis var. megalostipa (Spruce) Gradst. comb. nov., Lejeunea killipii (Herzog) Gradst. comb. nov., Lejeunea mojandae Gradst. sp. nov., Lejeunea schultesii Gradst. sp. nov., Lejeunea yasuniensis Gradst. & C.Bastos sp. nov., Marchesinia brachiata var. robusta (Mitt.) Gradst. & Uribe comb. nov., Myriocoleopsis tixieri Pócs & Gradst. nom. nov., Radula bischlerae Gradst. & Ilkiu-Borges sp. nov. y Xylolejeunea pellucidissima (Spruce) Gradst. comb. nov.

Abbreviations

Departments of Colombia Ama Amazonas Ant Antioquía Ara Arauca Bol Bolívar Boy Boyacá Cal Caldas Caq Caquetá Cas Casanare Cau Cauca Ces Cesar Cho Chocó Cor Córdoba Cun Cundinamarca Gua Guainía Hui Huila Mag Magdalena Met Meta Nar Nariño NSa Norte de Santander Put Putumayo Qui Quidío Ris Risaralda SAP San Andrés y Providencia San Santander Suc Sucre Tol Tolima Val Valle Vau Vaupés Vic Vichada xxiii

xxiv

Abbreviations

Provinces of Ecuador Az Azuay Bo Bolivar Ca Carchi Cañ Cañar Ch Chimborazo Co Cotopaxi EO El Oro Es Esmeraldas Ga Galápagos Islands Gu Guayas Im Imbabura Lo Loja LR Los Ríos Ma Manabí MS Morona Santiago Na Napo Or Orellana Pa Pastaza Pi Pichincha Su Sucumbíos Tu Tungurahua ZC Zamora Chinchipe

Further Abbreviations !

collection seen; used for types to indicate that a type specimen has been examined a.o. among others aff. affinity, related to alt. altitude elevation auct. auctorum, according to certain authors; used for misused names that have been applied by certain authors to a different species than the type specimen ca. circa, approximately cf. confer, compare; used for a preliminary, tentative identification of a species comb. combinatio, combination; referring to the association of a species epithet with a particular genus name c. andr. cum androeciis, with male gametangia

Abbreviations

c. gyn. cum gynoeciis, with female gametangia c. per. cum perianthiis, with perianths c. spor. cum sporis, with spores, with sporophytes det. determinavit, identified by e.g., exempli gratia, for example et al. et alii, and others fo. forma, form herb. herbarium incl. including ICN International Code of Nomenclature for Algae, Fungi and Plants l.c. loco citato, as cited above leg. legit, collected by loc. locality nom. nomen, name nom. illeg. illegitimate name nom. inval. invalid name nov. novum, nova, new nr. numero, number p.p. pro parte; partly s.l. sensu lato, in a broad sense s.n. sine numero, without number s.str. sensu stricto, in a narrow sense sp. species (singular) spp. species (plural) subfam. subfamily subsp. subspecies subg. subgenus syn. synonym var. variety

xxv

Introduction

Contents  rief History of Exploration B Underexplored Areas Format of the Book

   2    5    6

Colombia and Ecuador, located in northwestern South America in heart of the northern Andes, have a very rich flora of liverworts and hornworts. About 850 species are recognized in this book, including about 720 species in Colombia (of which 705 liverworts) and about the same numbers in Ecuador, making it the richest area in tropical America for liverworts and hornworts (Gradstein et al. 2001). The area has about 70 percent of the species of liverworts and hornworts occurring in the neotropics and almost all the families. The rich flora of Colombia and Ecuador coincides with the presence of an extensive and complex Andean cordilleran system, made up of two ranges of snow-capped volcanos in Ecuador rising to its highest peak on Mt. Chimborazo (6268 m), and three cordilleras in Colombia separated by the deep valleys of the Magdalena and Cauca rivers. The highest elevation in Colombia is reached on the Sierra Nevada de Santa Marta (5775 m), an isolated mountain massive in the north with a distinct vascular flora (Bernal et al. 2016). The Amazonian rainforests bordering Venezuela, Brazil, and Peru, and the lush rainforests along the Pacific coast — with an annual rainfall over 10,000 mm, the wettest area in the Western Hemisphere — further add to the extraordinarily floristic richness of Colombia and Ecuador. The literature dealing with liverworts and hornworts of Colombia and Ecuador is extensive and includes more than 500 titles. Both countries have annotated checklists (Gradstein and Hekking 1979; Uribe and Gradstein 1998; Gradstein and Uribe 2016a, b; León-Yánez et al. 2006) as well as a Red Book of rare and endangered species (Linares and Uribe 2002; Gradstein and León-Yánez 2011). However, there is no modern work providing keys, descriptions, and illustrations for the species of liverworts and hornworts of Colombia and Ecuador. Richard Spruce’s masterly Hepaticae Amazonicae et Andinae (Spruce 1884–1885), a 600-page monograph © Springer Nature Switzerland AG 2021 S. R. Gradstein, The Liverworts and Hornworts of Colombia and Ecuador, Memoirs of the New York Botanical Garden 121, https://doi.org/10.1007/978-3-030-49450-6_1

1

2

Introduction

describing about 560 species from Amazon Basin and the Andes of Ecuador and Peru, is out of date, and Margaret Fulford’s Manual of the leafy Hepaticae of Latin America (Fulford 1963, 1966, 1968, 1976) is incomplete. Many of the lowland and lower montane species of Colombia and Ecuador can be keyed with The Hepaticae and Anthocerotae of Brazil (Gradstein and Costa 2003), but the high-Andean flora is not treated in this work. Because of the lack of a comprehensive identification tool, I have written this book.

Brief History of Exploration Colombia  Hepatic collecting activities in Colombia have been summarized in the checklists (Gradstein and Hekking 1979; Uribe and Gradstein 1998; Gradstein and Uribe 2016a, b). The first liverwort records from Colombia appeared in the Flora de la Reál Expedición Botánica del Nuevo Reino de Granada of José Celestino Mutis, which contains illustrations of, a.o., Jensenia erythropus, Plagiochila sp. and Frullania sp. (Aguirre and Calonge 1985). Hooker (1819–1820) and Kunth (1822) described several liverwort species collected by Alexander von Humboldt and Aimée Bonpland, including Porella crispata and Symbiezidium transversale from Mt. Quindio. The first major contribution was by Gottsche (1864) who reported 120 species, collected mainly by the German physician, Alexander Lindig, in Cundinamarca and many of them described as new to science. Gottsche’s herbarium was destroyed with the bombing of the Berlin herbarium but duplicates of Lindig’s liverwort collections can be found in the herbaria of Geneva, Helsinki, London, New York, Paris, and Stockholm. Other nineteenth-century collections, e.g., those made by Gustav Wallis in Antioquía, Caldas, Santander, and Tolima, were studied by Jack and Stephani (1892) and by Stephani (1898–1924), who incorporated all early liverwort and hornwort records from Colombia in his Species Hepaticarum. Later publications on Colombian liverworts and hornworts include those of Herzog (1932, 1934, 1955) on the collections of K.  Troll, E.  P. Killip, and A. C. Smith; by Bischler (e.g., 1962d, 1964, 1969) on her collections made throughout the country during the years 1956–1959; by Robinson (1967) on the J. Cuatrecasas and R. King collections; and by Winkler (1970, 1976) on his collections from the Chocó and the Sierra Nevada de Santa Marta. Fulford (1963, 1966, 1968, 1976) published many new records from Colombia in her Manual of the leafy Hepaticae of Latin America, e.g., based on the collections made by J. Cuatrecasas in the Andean region and by R. E. Schultes and I. Cabrera in the Colombian Amazon. Extensive collecting was done by Antoine M. Cleef in the páramos of the Eastern Cordillera in the 1970s (Cleef 1981), and these materials were the basis for a series of papers in Studies on Colombian Cryptogams (e.g., Gradstein et al. 1977; van der Gronde 1980; van Reenen 1982; Kuwahara 1982; Meenks 1987). Important hepatic collecting was also done in the 1970s and 1980s by J.  Aguirre, S.  R. Gradstein, G. B. A. van Reenen, J. Wolf, and others in the framework of the Dutch-Colombian

Brief History of Exploration

3

ECOANDES project (van der Hammen 1983–2008), and these activities yielded several papers dealing with bryophyte ecology and biogeography (e.g., van Reenen and Gradstein 1983, 1986; Gradstein et al. 1989, 1996; Wolf 1993; Gradstein 1995). In addition, much collecting was done by J. Aguirre and his students in the páramos of Cundinamarca and elsewhere (e.g., Santana and Aguirre 1986; Castillo et  al. 1994; Aguirre and Ruiz 2001), by R.  M. Schuster for study of oil bodies of Colombian Lejeuneaceae (Schuster 1992c), by S. P. Churchill and his associates in the framework of the “Moss Flora of Colombia” project (Churchill and Linares 1995), and by S. R. Gradstein together with J. Aguirre and J.-P. Frahm in the Chocó biogeographic region (Rangel and Gradstein 2004; Gradstein and Schäfer-Verwimp 2011; Gradstein and Reiner-Drehwald 2017a, b; Reiner-Drehwald and Gradstein 2018). A useful key to the genera of Colombian liverworts was published by Uribe and Aguirre (1997). In recent years hepaticological studies have been carried out mainly by Colombian bryologists in many parts of the country, including the Andean region (e.g., Álvaro Alba et  al. 2007; Barbosa et  al. 2007; Ramírez et  al. 2008; Corrales et  al. 2010; Benavides and Gutiérrez 2011; Gradstein et  al. 2016), the Colombian Amazon (Pinzón et  al. 2003; Benavides et  al. 2006; Campos et  al. 2014, 2015), and the Chocó (Vasco et al. 2002; Benavides and Sastre-de Jesús 2011). The floristic novelties of these studies were incorporated in the latest checklist (Gradstein and Uribe 2016a, b), but numerous collections in the national herbarium (COL) and elsewhere remain without identification and are awaiting study. A new key to the genera was prepared by Gradstein (2016b). Mainland of Ecuador  Hepatic collecting activities have been reviewed in the checklist of Ecuador (León-Yánez et al. 2006). The earliest record of a liverwort species seems to have been Leptolejeunea elliptica gathered by an unknown collector near Guayaquil (Lehmann 1833; Bischler 1969). William Jameson, a British physician and botanist who lived in Quito from 1822 until his death in 1872, made the first important liverwort collections in the country. There has been considerable confusion about the origin of Jameson’s specimens, which are frequently cited as being from Peru or Colombia. All his collections, however, are from Ecuador (León-­ Yánez et al. 2006). Lindenberg (1839–1843) described Plagiochila laetevirens and P. procera (= P. adianthoides) collected by Jameson in Baños, whereas Taylor (1846, 1847, 1848) described many new liverwort species based on Jameson’s collections, including common Andean taxa such as Brachiolejeunea laxifolia, Frullania cuencensis, Herbertus pensilis, Lepicolea pruinosa, Plagiochila aerea, P. dependula, P. deflexirama, P. ensiformis, P. fuscolutea, P. pachyloma, Porella brachiata, and P. squamuligera. The most important contribution was made by Richard Spruce. Spruce resided in Ecuador from 1857 to 1863, mainly in Baños, Ambato, and Guayaquil, and collected a wealth of bryophytes mainly in the provinces of Pastaza and Tungurahua. He described 225 species of liverworts and hornworts from the country, many of them new to science and including rare endemic taxa such as Myriocolea irrorata (≡ Colura irrorata) and Spruceanthus theobromae (Hällingbäck and Hodgetts

4

Introduction

2000; Kautz and Gradstein 2001; Gradstein et al. 2004). His Hepaticae Amazonicae et Andinae (Spruce, 1884–1885) was the first major study describing tropical liverworts based on direct observation in the field. It laid the foundation for the modern classifications of Frullaniaceae and Lejeuneaceae (Thiers 1984). After Spruce, liverworts and hornworts of Ecuador were studied by Stephani (1898–1924) based on miscellaneous collections, including, e.g., those made by Abbé Michel Allioni during 1909–1910  in the surroundings of Gualaquiza and Bomboiza (Morona Santiago; but often cited as “Azuay”, see below) and by Herzog (1952, 1957) and Arnell (1961) based on the large collections made by G. Harling and other Scandinavian botanists. More recently, liverworts from throughout the country were studied by Schäfer-Verwimp et al. (2006, 2013a); from cacao plantations by Kautz and Gradstein (2001) and Andersson and Gradstein (2005); from páramo by Gradstein (1998, 1999), M. Burghardt, and others; from Parque Nacional Podocarpus and the San Francisco Biological Reserve by Nöske et  al. (2003), Parolly et al. (e.g., 2004), M. Nebel and others; from the Condor region by Churchill et  al. (2009), Gradstein and Benitez (2014b) and Gradstein et  al. (2019b); from Polylepis forests by Gradstein and León-Yánez (2018); etc. Exploration of the Río Nangaritza and tributaries of the Pastaza River yielded several unusual rheophilous species new to science (Gradstein and Reiner-Drehwald 2007; Gradstein et al. 2011; Crandall-Stotler and Gradstein 2017) and led to the rediscovery of Myriocolea (Gradstein et al. 2004; Gradstein and Benitez 2014a). Numerous additions to the flora have been published (e.g., Benitez and Gradstein 2011; Benitez et al. 2012; Schäfer-Verwimp et  al. 2013a; Gradstein and Benitez 2017; Burghardt 2019; Gradstein et al. 2019b), and the 2006 checklist has now become outdated. The Reserva Biológica San Francisco, situated between Zamora and Loja at the northern limit of the Parque Nacional Podocarpus, is the best explored area in Ecuador for bryophytes and probably in the Andes as a whole. The reserve, which contains about 1000 ha of pristine montane forest and páramo, has been the location of a large-scale interdisciplinary project financed by the German Research Foundation (Beck et  al. 2007; Gradstein et  al. 2007b). Bryophyte work focused mainly on phytosociology and ecology (e.g., Parolly and Kürschner 2004a, b, 2005; Nöske 2005; Mandl et al. 2009), but general collecting was also done. A staggering number of liverwort species (357), about half the total number known from Ecuador, has been recorded from the reserve and the number is still rising (Gradstein et al. 2007a; Schäfer-Verwimp et al. 2013a; Gradstein and Benitez 2017). A reason for the high number of species recorded is undoubtedly the large number of bryologists who conducted fieldwork in the reserve and studied the collections. As Churchill et  al. (2009) remarked: “One could readily predict similar diversity numbers throughout the montane forest of the tropical Andes employing such expertise. This study provides a basis for comparing other localities of similar vegetation and elevational range.” Galápagos Islands  These islands, which are part of the territory of Ecuador, have been well studied and 126 species of liverworts and hornworts are recorded, including 10 endemic species (Gradstein and Ziemmeck 2016, with updates). Charles

Underexplored Areas

5

Darwin was the first to gather liverworts on Galápagos during his visit to the archipelago in 1835. Six species were recorded (Hooker 1847), including Bryopteris filicina, Frullania aculeata, F. peruviana, Omphalanthus filiformis, and two species of Frullania (F. tamarisci, F. vaginata) whose presence on the islands is unlikely and has been rejected (León-Yánez et al. 2006). The identities of the latter two species have not been verified and should be checked. Further early publications on Galápagos hepatics include the papers by Ångström (1873) on the collections of Swedish botanist N.  J. Andersson and by Evans (1902) on various nineteenth-­ century collections. Evans recorded the first hornwort from Galápagos, Nothotylas orbicularis, which was subsequently redescribed as N. galapagensis endemic to the archipelago (Howe 1934). Further species were recorded by Howe (1934), Clark (1953), Arnell (1961), and Weber (1975). Weber (1966) summarized the state of knowledge of the bryophyte flora of the islands and listed 68 species of liverworts and 3 hornworts. During my 1976 visit to the islands, several hundreds of liverworts and hornworts were gathered throughout the archipelago. A series of papers appeared based on the collections (Inoue 1977; Jovet-Ast 1978; Gradstein 1978, 1986; Kuwahara 1979; Inoue and Gradstein 1980; Onraedt 1982; Gradstein and Weber 1982; Weber and Gradstein 1984; Yamada and Gradstein 1991; Uribe 2004), and a new checklist was prepared together with an analysis of the bryogeography of the islands (Gradstein and Weber 1982; Gradstein 2009). Collections made by H. van der Werff, L. H. Pike, O. Hammann, U. Eliasson, and others were also incorporated in these publications. After the 1976 exploration little collecting of liverworts and hornworts was done on the islands for many years. In 2006 a renewed inventory of bryophytes (and lichens) was undertaken coordinated by F. Bungartz, and hundreds of hepatic collections were made. No new records of liverworts and hornworts were published based on these collections, however.

Underexplored Areas Even though thousands of hepatic collections have been made, large parts of Colombia and Ecuador may still be considered underexplored. In Colombia, the department of Cundinamarca and the páramos of the Eastern Cordillera are probably the best explored areas for liverworts. Much less attention has been paid to the Central and Western Cordilleras, the Chocó region, Amazonia, the Caribbean region, and the Sierra Nevada de Santa Marta. Future liverwort explorations should pay attention to these underworked areas. In Ecuador, the best explored areas are probably the province of Pichincha, the Galápagos Islands, the Pastaza River valley, the easily accessible sites in Parque National Podocarpus (Cajanuma, Bombuscara), and, especially, the San Francisco Biological Reserve. All other parts of the country are underexplored. Hornworts have been very little studied and particularly need work.

6

Introduction

Format of the Book This book provides keys, descriptions, and illustrations for about 850 species (in 47 families and 148 genera), including about 830 species of liverworts and 20 of hornworts. The format of the book is somewhat similar to that of The Hepaticae and Anthocerotae of Brazil (Gradstein and Costa 2003), but the latter work dealt with fewer species, lacked species descriptions, and about one fourth of the species of Brazil were not treated. In the present work, brief morphological descriptions and discussions are provided for all species from Ecuador and Colombia (a few obscure or doubtful taxa excepted for which the types could not be located or were unavailable). The text on the species varies in length and is more detailed for groups that have not been well studied and are poorly known. No attempt has been made to provide full descriptions for every species; such an approach would have been unrealistic given the large number of species occurring in Colombia and Ecuador. Generic treatments usually follow the latest taxonomic revisions and monographs. Important titles from the last 20 years include the treatments of Pycnolejeunea (He 1999), Kymatocalyx (Gradstein and Váňa 1999), Lejeuneaceae of Misiones (e.g., Reiner-Drehwald 1998, 2000), Crossotolejeunea (Reiner-Drehwald and Goda 2000), neotropical Plagiochila (Heinrichs 2002a a.o; Gradstein 2016a), Ceratolejeunea (Dauphin 2003), Echinocolea (Ilkiu-Borges 2005), Amphilejeunea (Reiner-Drehwald 2005a), neotropical Marchantiidae (Bischler-Causse et al. 2005), neotropical Porella (So 2005), neotropical Herbertus (Feldberg and Heinrichs 2006), neotropical Metzgeria (Costa 2008), Frullania subg. Meteoriopsis (Uribe 2008), Inflatolejeunea and Macrolejeunea (Reiner-Drehwald and Schäfer-Verwimp 2008a), Balantiopsidaceae of Colombia (Campos and Uribe 2012), Rectolejeunea (Reiner-Drehwald and Grolle 2012), Odontoschisma (Gradstein and Ilkiu-Borges 2015), Prionolejeunea (Ilkiu-Borges 2016), Bazzania (Gradstein 2017b), Anastrophyllum of Colombia (Álvaro and Uribe 2017), neotropical Cheilolejeunea (Bastos 2017), Stephaniellaceae (Juárez-Martínez and Delgadillo 2017), and Andean Riccardia (Gradstein and Reeb 2018b). Where taxonomic treatments were lacking, incomplete, or outdated, a revision of the group was undertaken, for example, for Acrobolbus, Frullania p.p., Bazzania, Lejeunea p.p., Lophocolea, Plagiochila p.p., and Riccardia. An exception are the hornworts. Hornworts of Colombia and Ecuador have been very little studied (see Villarreal et al. 2012), and the available collections are few and often poorly preserved. For this reason, I had initially considered omitting the hornworts from this book. However, as they are traditionally treated in liverwort manuals, a chapter on the hornworts of Colombia and Ecuador has been added. Species are defined as monophyletic groups of populations characterized by two or more independent, diagnostic morphological characters, varieties, and subspecies by a single diagnostic morphological character. Subspecies differ from varieties by being allopatric, geographically or ecologically separate taxa (Gradstein 1975; Gradstein and Ilkiu-Borges 2015). Taxa are accepted when they can be keyed. I realize that a considerable number of species accepted in this book are characterized by

Format of the Book

7

only one distinct, diagnostic character. Theoretically, these should be varieties or subspecies. However, since in most cases nothing is known about their phylogenetic status I have maintained them at species rank, except when there was clear evidence for reducing them to varietal or subspecific rank, such as in the case of Frullania sabanetica (≡ Frullania intumescens var. sabanetica), Marchesinia robusta (≡ M. brachiata var. robusta), and a few other taxa. Taxa without diagnostic morphological characters that cannot be keyed, such as the genetic races Acrobolbus ruwenzorensis (= A. laxus), Ceratolejeunea calabriensis (= C. cornuta), C. diversicornua (= C. coarina), and Symphyogyna hymenophyllum (= S. podophylla), have not been accepted. For each species and selected infraspecific taxa, information is provided on habitat, elevational range, and distribution within Colombia (by department) and Ecuador (by province). In addition, the world distribution is briefly described based on the literature and GBIF (https://www.gbif.org/species/). Information on distribution within Colombia and Ecuador has been taken from the checklists (Gradstein and Hekking 1979; Uribe and Gradstein 1998; Gradstein and Uribe 2016a, b; León-­ Yánez et al. 2006) with additions and corrections. The identifications of the records from Colombia and Ecuador have not all been checked; such a task would have been enormous and would have fallen beyond the scope of the book. The published records may therefore contain errors and must be used with care, especially in groups that have not been monographed. It should also be realized that some departments or provinces were established rather recently, and this may be a further source of error. In Ecuador, the provinces of Morona Santiago and Zamora Chinchipe were established in 1953, Napo and Pastaza in 1959, Sucumbios in 1989, and Orellana in 1998. As a result, older records from Ecuador are sometimes referred to provinces to which the localities previously belonged (León-Yánez et  al. 2006). The early twentieth-century collections of M.  Allioni from Morona Santiago (Gualaquiza, Bomboiza), for example, are frequently referred to Azuay which at that time included the territory of Morona Santiago. Important new records are documented by means of a specimen citation or a reference. Additionally, a reference is usually given when the taxon is known from only one or two locations in the country. Names of taxa and authors abbreviations generally follow the World Checklist (Söderström et al. 2016) except for new taxonomy, a few spelling corrections and the use of “ex” in author citations. The latter is omitted because the use of “ex” is not obligatory by the Code. Instead, only the validating author is cited. For taxon names with four or more authors, the author abbreviation “et  al.” is used. Synonyms include names used for Colombia and Ecuador and other relevant ones, but no effort has been made to cite all the homotypic synonyms which are often numerous, especially in Lejeuneaceae. Almost 100 new synonyms are proposed in this book. A reference to the type, or to the original description and illustration when the type could not be studied, is given for new synonyms. Holotypes are indicated when mentioned in the original publication. Illustrations are given for the genera and about half of the species. They were mostly taken from my previous books published in the Memoirs (Guide to the Bryophytes of Tropical America, The Hepaticae and Anthocerotae of Brazil, Guide

8

Introduction

to the Plants of Central French Guiana Part 4) or redrawn from other sources. The drawings emphasize important characters for identification and are not exhaustive. A glossary of technical terms is included but an introductory chapter on the morphology of liverworts and hornworts has been omitted. For this, the Guide to the Bryophytes of Tropical America, The Hepaticae and Anthocerotae of Brazil, and other hepatic manuals may be consulted. For an introduction on the morphology of liverworts and hornworts in Spanish, see León-Yánez et al. (2014). In spite of all the work done on the liverworts of Colombia and Ecuador, our knowledge of the exceptionally rich flora of these two countries remains incomplete. Many species new to the countries, or possibly new to science, are still to be discovered; in addition, numerous herbarium collections are awaiting identification or verification. Much work remains to be done. I hope that this book will stimulate work on the liverworts and hornworts of Colombia and Ecuador and of the wider Andean region in general.

Diversity and Endemism

Contents Phytogeographic Elements Regional Diversity

 11  16

With about 850 accepted species of liverworts and hornworts (720  in Colombia, 720  in Ecuador) and an endemism rate of about 10 percent, Colombia and Ecuador count among the most speciose regions on Earth. Almost 40 percent of the species belong to a single family, Lejeuneaceae, which shows enormous diversification in the area with about 325 species in 48 genera. Other important families are Lepidoziaceae (66 spp.), Plagiochilaceae (65 spp.), and Frullaniaceae (54 spp.) (Table 1). Lejeunea is the largest genus with 66 species, followed by Plagiochila (65 spp.). Since the appearance of the latest checklists, at least 35 species have been reported new to Colombia (e.g., Gradstein et  al. 2016, 2018c) and about 120 to Ecuador (e.g., Schäfer-Verwimp et al. 2013a; Gradstein and Benitez 2017; Burghardt 2019), and further new ones are included in this book. In spite of this, the number of species recorded from the two countries has remained almost unchanged, due largely to the large number of species that proved to be synonyms. As noted by Churchill (2009), the redundancy among published species names of tropical bryophytes is enormous. About 100 new synonyms are proposed in this book. Many of these were already listed as dubious species in recent checklists, although some of these have proven to be good taxa, for example, Aneura latissima and Lophocolea tenerrima from Ecuador (Dauphin et  al. 2018) and Lejeunea elongella from Colombia (M. E. Reiner-­Drehwald, pers. comm.). Some unverified species records of Colombia and Ecuador still remain, especially in the genera Lejeunea and Plagiochila, but their number has become limited. Many of them are dubious species described in the nineteenth century of which the type is lost. Colombia has not more species than Ecuador even though the country is almost four times larger. This supports the notion that richness is largely driven by habitat © Springer Nature Switzerland AG 2021 S. R. Gradstein, The Liverworts and Hornworts of Colombia and Ecuador, Memoirs of the New York Botanical Garden 121, https://doi.org/10.1007/978-3-030-49450-6_2

9

10 Table 1  The largest families and genera of liverworts of Colombia and Ecuador

Diversity and Endemism Family (no. genera/species) Lejeuneaceae (48/330) Lepidoziaceae (12/66) Plagiochilaceae (1/65) Frullaniaceae (1/54) Radulaceae (1/33) Metzgeriaceae (1/33) Lophocoleaceae (6/32) Aneuraceae (3/19) Jamesoniellaceae (1/18) Calypogeiaceae (2/17) Cephaloziaceae (6/17) Scapaniaceae (8/15)

Genus (no. species) Lejeunea (66) Plagiochila (65) Frullania (54) Radula (33) Metzgeria (33) Cololejeunea (32) Cheilolejeunea (30) Bazzania (26) Drepanolejeunea (25) Syzygiella (18) Diplasiolejeunea (18) Ceratolejeunea (18)

diversity — which is rather similar in Colombia and Ecuador — and not by surface area (e.g., Lack 1973; Riclefs and Lovette 1999; Patiño et al. 2013). The composition of the floras of the two countries is different, however, and about 30 percent of the species are only known from one of the two countries. About 140 species are known from Colombia but not from Ecuador, whereas about the same number are found in Ecuador but not in Colombia. The differences in species composition of the two countries probably reflect insufficient exploration or problems in identification. Lepidolejeunea cuspidata and Aneura latissima, for example, two species known from Ecuador, may have been recorded from Colombia as Lepidolejeunea involuta and Aneura pinguis, respectively. Blackwater rivers with granite rock should be searched in Colombia for rare rheophilous taxa such as Colura irrorata, Lejeunea topoensis, and Myriocoleopsis gymnocolea, which are characteristic of this type of habitat in Ecuador. Other species known from Ecuador that may be found in Colombia include Frullania aculeata, a subendemic species (one record from Panama) that is remarkably common in Ecuador, and Frullania planifolia, Haplomitrium blumei, various epiphyllous species of Cololejeunea and Diplasiolejeunea, Lejeunea rionegrensis, Lophocolea tenerrima, Microlejeunea globosa (dry montane forest!), Plagiochila ecuadorica, Radula yanoella, Reinerantha foliicola, Xylolejeunea pellucidissima, and Lobatiriccardia verdoornioides. On the other hand, species known from Colombia that should occur in Ecuador include characteristic Amazonian taxa such as Archilejeunea crispistipula, Cheilolejeunea neblinensis, C. urubuensis, Lejeunea immersa, Micropterygium pterygophyllum, Odontoschisma portoricense, Pteropsiella serrulata, Schusterolejeunea inundata, Telaranea pecten, Thysananthus plicatiflorus, and Zoopsidella integrifolia; Chocó elements such as Fuscocephaloziopsis pulvinata, Lepidolejeunea grossepapulosa, Luteolejeunea herzogii, and Prionolejeunea magnistipula (to be looked for in Esmeraldas); and páramo taxa such as Harpalejeunea grandis, Odontoschisma cleefii, Gymnomitrion setaceum, Radula sonsonensis and several members of Stephaniellaceae. Further examples are mentioned in the text of this book; plenty of interesting species remain to be found in the two countries.

Phytogeographic Elements

11

Phytogeographic Elements Five phytogeographic elements are here recognized in the liverwort and hornwort flora of Colombia and Ecuador: the neotropical element, the Afro-American element, the pantropical element, the temperate element, and the endemic element (Table 2). Neotropical Element This is the largest phytogeographic category and includes species that are restricted to tropical America but not endemic to Colombia and/or Ecuador. More than 600 species, or about 72 percent of the species of Colombia and Ecuador, belong to this category. Most of them are widely distributed in the neotropics, but about one third have more restricted ranges and occur in the Andes with limited distributions elsewhere, e.g., in Central America, the West Indies, or along the Atlantic coast of Brazil (Gradstein and Costa 2003). Since the descriptions of the ranges of the neotropical species are frequently altered as a result of new explorations (e.g., Schäfer-Verwimp 2014), no attempt is made here to further subdivide the category of neotropical species. Afro-American and Pantropical Elements  This group contributes 14 percent to the flora (120 spp.) and contains species that are also recorded from the paleotropics. Some of them have ranges extending to Africa, the Afro-American element (48 spp.), but the majority are pantropical taxa occurring in Africa and Asia, and some of them range into warm-temperate regions, e.g., Dumortiera hirsuta, Lejeunea flava, Lepidozia cupressina, Plagiochila bifaria, and Pseudomarsupidium decipiens. Although being much fewer than neotropical ones, the number of Afro-­ American and pantropical species has considerably increased in recent years based on the results of taxonomic and floristic work (Gradstein 2013b). Temperate Species  This is a small but interesting phytogeographic group made up of species which have their main distributions in temperate regions and probably originated there. They presumably migrated to the northern Andes in the late Tertiary or the Quaternary, after the upheaval of the Andean cordilleras, and three species, Solenostoma hyalinum, Schistochilopsis incisa, and Diplophyllum obtusatum, are considered to have been introduced by man (Gradstein and Váňa 1987). Temperate taxa include 37 species (Tables 2 and 3). They occur in the Andean region and at higher elevation on the Galápagos Islands, and about half of them are only found above 2000  m. Three categories are recognized: north-temperate Table 2 Phytogeographic elements in the flora of liverworts and hornworts of Colombia and Ecuador

Phytogeographic element Neotropical Afro-American and pantropical Temperate Endemic

Number of species 605 120 37 88

Percentage 72% 14% 4% 10%

12

Diversity and Endemism

Table 3  Temperate species Species North-temperate Anthelia juratzkana Blepharostoma trichophyllum Cephalozia bicuspidata Diplophyllum obtusatum Fossombronia pusilla Fuscocephaloziopsis pleniceps a Isopaches bicrenatus Lepidozia reptans Lophocolea bidentata Marchantia paleacea Metzgeria conjugata Nowellia curvifolia Odontoschisma denudatum Pallavicinia lyellii a Riccia huebeneriana Scapania cuspiduligera Schistochilopsis incisa Solenostoma hyalinum Solenostoma sphaerocarpum Sphenolobus minutus Syzygiella autumnalis Tritomaria capitata South-temperate Bazzania peruviana Clasmatocolea vermicularis Gymnocoleopsis cylindriformis Jensenia difformis Lophocolea muricata Marchantia beteroana Pseudocephalozia quadriloba Symphyogyna podophylla Syzygiella sonderi Pantemperate Cephaloziella divaricata Lunularia cruciata Marchantia polymorpha Phaeoceros carolinianus Ricciocarpos natans Targionia hypophylla =new records from the northern Andes

a

Elevation in Colombia and Ecuador/m 4300 3000–4300 ca. 3800 2650–3400 1100–3400 3500–3700 4300–4500 2600–3100 600–4450 2300–3500 2300–3820 900–3200 3500–4000 500–2500 3450 4300 2500–4500 1000–4000 1800–4100 4500 (2300–)3300–3700 3550 ? 1800–4500 3300–4460 1500–3500 1000–3400 (650–)2000–4200 3200–4100 500–3900 2000–4300 (400–)3000–3500 ca. 2000–3000 1600–4000 700–3500 ca. 2500 630–3800

Phytogeographic Elements

13

species, distributed mainly in the temperate regions of the Northern Hemisphere; south-­temperate species, occurring mainly in the temperate regions of the Southern Hemisphere; and pantemperate species, distributed widely in temperate regions of both Hemispheres (Table 3). North-temperate species are the largest group (22 spp.) and were mapped by Gradstein and Váňa (1987); about half of them are restricted to páramo (Table  4). A few of them, such as Fuscocephaloziopsis pleniceps and Tritomaria capitata, have also been found in the temperate regions of the Southern Hemisphere, but their main distribution is in the north. South-temperate liverworts are fewer (9 spp.), as already noted by Gradstein et al. (1989). Griffin et al. (1982) presented a list of south-temperate species of Colombia, but two of them, Colura naumannii and Pleurozia paradoxa, have main ranges in the tropical Andes and do not belong in this category. Since the appearance of the reviews by Griffin et  al. (1982) and Gradstein and Váňa (1987), very few temperate species have been added to the flora of the tropical Andes. Isopaches bicrenatus was newly found in Colombia (Gradstein et al. 2018c), Riccia huebeneriana was discovered in Ecuador (this book), and Flores et al. (2017) recorded the Holarctic Cephaloziella hampeana from the Andes of northern Argentina. Endemic Species  About 90 species are only known from Colombia and Ecuador. Endemism in Ecuador is 6% (44 species and subspecies) and in Colombia 3.5% (26 species and subspecies) (Tables 5 and 6). The higher endemism in Ecuador may be explained by the contribution of the Galápagos Islands to endemism (10 spp.; Table 6) and the intensive recent explorations in Ecuador, which resulted in a large number of species new to science (Schäfer-Verwimp et al. 2013a; Crandall-Stotler and Gradstein 2017; Gradstein and Benitez 2017, 2018; Gradstein and León-Yánez 2018; Gradstein et al. 2018b; this book). In spite of the new discoveries, the total number of endemic species has remained unaltered since the appearance of the latest lists (Gradstein and León-Yánez 2011; Gradstein and Uribe 2016a, b), due to the fact that about 30 species previously considered endemic to Colombia or Ecuador proved to be ill-defined or were found elsewhere. Ill-defined endemics, which have now fallen into synonymy, include Acrobolbus caducifolius, Metzgeria metaensis, Prionocolea viridissima, Syzygiella colombiana, and Zoopsidella dichotoma from Colombia and Frullania squamuligera, Gongylanthus oniscoides, Lejeunea megalostipa, Plagiochila inouei, and P. spinifera from Ecuador. Among those found elsewhere are the putative Colombian endemics Frullania dulimensis (recorded from Jamaica), Lejeunea inflexiloba, Leptoscyphus cleefii, Metzgeria bischlerae, and Plagiochila cuatrecasii (reported from Ecuador), and Odontoschisma cleefii, which was found in Peru. Putative endemic species from Ecuador which proved to be more widespread include Calypogeia oblata, Cololejeunea cingens, C. ecuadoriensis, Frullania formosa, Lepidozia subdichotoma, and Heteroscyphus polychaetus. The data clearly indicate that the recognition of endemic species in the neotropics is still hampered by the paucity of adequate information on the distribution of the species. More data are needed, both on distribution, necessitating more collecting in unknown places, and on taxonomy, necessitating revisionary work on neotropical groups (Knapp 2002).

14 Table 4 Species characteristic of (sub)páramo

Diversity and Endemism Acrobolbus setaceusa Adelanthus aureomarginatusa Adelanthus lindenbergianus Anastrophyllum auritum Anastrophyllum nigrescensa Anastrophyllum stellatuma Anastrophyllum tubulosum Anthelia juratzkana Bazzania placophyllaa Blepharostoma trichophyllum Cephalozia bicuspidata Cheilolejeunea auriferaa Cheilolejeunea paramicola Cheilolejeunea quinquecarinata Colura ornithocephala Colura stotleria Diplasiolejeunea erostrataa Diplasiolejeunea grandirostrataa Drepanolejeunea andinaa Drepanolejeunea auritaa Drepanolejeunea subvittataa Fossombronia peruviana Fuscocephaloziopsis pleniceps Gongylanthus liebmannianus Gongylanthus limbatus Gymnocoleopsis cylindriformis Gymnomitrion atrofiluma Gymnomitrion bolivianum Gymnomitrion setaceuma Gymnomitrion truncatoapiculatuma Harpalejeunea grandisa Isopaches bicrenatus Isotachis lopeziia Isotachis obtusaa Jensenia florschuetziia Kurzia brasiliensis Leiomitra robustaa Lejeunea inflexiloba Lejeunea schultesiia Leptoscyphus autoicusa Leptoscyphus cleefiia Leptoscyphus hexagonusa Leptoscyphus obcordatus Leptoscyphus sotiauxiia Lobatiriccardia verdoornioidesa (continued)

15

Phytogeographic Elements Table 4 (continued)

Lophocolea fragmentissimaa Lophonardia jamesoniia Marsupella emarginata Marsupella miniata Metzgeria bischleraea Metzgeria chilensis Metzgeria cleefiia Myriocoleopsis tixieria Nanomarsupella xenophyllaa Noteroclada confluens Odontoschisma cleefiia Odontoschisma denudatum Odontoschisma engeliia Plagiochila buchtiniana Plagiochila cleefiia Plagiochila cuatrecasiia Plagiochila dependula Plagiochila guevaraia Plagiochila pautaphilaa Plagiochila priceanaa Plagiochila revolvensa Platycaulis renifoliaa Pseudocephalozia quadriloba Riccia ecuadoriensisa Radula bischleraea Riccardia aberransa Riccardia cataractarum Riccia huebeneriana Riccia lamellosa Ruizanthus venezuelanusa Scapania cuspiduligera Spenolobus austroamericanusa Sphenolobus minutus Stephaniella paraphyllina Stephaniella rostrata Stephaniella uncigera Stephaniellidium sleumeri Symphyogyna rubescensa Syzygiella autumnalis Syzygiella burghardtiia Syzygiella rubricaulis Syzygiella setulosa Triandrophyllum eophylluma Tritomaria capitata =endemic to páramo

a

16

Diversity and Endemism

Table 5  Endemic species and subspecies of Colombia Species Liverworts Bazzania denticulifera Bazzania serrata Cheilolejeunea schiavoneana Colura stotleri Cyclolejeunea glimeana Dibrachiella bischleriana Diplasiolejeunea columbica Drepanolejeunea aurita Drepanolejeunea orthophylla subsp. saxicola Frullania bogotensis Harpalejeunea grandis Kurzia capillaris subsp. paramicola Leiomitra robusta Lejeunea killipii Lejeunea ryszardii Lejeunea schultesii Metzgeria cleefii Micropterygium longicellulatum Odontoschisma soratamum Prionolejeunea magnistipula Pycnolejeunea chocoensis Radula aguirrei Radula bischlerae Radula sonsonensis Rectolejeunea halinae Hornworts Nothoceros renzagliensis

Distribution

Habitat

Chocó Boyacá, Casanare, Santander Risaralda Boyacá

Lowland rainforest Montane cloud forest Submontane rainforest Sphagnum bog in superpáramo Coastal rainforest Lowland rainforest Upper montane scrub Páramo Montane rainforest environment Montane rainforest

Chocó Chocó Cundinamarca, Huila Cauca/Huila Quindío Boyacá, Cundinamarca, Santander Tolima Boyacá, Cundinamarca, Risaralda Boyacá, Cundinamarca

Páramo Páramo

Cundinamarca Cauca Chocó

Montane cloud forest and páramo Lowland rainforest Lower montane rainforest Páramo Subalpine dwarf forest Montane forest

Amazonas Chocó, Valle Chocó Cauca Boyacá Antioquía, Boyacá, Cauca, Meta, Risaralda, Tolima Risaralda

Lowland rainforest Coastal rainforest Coastal rainforest Coastal rainforest Páramo Montane cloud forest and páramo Submontane rainforest

Nariño

Upper montane forest

Antioquía, Boyacá, Chocó, Cundinamarca, Quindío, Risaralda Valle Quindío

Regional Diversity Biogeographic regions of Colombia were described by Rangel (1995) and Bernal et al. (2016), those of Ecuador in Jørgensen and León-Yánez (1999). The three main biogeographic regions of Colombia and Ecuador are the Andean region, the Amazon region and the Pacific coastal region. Due to their humid climate, these regions harbor most of the liverwort and hornwort species of the two countries. Other major

Regional Diversity

17

Table 6  Endemic species and subspecies of Ecuador Species Liverworts Cololejeunea kuciana

Distribution

Habitat

Zamora Chinchipe

Cololejeunea pterocolea

Morona Santiago

Cololejeunea stotleriana

Zamora Chinchipe

Colura andoi Colura irrorata

Galápagos Pastaza, Zamora Chinchipe

Diplasiolejeunea erostrata Diplasiolejeunea grandirostrata Diplasiolejeunea lanceolata Diplasiolejeunea mayaykuensis Fossombronia jostii Frullania blepharozia

Loja, Zamora Chinchipe Zamora Chinchipe Esmeraldas, El Oro Zamora Chinchipe Pastaza Chimborazo, Pichincha, Zamora Chinchipe Galápagos Galápagos Galápagos

Montane rainforest, epiphyllous Submontane rainforest, epiphyllous Submontane blackwater rivers, epiphyllous Evergreen woodland Lower montane blackwater rivers Subpáramo Páramo Submontane rainforest Cultivated land, submontane Calcareous rock along river Upper montane environment

Frullania darwinii Frullania gradsteinii Frullanoides densifolia subsp. grandidentata Fuscocephaloziopsis subintegra Lejeunea flagellaris

Leptoscyphus leoniae

Zamora Chinchipe Chimborazo, Galápagos, Morona Santiago Pichincha Chimborazo Los Ríos Orellana Azuay, Loja, Napo, Zamora Chinchipe Loja, Pichincha

Leptoscyphus revolutus Leptoscyphus sotiauxii Lobatiriccardia oberwinkleri

Pichincha Napo Loja, Zamora Chinchipe

Lobatiriccardia verdoornioides Metzgeria grandiflora Plagiochila macrifolia Plagiochila priceana

Azuay, Pichincha Galápagos Chimborazo Azuay, Cotopaxi, Pichincha

Radula eggersii

Loja, Zamora Chinchipe

Lejeunea mojandae Lejeunea suffruticola Lejeunea xiphophylla Lejeunea yasuniensis Leptoscyphus autoicus

Evergreen woodland Semi-deciduous woodland

Upper montane elfin forest Evergreen woodlands Polylepis forest Montane scrub Submontane rainforest Lowland rainforest Upper montane forest and páramo Upper montane forest and Polylepis forest Upper montane forest Páramo Wet rock in upper montane forest Páramo Evergreen woodland Upper montane forest Polylepis reticulata forest, páramo Isolated trees in pasture (continued)

18

Diversity and Endemism

Table 6 (continued) Species Radula galapagona Radula sinskeana Riccardia verticillata Riccia howellii Riccia ianthina Riccia limicola Spruceanthus theobromae Syzygiella burghardtii Syzygiella ciliata Triandrophyllum eophyllum Hornworts Anthoceros simulans Notothylas galapagensis Phaeoceros pichinchensis

Distribution Galápagos Pichincha Morona Santiago Galápagos Galápagos Galápagos Guayas, Los Ríos Loja/ Zamora Chinchipe Pichincha, Zamora Chinchipe Napo/Pichincha

Habitat Evergreen woodland Submontane forest Montane forest Dry scrub Dry volcanic rock Moist soil at pool Old cacao plantations Subpáramo scrub Upper montane forest and scrub Superpáramo

Galápagos Galápagos Pichincha

Dry scrub Semi-deciduous woodland Rock in river, upper montane

regions are the Caribbean coastal region and “Llanos orientales” of Colombia, each with a rather dry climate and with a relatively poor liverwort and hornwort flora, and the Galápagos Islands. Andean Region  This region, defined here as the mainland territory above 500 m, is by far the richest area for liverworts and hornworts. Hepatic diversity in the Andean region has been described by Gradstein (1995, 1998). Almost 800 species have been recorded here or about 95 percent of the hepatic flora of Colombia and Ecuador. Many of them are widespread in the region, and very high species numbers can be found in relatively small areas of well-preserved Andean vegetation. More than 350 liverwort species or about 45% of all species of the Andean region have been recorded from the San Francisco Biological Reserve in southern Ecuador, with a surface area of about 10  km2 probably the best explored area in the Andes (Gradstein et al. 2007a; Schäfer-Verwimp et al. 2013a). The extraordinary biological richness of the Andean region is due to the great climatic and elevational variation of the area as well as historical factors (e.g., van der Hammen 1983–2008; Young et al. 2002). Unfortunately, many of the rich Andean forests has been damaged or cleared and put into cultivation for cattle grazing or altered into coffee plantations or potato fields. About 12 percent (106 spp.) of the liverworts and hornworts found in the Andean region are endemic to the northern Andes, the cordilleran area ranging from Venezuela to northern Peru (Table 7). Among them are three genera, Chaetocolea, Platycaulis, and Reinerantha, each containing a single species. Many further endemic genera have been described from the Andean region, and the area has been considered a hotspot for generic endemism in liverworts (Schuster 1990). However,

Regional Diversity Table 7  Species and subspecies endemic to the northern Andes

19 Acrobolbus setaceus Adelanthus aureomarginatus Archilejeunea nebeliana Bazzania placophylla Bazzania serrata Blepharolejeunea saccata subsp. harlingii Calypogeia oblata Ceratolejeunea pungens Chaetocolea palmata Cheilolejeunea asperrima Cheilolejeunea choachina Cheilolejeunea paramicola Cheilolejeunea schiavoneana Cololejeunea kuciana Cololejeunea pterocolea Cololejeunea stotleriana Colura irrorata Colura stotleri Diplasiolejeunea columbica Diplasiolejeunea erostrata Diplasiolejeunea grandirostrata Diplasiolejeunea lanceolata Diplasiolejeunea mayaykuensis Diplasiolejeunea papilionacea Drepanolejeunea andina Drepanolejeunea aurita Drepanolejeunea cutervoensis Drepanolejeunea orthophylla subsp. saxicola Drepanolejeunea pterocalyx Drepanolejeunea ramentiflora Drepanolejeunea subvittata Drepanolejeunea urceolata Fossombronia jostii Frullania blepharozia Frullania bogotensis Frullania formosa Frullania haematocysta Frullania jelskii Frullania lobatohastata Frullania meridana Frullania ringens Fuscocephaloziopsis subintegra (continued)

20 Table 7 (continued)

Diversity and Endemism Harpalejeunea grandis Heteroscyphus polychaetus Heteroscyphus thraustus Jensenia florschuetzii Kurzia capillaris subsp. paramicola Leiomitra robusta Lejeunea chimborazensis Lejeunea drehwaldii Lejeunea elongella Lejeunea flagellaris Lejeunea galeata Lejeunea inflexiloba Lejeunea killipii Lejeunea meridensis Lejeunea mojandae Lejeunea prionoides Lejeunea ryszardii Lejeunea schultesii Lejeunea suffruticola Lejeunea xiphophylla Lepidolejeunea auriculata Lepidozia alstonii Leptoscyphus autoicus Leptoscyphus cleefii Leptoscyphus jackii Leptoscyphus leoniae Leptoscyphus revolutus Leptoscyphus sotiauxii Lobatiriccardia oberwinkleri Lobatiriccardia verdoornioides Metzgeria bischlerae Metzgeria cleefii Metzgeria jamesonii Micropterygium longicellulatum Neesioscyphus allionii Neolepidozia rectangularis Nothoceros renzagliensis Odontoschisma cleefii Odontoschisma engelii Odontoschisma soratamum Phaeoceros pichinchensis Plagiochila cleefii Plagiochila cuatrecasii (continued)

Regional Diversity Table 7 (continued)

21 Plagiochila ensiformis Plagiochila guevarai Plagiochila macrifolia Plagiochila pachyloma Plagiochila paraphyllina Plagiochila pautaphila Plagiochila priceana Plagiochila revolvens Platycaulis renifolia Radula bischlerae Radula eggersii Radula sonsonensis Rectolejeunea halinae Reinerantha foliicola Riccardia aberrans Riccardia hans-meyeri Riccardia verticillata Syzygiella burghardtii Syzygiella ciliata Temnoma chaetophyllum Triandrophyllum eophylla Endemic genera in bold face

most of these so-called endemic genera proved to be synonyms or have wider ranges and extend northwards to Costa Rica (e.g., Lindigianthus, Ruizanthus). Most species of the Andean region occur in the moist Andean forests, including rainforests and cloud forests, and in the treeless páramo. Few species are found in dry forests, such as Microlejeunea globosa and Frullania pluricarinata, which are taxa with main distributions in subtropical regions including southern USA, Mexico, northern Argentina, and Central Chile. The páramos of the Andean region are of major floristic interest due to the occurrence of a mixture of temperature and tropical taxa and the high rate of endemism (Luteyn 1999; Gradstein 1998). Páramos occur above the forest line, which varies in the northern Andes between about 3000 and 4000  m depending on climatic, edaphic, and human factors, and extend up to the perennial snowline at about 4800  m or higher. They harbor a wide variety of habitats, including grasslands, moorlands, bogs, river valleys, scrubby vegetations, and various type of rock, and are frequently inhabited by large, pachycaul plants with giant apical leaf rosettes of the genus Espeletia (Cleef 1981; Luteyn 1999). Great ecological differences may be seen between one páramo and the next, and some páramos are very humid, others rather dry. The ecological differentiation of páramos and their island-like distribution must have enhanced the rich flora of the páramo environment. About 10 percent of the liverworts of Colombia and Ecuador are characteristic of páramo, and more than half of them are endemic here (Table  4). Twelve are temperate species; the

22

Diversity and Endemism

remaining ones are tropical taxa. The liverwort families Arnelliaceae, Antheliaceae, Gymnomitriaceae, Pelliaceae, Scapaniaceae, and Stephaniellaceae are largely restricted to páramo, while Lejeuneaceae with about 15 species are poorly represented. Hornworts are very scarce in páramo, and two species, Nothoceros vincentianus and Phaeomegaceros fimbriatus, have been recorded. None of them is limited to páramo. Amazon Region  This region includes the vast lowland area east of the Andes drained by the Amazon River system, extending in the southeast of Colombia and the east of Ecuador. The region has a humid climate with annual rainfalls over 2000 mm and a short dry season and is largely covered by rainforest. Some of the largest protected rainforest areas on Earth, such as Parque Nacional Chiribiquete in Colombia and Parque Nacional Yasuni in Ecuador, are in this region. About 250 liverwort species and one hornwort, Dendroceros crispus, have been recorded (Pinzón et al. 2003; Benavides et al. 2006; Mota de Oliveira and ter Steege 2013; Schäfer-Verwimp et al. 2013a; Campos et al. 2014, 2015; this book), including 22 species and 3 genera (Pteropsiella, Schusterolejeunea, Verdoornianthus) endemic to the wider Amazon region (Table 8). The region is the center of diversity of the genus Archilejeunea. Species richness and composition of the Amazon rainforests seem rather uniform, but a tendency toward higher richness is seen in areas with high humidity, such as in Yasuni National Park (Mota de Oliveira and ter Steege 2013) or in locations close to the Andes (Campos et al. 2015). Lejeuneaceae are the most important family in the region and include more than half of all bryophyte species. The Colombian Amazon region holds important sandstone rock outcrops of the Guiana shield, such as the Serranía de Chiribiquete, Serranía de la Lindosa, and the Serranía de la Macarena. Although being much lower than the tepuis of Venezuela (Guayana Highland), the flora and vegetation of these low mountains resembles that of the tepuis (Cortés et al. 1998; Cleef and Arbelaez Velasquez 2005), and the summit area of Serranía de la Macarena (1600 m) is covered by low forest with great abundance of bryophytes (Galeano 2016). Nevertheless, there are no publications describing the liverwort diversity of these tabletop areas. In Ecuador, tepui-like mountains are present in the Cordillera del Condor situated in the south of the country, between the eastern Andean cordillera and the Amazon Basin, at the border with Peru. Although physiographically and floristically allied to the Guayana Highlands, the sandstone rocks of the Condor region are of more recent age than those of the Guiana shield. Recent studies indicate that the bryophyte flora of the Cordillera del Condor is closely similar to that of the Andean region and has many species in common with San Francisco Biological Reserve (Churchill et al. 2009). One liverwort species, Micropterygium duidae, is only known from the Condor region and the Guayana Highland (Gradstein and Benitez 2014c). Pacific Coastal Region  The Pacific coastal region, also called Chocó biogeographic region, is one of the world’s major hotspots of biodiversity (Myers et al. 2000). The exceptional biological richness of the region is due to its separation from the rest of South America after the uplift of the Andes, allowing for many species to evolve in isolation, and to its very humid climate with annual rainfall up to

Regional Diversity

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Table 8  Species endemic to Amazonia (including the Guianas)

Calypogeia densifolia Calypogeia tenax Allorgella schnelliia Archilejeunea badia Archilejeunea crispistipula Cheilolejeunea neblinensis Cheilolejeunea urubuensis Cololejeunea bekkerae Cololejeunea surinamensisa Cyclolejeunea folioruma Diplasiolejeunea buckii Leptolejeunea obfuscataa Lejeunea yasuniensis Micropterygium leiophylluma Micropterygium parvistipuluma Odontoschisma soratamum Pallavicinia sp. Pictolejeunea sprucei Pteropsiella serrulata Riccia crassifrons Schiffneriolejeunea amazonica Schusterolejeunea inundata Telaranea pecten Thysananthus innovans Verdoornianthus griffinii Verdoornianthus marsupiifolius Vitalianthus aphanellus Xylolejeunea pellucidissima Endemic genera in bold face =extending to the Pacific coastal region

a

12,000  mm. The high rainfalls prevail in Colombia and in the north of Ecuador, further to the south the climate becomes dry. Except for the dry south and the deforested coastal part of Ecuador, the region is covered by evergreen rainforests with a rich hepatic flora. More than 200 species have been registered (e.g., Winkler 1970; Mägdefrau 1983; Vasco et  al. 2002; Rangel and Gradstein 2004; Andersson and Gradstein 2005; Benavides and Sastre-de Jesús 2011; Gradstein and Reiner-­ Drehwald 2017a), and many more may probably be discovered. The Pacific coastal region is a center of diversity of the genus Prionolejeunea (Ilkiu-Borges 2016) and the pantropical Dibrachiella, and eleven species and two genera, Fulfordianthus and Luteolejeunea, are endemic or subendemic to the region (Table 9). Several of these endemic taxa are very common in the area in spite of

24 Table 9  Species endemic to the Pacific coastal region

Diversity and Endemism Bazzania denticulifera Cyclolejeunea glimeana Dibrachiella bischleriana Fulfordianthus pterobryoidesa Luteolejeunea herzogii Prionolejeunea magnistipula Prionolejeunea recurvulaa Pycnolejeunea chocoensis Radula aguirrei Spruceanthus theobromae Symbiezidium dentatuma (Sub)endemic genera in bold face a =extending to western Amazonia

their absence elsewhere. A conservation icon in the flora is Spruceanthus theobromae, the single New World representative of the palaeotropic genus Spruceanthus and known only from old cacao plantations in the province Los Ríos (Kautz and Gradstein 2001). An unusual aspect of the flora of the Chocó rainforests is the high number of montane species found here at sea level (Gentry 1986; Gradstein 1995). This has been explained by the very humid, foggy climate, and somewhat lower temperatures in the region as compared to the east side of the Andes. The climatic and floristic peculiarities of the Chocó forests are suggestive of the presence of lowland cloud forest, a forest type described from French Guiana characterized by morning fog and a highly diverse liverwort flora rich in montane taxa (Gehrig-­ Downie et  al. 2013). The Pacific coastal region is still underexplored, and more fieldwork is necessary to assess the diversity of its hepatic flora. Caribbean Coastal Region and Llanos Orientales  The coastal lowlands in the north of Colombia and the Llanos orientales, the drainage area of Orinoco River in the east of the country, are relatively dry regions with a rainfall of mostly less than 2000 mm per year and a prolonged dry season of about 6 months. These areas are largely covered by savannas, deciduous woodlands, scrub, and semi-desert vegetation. Few liverworts and hornworts are found in these dry lowland habitats (Table 10). They include various species of the genus Riccia and the hornwort genus Notothylas, which are characteristic of open soil and rock. Epiphytes include members of the genus Frullania, especially F. ericoides, F. gibbosa, and F. rio-­janeirensis, and a few species of Lejeunea, especially L. laetevirens. Frullania corticalis is characteristic of mangrove. Some epiphytes of dry forest and savanna may occur also in the outer canopy of evergreen rainforests, such as Acrolejeunea spp. and Frullania spp., and this has been described as the “canopy effect” (Cornelissen and Gradstein 1990). The liverworts and hornworts of the dry lowland regions have been little studied and need more work.

Regional Diversity Table 10  Species of dry lowland environments

25 Acanthocoleus aberrans Asterella pringlei Brachiolejeunea phyllorhiza Cololejeunea minutissima Cylindrocolea rhizantha Frullania ericoides Frullania gibbosa Frullania nodulosa Frullania obcordata Frullania rio-janeirensis Lejeunea laetevirens Lejeunea setiloba Lejeunea trinitensis Lopholejeunea subfusca Metzgeria myriopoda Notothylas spp. Riccia spp.

Galápagos Islands  The Galápagos Islands (Ecuador) are a group of young volcanic islands along the Equator in the eastern Pacific Ocean, about 1000 km west of the mainland of Ecuador. There are some 45 islands covering almost 8000 km2, but most of these are small and low and are covered with sparse, dry vegetation. Eight islands exceed 400 m in elevation (reaching to 1700 m on Isabela) and are covered with more or less luxuriant vegetation rich in bryophytes in the humid uplands; they include Floreana, Isabela, Pinta, Pinzón, San Cristóbal, San Salvador, and Santa Cruz. Liverworts and hornworts of the Galápagos Islands have been rather well studied, and 113 species of liverworts and hornworts are recorded (Gradstein 2009; Gradstein and Zimmeck 2016). Frullania is the largest genus with sixteen species, followed by Lejeunea (9 spp.), Plagiochila (8), Radula (7), Riccia (6), and Cheilolejeunea (5). Most of the species occur on several islands and copiously produce spores and/or asexual dispersal devices. Those known from only one island are usually sterile and lack specialized means of dispersal (Gradstein and Weber 1982). Ten species and one subspecies of liverworts and hornworts are endemic to the islands; one further species, Lejeunea flagellaris, is restricted to Galapágos and the mainland of Ecuador (Table 6). The relatively low rate of hepatic endemism (10%) as compared to flowering plants (50%) is explained by the high dispersal capacity of spore plants (see Patiño et al. 2013). Interestingly, the majority of the endemic species (60%) occur in rather dry environments. The elevated endemism in the dry areas is explained by the continued availability and expansion of dry regions on the islands during the Pleistocenic glaciations, when climates were drier than at present, offering more time for natural selections and evolution in dry regions than in moist regions (Johnson and Raven 1973; Porter 1979).

Classification

Contents Marchantiophyta Haplomitriopsida Marchantiopsida Jungermanniopsida Anthocerotophyta

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The classification of liverworts and hornworts in this book is according to the Catalogue of the Plants and Lichens of Colombia (Gradstein and Uribe 2016a, b) with updates. The classification of families into orders and (sub)classes follows Crandall-Stotler et al. (2009) except for Pleuroziaceae, which are placed in a separate subclass, Pleuroziidae, following Frey (2009). The arrangement of families, genera, and species is alphabetical.

Marchantiophyta (Liverworts)

Haplomitriopsida Haplomitriaceae Haplomitrium Nees

© Springer Nature Switzerland AG 2021 S. R. Gradstein, The Liverworts and Hornworts of Colombia and Ecuador, Memoirs of the New York Botanical Garden 121, https://doi.org/10.1007/978-3-030-49450-6_3

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Marchantiopsida Aytoniaceae Asterella P.Beauv. Plagiochasma Lehm. & Lindenb. Cleveaceae Clevea Lindb. Sauteria Nees Corsiniaceae Cronisia Berk. Cyathodiaceae Cyathodium Kunze Dumortieraceae Dumortiera Nees Lunulariaceae Lunularia Adans. Marchantiaceae Marchantia L. Monocleaceae Monoclea Hook. Ricciaceae Riccia L. Ricciocarpos Corda Targioniaceae Targionia L.

Jungermanniopsida Pelliidae Fossombroniaceae Fossombronia Raddi

Classification

Marchantiophyta

Pallaviciniaceae Jensenia Lindb. Pallavicinia Gray Symphyogyna Nees & Mont. Pelliaceae Noteroclada Hook.f. & Wilson Metzgeriidae Aneuraceae Aneura Dumort. Lobatiriccardia (Mizut. & S.Hatt.) Furuki Riccardia Gray Metzgeriaceae Metzgeria Raddi Pleuroziidae Pleuroziaceae Pleurozia Dumort. Jungermanniidae 1. Jungermanniales Acrobolbaceae Acrobolbus Nees Lethocolea Mitt. Adelanthaceae Adelanthus Mitt. Pseudomarsupidium Herzog Antheliaceae Anthelia (Dumort.) Dumort. Arnelliaceae Gongylanthus Nees

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Balantiopsidaceae Isotachis Mitt. Neesioscyphus Grolle Ruizanthus R.M.Schust. Calypogeiaceae Calypogeia Raddi Mnioloma Herzog Cephaloziaceae Alobiella (Spruce) Schiffn. Alobiellopsis R.M.Schust. Cephalozia (Dumort.) Dumort. Fuscocephaloziopsis Fulford Nowellia Mitt. Odontoschisma (Dumort.) Dumort. Cephaloziellaceae Cephaloziella (Spruce) Schiffn. Cephaloziopsis (Spruce) Schiffn. Cylindrocolea R.M.Schust. Gymnocoleopsis (R.M.Schust.) R.M.Schust. Kymatocalyx Herzog Gymnomitriaceae Gymnomitrion Corda Marsupella Dumort. Nanomarsupella A.Hagborg, L.Söderstr. & von Konrat Herbertaceae Herbertus Gray Triandrophyllum Fulford & Hatcher Jamesoniellaceae Syzygiella Spruce Jungermanniaceae Jungermannia L. Lepicoleaceae Lepicolea Dumort. Lepidoziaceae Bazzania Gray

Classification

Marchantiophyta

Kurzia G.Martens Lepidozia (Dumort.) Dumort. Micropterygium Gottsche Monodactylopsis (R.M.Schust.) R.M.Schust. Mytilopsis Spruce Neolepidozia Fulford & J.Taylor Paracromastigum Fulford & J.Taylor Pseudocephalozia R.M.Schust. Pteropsiella Spruce Telaranea Schiffn. Zoopsidella R.M.Schust. Lophocoleaceae Clasmatocolea Spruce Cryptolophocolea L.Söderstr. Heteroscyphus Schiffn. Leptoscyphus Mitt. Lophocolea (Dumort.) Dumort. Platycaulis R.M.Schust. Plagiochilaceae Plagiochila (Dumort.) Dumort. Pseudolepicoleaceae Blepharostoma (Dumort.) Dumort. Chaetocolea Spruce Temnoma Mitt. Scapaniaceae Anastrophyllum (Spruce) Steph. Diplophyllum (Dumort.) Dumort. Isopaches H.Buch Lophonardia R.M.Schust. Scapania (Dumort.) Dumort. Schistochilopsis (N.Kitag.) Konstant. Sphenolobus (Lindb.) Berggr. Tritomaria Loeske Solenostomataceae Nardia Gray Solenostoma Mitt. Stephaniellaceae Stephaniella J.B.Jack Stephaniellidium Grolle

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Trichocoleaceae Leiomitra Lindb. 2. Porellales Frullaniaceae Frullania Raddi Jubulaceae Jubula Dumort. Lejeuneaceae Acanthocoleus R.M.Schust. Acrolejeunea (Spruce) Steph. Allorgella Tixier Anoplolejeunea (Spruce) Schiffn. Archilejeunea (Spruce) Steph. Blepharolejeunea S.W.Arnell Brachiolejeunea (Spruce) Schiffn. Bromeliophila R.M.Schust. Bryopteris (Nees) Lindenb. Caudalejeunea Schiffn. Ceratolejeunea (Spruce) J.B.Jack & Steph. Cheilolejeunea (Spruce) Steph. Cololejeunea (Spruce) Steph. Colura (Dumort.) Dumort. Cyclolejeunea A.Evans Dibrachiella (Spruce) X.Q.Shi, R.L.Zhu & Gradst. Dicranolejeunea (Spruce) Schiffn. Diplasiolejeunea (Spruce) Schiffn. Drepanolejeunea (Spruce) Steph. Frullanoides Raddi Fulfordianthus Gradst. Harpalejeunea (Spruce) Schiffn. Lejeunea Lib. Lepidolejeunea R.M.Schust. Leptolejeunea (Spruce) Steph. Lindigianthus Kruijt & Gradst. Lopholejeunea (Spruce) Steph. Luteolejeunea Piippo Marchesinia Gray Metalejeunea Grolle Microlejeunea (Spruce) Steph. Myriocoleopsis Schiffn. Neurolejeunea (Spruce) Schiffn.

Classification

Anthocerotophyta

Odontolejeunea (Spruce) Schiffn. Otigoniolejeunea (Spruce) Schiffn. Pictolejeunea Grolle Prionolejeunea (Spruce) Schiffn. Pycnolejeunea (Spruce) Schiffn. Rectolejeunea A.Evans Reinerantha Gradst. & R.L.Zhu Schiffneriolejeunea Verd. Schusterolejeunea Grolle Spruceanthus Verd. Stictolejeunea (Spruce) Schiffn. Symbiezidium Trevis. Thysananthus Lindenb. Verdoornianthus Gradst. Xylolejeunea X.L.He & Grolle Porellaceae Porella L. Radulaceae Radula Dumort.

Anthocerotophyta (Hornworts) Anthocerotaceae Anthoceros L. Dendrocerotaceae Dendroceros Nees Nothoceros (R.M.Schust.) J.Haseg. Phaeomegaceros R.J.Duff et al. Leiosporocerotacaceae Leiosporoceros Hässel Notothyladaceae Notothylas A.Gray Phaeoceros Prosk.

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Keys and Descriptions

Contents Introductory Key����������������������������������������������������������������������������������������������������������������������   37 Liverworts��������������������������������������������������������������������������������������������������������������������������������   37 Class Haplomitriopsida������������������������������������������������������������������������������������������������������������   38 Class Marchantiopsida ������������������������������������������������������������������������������������������������������������   41 Haplomitriaceae ����������������������������������������������������������������������������������������������������������������������   39 Aytoniaceae������������������������������������������������������������������������������������������������������������������������������   42 Cleveaceae ������������������������������������������������������������������������������������������������������������������������������   46 Corsiniaceae����������������������������������������������������������������������������������������������������������������������������   48 Cyathodiaceae��������������������������������������������������������������������������������������������������������������������������   48 Dumortieraceae������������������������������������������������������������������������������������������������������������������������   51 Lunulariaceae��������������������������������������������������������������������������������������������������������������������������   52 Marchantiaceae������������������������������������������������������������������������������������������������������������������������   53 Monocleaceae��������������������������������������������������������������������������������������������������������������������������   58 Ricciaceae��������������������������������������������������������������������������������������������������������������������������������   59 Targioniaceae ��������������������������������������������������������������������������������������������������������������������������   67 Class Jungermanniopsida��������������������������������������������������������������������������������������������������������   68 Subclass Pelliidae��������������������������������������������������������������������������������������������������������������������   69 Fossombroniaceae��������������������������������������������������������������������������������������������������������������������   70 Pallaviciniaceae������������������������������������������������������������������������������������������������������������������������   77 Pelliaceae ��������������������������������������������������������������������������������������������������������������������������������   85 Subclass Metzgeriidae ������������������������������������������������������������������������������������������������������������   85 Aneuraceae������������������������������������������������������������������������������������������������������������������������������   87 Metzgeriaceae�������������������������������������������������������������������������������������������������������������������������� 105 Subclass Pleuroziidae�������������������������������������������������������������������������������������������������������������� 121 Pleuroziaceae �������������������������������������������������������������������������������������������������������������������������� 122 Subclass Jungermanniidae ������������������������������������������������������������������������������������������������������ 123 Order Jungermanniales������������������������������������������������������������������������������������������������������������ 124 Acrobolbaceae�������������������������������������������������������������������������������������������������������������������������� 130 Adelanthaceae�������������������������������������������������������������������������������������������������������������������������� 136 Antheliaceae���������������������������������������������������������������������������������������������������������������������������� 142

© Springer Nature Switzerland AG 2021 S. R. Gradstein, The Liverworts and Hornworts of Colombia and Ecuador, Memoirs of the New York Botanical Garden 121, https://doi.org/10.1007/978-3-030-49450-6_4

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Keys and Descriptions

Arnelliaceae ���������������������������������������������������������������������������������������������������������������������������� 143 Balantiopsidaceae�������������������������������������������������������������������������������������������������������������������� 145 Calypogeiaceae������������������������������������������������������������������������������������������������������������������������ 152 Cephaloziaceae������������������������������������������������������������������������������������������������������������������������ 160 Cephaloziellaceae�������������������������������������������������������������������������������������������������������������������� 172 Gymnomitriaceae�������������������������������������������������������������������������������������������������������������������� 179 Herbertaceae���������������������������������������������������������������������������������������������������������������������������� 183 Jamesoniellaceae���������������������������������������������������������������������������������������������������������������������� 191 Jungermanniaceae�������������������������������������������������������������������������������������������������������������������� 201 Lepicoleaceae�������������������������������������������������������������������������������������������������������������������������� 203 Lepidoziaceae�������������������������������������������������������������������������������������������������������������������������� 204 Lophocoleaceae������������������������������������������������������������������������������������������������������������������������ 243 Plagiochilaceae������������������������������������������������������������������������������������������������������������������������ 266 Pseudolepicoleaceae���������������������������������������������������������������������������������������������������������������� 320 Scapaniaceae���������������������������������������������������������������������������������������������������������������������������� 324 (Anastrophyllaceae, Jungermanniaceae p.p., Lophoziaceae)������������������������������������������ 324 Solenostomataceae������������������������������������������������������������������������������������������������������������������ 338 Stephaniellaceae���������������������������������������������������������������������������������������������������������������������� 342 Trichocoleaceae ���������������������������������������������������������������������������������������������������������������������� 346 Order Porellales ���������������������������������������������������������������������������������������������������������������������� 349 Frullaniaceae���������������������������������������������������������������������������������������������������������������������������� 350 General Key �������������������������������������������������������������������������������������������������������������������� 352 Key 1. Frullania subg. Diastaloba s.l.������������������������������������������������������������������������������ 353 Key 2. Frullania subg. Chonanthelia�������������������������������������������������������������������������������� 353 Key 3. Frullania subg. Meteoriopsis sect. Meteoriopsis�������������������������������������������������� 355 Key 4. Frullania subg. Meteoriopsis sect. Intumescentes and sect. Obtusilobae �����������������  356 Jubulaceae�������������������������������������������������������������������������������������������������������������������������������� 383 Lejeuneaceae���������������������������������������������������������������������������������������������������������������������������� 384 Classification of the genera of Lejeuneaceae of Colombia and Ecuador������������������������ 385 Synoptic key to the subfamilies and tribes of Lejeuneaceae ������������������������������������������ 386 Artificial key to the genera of Lejeuneaceae of Colombia and Ecuador ������������������������ 386 Key 1. Genera of Lejeuneaceae with undivided underleaves������������������������������������������ 386 Key 2. Genera of Lejeuneaceae with bifid underleaves �������������������������������������������������� 391 Porellaceae ������������������������������������������������������������������������������������������������������������������������������ 614 Radulaceae ������������������������������������������������������������������������������������������������������������������������������ 618 Hornworts�������������������������������������������������������������������������������������������������������������������������������� 640 (Anthocerotophyta - Anthocerotae) �������������������������������������������������������������������������������� 640

Introductory Key

37

Introductory Key 1. Plants with stem and leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants thalloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Leaves with a midrib (sometimes very short, restricted to the leaf base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mosses (Bryophyta) 2. Midrib lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaves in 2–3 rows . . . . . . . . . . . . . . . . . . . . . . . Liverworts (Marchantiophyta) 3. Leaves not in 2–3 rows (except Fontinalis, Phyllogonium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mosses (Bryophyta) 4. Thallus only 1 cell layer thick (except in the area of the archegonia); midrib lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Thallus more than 1 cell layer thick, at least in the middle; midrib present or lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Thallus pale green, narrowly linear, dichotomous, with colorless hairs on the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metzgeria 5. Thallus dark green to brown, different . . . . . . . . . . . . . . . . . . Fern prothallium 6. Thallus cells with one to two large chloroplasts. Capsule narrowly ellipsoidal to linear, green, turning black after dehiscence, dehiscing gradually (over a period of weeks or month) by 2 valves, from the apex downward . . . . . . . . . . . . . . . . . . . . . . Hornworts (Anthocerotophyta) (p. 640) 6. Thallus cells with numerous small chloroplasts. Capsule globose to ellipsoidal(cylindrical), black, dehiscing at once by (1–)4 valves (irregular or by a small operculum in some thalloid liverworts). . . . . . . Liverworts (Marchantiophyta)

LIVERWORTS (Marchantiophyta - Hepaticae) Plants (= gametophytes) leafy or thalloid, growing by means of a two- to four-sided apical cell. Leaves, when present, in 2–3 straight rows, the third row of leaves ventral and usually reduced in size, and sometimes absent, leaf usually only 1 layer of cells thick, occasionally more than 1 layer of cells thick in the lower half, without midrib, undivided or divided into segments. Thallus, when present, several cells thick at least in the middle, with or without midrib, inner tissue homogeneously green or differentiated into a green upper layer and a colorless to brown or reddish lower layer. Cells usually with numerous small chloroplasts and with oil bodies, pyrenoids lacking, cell walls often with trigones. Rhizoids unicellular (very rarely of several cells). Antheridia spherical, stalked, one to several in axils of leaf-like bracts, in thallus cavities, or on the thallus surface. Archegonia flask-shaped, one to several surrounded by leaf-like bracts (leafy liverworts) or an involucre. Sporophytes consisting of foot, seta, and capsule, enveloped until maturity in a calyptra and usually surrounded by additional protective structures: a perianth, perigynium, or marsupium in leafy liverworts and an involucre or pseudoperianth in thalloid liverworts. Setae colorless and rather fragile, elongating after spore maturation. Capsules globose to cylindrical, containing spores and elaters, capsule wall without stomata, columella and peristome absent; dehiscence

38

 

Keys and Descriptions

of capsule usually by means of (1–)4 valves, rarely irregular or with an operculum. Protonema very small, thalloid, normally giving rise to only one gametophyte. About 5000–6000 species worldwide, ca. 830 species in Colombia and Ecuador, in 141 genera and 43 families. Key to the classes of liverworts of Colombia and Ecuador 1. Plants with stem and leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants thalloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Leaves divided into segments . . . . . . . . . . . . . . . . Jungermanniopsida (p. 68) 2. Leaves not divided into segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaves in 3 ± equal rows, leaf insertion transverse. (Plants growing erect from a leafless rhizome, rhizoids absent). . . . . . . . . . . . . . Haplomitriopsida (p. 37) 3. Leaves not in 3 equal rows, leaf insertion incubous or succubous (rarely transverse) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jungermanniopsida (p. 68) 4. Thallus with air chambers (cross section) and with numerous small pores on dorsal surface. Thallus underside with scales, at least near the tip. Antheridia and archegonia usually born in receptacles . . . . . . . . . Marchantiopsida (p. 38) 4. Thallus lacking air chambers and pores. Thallus underside with or without scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Upper surface of the thallus with a median groove. Plants often growing in rosettes (loosely forked in aquatic plants). Sporophytes embedded in the thallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marchantiopsida: Ricciaceae (p. 58) 5. Upper surface of the thallus without median groove. Plants not growing in rosettes. Sporophytes not embedded in the thallus . . . . . . . . . . . . . . . . . . . . . 6 6. Rhizoids papillose. Thallus large, 1–3  cm wide, surface uniformly green. Gametangia produced in rounded receptacles with black hairs at the margin . . . . . . . . . . . . . . . . . . . . . . . Marchantiopsida: Dumortieraceae (p. 50) 6. Rhizoids smooth (exceptionally a few papillose rhizoids present). Thallus small or large. Gametangia not produced in receptacles with black hairs . . . . . . . . 7 7. Thallus large, (0.5–)1–3  cm wide, without midrib, margins crispateundulate, surface with numerous small whitish dots (fresh material). Antheridia produced in rounded or elongate receptacles on the thallus surface . . . . . . . . . . . . . . . . . . . . . . . Marchantiopsida: Monocleaceae (p. 57) 7. Thallus smaller, with or without midrib, margins plane or undulate, surface without small whitish dots. Antheridia not produced in receptacles on the thallus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jungermanniopsida (p. 68) Class HAPLOMITRIOPSIDA (Basal Liverworts) Plants foliose, erect, or creeping. Stems with central strand. Leaves transverse or succubous, originating from 1 initial cell, in 2 or 3 rows, usually undivided. Cells thin-walled, without trigones, with or without oil bodies. Rhizoids smooth or lacking. Antheridia and archegonia on the axis, in leaf axils, archegonia positioned laterally (anacrogynous), not terminally. Spermatozoid body massive. Sporophyte

Marchantiopsida

39

surrounded by a fleshy calyptra. Seta long, thick. Capsules opening with 2–4 valves, capsule wall 1–5-layered. Spores small, surface not ornamented. Containing two genera in two families, the worldwide genus Haplomitrium (Haplomitriaceae) and the Asian-Australasian genus Treubia (Treubiaceae).

Haplomitriaceae A family with a single genus. Haplomitrium Nees Calobryum Nees Plants foliose, with leaves in 3 ± equal rows, light green, 1–5 cm high, growing upright from a whitish leafless rhizome, rhizoids lacking. Stems pale, with a central strand. Branches lateral, without collar, originating from the epidermis. Leaves undivided, transverse, ovate-orbicular to broadly reniform, with very narrow insertion, margins ± entire, basal part more than 1 cell thick (2–4-stratose). Cells very large, ca. 50–100 μm long, quadrate to rectangular, arranged in rows, uniformly thin-walled, without trigones, cuticle smooth; oil bodies small, homogeneous, many per cell. Underleaves similar to leaves or slightly smaller. Dioicous. Antheridia and archegonia borne in discs terminal on the stem, surrounded by bracts and bracteoles in one or a few series. Sporophyte surrounded by a fleshy calyptra; perianth lacking. Seta thick, of numerous cells (cross section). Capsules cylindrical, wall only 1 cell layer thick. A genus of seven species worldwide, one in tropical America. Haplomitrium is readily recognized by (1) upright stems with stoloniform base and with 3 ± equal rows of undivided, transversally inserted leaves (underleaves ± similar to leaves); (2) basal part of leaves more than 1 cell thick; (3) leaf cells very large, ca. 50–100 μm long, thin-walled; and (4) rhizoids absent. Literature: Bartholomew-Began (1991), Gradstein et al. (2001) 1. Haplomitrium blumei (Nees) R.M.Schust.  Scalia andina Spruce (Calobryum andinum [Spruce] Steph.)

Fig. 1A–C

On rotten wood and humus in more or less undisturbed, lower montane rainforests at 1200–1500 m and in mossy Polylepis forest and páramo at ca. 3800–4000 m. Ecuador: Ca (páramo El Ángel), Co (Burghardt et al., QCA), Tu (Spruce 1885); to be expected in Colombia. General distribution: pantropical but rare in tropical America (tropical Andes, Lesser Antilles, Costa Rica). Haplomitrium blumei (sometimes erroneously spelled “blumii”) is rare in the Andes where it is known from a few collections from Ecuador, northern Peru, and Bolivia (Spruce 1885; Frey 1987; León-Yánez et al. 2006; Churchill et al. 2015). Most specimens are from lower montane rainforest but some are from páramo. The species is readily recognized by (1) stems growing upright, stem base whitish, stoloniform, without rhizoids, with three subequal rows of undivided, ovate-orbicular leaves; (2) leaf

40

 

Keys and Descriptions

Fig. 1 (A–C) Haplomitrium blumei. (A) Habit with gynoecia and sporophyte. (B) Portion of shoot, ventral view, 10×. (C) Midleaf cells. (D–G) Asterella macropoda. (D) Habit with female receptacles, dorsal view. (E) Female receptacle with three pseudoperianths, ventral view. (F) Thallus with 2 rows of scales, ventral view. (G) Scale. (H) Plagiochasma rupestre, habit with two female receptacles, dorsal view. (I–L) Clevea spathysii. (I) Habit with female receptacles, dorsal view. (J) Habit with female receptacle and sporophyte, dorsal view. (K) Portion of cross section of thallus. (L) Pore. (M and N) Sauteria berteroana. (M) Habit with female receptacles, sporophytes and androecia, dorsal view. (N) Pore. (B from Fulford 1963; J from Bischler-Causse et  al. 2005; rest from Gradstein et al. 2001)

Haplomitriaceae

41

cells large, ca. 60–80 μm long, thin-walled, arranged in straight rows; and (3) mature capsule cylindrical, on a long, thick seta, dehiscing by 2 valves, wall 1 cell thick. Class MARCHANTIOPSIDA (Complex Thalloid Liverworts) Plants thalloid, small to very large, prostrate, simple or forked, ventral or apical innovations sometimes present. Thallus with or without a midrib, dorsal surface often reticulate and with small pores; in cross section usually differentiated into 2 layers: a green upper layer composed of air chambers and photosynthetic tissue and a colorless to red or brown lower layer without air chambers and without photosynthetic tissue; sometimes thallus undifferentiated, homogeneous. Thallus usually with scattered oil cells (containing one large oil body and lacking chlorophyll); green cells without oil bodies. Ventral scales usually present. Rhizoids of two types: with smooth walls and with papillose (= pegged) walls. Antheridia and archegonia in dorsal cavities, in cushions or on stalked receptacles, archegonia often surrounded by an involucre. Sporophytes on a stalked receptacle or ± immersed in the thallus, protected by a thin calyptra and sometimes by an involucre (pseudoperianth). Setae short or lacking, rarely long (Monoclea). Capsules spherical to cylindrical, opening irregularly, by an operculum or by valves, capsule wall 1-layered. Spores usually large, 50–150 μm in diameter, and with a conspicuously ornamented surface (spores small and not ornamented in Lunularia, Marchantia, and Monoclea). Elaters present or lacking. Vegetative reproduction rare, by gemmae in Lunularia and Marchantia. About 350 species worldwide, 36 species in Colombia and Ecuador, in 13 genera and 10 families. Key to the families of Marchantiopsida of Colombia and Ecuador 1. Upper surface of the thallus with distinct pores, appearing as tiny, colorless or whitish dots (do not confuse with the oxalate crystals in the epidermis of Monoclea, which also appear as whitish dots). Air chambers present (cross section) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Pores lacking or indistinct. Air chambers absent or inconspicuous . . . . . . . . 10 2. Gemma receptacles present on dorsal surface of thallus. . . . . . . . . . . . . . . . . 3 2. Gemma receptacles absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Gemma receptacles cup-shaped. . . . . . . . . . . . . . . . . . . . . . . . Marchantiaceae 3. Gemma receptacles moon-shaped . . . . . . . . . . . . . . . . . . . . . . . . Lunulariaceae 4. Thallus usually with a median groove and often growing in (partial) rosettes, sometimes floating on water. Sporophyte embedded in the thallus . . Ricciaceae 4. Thallus without median groove, usually not growing in rosettes. Sporophyte not embedded in the thallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Thallus very thin, 2–3-layered, translucent, iridescent. Ventral scales lacking (except at thallus apex). Sporophytes on the ventral thallus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyathodiaceae 5. Thallus thicker, not translucent and not iridescent. Ventral scales present. Sporophytes not on the ventral thallus surface . . . . . . . . . . . . . . . . . . . . . . . . 6

42

 

Keys and Descriptions

6. Thallus completely black when dry, margins and underside dark purplishblack. Sporophyte in a dark, swollen, mussel-like involucre below the thallus apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Targioniaceae 6. Thallus not completely black when dry, margins and underside green or tinged with purple. Sporophyte not in a mussel-like involucre below the thallus apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Pores compound, made up of several layers of cells (cross section). Air chambers with green filaments. Male receptacle stalked . . . . . . . . . Marchantiaceae 7. Pores simple, of only 1 layer of cells. Air chambers without filaments. Male receptacle without stalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Underside of female receptacle with large, whitish or purplish, conical pseudoperianths, splitting into ribbon-like segments with age. Pores surrounded by several rings of cells. Thallus deep green to dark green. . . . . . . . . Aytoniaceae: Asterella 8. Pseudoperianths lacking. Pores surrounded by only one ring of differentiated cells. Thallus light green to bluish-green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Ventral scales ovate, in 2 rows, each scale terminating in 1–3 lanceolate appendages. Female receptacles arising from the dorsal thallus surface (not from the apex). Capsule opening by an operculum. Thallus light green to bluish-green, indistinctly or distinctly reticulate. . . . . . . . . . . . . . Aytoniaceae: Plagiochasma 9. Ventral scales lanceolate, in several ill-defined rows, each scale terminating in a linear-lanceolate appendage (appendage not well defined). Female receptacles arising from the thallus apex or from the dorsal thallus surface. Capsule opening by several irregular valves. Thallus light green (never bluish-green), distinctly reticulate. . . . . . . . . . . . . . . . . . . . . . Cleveaceae (and Corsiniaceae) 10. Plants small, thallus segments less than 0.5 cm wide, often in (partial) rosettes. Thallus surface often with a median groove . . . . . . . . . . . . . . . . . . Ricciaceae 10. Plants larger, thallus segments (0.5–)1–3 cm wide, never in rosettes. Thallus surface without median groove . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Thallus very light green. Ventral scales present, purplish, in two well defined rows. Female receptacles arising from the thallus surface (not from the apex). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aytoniaceae: Plagiochasma 11. Thallus dark green. Ventral scales absent or very small, colorless. Female receptacles arising from the thallus apex or absent . . . . . . . . . . . . . . . . . . . . 12 12. Thallus margins undulate-crispate. Thallus surface with numerous small whitish dots (consisting of oxalate crystals). Thallus without midrib. Rhizoids mostly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monocleaceae 12. Thallus margins plane. Thallus surface without small whitish dots. Thallus usually with a narrow midrib formed by rhizoids (on the ventral thallus surface). Rhizoids mostly papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dumortieraceae

Aytoniaceae Thallus medium-sized, with simple pores, the pores surrounded by one to several rings of differentiated cells. Air chambers usually in more than 1 layer, the walls green, filaments lacking. Ventral scales large, in two rows. Antheridia in irregular

Aytoniaceae

43

groups embedded in the dorsal surface of the thallus or in sessile receptacles; archegonia in receptacles. Female receptacles stalked after fertilization, stalk with or without a furrow, receptacle ± lobed, each segment with 1(−several) archegonia in a cavity surrounded by a two-lipped involucre. Fertilized archegonium and sporophyte sometimes surrounded by a large pseudoperianth (Asterella). Sporophyte with a very short seta. Capsules opening by an operculum. Spores usually large (ca. 50–150 μm). Vegetative reproduction lacking. A family of five genera worldwide, two in Colombia and Ecuador. The main characters of Aytoniaceae are (1) thallus firm, with simple pores; (2) air chambers usually in more than 1 layer, without filaments; (3) male receptacles sessile on the thallus surface (not stalked); (4) female receptacle stalked; and (5) capsule opening by an operculum. 1. Female receptacle arising from the tip of the thallus (or at the tip of a short ventral branch). Sporophyte surrounded by a large, conical, membranaceous, whitish or purplish pseudoperianth, hanging down from the underside of the female receptacle and splitting into ribbon-like segments. Thallus light green to glossy dark green to reddish. Pores distinct, surrounded by several rings of cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asterella 1. Female receptacle arising from the thallus surface, away from the tip. Sporophyte surrounded by a fleshy, green involucre opening to the lateral side of the receptacle (large pseudoperianth lacking). Thallus light green to pale bluish-green. Pores indistinct, surrounded by a one ring of cells (in P. rupestre) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plagiochasma Asterella P.Beauv. Fimbriaria Nees Thallus green to reddish-purple, medium-sized, 1.5–9 mm wide. Dorsal surface of thallus reticulate and with simple pores; epidermis cells with or without trigones. Thallus underside usually reddish-purple, with large violet scales in 2 rows, each scale with 1–4 appendages. Air chambers in several layers, without filaments. Monoicous or dioicous. Antheridia in irregular cushions on the thallus or on short lateral branches. Female receptacles arising from apex of the main thallus or small branches on a stalk with one furrow; receptacle usually papillose above and ± 4-lobed in the species from Colombia and Ecuador, each lobe with a small group of archegonia underneath surrounded by a cup-shaped involucre. Each fertilized archegonium (and sporophyte) surrounded by a large pseudoperianth, splitting above into 8–16 narrow segments. Sporophyte as in the family. About 50 species worldwide, 3 in Colombia and Ecuador, in 2 subgenera, subg. Phragmoblepharis Grolle and subg. Saccatae (Grolle) D.G.Long (see Long 2005). The most characteristic features of Asterella are the large, conical, whitish pseudoperianths hanging down from the underside of the female receptacle. When the sporophyte is mature, the pseudoperianth splits along its entire length into numerous narrow, ribbon-like segments, which remain connected at the tip after capsule dehiscence in the species from Colombia and Ecuador. The dorsal surface of the female receptacle is usually papillose (covered by small to large turbercles).

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Keys and Descriptions

Literature: Long (2005) 1. Thallus large, 5–9 mm wide. Female receptacle arising from the main thallus, stalk 10–40 mm long. Plants on moist soil and rock in montane forests above 2000 m, common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. macropoda 1. Thallus smaller, 2–5 mm wide. Female receptacle arising from the thallus apex or from a short ventral branch, stalk 10–25 mm long. Plants on road banks in open, rather dry habitats below 2000 m (not in forest), rare . . . . . . . . . . . . . . 2 2. Female receptacle arising from a short ventral branch (not from the main thallus), deeply 4-lobed, stalk 10–25  mm long. Spores yellow to red-brown, winged, with large areoles. Dry thallus strongly inrolled, becoming black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lateralis 2. Female receptacle arising from the main thallus, unlobed or shallowly four-lobed, stalk 10–15 mm long. Spores black or dark-brown, wingless, without large areoles. Dry thallus not strongly inrolled, not becoming black . . . . . . . . . . . . . . A. pringlei 1. Asterella lateralis M.Howe Fimbriaria elegans Spruce, nom. illeg. On earth banks and rock along roads, in rather open, dry environments, 1500–1800  m. Colombia: Cau (Robinson 1967), Qui (Gradstein and López Gonzales 2018a). Ecuador: Tu (Baños, R.  Spruce). General distribution: Central America, northern Andes. Asterella lateralis (subg. Phragmoblepharis) is a rare Central American species that has been found two times in Colombia and once in Ecuador. The species is distinguished by the deeply 4-lobed female receptacles which originate from a short ventral branch (not from the main thallus) and by the yellow to red-brown spores (80–130 μm in diam.) with a wing and areolate distal surface. The thallus is xeromorphic and becomes inrolled and blackish when dry (like in Targionia). 2. Asterella macropoda (Spruce) A.Evans Fimbriaria macropoda Spruce Fimbriaria canelensis Spruce

Fig. 1D–G

On moist soil in humid montane forests and páramo, often near streams or waterfalls, 2000–4000  m. Colombia: Ant, Boy, Cau, Cun, Nar, Qui, Ris, San, Tol. Ecuador: Az, Co, Lo, Pi, Tu, ZC. General distribution: tropical America. Asterella macropoda (subg. Phragmoblepharis) is a common Andean species that is recognized by (1) thallus rather large, 5–9 mm wide, not inrolling when dry, dorsal surface glossy dark green, reticulate, ventral surface deeply violet; (2) female receptacles 4-lobed, stalk 1–4 cm long, pseudoperianths conical, whitish to purplish, deeply ribbed, tips remaining connected after capsule dehiscence; and (3) spores reddish-brown, 80–105 μm, with a narrow wing and with large areoles (5–6 areoles across the surface). 3. Asterella pringlei Underw. Fimbriaria bakeri Steph. On soil over rock, on steep and rather dry, cut roadbank, 200–850 m. Colombia: Mag (Gradstein et al. 2016). General distribution: Mexico, Guatemala, northern Colombia.

Aytoniaceae

45

Asterella pringlei (subg. Saccatae) is a rare Mesoamerican species that has been found in the foothills of the Sierra Nevada de Santa Marta (Gradstein et al. 2016). The species is distinguished by the rather small, pale green thalli (3–5 mm wide) with almost unlobed female receptacles on very short stalks (10–15 mm long) arising from the apex of the main thallus (not from small branches), and by the blackish, wingless spores without areoles. Excluded Records Asterella lindenbergiana (Nees) S.W.Arnell  – Colombia (Orrego and Uribe 2004). Holarctic species, not in the tropics (Long 2005). The Colombian specimens are probably misidentified but could not be examined. Asterella saccata (Wahlenb.) A.Evans  – Colombia (Orrego and Uribe 2004). Northern temperate species of Europe and Asia, not in the tropics (Long 2005). The Colombian specimen is probably misidentified but could not be examined. Asterella venosa (Lehm. & Lindenb.) A.Evans – Colombia (Orrego and Uribe 2004). Subtropical species (Mexico, Southern Brazil, etc.), not in the equatorial tropics (Long 2005). The Colombian specimens are probably misidentified but could not be examined. Plagiochasma Lehm. & Lindenb. Thallus light green or pale bluish-green, often tinged with purple, small- to mediumsized, 3–5 mm wide. Dorsal surface of thallus reticulate or not, pores large or very small and inconspicuous; epidermis cells with or without trigones. Thallus underside with small or large, ovate, violet scales in 2 rows, each scale with 1–3 appendages. Air chambers in several layers, without filaments. Monoicous. Antheridia in sessile receptacles. Female receptacles arising from the middle of the thallus surface away from the apex, without a furrow; divided into one to four shallow segments, each segment with a strongly swollen, fleshy, (sub)globose involucre larger than the segment; involucre opening along a median slit into 2 valves and holding unfertilized archegonia and one sporophyte. Pseudoperianth lacking. Sporophyte as in the family. A genus of 16 species worldwide, one in Colombia and Ecuador. For main features of the genus, see under the species. Literature: Bischler-Causse et al. (2005) 1. Plagiochasma rupestre (J.R.Forst. & G.Forst.) Steph. Aitonia lanigera Spruce Aitonia subplana Spruce

Fig. 1H

On dry and moist volcanic rock and on loamy soil over rock, in open or partially shaded places in rather mesic woodlands, (50–)800–3200  m (on the Galápagos Islands at 50–1100 m). Colombia: Cun, Mag, San, Tol. Ecuador: Co, Ga, Lo, Tu. General distribution: widespread in tropical and Mediterranean regions. Plagiochasma rupestre is a rather common montane species that is readily distinguished by (1) thallus light green to pale bluish-green, ca. 5 mm wide, surface finely and rather indistinctly reticulate, pores inconspicuous, surrounded by a single

46

 

Keys and Descriptions

ring of 4–6 cells; (2) scales ovate, in 2 rows, with 1–2(−3) appendages; and (3) female receptacles arising from the dorsal thallus surface, not from the apex, each receptacle with one to four large, fleshy, subglobose involucres, which are projecting sideward and open with 2 valves, each involucre holding one sporophyte.

Cleveaceae Thallus small- to medium-sized, reticulate and with simple pores, the pores surrounded by only one ring of differentiated cells, pores sometimes star-shaped due to the thickening of the radial walls of the surrounding cells. Air chambers in 1–4 layers, empty, without filaments. Ventral scales in several rows, narrowly lanceolate, with filiform appendage. Antheridia in irregular or regular groups embedded in the dorsal surface of the thallus, archegonia in receptacles. Female receptacles stalked after fertilization, stalk short, with or without furrow, receptacle deeply lobed, dorsally with simple pores, each segment with archegonia surrounded by a 2-valved or tubular involucre. Pseudoperianth lacking. Sporophyte with a very short seta. Capsules opening by several irregular valves, operculum lacking. Spores large (ca. 40–80 μm), surface with large, rounded protuberances (areoles lacking). Vegetative reproduction lacking. A small family of three genera, two in Ecuador and Colombia, both very rare. Cleveaceae are closely related to Aytoniaceae, but the capsules open by valves instead of an operculum and pseudoperianths are lacking. This treatment follows Bischler-Causse et al. (2005) and Rubasinghe (2011). 1. Female receptacle arising from the thallus apex, involucre and sporophyte hanging down from underneath the receptacle. Pores star-shaped. Ventral scales with oil cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sauteria 1. Female receptacle arising from the thallus surface, involucre and sporophyte standing up obliquely from the side of the receptacle. Pores not star-shaped. Ventral scales without oil cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clevea Clevea Lindb. Athalamia W.Falc. Thallus light green, sometimes tinged with purple, small, ca. 1–1.5  cm long, 2–5 mm wide, forked; dorsal surface of thallus distinctly reticulate and with simple pores, the pores not star-shaped in the neotropical species; thallus underside with long, purplish, linear-lanceolate scales in several irregular rows, scales without oil cells, apex tapering-acuminate. Air chambers in 1–4 layers, without filaments. Monoicous or dioicous. Female receptacles 2–8-lobed, arising singly or several in a row along the midline of the thallus surface, stalk short, to 5 mm long, without a furrow, receptacle with two to three large, tubular, outwardly projecting involucres, each involucre 2-valved, opening by a long median slit. Sporophyte as in the family. A small genus of two species, one in Ecuador and Colombia (very rare). For main characters of Clevea, see under the species.

Cleveaceae

47

Literature: Bischler-Causse et al. (2005), Rubasinghe (2011) 1. Clevea spathysii (Lindenb.) Müll.Frib. Clevea andina Spruce (Athalamia andina [Spruce] S.Hatt.) Targionia robusta Steph.

Fig. 1I–L

On moist, loamy soil and rock along rivers, 2000–2800  m. Colombia: Qui (Gradstein and López Gonzales 2018b). Ecuador: Pa, Tu (Spruce 1885). General distribution: mainly in Mediterranean and subtropical regions of the Northern and Southern Hemisphere; a few records from the tropical Andes and one from Africa. Clevea spathysii is an amphitropical species that is very rare in the tropics (Rubasinghe 2011; Gradstein 2017a). In the northern Andes, the species has only been found in the mid-nineteenth century in Ecuador and in 2018  in Colombia (Gradstein and López Gonzales 2018b). It resembles Plagiochasma rupestris by the light green thallus with female receptacles arising from the dorsal surface (not from the thallus apex) but clearly differs in the distinctly reticulate thallus surface, the narrowly lancolate scales (numerous), the capsule opening by several irregular valves instead of an operculum, and the spores with large, rounded protuberances instead of areoles. Sauteria Nees Thallus dark green, small, to 1.2 cm long and 2.5 mm wide, forked; dorsal surface of thallus finely reticulate and with small pores, the pores star-shaped; thallus underside with reddish-black scales in irregular rows, reaching to the margin of the thallus, apex of scales sometimes with a ciliate appendage. Air chambers narrow-rectangular, in 1(−2) layers, without filaments. Monoicous. Female receptacles arising from the thallus apex, on a very short stalk (to 5 mm long) with one furrow, receptacle convex, 3–5-lobed, each lobe with a 2-valved involucre underneath. Sporophyte as in the family. A small genus of two species, one in Ecuador (Galápagos Is.). For main characters of Sauteria, see under the species. Literature: Bischler-Causse et al. (2005) 1. Sauteria chilensis (Mont.) Grolle Sauteria berteroana Mont.

Fig. 1M, N

On gravelly, volcanic soil in cool and dry, open, treeless environment, together with Plagiochasma rupestris and Targionia hypophylla, ca. 1200 m. Ecuador: Ga (Cerro Azul; Gradstein 2009). General distribution: Central Chile, northern Argentina, Peru, Galápagos Islands. Sauteria chilensis is a southern subtropical species that has been found once at high elevation in the Galápagos Islands. It is distinguished by the small, dark green, reticulate thallus (1.5–2.5 mm wide) with distinctly star-shaped pores, small ventral scales terminating in a ciliate appendage, and 3–5-lobed female receptacles with swollen, 2-valved involucres on the underside, beneath the lobes.

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Keys and Descriptions

Corsiniaceae A small family with two genera, one in Colombia. Cronisia Berk. Thallus medium-sized, light green, underside red to purplish or purplish-brown, to 3 cm long, 2–6 mm wide, forked. Dorsal surface of thallus finely reticulate, with simple pores surrounded by one ring of cells, the pores sometimes elevated; epidermis cells thin-walled, often decaying. Air chambers in 1 layer, with or without filaments. Thallus underside with scales in 2 rows, each scale with filiform appendages. Monoicous. Stalked receptacles lacking. Antheridia in linear receptacles along the median line of the thallus. Archegonia and sporophytes in shallow depressions along the median line of the thallus and covered by a conical involucre. Calyptra smooth. Pseudoperianth lacking. Seta very short. Capsules cleistocarpous, opening by decay of the capsule wall. Spores large (ca. 60–100 μm). Elaters present, short. A small neotropical genus (2 spp.), one species in Colombia (very rare). For main features of Cronisia, see under the species. Literature: Bischler-Causse et al. (2005) 1. Cronisia weddellii (Mont.) Grolle Funicularia bischleriana Jovet-Ast

Fig. 2A

On moist soil over rock along river, 350 m. Colombia: Meta (foot of Serranía de la Macarena; Bischler-Causse et  al. 2005). General distribution: scattered in rather dry regions of tropical America. Cronisia weddellii is a rare neotropical species of dry areas that had been found by H. Bischler along the rapids of the Río Guayabero, at the foot of the Serranía de la Macarena. The species is recognized by (1) thallus light green with purplish underside, rather small (2–5 mm wide), reticulate; (2) ventral scales with 1–6 filiform appendages; and (3) archegonia and sporophytes on the dorsal surface of the thallus, in shallow depressions along the median thallus line and covered by a conical involucre.

Cyathodiaceae A family with a single genus. Cyathodium Kunze Thallus very delicate, 2–3-layered, pale green to yellowish-green (rarely glaucous green), not tinged with purple, sometimes luminescent and with a bright metallic green luster, to 1.5 cm long and 1–4 mm wide, ± obcuneate with a narrow base and a broad, truncate apex, sometimes with a darker-colored midrib, usually growing in dense, flat mats. Dorsal surface of thallus reticulate and with large whitish pores surrounded by a few rings of cells (rings sometimes absent), epidermal cells

Cyathodiaceae

49

Fig. 2 (A) Cronisia weddelii, habit with female receptacles, dorsal view. (B and C) Cyathodium cavernarum. (B) Habit with male and ventral receptacles, dorsal view. (C) Spore. (D) Cyathodium foetidissimum, spores. (E and F) Dumortiera hirsuta. (E) Habit with male recetacle, dorsal view. (F) Habit with female receptacles and sporophytes, dorsal view. (G–I) Lunularia cruciata. (G) Habit with gemma cups, dorsal view. (H) Pore in cross section. (I) Portion of thallus with 2 rows of scales, ventral view. (J–N) Marchantia chenopoda. (J) Habit with female receptacles, dorsal view. (K) Habit with male receptacle, dorsal view. (L) Portion of cross section of thallus. (M) Pore in cross section. (N) Pegged rhizoid. (D redrawn from Wigginton 2004; rest from Gradstein et al. 2001)

50

 

Keys and Descriptions

thin-walled. Ventral scales present near thallus apex (in 2 rows) or fully absent. Rhizoids smooth or papillose. Air chambers in 1 layer, without filaments. Monoicous or dioicous. Antheridia on very small lateral branches. Archegonia 1–30 per thallus, on the ventral thallus surface. Sporophytes in a short row on the thallus underside, each sporophyte surrounded by an involucrum. Setae reduced. Capsules opening by an operculum. Spores ca. 30–55 μm, variously ornamented. Elaters few to many, with 2–4 spirals. Vegetative reproduction by tubers. A pantropical genus (ca. 12 spp.), two species in Colombia and Ecuador (rare). The main features of Cyathodium are (1) thallus green, very delicate, 2–3-layered, usually obcuneate in shape, with or without midrib, sometimes with a bright metallic green luster; (2) epidermis thin-walled, with large pores; (3) air chambers in 1 layer, without filaments; and (4) sporophytes on the thallus underside near the apex, surrounded by an involucrum. Literature: Salazar Allen (2005) 1. Rhizoids with smooth walls. Spores surface covered by short spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cavernarum 1. Rhizoids with papillose walls. Spore surface with rounded turbercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. foetidissimum 1. Cyathodium cavernarum Kunze

Fig. 2B, C

On earth walls and cut road banks, ca. 750 m. Colombia: Mag (Gradstein et al. 2016), Boy. General distribution: pantropical; in the neotropics known from Mexico, Cuba, Costa Rica, Panama, northern Colombia, and Brazil. Cyathodium cavernarum is a pantropical species that is uncommon in the neotropics. In Colombia the species is only known from the foothills of the Santa Nevada de Santa Marta (Gradstein et al. 2016). The species is not yet known from Ecuador. Cyathodium cavernarum is recognized by the glossy light green thalli, sometimes with a bright metallic green luster, without midrib and with smoothwalled rhizoids. The plants are monoicous and often fertile. Antheridia are produced on minute lateral thallus branches, and the sporophytes are on the underside of the thallus, near the apex. The mature spores are dark brown and densely spinose. 2. Cyathodium foetidissimum Schiffn.

Fig. 2D

On bare soil along trails, 50–2100  m. Colombia: Hui (Gradstein et  al. 2016). Ecuador: Bo (Pital near Caluma, trail to Cerro Samama, 350  m, Gradstein 9915, GOET), Gu (near Buenos Aires, Holm-Nielsen et al. 2591, AAU, PC, ster.), LR (Salazar Allen 2005). General distribution: pantropical but rare in tropical America (Costa Rica, Colombia, western Ecuador), also in southern Europe (Ducktt and Ligrone 2006). Cyathodium foetidissimum is readily distinguished from C. cavernarum by the densely papillose rhizoids (rhizoids smooth in C. cavernarum). Fertile material differs furthermore by the antheridia produced in small cushions near the thallus apex (not on small branches) and the spores covered by large, rounded turbercles (not by spines). The plants are monoicous, and sporophytes are produced in a short row on the thallus underside, near the thallus apex. Small, yellowish tubers may be produced on the ventral thallus surface.

Dumortieraceae

51

Cyathodium foetidissimum resembles Riccia membranacea in the thin and broad thallus, which is widened toward the apex, but the latter species clearly differs from C. foetidissimum by the sporophytes situated inside the thallus (not on the ventral surface), spores covered by short spines and absence of elaters. The two species may grow mixed (e.g., Gradstein 9915).

Dumortieraceae A family with a single genus. Dumortiera Nees Thallus uniformly deep green, large, 5–20 cm long, 1–3 cm wide, forked, with a narrow, colorless midrib consisting of rhizoids. Dorsal surface of thallus not reticulate, without pores and without epidermis, sometimes with papilliform cells in groups on the surface. Thallus margins often with hairs. Thallus underside green, with small, colorless scales in 2 rows and papillose and smooth rhizoids forming a narrow, false midrib and often radiating to the thallus margins. Air chambers lacking or vestigial. Monoicous. Male receptacle on a short stalk, circular and ± unlobed, margins with stiff, bristle-like hairs. Female receptacles on an elongate, 3–6  cm long stalk with two furrows, becoming 8–10-lobed after fertilization, margins with stiff, bristle-like hairs. Archegonia in groups on the ventral side of the segments, each group surrounded by a tubular involucre. Pseudoperianths lacking. Setae short. Capsules opening irregularly by valves. Spores small, 20–35 μm. Sporophyte otherwise as in the family. Vegetative reproduction lacking. A genus with one or several species, one in Colombia and Ecuador. For recognition of Dumortiera, see under the species. Literature: Bischler-Causse et al. (2005), Forrest et al. (2011) 1. Dumortiera hirsuta (Sw.) Nees Dumortiera hirsuta var. irrigua (Wilson) Spruce Dumortiera hirsuta var. trichopus Spruce

Fig. 2E, F

On moist rock, soil, and rotten wood in humid montane forests, often in or near running water, often together with Monoclea gottschei, (200–)500–3300  m. Colombia: Cal, Ces, Cho, Cun, Mag, Met, Nar, Qui, Ris, San, Tol, Val. Ecuador: Co, EO, ES, Ga, MS, Na, Pa, Pi, Tu, ZC. General distribution: widepread in tropical and oceanic warm-temperate regions of the world. Dumortiera hirsuta is a widespread hygrophilous species that is common in the Andean region. The species is genetically diverse and may split into several taxa (Forrest et al. 2011). Dumortiera hirsuta is readily distinguished by the large, flat, homogeneously green thalli without pores and without air chambers and usually with a (false) midrib formed by rhizoids. Fertile plants are recognized by the rounded receptacles on the thallus surface with dark, bristle-like hairs on the margins; both the male and the female receptacles are stalked (like in Marchantia).

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Keys and Descriptions

Sterile plants can be confused with Monoclea, which often grows together with Dumortiera. Monoclea can be readily separated from Dumortiera by the undulatecrisped thallus margins, the thallus surface with numerous whitish dots, the absence of a midrib, and the predominance of smooth rhizoids. A few rhizoids with papillose walls may be present in Monoclea, but they are very scarce and the walls are much less densely papillose than in Dumortiera.

Lunulariaceae A family with a single genus. Lunularia Adans. Thallus bright green, 1.5–4 cm long, 0.5–1 cm wide. Dorsal surface of thallus reticulate and with simple pores, the pores surrounded by three to four rings of cells; epidermis cells fully thin-walled (in plants growing in greenhouses) or with trigones. Thallus underside green, rarely purplish, scales very broad, colorless, in 2 rows, each scale with a rounded appendage. Air chambers in a very narrow layer, with numerous green filaments. Dioicous, mostly sterile and with gemmae. Antheridia and archegonia in receptacles. Male receptacles ellipsoid, at the thallus apex or on branches. Female receptacles 4-lobed, each segment with a long, tubular involucre enveloping the archegonia and sporophytes, receptacle becoming longly stalked at spore maturity, stalk hairy and very fragile, without a furrow. Pseudoperianths lacking. Sporophytes with a short seta. Capsules opening by an operculum and 4 valves. Spores very small, 14–22 μm. Vegetative reproduction by discoid gemmae borne on the thallus surface on the inner side of moon-shaped receptacles with entire margins. A genus with a single species; for recognition of Lunularia, see under the species. Literature: Bischler-Causse et al. (2005) 1. Lunularia cruciata (L.) Dumort.

Fig. 2G–I

On loamy soil in montane environments, usually in man-made habitats such as gardens, greenhouses, nurseries, and roadsides, occasionally growing naturally along trails and rivers, ca. 2000–3000  m. Colombia: Boy, Cun, Mag (Gradstein et  al. 2016), Qui (Gradstein & Lopez s.n., HUQ). Ecuador: Lo, Pi (Botanical Garden, Quito) (Benitez et al. 2012). General distribution: widespread in subtropical and warm-temperate regions, less common in the tropics. Lunularia cruciata is readily recognized by the large, bright green thalli with gemmae born in lunate (not cup-shaped) receptacles on the thallus surface. The species has been little collected in Colombia and Ecuador and has probably been overlooked. The species grows mostly in gardens and greenhouses but may also occur in natural environments (e.g., along the trail to Montserrate, Bogotá; Gradstein et  al. 2016). Lunularia cruciata is commonly introduced via gemmae in soil of garden plants.

Marchantiaceae

53

Marchantiaceae A family of three genera worldwide, one in Colombia and Ecuador. Marchantia L. Thallus light green to rather dark green to glaucous, underside often purplish; plants medium to large, 3–20(−30) cm long, 0.3–2 cm wide, forked, with or without midrib (cross section). Dorsal thallus surface reticulate and with numerous pores, the pores barrel-shaped, consisting of 4–7 layers of cells (cross section), each pore connected with a small air chamber. Thallus margins smooth or with small scales protruding beyond the margin. Thallus underside green or purple, with large colorless to violet scales in 4–10 rows, median scales with an appendage. Air chambers in 1 narrow layer, with green filaments. Dioicous. Male and female receptacles on long stalks with two furrows, shallowly or deeply lobed, the lobes flat or terete, margin of the receptacles without hairs. Archegonia in rows on the ventral side of the female receptacle, each row of archegonia surrounded by an involucre and each fertilized archegonium (and young sporophyte) surrounded by a pseudoperianth. Setae short. Capsules opening irregularly by valves. Spores small, 10–35 μm. Vegetative reproduction by discoid gemmae born in rounded, cup-shaped receptacles on the dorsal surface of the thallus; margins of the receptacles usually longly ciliate. A genus of about 50 species worldwide, 8 in Colombia and Ecuador, in 2 subgenera, subg. Marchantia and subg. Chlamidium ((Corda) Bischl. (see BischlerCausse et  al. 2005). The principal characteristics of Marchantia are (1) gemmae produced in cup-shaped receptacles on the thallus; (2) pores barrel-shaped, consisting of 4(−7) layers of cells (cross section); (3) ventral scales in 4–10 rows, scales sometimes reaching the thallus margin; (4) air chambers with filaments, in 1 layer; and (5) male and female receptacles stalked; in other neotropical genera of Marchantiidae, male receptacles are usually sessile (exception, Dumortiera). Literature: Bischler-Causse et al. (2005) 1. Scales extending over more than 3/4 of the width of the ventral thallus surface, often reaching the thallus margin. Outer surface of gemma cups papillose. Female receptacle very deeply divided into 9–11 narrow, terete lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (subg. Marchantia) 2 1. Scales extending over less than 2/3 of the width of the ventral surface. Outer surface of gemma cups smooth (papillose in M. paleacea). Female receptacle shallowly or rather deeply divided into 5–11 flat lobes. . . . . . (subg. Chlamidium) 4 2. Thallus margins irregularly lobulate. . . . . . . . . . . . . . . . . . . . . . M. berteroana 2. Thallus margins entire or crenulate (but sometimes seemingly lobulate due to ventral scales protruding beyond the margin). . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Ventral thallus surface green or somewhat purplish (not reddish), dorsal surface often with a dark median band. Ventral scales hyaline or light reddish, appressed to thallus surface. Lobes of female receptacle papillose. Male receptacle shallowly lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. polymorpha

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Keys and Descriptions

3. Ventral thallus surface reddish or black, dorsal surface without dark median band. Scales usually dark reddish, curved away from thallus surface. Lobes of female receptacle smooth. Male receptacle deeply lobed . . . . . . . . . M. plicata 4. Outer surface of gemma cups papillose. Appendage of scales with numerous oil cells. Male receptacle unlobed or very shallowly lobed. Female receptacle deeply lobed. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. paleacea 4. Outer surface of gemma cups smooth. Appendage of scales without oil cells (rarely with 1–2 oil cells). Male receptacle clearly palmately lobed. Female receptacle shallowly or deeply lobed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Thallus margin cells thin-walled, colorless or light red, rarely purplish. Appendage of ventral scales sharply toothed. Female receptacle deeply lobed, to 1/3–2/3 the diameter of the receptacle. Plants rare . . . . . . . . . . . . . . . . . . . 6 5. Thallus margin cells thick-walled, usually purplish, occasionally colorless or light red. Appendage of ventral scales entire or crenulate. Female receptacle shallowly lobed, to maximally 1/4 the diameter of receptacle . . . . . . . . . . . . 7 6. Thallus without black median line. Thallus margin coloress. Margin of gemma cups ciliate. Female receptacle lobed to about half the diameter of the receptable, lobes not strongly narrowed to the base . . . . . . . . . . . . . . . . . . M. inflexa 6. Thallus usually with a black median line. Thallus margin purplish. Margin of gemma cups crenulate. Female receptacle more deeply lobed, to 2/3× diameter of receptacle (or more), lobes strongly narrowed to the base, the base stalklike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. papillata 7. Appendage of scales with acute-acuminate apex. Female receptacle 5-lobed, apex of lobes rounded. Dorsal surface of male receptacle papillose. Plants very common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. chenopoda 7. Appendage of scales with rounded to subacute apex. Female receptacle 7–11lobed, apex of lobes truncate to emarginate. Dorsal surface of male receptacle smooth. Plants uncommon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. breviloba 1. Marchantia berteroana Lehm. & Lindenb. Marchantia discoidea Bonner On moist soil and rock in well-illuminated areas, often on road banks, 2000–4200 m; on the Galápagos Islands in summit areas in pampa at lower elevation, 650–800 m. Colombia: Ant, Boy, Cau, Cun, Met, Nar, Qui, San. Ecuador: Az (S. León-Yánez, QCA), Ga, ZC. General distribution: temperate regions of the Southern Hemisphere and on tropical high mountains. Marchantia beteroana (subg. Marchantia) is a southern-temperate species close to M. plicata and M. polymorpha but differing in having an irregularly hirsute thallus margin (margin entire or crenulate in M. plicata and M. polymorpha). The lobulation of the thallus margin should not be confused with ventral scales protruding beyond the thallus margin. From M. plicata the species differs furthermore in the pale-colored ventral thallus surface (reddish to black in M. plicata) and from M. polymorpha in the lack of a dark median band on the dorsal surface of the thallus.

Marchantiaceae

2. Marchantia breviloba A.Evans

55

Fig. 3A

On moist soil and rock on road banks and along trails in montane rainforest areas, 1000–3900 m. Colombia: Boy, Cun, Nsa, San. Ecuador: Pi. General distribution: scattered in tropical America, uncommon. Marchantia breviloba (subg. Chlamidium) is a rather rare species that is close to M. chenopoda; for differences see the key and under M. chenopoda. 3. Marchantia chenopoda L. Marchantia columbica Bonner Marchantia ecuadorensis S.W.Arnell

Fig. 2J–N

On moist soil and rock in open or shaded places, often on road banks, in meadows, and along running water, 150–3000 m. Colombia: Ant, Boy, Cau, Cal, Ces, Cho, Cun, Hui, Mag, Met, Nar, Nsa, Put, Qui, Ris, San, Tol. Ecuador: EO, Es, Ga, Gu, Lo, MS, Na, Pa, Tu, ZC.  General distribution: very common and widespread in the mountains of tropical America. Marchantia chenopoda (subg. Chlamidium) is a common and variable neotropical species that is recognized by (1) thallus margin cells thick-walled, often purplish (occasionally colorless); (2) ventral scales in (2–)4 rows, appendage of scales acute to acuminate and with entire or crenulate margins; (3) gemma cups with longly ciliate margin (cilia 3–5 cells long); (4) male receptacles deeply palmately lobed, with 2–9 flat lobes, dorsal surface papillose; and (5) female receptacle shallowly and asymmetrically 5-lobed, apex of the lobes broadly rounded. The species is close to M. breviloba, but in the latter species, the apex of the appendage of the scales is rounded to acute, the female receptacles are ± symmetrically lobed with 7–11 truncate to hirsuted lobes, and the dorsal surface of the male receptacles is smooth. 4. Marchantia inflexa Nees & Mont. On soil and soil over rock (limestone, volcanic) in humid and usually shaded places in rainforest or at water edges, more rarely on road banks, from sea level to 2300 m. Colombia: San, SAP (Bischler-Causse et al. 2005). General distribution: West Indies, Central America, northern Andes (Colombia, Venezuela). Marchantia inflexa (subg. Chlamidium) is a Caribbean species that has been found twice in Colombia; the record from Chocó (Vasco et al. 2002) needs verification. The species somewhat resembles M. breviloba but differs in (1) appendage of scales sharply toothed; (2) margin of gemma cups with shorter teeth (1–2 cells long); and (3) female receptacles more deeply lobed, to 0.3–0.6× receptacle diameter. 5. Marchantia paleacea Bertol. 

Fig. 3B

On soil, 2300–2500  m. Colombia: Cun, San (Bischler-Causse et  al. 2005). Widely distributed in subtropical and Mediterranean areas of the Northern Hemisphere, rare in the tropics; in tropical America mainly in Mexico and the West Indies, reaching southward to Venezuela and Colombia. Marchantia paleacea (subg. Chlamidium) is a rare subtropical species that has been recorded twice from Colombia and is not known from Ecuador. The species is readily recognized by (1) gemma cups with papillose outer surface and lobed-ciliate margin; (2) ventral scales in 4 rows, covering about half the width of the thallus; (3)

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Keys and Descriptions

Fig. 3 (A) Marchantia breviloba, female receptacle. (B) Marchantia paleacea, male receptacle. (C–F) Marchantia polymorpha. (C) Habit with female receptacle (c = gemma cup). (D) Habit with male receptacle. (E) Gemma cup. (F) Thallus, ventral view, showing scales up to the thallus margin. (G) Marchantia plicata, female thallus. (H and I) Monoclea gottschei subsp. gottschei. (H) Habit with sporophyte. (I) Habit with male receptacles. (J and K) Ricciocarpos natans. (J) Habit, dorsal view. (K) Cross section of female thallus. (L and M) Targionia hypophylla. (L) Habit with sporophyte, lateral view. (M) Ventral scale. (A, B from Bischler-Causse et al. 2005; rest from Gradstein et al. 2001)

Marchantiaceae

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appendage of scales with numerous oil cells; (4) thallus margin cells thin-walled; (5) male receptacles very hirsute lobed (like in M. polymorpha); and (6) female receptacles deeply divided into 7–11 flat lobes. 6. Marchantia papillata Raddi Marchantia subandina Spruce On soil and soil over rock (limestone, volcanic), often near water edges, from sea level to ca. 1000  m. Ecuador: without locality (Bischler-Causse et  al. 2005). General distribution: mostly in subtropical South America (southern Brazil, Paraguay, northern Argentina, Chile), northward to Ecuador and the central Amazon Basin (Manaus); a separate subspecies occurs in Asia. Marchantia papillata (subg. Chlamidium) is a southern subtropical species that has been collected once in the northern Andes. The species is close to M. inflexa and the two have been considered conspecific. However, they differ in the usual presence of purplish margins and a black median line on the thallus in M. papillata and in the shape of the female receptacle, which is much more deeply divided (to near the base) in M. papillata, with the basal part of the receptacle lobes being very narrow, stalk-like. Moreover, the margins of the gemma cups are crenulate in M. papillata and short-ciliate in M. inflexa. The two species have very different geographical distributions, M. papillata occurring in southern subtropical America, south of the Equator, while M. inflexa is distributed in the West Indies, Central America, and the northern Andes (Bischler-Causse et al. 2005). 7. Marchantia plicata Nees & Mont. Marchantia lamellosa Hampe & Gottsche

Fig. 3G

On soil over volcanic or calcareous rock, in stonework, in seepage areas, on road banks, in marshes in páramo, and along running water, in exposed or shaded locations, (500–)1800–4150 m. Colombia: Boy, Cal, Ces, Cho, Cun, Mag, Met, Nar, Qui, San, Tol. Ecuador: Az, Bo, Ch, Co, Im, Lo, MS, Na, Pa, Pi, Tu. General distribution: tropical Andes, Central America (?). Marchantia plicata (subg. Marchantia) is a common high-Andean species that grows scattered or in mats and may form rosettes up to 75 cm in diameter. It is considered the world’s largest terrestrial liverwort (Gradstein 1989b). The species is close to M. polymorpha but differs in (1) thalli with reddish or black underside, dorsal thallus surface without dark median band; (2) scales ± curving away from the ventral thallus surface; (3) male receptacles rather deeply lobed; and (4) female receptacles with ± smooth lobes. 8. Marchantia polymorpha L.

Fig. 3C–F

On moist soil in ruderal habitats, often near human settlements, in plantations, gardens, and near greenhouses, 1600–4000(−4700?) m. Colombia: Boy, Cau, Ces, Cun, Mag, Nar, Nsa, Qui, San. Ecuador: Ga, Tu. General distribution: subcosmopolitan. Marchantia polymorpha (subg. Marchantia) is a widespread, subcosmopolitic species that usually grows in ruderal habitats on nitrate-rich soil. The species is recognized by (1) ventral thallus surface green or somewhat purplish (not reddish or black), dorsal surface often with a dark median band; (2) ventral scales appessed to the ventral thallus surface, often protruding beyond the thallus margin; (3) lobes of female receptacle papillose; and (4) male receptacles very shallowly lobed.

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Keys and Descriptions

Monocleaceae A family with a single genus. Monoclea Hook. Plants thalloid, pure green, glossy, small to very large, 1–10(−20) cm long, 0.5–3 cm wide, prostrate, forked. Thallus without a midrib, in cross section homogeneous, without differentiation of air chambers; upper thallus surface densely spotted by whitish or darkish dots in fresh material; margins usually crisped-undulate. Oil bodies large, colorless or brown, occurring singly and scattered in inner cells of the thallus, especially in the upper and lower subepidermis. Ventral scales lacking. Rhizoids mostly with smooth walls, a few slightly papillose rhizoids present. Dioicous. Gametoecia within dorsal cavities. Antheridia in swollen receptacles along the median portion of the thallus, the receptacles rounded to linear or furcate. Archegonia beneath an elongate swelling (= involucre) along the median portion of the thallus, near the apex. Sporophytes arising one to three per involucre, when young surrounded by a thin calyptra; pseudoperianth absent. Setae long and thick, to 5 cm. Capsules cylindrical, opening by only 1 valve, wall 1-layered, with two sets of anastomosing thickening bands in the radial cell walls. Spores very small, 15–20 μm. Elaters very long, with 2 spirals. A genus of two species, one in New Zealand and one in Latin America (with two subspecies). For recognition of Monoclea, see under the species. Literature: Gradstein (2005, 2017c) 1. Monoclea gottschei Lindb. subsp. gottschei  Monoclea gottschei subsp. elongata Gradst. & Mues

Fig. 3H, I

On moist rock, soil, rotten wood, and trunk bases in montane forests, in or near running water, and always in shaded locations, often growing together with Dumortiera hirsuta, 150–4000 m. Colombia: Ama, Boy, Cas, Ces, Cho, Cun, Hui, Mag, Met, Nar, Put, Qui, Ris, San, Tol, Vau. Ecuador: EO, Lo, LR, MS, Na, Pa, Pi, Tu, ZC. General distribution: widespread in tropical America; a separate subspecies occurs in warm-temperate regions of Chile and Argentina. Monoclea gottschei is a common hygrophilous species that is readily distinguished by the large green thallus with undulate to crisped margins and the thallus surface densely spotted by minute whitish dots. The dots, which are particularly well visible in fresh material, are crystal druses (not pores). A midrib is lacking. Sporophytes, often in pairs or even triplets, arise near the thallus apex from beneath an elongate swelling of the thallus and possess a long seta and a cylindrical capsule, which opens along a longitudinal slit (hence the name “Monoclea”). The antheridia are developed in elongate receptacles on the thallus surface. Based on the shape of the mature male receptacle, M. gottschei is divided into two subspecies: Monoclea gottschei subsp. gottschei in tropical America and characterized by elongate male receptacles and Monoclea gottschei subsp. australis Gradst. in warm-temperate regions of Argentina and Chile and with orbicular receptacles (Gradstein 2017c).

Ricciaceae

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The shape of the male receptacle must be observed on mature receptacles; young male receptacles may be orbicular in both subspecies. Monoclea gottschei frequently grows mixed with Dumortiera hirsuta, and sterile thalli of the two species can easily be confused; for differences see under D. hirsuta.

Ricciaceae Thallus forming rosettes or gregarious patches, forked. Dorsal surface of thallus frequently with a median groove, usually not reticulate; pores present or absent, when present inconconspicuous and simple, surrounded by a ring of cells or not. Air chambers present or ± lacking, in one to several layers, without filaments. Ventral scales scattered or in 1–2 rows or absent. Gametoecia sunken in the thallus with only the necks projecting beyond the surface; receptacles lacking. Involucre and pseudoperianth lacking. Sporophytes embedded in the thallus, consisting merely of a globose capsule, lacking foot and seta. Capsule cleistocarpous, wall disintegrating at the time of spore maturation. Spores large, 50–200 μm, surface variously ornamented. Elaters lacking. Vegetative reproduction usually lacking, occasionally by tubers. A family of two genera, the large genus Riccia and Ricciocarpos with a single species. The species usually grow as pioneers on naked soil in rather dry areas, sometimes floating on water. The main characters of the family are (1) thallus small, forked, often fleshy and growing in rosettes, dorsal surface frequently with a median groove; (2) air chambers and pores frequently absent or rudimentary; (3) gametoecia and sporophyte embedded in the thallus; (4) seta and elaters lacking; (5) capsule dehiscence by disintegration of the capsule wall; and (6) spores large. 1. Plants floating on water (rarely on wet soil). Pores present, surrounded by a ring of cells. Ventral scales large, long-ligulate, with oil cells. Dorsal thallus surface ± dark green, reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ricciocarpos 1. Plants on soil, rarely in water. Pores absent or present, when present without ring of cells. Ventral scales absent or small, not long-ligulate, without oil cells. Dorsal thallus surface pale green to glaucous green, reticulate or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Riccia Riccia L. Thallus pale green to glaucous green, often tinged with red or purple, small, thallus segments 0.5–4 mm wide, forked, often forming rosettes. Dorsal surface of thallus usually with a median groove, not or slightly reticulate; pores lacking or very small and inconspicuous; dorsal epidermis without oil cells, sometimes decayed and then thallus surface with openings. Thallus margins acute or obtuse, sometimes with cilia or with scales projecting beyond the margin, the flanks of the thallus usually steepsloping. Air chambers in 1–3 layers or ± lacking, when lacking the upper tissue of the thallus forming dense vertical rows. Ventral scales small or large, when large sometimes projecting beyond the thallus margin, variously colored, in 1–2 rows, without oil cells. Monoicous or dioicous. Gametoecia and sporophyte as in the family.

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Keys and Descriptions

About 150 species worldwide, 15 in Colombia and Ecuador, in 3 subgenera, subg. Leptoricca R.M.Schust., subg. Riccia, and subg. Ricciella sect. Riccia (A.Braun) Bisch. (see Bischler-Causse et al. 2005); four species are endemic to the Galápagos Islands. For main features of Riccia, see under the family. The species of Riccia have been little collected in Colombia and Ecuador and deserve more attention. They are mostly found in open, cultivated areas and should be much more common than is currently known. Mature spores are sometimes necessary for species identification. Literature: Jovet-Ast (1978), Bischler-Causse et al. (2005) 1. Thallus linear, 0.3–0.6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Thallus wider . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Spores 80–95 μm, with 7–9 areoles across the outer face. Plants endemic to the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. limicola 2. Spores smaller, 60–75 μm, with 4–7 areoles across the outer face. Plants widespread, but not known from the Galápagos Islands. . . . . . . . . . . . . . R. fluitans 3. Thallus only 2–3 cell layers thick, thallus lobes conspicuously widened toward apex. Spores 40–65  μm in diameter, densely covered by spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. membranacea 3. Thallus more than 3 cell layers thick, lobes not conspicuously widened toward apex. Spores 50–125 μm, not covered by spines. . . . . . . . . . . . . . . . . . . . . . . 4 4. Thallus with large whitish scales extending beyond the lobe margins. Plants found once in superpáramo of the Sierra Nevada de Santa Marta, 4300 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. lamellosa 4. Thallus not with large whitish scales extending beyond lobe margins . . . . . . 5 5. Thallus with numerous whitish dots. Plants endemic to the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. albopunctata 5. Thallus without whitish dots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Thallus with papilla-like cells protruding from the thallus margins and apex. [Thallus glaucous green, margins dark violet. Spores wingless. Outer spore surface with 16–25 areoles across the diameter] . . . . . . . . . . . . . . . R. iodocheila 6. Thallus without papilla-like cells protruding from margins and apex . . . . . . . . . 7 7. Epidermis cells persistent, not disintegrating (cross section). Thallus surface irregularly reticulate. Thallus margins with thin, translucent wings. Thallus with large air chambers in (1–)2 layers (cross section). Sporangia bulging ventrally. Spores 50–70 μm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Epidermis cells disintegrating. Thallus surface not reticulate. Thallus margins with or without translucent wings. Thallus without large air chambers. Sporangia not bulging ventrally. Spores larger, 70–130 μm . . . . . . . . . . . . . . 9 8. Thallus lobes 0.7–1.2 mm wide. Spores without or with a rudimentary wing. Plants of Amazonian lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . . R. crassifrons 8. Thallus lobes broader, 1.5–3  mm wide. Spores with a conspicuous, broad, yellow-brown wing. Plants of páramo . . . . . . . . . . . . . . . . . . . R. huebeneriana 9. Subepidermal cells thick-walled (cross section). Spores winged. Plants widespread, bipolar, recorded from high elevations in Peru and Costa Rica; not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . R. sorocarpa Bisch. 9. Subepidermal cells thin-walled (cross section). Spores wingless (winged in R. australis, couplet 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

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10. Cells in the dorsal half of the thallus with two thickened vertical strips in the wall (cross section). Plants of dry Acacia woodlands along the Caribbean coast. . . . 11 10. Thickened vertical strips lacking, all thallus cells thin-walled . . . . . . . . . . . 12 11. Thallus scales extending beyond the thallus margin, clearly visible in dorsal view, dark violet. Thallus lobes 2–3  mm wide. Inner spore surface without areoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. vitalii 11. Thallus scales not extending beyond the thallus margin, not clearly visible in dorsal view, pink or violet. Thallus lobes 1–2(−2.3) mm wide. Inner spore surface with areoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. weinionis 12. Green thallus tissue in only 2 cell layers (cross section). Spores with a broad wing. Plants very rare in the Andes. . . . . . . . . . . . . . . . . . . . . . . . . R. australis 12. Green thallus tissue in more than 2 cell layers (cross section). Spores without wing. Plants common in lowland areas (to ca. 1000 m) . . . . . . . . . . . . . . . . 13 13. Thallus lobes very narrow, 0.7–1 mm wide. Green thallus tissue in 2–4 cell layers (cross section). Thallus underside violet brown. Plants endemic to the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. howellii 13. Thallus lobes broader. Green thallus tissue in 6–10 cell layers. Thallus underside green or violet brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Ventral scales pink or pink-violet, bordered by 2 rows of hyaline cells. Outer spore surface with 12–16 areoles across the diameter. Plants endemic to the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. ianthina 14. Ventral scales hyaline or brown-violet, not bordered by 2 rows of hyaline cells. Outer spore surface with less than 12 areoles across the diameter. Plants widespread. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15. Outer spore surface with (8–)9–11 areoles across the diameter. Thallus with a 3–4-cell-wide hyaline margin. Ventral scales hyaline. Plants dioicous, occurring in rather dry lowland areas. . . . . . . . . . . . . . . . . . . . . . R. planobiconvexa 15. Outer spore surface with 4–7 areoles across the diameter. Thallus without hyaline margin. Ventral scales brown-violet. Plants monoicous, occurring in rather humid lowland areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. subplana 1. Riccia albopunctata Jovet-Ast On volcanic rock and soil, 1100  m. Ecuador: Ga (Isabela: volcán Alcedo; Bischler-Causse et al. 2005). General distribution: southeastern Brazil, northern Argentina (Misiones), Galápagos Islands. Ricca albopunctata (subg. Riccia) is an uncommon neotropical species that is distinguished by the presence of scattered dead cells with whitish contents (“idioblasts”) inside the thallus, visible as whitish dots on the thallus, and the large spores (100–125 μm) with areolate outer surface and a narrow wing. 2. Riccia australis Steph. On moist soil in open places, from sea level to 1800 m. Ecuador: Pi (BischlerCausse et  al. 2005). General distribution: subtropical South America, Peru (Amazonas), northern Andes (Ecuador, Venezuela). Riccia australis (subg. Riccia) is a tropical South American species with a scattered distribution, being common in southeastern Brazil but rare elsewhere. Characteristics are the rather flat, 5–6× wider than high thalli with green dorsal tissue in 2 layers (cross

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Keys and Descriptions

section), small hyaline or violet scales, and large, yellowish-brown spores (95–120 μm) with a broad wing and with papillose-areolate inner and outer surfaces. 3. Riccia crassifrons Spruce

Fig. 4A, B

On soil on periodically inundated river banks, from sea level to ca. 1000  m. Colombia (new): Vau (Río Apoporis, Raudal Yayacopi and vicinity, in mud between rocks of falls, ca. 800 ft., 16 Feb 1952, Schultes & Cabrera 15388, ster., CINC, PC; ibid., Río Ricapuyá, on flat, exposed clay bank, 25 Sept 1952, Schultes & Cabrera 17660, abundantly fertile, c. spor., CINC, PC). Ecuador: MS (Bomboiza, Pérez et  al. s.n., QCA), ZC (Benitez and Gradstein 2011). General distribution: Amazonia (Venezuela, Brazil, Colombia, Ecuador, Peru). Riccia crassifrons (sect. Ricciella) is a rare Amazonian species that is recognized by the 0.7–1.2-mm-wide thalli with a reticulate dorsal surface and with the dorsal thallus edges expanded as thin, translucent wings. The thallus in cross section has air chambers in 2 layers and ventral tissue in 2–3 layers. The sporangia are bulging ventrally, and the spores are 50–70 μm in diameter, reddish-brown, without or with a rudimentary wing and with 6–8 complete areoles across the outer and inner spore surface; the areoles on outer and inner surface are similar in size. 4. Riccia fluitans L. Riccia stenophylla Spruce (fide Rabeau et al. submitted)

Fig. 4N

On moist sandy or clayey soil, mostly in cultivated areas, in gardens, plantations, on trails and at springs, rarely floating on stagnant water, 400–2000 m. Colombia: Cho (road Tutunendo to El Carmen, Gradstein 9033, COL, GOET), Mag (Gradstein et al. 2016), Met (Schultes 11595, CINC, PC). Ecuador: MS, Tu, ZC. General distribution: widespread in the Holarctic region, scattered in the tropics, reaching southward to South Africa (Rabeau et al. submitted). Riccia fluitans (sect. Ricciella) is a widespread species that has been little collected in Colombia and Ecuador and should be more common. It is distinguished by the very narrow, linear thallus lobes (0.3–0.6 mm wide), the reticulate dorsal surface of the thallus with discrete pores, and the presence of air chambers in 1–2 layers. The spores are relatively small, 60–75 μm in diameter, with 4–7 complete or incomplete areoles across the diameter on the outer surface. The species is monoicous or dioicous and is often sterile in Colombia and Ecuador. Riccia fluitans was called R. stenophylla in tropical America (Bischler-Causse et al. 2005), but the latter species, originally described from Paraguay, is a synonym of R. fluitans according to Rabeau et al. (submitted). 5. Riccia howellii M.Howe

Fig. 4C, D

On soil in dry coastal vegetation, often along trails, 50–250 m. Ecuador: Ga (Bischler-Causse et  al. 2005). General distribution: only known from the Galápagos Islands where it is widespread. Riccia howellii (subg. Riccia) is a common Galápagos endemic that is distinguished by small thalli (lobes 0.7–1 mm wide) with a conspicuous hyaline border and with the green thallus tissue in 2–4 cell layers (cross section) and large wingless spores (95–130 μm in diameter). The species is close to R. planobiconvexa, but the latter species has broader thallus lobes (1.5–3 mm wide) and thicker green thallus tissue (8–10 cell layers thick) and is dioicous (R. howellii is monoicous).

Fig. 4 (A and B) Riccia crassifrons. (A) Habit with sporophytes, dorsal view. (B) Cross section of thallus. (C and D) Riccia howellii. (C) Habit with sporophytes, dorsal view. (D) Cross section of thallus. (E–G) Riccia iodocheila. (E) Portion of thallus, dorsal view. (F) Cross section of thallus. (G) Scale margin, showing large papilliform cells. (H and I) Riccia lamellosa. (H) Habit, dorsal view. (I) Cross section of thallus. (J–L) Riccia membranacea. (J) Habit with sporophytes, dorsal view. (K) Cross section of thallus. (L) Cross section of thallus with sporophyte. (M) Riccia planobiconvexa, cross section of thallus. (N) Riccia fluitans, habit. (O–Q) Riccia vitalii. (O) Portion of thallus. (P) Vertical strips of thickening in the dorsal half of the thallus (cross section). (Q) Cross section of thallus. (All from Bischler-Causse et al. 2005)

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6. Riccia huebeneriana Lindenb. On soil along rivulet in páramo, at the edge of remnant Polylepis forest, ca. 3650  m. Ecuador (new): Na (páramo de Oyacachi, Gradstein et  al. 12791, det. C. Reeb, QCA, PC). General distribution: widespread in the Holarctic region; new to Ecuador. Riccia huebeneriana (sect. Ricciella) is a widespread north-temperate species that is recorded here from the first time from the tropics. Its occurrence in the páramo region reflects the preference of the species for cool climates; indeed, amost all temperate species of Colombia and Ecuador are restricted to páramo (Table  3). Characteristic features of the species are (1) thallus large and very flat, forming rosettes of 1–2 cm in diameter, lobes 1.5–3 mm wide, dorsal thallus surface glaucous green when fresh, pale green to whitish when dry, irregularly reticulate, pores inconspicuous, thallus edges with 2–3 cells wide, translucent wings made up of 1 layer of very large air chambers; (2) ventral thallus surface green, scales absent; (3) thallus cells thin-walled, epidermis cells strongly convex, not disintegrating; (4) thallus in cross section with large air chambers in (1–)2 layers and with small-celled ventral tissue in 4–5 layers; (5) sporangia numerous per thallus, bulging ventrally and dorsally; (6) mature spores ca. 65 μm in diameter, brown with a broad yellow-brown, crenate wing; and (7) outer and inner spore surfaces with 5–7 areoles across the surface, the areoles mostly complete, few incomplete ones, margins of areoles brown. The Ecuadorian population of R. huebeneriana stands out by the abundance of sporangia, which occur throughout the thallus and, when mature, are prominent both dorsally and ventrally. Riccia huebeneriana is very similar to R. crassifrons; both species have air chambers in 2 layers and hyaline thallus wings and lack ventral scales. Also, spore size and the number of areoles across the distal and proximal surface are the same in the two species. Riccia crassifrons differs, however, in the much smaller thalli (lobes maximally 1.2 mm wide), which do not form rosettes, the ventral tissue in only 2–3 layers and the spores without wing. Moreover, R. crassifrons is a lowland species. 7. Riccia ianthina Jovet-Ast On volcanic rock near a pool, 650  m. Ecuador: Ga (San Salvador; BischlerCausse et al. 2005). General distribution: only known from the type from the Galápagos Islands. Riccia ianthina (subg. Riccia) is a very rare Galápagos endemic that is recognized by the rather narrow thalli, 1.5–3× wider than high, with green tissue in 6–9 layers (cross section), pink or pink-violet ventral scales that are bordered by 2 rows of hyaline cells, and large, wingless spores (95–110 μm) with 12–16 areoles across the inner and outer surfaces. The edges of the areoles are covered by small turbercles. 8. Riccia iodocheila M.Howe Fig. 4E–G Riccia atromarginata Levier subsp. iodocheila (M.Howe) R.M.Schust. On soil in dry coastal vegetation, sea level to 100 m. Ecuador: Ga (San Cristóbal; Bischler-Causse et al. 2005). General distribution: Galápagos Islands (type) and subtropical regions of southwestern USA, southern Brazil, and northern Argentina. Riccia iodocheila (subg. Riccia) is an amphitropical New World species that occurs disjunct on the Galápagos Islands (Gradstein 2017a). Characteristics are (1)

Ricciaceae

65

scattered papillae on the thallus surface, especially near apex and on thallus margins; (2) thallus surface bluish-green, flanks dark violet; and (3) spores wingless, 70–100 μm, outer surface with numerous (16–25) small areoles across the diameter. 9. Riccia lamellosa Raddi Riccia indusiata S.Winkl.

Fig. 4H, I

On soil in superpáramo, 4200  m. Colombia: Mag (Winkler 1976 as R. indusiata). General distribution: widespread in Mediterranean-subtropical regions of the world and occasionally at high elevation (above 3000 m) in tropical South America and Africa (Gradstein 2017a). Riccia lamellosa (subg. Riccia) is a widespread amphitropical species that is readily distinguished by the numerous large, whitish scales that extend beyond the lobe margins. The occurrence of R. lamellosa at 4200 m in the Sierra Nevada de Santa Marta, where it was discovered by Winkler (1976) and described as R. indusiata, is remarkable. 10. Riccia limicola Jovet-Ast On moist soil on bank of pools in the evergreen woodland zone and pampa, ca. 400–700  m. Ecuador: Ga (Santa Cruz; Bischler-Causse et  al. 2005). General distribution: only known from the Galápagos Islands. Riccia limicola (sect. Ricciella) is only known from the Galápagos Islands where it was first recorded as R. fluitans. The species is morphologically similar to the widespread R. fluitans, but the spores in R. limicola are larger (80–95 μm) and the areoles on the outer surface more numerous (7–9 across the diameter). 11. Riccia membranacea Gottsche & Lindenb. Riccia amazonica Spruce Riccia echinatispora Schiffn.

Fig. 4J–L

On moist soil in open places, along trails, in gardens, on the bank of ponds, etc., 200–1800  m. Colombia: San. Ecuador: Bo (Gradstein 9914, GOET), Pi, ZC (Gradstein 12104, PC). General distribution: tropical America, tropical Africa. Riccia membranacea (subg. Leptoriccia) is a common Afro-American species that differs from all other neotropical Riccia species by the thin, 2–3 cells thick thallus with lobes widened toward the apex and small brownish spores (40–65  μm) which are densely covered by sharp spines. None of the other neotropical Riccia species has spinose spores (Bischler-Causse et al. 2005). In its thin thallus widened toward the apex and spinose spores, R. membranacea resembles Cyathodium but is readily separated from Cyathodium by the sporangia inside the thallus (not on the lower surface) and the lack of elaters. 12. Riccia planobiconvexa Steph.

Fig. 4M

On exposed soil in rather dry lowland areas, from sea level to ca. 1000  m. Colombia: Mag (Gradstein et  al. 2018c). Ecuador: Ga (Cerro Azul; BischlerCausse et  al. 2005). General distribution: Costa Rica, northern Colombia, Ecuador (Galápagos Is.), subtropical South America. Riccia planobiconvexa (subg. Riccia) is a rather robust South American species with green, dioicous thalli with 3–4-cell-wide hyaline borders and thallus lobes to 3  mm wide and 3–5× wider than high, forming large, gregarious patches. The

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thallus cells are uniformly thin-walled, the epidermis cells are mostly disintegrated, the ventral scales are usually inconspicuous and hyaline, and the spores are dark reddish-brown, wingless, 70–130 μm, with 8–11 areoles across the diameter on the outer surface. The species has been very little collected and should be more common. Riccia planobiconvexa may be confused with R. subplana, but the latter species differs in having spores with fewer areoles (4–7) across the outer surface, thalli without hyaline borders and brownish-violet scales. Moreover, R. subplana is monoicous. Riccia planobiconvexa is also close to R. howellii from the Galápagos Islands; for differences see under the latter species. 13. Riccia subplana Steph. On exposed soil in lowland areas, from sea level to ca. 1000 m. Ecuador (new): MS (Bomboiza, Pérez, Gradstein & Deleg 11562, QCA, PC), Or (Yasuni National Park, Gradstein, Deleg & Bader 12810 (cf.), 12813 (cf.), HUTPL, PC, without spores), Su (Deleg s.n. (cf.), HUTPL, without spores); to be expected in Colombia. General distribution: Central America, Lesser Antilles, northern South America (Bischler-Causse et al. 2005). Riccia subplana (subg. Riccia) is close to R. planobiconvexa but differs from the latter by spores with fewer areoles (4–7) across the outer surface, thallus monoicous and without hyaline borders, and scales brown-violet. The species has been little collected and should be more common. 14. Riccia vitalii Jovet-Ast

Fig. 4O–Q

On soil in dry coastal cactus-Acacia woodland and in gardens, 50–1000  m. Colombia: Mag (Gradstein et al. 2016). Ecuador (new): MS (Gradstein & Pérez, QCA). General distribution: Costa Rica, northern Colombia, eastern Brazil, Paraguay. Riccia vitalii (subg. Riccia) is a characteristic species of dry caatinga in eastern Brazil that has been found once in Colombia (Magdalena). The species is recognized by the narrow, 1–1.5-mm-wide thallus lobes with a dark green upper side and large, dark violet scales, which extend beyond the thallus margin and are bordered by large, hyaline cells. Dry thalli are inrolled and dark purplish due to the scales covering the inrolled thallus. A further characteristic of the species is the occurrence of long, paired, vertical strips of thickening in the dorsal half of the thallus (cross section), running closely parallel through the cells (see Bischler-Causse et al. 2005, Fig. 54N). The latter feature is otherwise only seen in R. weinionis (see below) and the rare Riccia ridleyi A.Gepp from Fernando Noronha (Brazil), Peru, and northern Venezuela. In the latter two species, however, the scales do not extend beyond the thallus margins, and the inner and outer spore surfaces are areolate. In R. vitalii, the inner spore surface lacks complete areoles (Bischler-Causse et al. 2005). 15. Riccia weinionis Steph. On soil in dry coastal cactus-Acacia woodland, 50 m. Colombia: Mag (Winkler 1976). General distribution: tropical America. Riccia weinionis (subg. Riccia) is a widespread neotropical species that is recognized by (1) presence of vertical strips of thickening in the dorsal half of the thallus (cross

Targioniaceae

67

section), running closely parallel through the cells (see R. vitalii: Fig. 4P, Q); (2) ventral scales not extending beyond the thallus margins; and (3) outer spore surfaces areolate. Riccia weinionis is closely related to R. vitalii and grows in the same habitat. In R. vitalii, however, the spores are not areolate, and the ventral scales are clearly extending beyond the thallus margin. Like R. vitalii, R. weinionis is undercollected and should be looked for in dry lowland areas with scrubby vegetation. Ricciocarpos Corda Thallus yellow-green to dark green, often with purple margins, medium-sized, to 1 cm long, the segments 3–6 mm wide, usually forming rosettes floating on water; dorsal surface of thallus with a median groove, finely reticulate and with small pores, dorsal epidermis with scattered oil cells. Ventral scales large, ligulate, violet, in several diffuse rows, with oil cells. Air chambers large, in several layers (almost the entire thallus composed of air chambers). Monoicous. Gametoecia and sporophyte as in the family. A genus with a single species. For recognition see under the species. Literature: Bischler-Causse et al. (2005) 1. Ricciocarpos natans (L.) Corda

Fig. 3J, K

Floating on stagnant water, occasionally terrestrial on muddy banks of ponds and lakes that have dried up, ca. 2500 m. Colombia: Cun (near Bogotá; Gradstein and Uribe 2016a). General distribution: widespread in temperate regions of the Northern and Southern Hemisphere, scattered at high elevation in the tropics. Ricciocarpos natans is a subcosmopolitan species that is known from one locality in Colombia. The species is aquatic and is readily distinguished by the dark green thalli (to 1 cm long, 3–6 mm wide) with purplish margins, floating on water. The dorsal thallus surface is distinctly reticulate with small whitish pores, and the ventral surface has large, ligulate, violet scales with numerous oil cells.

Targioniaceae A family with a single genus. Targionia L. Thallus leathery, narrowly oblong to ligulate, deep green, margins and ventral surface black or blackish-purple, dry thalli entirely black and narrowly linear due to inrolling of the thallus, thalli to 4 cm long, 2–4 mm wide. Dorsal surface of thallus finely reticulate, with small whitish pores, the pores sometimes star-shaped, epidermal walls smooth or papillose, with bulging trigones. Thallus underside with dark purple scales in 2 rows, scales with oil cells and marginal slime papillae, each scale with a lanceolate appendage. Air chambers in 1 layer, with filaments. Rhizoids smooth and tuberculate. Monoicous or dioicous. Antheridia sunken in the thallus

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Keys and Descriptions

surface or in small receptacles on short branches. Sporophytes 1–3 in a large, purplish, mussel-like involucre, projecting forward from beneath the thallus apex and opening by 2 valves. Capsule opening by an operculum. Spores large, 50–100 μm. Vegetative reproduction probably by fragmentation of dry thalli. A small subtropical-Mediterranean genus (3 spp.), one species in Colombia and Ecuador. A further species, T. stellaris Hässel, has been recorded from the Galápagos Islands, but the material was misidentified and belongs to T. hypophylla (BischlerCausse et al. 2005). The main features of the genus Targionia are (1) thallus narrowly oblong to ligulate, leathery, deep green (becoming black when dry due to inrolling of the thallus), with simple pores; (2) thallus margins and underside blackish-purple; (3) air chambers in 1 layer, with filaments; and (4) sporophyte in a large, swollen, purplish involucre beneath the thallus apex; capsule opening by an operculum. Literature: Bischler-Causse et al. (2005) 1. Targionia hypophylla L.

Fig. 3L, M

On soil and rock in rather mesic areas, usually on road banks, in partial shade, 630–3800 m. Colombia: Mag, Met, Qui (Gradstein s.n., HUQ), San, Tol. Ecuador: Ch, Ga. General distribution: subcosmopolitan. Targionia hypophylla is readily distinguished by the generic characters given above. Class JUNGERMANNIOPSIDA (Foliose and Simple Thalloid Liverworts) Plants foliose or thallose, prostrate, ascending or erect rhizomes present or absent. Stems or thallus with or without central strand. Branches lateral or ventral, originating from leaf initial cells or from stem or thallus cells, when from stem originating from inner stem cells, not from the epidermis; collar present or lacking. Leaves (when present) in 2–3 rows: 2 lateral rows and 0–1 ventral row (= underleaves), each leaf originating from 1–3 initial cells, undivided or divided. Thallus (when present) homogeneous in cross section, air chambers and pores lacking; ventral scales absent. Cells with or without trigones, usually with oil bodies. Underleaves present or absent, when present usually smaller than the lateral leaves. Rhizoids usually present on the ventral side of the plant, always with smooth walls. Antheridia in leaf axils or on dorsal surface of thallus. Archegonia terminal on long or short shoots, in foliose plants surrounded by bracts and often by bracteoles, in thallose plants protected by an involucre or scale, rarely naked. Sporophytes surrounded by the calyptra and a perianth or fleshy perigynium. Setae well developed, 1–10 cm long, thin to very thick. Capsules globose to cylindrical, opening by (2–)4 valves, rarely irregularly, capsule wall 2–10-layered. Elaters present. The largest class of the liverworts, with ca. 5000 species worldwide, about 795 species in Colombia and Ecuador, in 127 genera and 32 families. The class is divided into four subclasses, Pelliidae, Metzgeriidae, Pleuroziidae, and Jungermanniidae.

Jungermanniopsida

69

Key to the subclasses of Jungermanniopsida of Colombia and Ecuador 1. Plants with stem and leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants thalloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Plants robust, to 10  cm long and 4–7  mm wide, reddish or purplish. Leaves deeply concave, strongly clasping the stem, with a white leaf border and undivided, or without white border and bifid (when bifid divided into a large ventral lobe and a smaller dorsal lobe, the smaller lobe usually turned into a sac). Underleaves absent. Perianth terete or plicate or both . . . Pleuroziidae (p. 121) 2. Plants different. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaves undivided, succubous. Leaf base several layers of cells thick. Cells thinwalled, without trigones. Rhizoids violet or colorless. Archegonia and antheridia on dorsal surface of stem. Plants on soil or rock . . . . . . . Pelliidae (p. 69) 3. Leaves divided into lobes or undivided, incubous, succubous, or transverse. Leaf base only 1 layer of cells thick. Cells thin-walled or thick-walled, often with trigones. Rhizoids not violet. Archegonia on apex of stem or short branch, antheridia in leaf axils. Plants on bark, living leaves, rotten wood, rock and soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jungermanniidae (p. 123) 4. Thallus unbranched or with a few short, ventral-intercalary branches. Thallus  margins, at least near the apex, with long, sausage-shaped slime papillae. Archegonia and antheridia on leafy branches. . . . . . . . . . . . . . . . . . . . . . . . Jungermanniidae: Pteropsiella (p. 238) 4. Thallus variously branched. Thallus margins without sausage-shaped slime papillae. Archegonia and antheridia on the thallus surface or in cavities, not on leafy branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Thallus midrib with a central strand. Gametoecia on the dorsal side of the midrib. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pelliidae: Pallaviciniaceae (p. 77) 5. Thallus midrib without central strand or midrib absent. Gametoecia on the ventral side of the midrib or at the thallus margin. . . . . . . . . Metzgeriidae (p. 86) Subclass Pelliidae (Simple Thalloid Liverworts I) Plants thalloid or foliose, growing with a two-sided apical cell, divided into two lateral merophytes, ventral merophyte absent. Thallus (when present) differentiated into a midrib and thinner lateral wings, midrib frequently with a central strand, wings undivided, lobed, or deeply dissected. Leaves (when present) undivided, originating from 1 initial cell, succubous, several layers of cells thick toward the base; underleaves lacking. Cells thin-walled, usually without trigones; oil bodies usually present. Rhizoids with smooth walls. Antheridia and archegonia usually on the surface of the axis, naked or surrounded by an involucre or scales; archegonia not arising from the apical cell of thallus (plants anacrogynous). Sporophytes surrounded by a calyptra and, usually, by an involucre. Setae well developed, 1–10 cm long, thin to very thick. Capsules globose to cylindrical, opening by 2–4 valves or irregularly, wall 2–6-layered.

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Contains 21 species in Colombia and Ecuador, in 5 genera and 3 families. A further genus, Hymenophyton (Hymenophytaceae) from Chile and Australasia  – resembling Jensenia (Pallaviciniaceae) but with gametoecia and sporophytes located on the ventral side of the thallus instead of on the dorsal side – has been recorded from Colombia and Costa Rica. The Costa Rican material was misidentified and belongs to Jensenia erythropus (Gradstein et al. 1994), but the identification of the specimen from Colombia (J. Weir, NY) was correct (Evans 1925). As no further records of Hymenophyton are known from Colombia, the material may have been mislabeled. Key to the families of Pellidae of Colombia and Ecuador 1. Plants thalloid, sometimes deeply divided into segments (Symphyogyna sect. Lobatae) but true leaves absent. (Thallus simple or dichotomous, with a thick midrib and broad wings. Midrib with a central strand). . . . . . Pallaviciniaceae 1. Plants with leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants 4–10  mm wide, deep green to glaucous. Leaf margins flat, entire. Rhizoids pale brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pelliaceae 2. Plants 2–3 mm wide, pale green, usually forming rosettes. Leaf margins undulate, usually sinuate or toothed. Rhizoids purplish . . . . . . . Fossombroniaceae

Fossombroniaceae by Barbara Crandall-Stotler and S. Robbert Gradstein A monogeneric family, containing only the genus Fossombronia (Stotler et al. 2016) Fossombronia Raddi Austrofossombronia R.M.Schust. Plants differentiated into stem and leaves, prostrate, simple or forked, often growing in rosettes. Leaves succubous, margins shallowly lobed and undulate, sometimes plicate, 2–6 cell layers thick at the base, with the multistratose portion usually extending to about 1/4 the leaf length. Central strand and ventral scales lacking. Cells large, thin-walled, with numerous minute, homogeneous oil bodies. Rhizoids usually deep violet. Gametangia scattered on the dorsal surface of the axis, usually naked, not covered by an involucre or scales. Sporophytes surrounded by a bell-shaped caulocalyx with a wide, open mouth. Setae rather short, thick. Capsules spheroidal, rarely ellipsoidal, opening irregularly, wall 2-layered. Spores ca. 35–50  μm in diameter, surface ornamented by lamellae, vermiculae, or cristae. Elaters with 2–3 spirals, free inside the capsule. Vegetative reproduction occasionally by apical tubers or by leafy buds sprouting from the stem or leaves (Fig. 5A–C).

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71

Fig. 5 (A–C) Fossombronia sp. (A) Habit with sporophyte. (B) Leaf. (C) Leaf cells. (D) Fossombronia peruviana, habit showing antheridia. (E–H) Jensenia spinosa. (E) Female thallus (i = involucre, p = pseudoperianth). (F) Male thallus. (G) Cross section of thallus, showing central strand. (H) Thallus margin with tooth. (I) Jensenia florschuetzii, teeth on thallus margin. (J–L) Pallavicinia lyellii. (J) Habit with gynoecia, dorsal view. (K) Habit with androecia, dorsal view. (L) Thallus margin with slime hair. (I redrawn from van der Gronde 1980; rest from Gradstein et al. 2001)

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Keys and Descriptions

About 80 species worldwide (Stotler et  al. 2016), 7 species in Colombia and Ecuador. The main characters of Fossombronia are (1) succubous leaves with lobedundulate margins and surface and thickened leaf bases; (2) violet rhizoids; (3) gametangia “naked” on the dorsal side of the stem, not covered by an involucre or scale; (4) sporophyte surrounded by a caulocalyx; (5) capsules spheroidal to short-ellipsoidal, dehiscence irregular; and (6) spores ca. 30–50  μm in diameter, surface coarsely sculptured. For species identification, material with mature spores is usually required. Literature: Freire (2002), Crandall-Stotler and Gradstein (2017), Crandall-Stotler et al. (2019) 1. Plants large, 5–7 mm wide, semi-aquatic, in mires and along rivers, dioicous. Leaf surface usually deeply folded (± plane in plants from very wet habitats). Seta more than 12 cells in diameter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants smaller, 1.5–4  mm wide, not semi-aquatic, on soil or rotten wood, monoicous or dioicous. Leaf surface not deeply folded. Seta less than 10 cells in diameter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Plants without any reddish or brownish pigmentation. Midleaf cells 40–70 × 35–40 μm. Capsules spheroidal. Spores 28–32 μm, outer surface vermiculate, without areoles. Plants on limestone rock in river, 1100 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. jostii 2. Plants with or without reddish or brownish pigmentation. Midleaf cells (50–) 70–120  ×  40–60  μm. Capsules longer than wide, ellipsoidal. Spores (31–) 35–50  μm in diameter, outer surface lamellate-reticulate, with complete or incomplete areoles. Plants from páramo and puna above 3000 m . . . . . . . . . . 3 3. Spores (31–)35–39  μm in diameter, outer surface lamellate-reticulate, with incomplete areoles. Thallus 3-6 cells thick in the lower 1/3, unistratose above. Plants usually with reddish or brownish pigmentation . . . . . . . . . F. peruviana 3. Spores larger, 40–50 μm in diameter, outer surface fully reticulate, with complete areoles. Thallus 2-3 cells thick in the lower 1/4 (unistratose above). Plants without any reddish or brownish pigmentation; recorded from Mexico and Venezuela, not yet known from Colombia and Ecuador. . . . . F. delgadilloana Crand.-Stotl. et al. 4. Leaves longer than wide to subquadrate. Leaf margin entire, sometimes bordered with tangentially elongated cells. Outer spore surface lamellate or reticulate. Plants monoicous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Leaves wider than long or as wide a long [when in doubt try both leads]. Leaf margin entire or toothed, never bordered. Outer spore surface lamellate or lamellate-reticulate. Plants monoicous or dioicous. . . . . . . . . . . . . . . . . . . . . 6 5. Tubers occasionally present at shoot apex. Outer spore surface lamellate, with 24–32 high, narrow lamellar projections around the margin . . . . F. fernandeziensis 5. Tubers never present. Outer spore surface ± regularly reticulate, with 4–6(−7) areoles across the diameter. Spore margin without or with rudimentary lamellar projections, more or less winged. Plants common . . . . . . . . . . F. porphyrorhiza 6. Tubers frequently present at shoot apex. Leaves conspicuously wider than long. Outer spore surface lamellate-reticulate, covered by long, simple or furcate lamellae and a few small central areoles. Spore margin with 15–20 short, conical lamellar projections. Plants dioicous . . . . . . . . . . . . . . . . . . . . F. lamellata

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6. Tubers absent. Leaves wider than long to subquadrate. Outer spore surface lamellate (occasionally with a few central areoles). Spore margin with 15–28 truncate, lamellar projections. Plants monoicous . . . . . . . . . . . . . . . . . . . . . . 7 7. Leaves conspicuously wider than long, with the margins entire and the basal portion 4–5 cell layers thick (cross section). Spore margin with 22–28 lamellar projections, elaters with 3 or more spiral bands . . . . . . . . . . . . . F. crassifolia 7. Leaves subquadrate, with the margins irregularly toothed and the basal portion 2–3 cell layers thick. Spore margin with fewer than 20 lamellar projections, elaters with 2(−3) spiral bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. pusilla 1. Fossombronia crassifolia Spruce

Fig. 6A, B

On moist soil in the (sub)montane belt, 500–2100  m. Ecuador: MS, Tu, ZC. General distribution: tropical America. Fossombronia crassifolia is an uncommon neotropical species that is recognized by (1) leaves wider than long, margin undulate-crispate, without border, basal portion of leaf 4–5 cell layers thick; (2) tubers absent; (3) plants monoicous, protandrous; (4) outer spore surface irregularly covered by broken lamellae that are here and there connected but are not forming small areoles, spore margin with 22–28 lamellar projections; and (5) elaters with 3–5 spirals. 2. Fossombronia fernandeziensis Steph. Fossombronia columbica S.W.Arnell, nom. inval.

Fig. 6C, D

On moist soil on earth banks in upper montane forest regions and páramo, 2350–3560  m. Colombia: Boy (as F. columbica nom. nud., Sierra Nevada del Cocuy, 3700  m, Grubb & Guymer B105, det. V.  Freire, BM), Cun, Nar, Tol (Gradstein et al. 2018c. Ecuador: Ch, Pi (Crandall-Stotler and Gradstein 2017). General distribution: tropical Andes (Bolivia to Colombia), Juan Fernández Islands. Fossombronia fernandeziensis is a rare Andean species that is characterized by (1) leaves distinctly longer than wide, plane or slightly undulate, margins entire, bordered by tangentially elongate cells; (2) tubers sometimes produced at shoot apex; (3) plants monoicous, antheridia and archegonia intermixed; (4) outer spore surface densely lamellate, margin with 24–32 high, narrow lamellar projections; and (5) elaters with 2 spirals. 3. Fossombronia jostii Crand.-Stotl. & Gradst. 

Fig. 6E, F

On soil over periodically inundated limestone rock in a river canyon in submontane rainforest, 1100 m. Ecuador: Pa (canyon of Río Anzu; Crandall-Stotler and Gradstein 2017). General distribution: only known from the type locality in Ecuador. Fossombronia jostii is a robust Ecuadorian endemic that habitually resembles F. peruviana. Both species share a semi-aquatic habitat; relatively large plant size (5–6 mm wide); fleshy stems with a dense mat of short, violet rhizoids on the ventral side; undulate-plicate leaves that are much wider than long, inserted up to the dorsal midline of the stem and 2–3 cells thick near the base; a massive seta that is

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Fig. 6  Fossombronia spores, distal views: SEM micrographs (A, C, E, G, I, K, M–O) and optical micrographs (B, D, F, H, J, L, P). (A and b) Fossombronia crassifolia. (C and D) Fossombronia fernandeziensis. (E and F) F. jostii. (G and H) Fossombronia lamellata Steph. (I and J) Fossombronia peruviana Gottsche. (K and L) Fossombronia porphyrorhiza (Nees) Prosk. (M–P) Fossombronia pusilla. (A from Lautner 01-41/42, GOET; B from the lectotype; C from Benavides 4064, F-ABLS; D from from the lectotype; E, F from the type; G, H from the lectotype; I, J from Stotler & Crandall-Stotler 4187, F-ABLS; K, L from Stotler & Crandall-Stotler 3993, F-ABSH; M from Cargill 451, F-ABSH; N–P from the lectotype of F. tenuifolia). Scale bars: A, K  =  7.0  μm; D = 8 μm; C, D = 6 μm; E = 4.8 μm; F = 5.5 μm; G, H, M, O = 7.5 μm; I = 5 μm; J = 5.8 μm; L = 14 μm; N = 8.2 μm; P = 7.2 μm

more than 12 cells in diameter; and dioicous sexuality. Fossombronia jostii differs from F. peruviana in several important respects, however (see Crandall-Stotler and Gradstein 2017): (1) plants glossy light green to glaucous green when fresh, without any trace of reddish or brownish pigmentation (with reddish or brownish pigmentation in F. peruviana); (2) cells in midleaf 40–70 × 35–40 μm ([50–]70–120 × 40–60 μm in F. peruviana); (3) caulocalyx without stalk (stalked in F. peruviana); (4) seta 13 cells thick (15–20 cells thick in F. peruviana); (5) capsules rounded (longer than

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wide in F. peruviana); (6) spores 28–32 μm in diameter ([31–]35–39 μm in F. peruviana); and (7) outer spore surface vermiculate (lamellate and with incomplete areoles in F. peruviana). The two species furthermore have very different elevational preferences: F. jostii occurs in submontane rainforest at 1100  m, whereas F. peruviana grows in alpine tundra (páramo, puna) above 3000 m. Fossombronia delgadilloana from high elevation in Mexico and Venezuela (Crandall-Stotler et al. 2019) also resembles F. jostii but differs in larger leaf cells, ellipsoidal capsules, larger spores (40–50  μm in diameter), and reticulate outer spore surfaces, with complete areoles. 4. Fossombronia lamellata Steph. Fossombronia tuberifera Goebel (fide Freire 2002)

Fig. 6G, H

On moist soil, also on rotten wood and rock, 100–3700  m. Colombia: Cho (Tutunendo, Gradstein 9039a, COL, GOET), Cun (páramo de Palácio, Cleef 3966, COL), Mag (Winkler 1976), Vau (Río Apoporis, Schultes & Cabrera 11967, 11973, 11974, “cf.”, CINC, PC). General distribution: scattered in tropical America. Fossombronia lamellata is a widespread neotropical species that is distinguished by (1) plants dioicous, leaves wider than long, margins plane or slightly undulate, entire, without a border; (2) tubers frequently produced at shoot apex; (3) outer spore surface with numerous subparallel, simple, or furcate lamellae, the lamellae anastomosing in the center and forming several small areoles (2–6), spore margin with 15–20 lamellar projections, often with an incomplete wing; and (4) inner spore surface covered by short lamellae or cristae (Freire 2002). 5. Fossombronia peruviana Gottsche & Hampe Figs. 5D, 6I, J Austrofossombronia peruviana (Gottsche & Hampe) Crand.-Stotl., Stotler & A.V.Freire Fossombronia herzogii Goebel Fossombronia ptychophylla Spruce On peaty soil along streamlets and in mires in páramo and puna, sometimes growing submerged, often forming extensive, pure mats, 3300–4100 m. Colombia: Boy, Cau, Cun. Ecuador: Ca, Ch, Co, Lo, MS, Na, Pi (Crandall-Stotler and Gradstein 2017). General distribution: tropical Andes (Venezuela to northern Chile). Fossombronia peruviana is a characteristic liverwort of mires and streamlets in the high Andes that is recognized by (1) large, light green plants (5–6 mm wide) with usually some reddish or brownish pigmentation; (2) densely imbricate, crispedplicate leaves, each leaf usually having four to six folds, and the numerous short, violet rhizoids; (3) seta relatively thick, 15–20 cells in diameter; (4) capsules longer than wide, ellipsoidal; and (5) outer spore surface lamellate and with incomplete areoles. The leaves are up to 4–6 cells thick in the lower third of the leaf in plants from rather dry habitats but thinner, maximally 3 cells thick, in plants from wet habitats. The leaves of plants from wet habitats are also less plicate, the leaf cells are a bit smaller on average (50–100 × 40–50 μm in midleaf), and the leaf margins may produce a few small teeth or lobes.

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Fossombronia peruviana may be confused with Noteroclada confluens (Pelliaceae), which occurs in the same habitat. However, the rhizoids in N. confluens are pale-colored (never violet), the leaves are planate (not crisped-plicate), the capsules are globose and produced on a very long seta (to 10  cm long), and the spores are green and very large, to 100 μm in diameter, with an unsculptured surface. Fossombronia peruviana is morphologically close to F. jostii and F. delgadilloana; for differences with F. jostii, see under the latter species. Fossombronia delgadilloana, recently described from Mexico and Venezuela (Crandall-Stotler et al. 2019), differs in thinner and fully planate leaves which are only 2–3 cells thick in the lower 1/4, furthermore absence of any reddish or brownish pigmentation, larger spores (40–50 μm in diameter), and reticulate outer spore surfaces, with complete areoles. 6. Fossombronia porphyrorhiza (Nees) Prosk. Androcryphia porphyrorhiza Nees Fossombronia brasiliensis Steph.

Fig. 6K, L

On moist soil and rock in rather open places, often on road banks or along rivers, from sea level to 2000(−2800) m. Colombia: Cun, Mag, Qui, Vau (Schultes & Cabrera 13874, CINC, PC). Ecuador: EO, Ga (Isabela, San Cristóbal, Santa Cruz), MS, Or (Parque Nac. Yasuni, abundant on road bank near Yasuni station, Oct 2018, Gradstein, Deleg & Bader 12811, QCA), ZC. General distribution: widespread in tropical and subtropical America. Fossombronia porphyrorhiza is the most common and widespread Fossombronia species in the neotropics. The species is distinguished by the rather regularly reticulate spores with 4–6(−7) areoles across the diameter on the outer spore surface and a more or less winged spore margin. The plants are monoicous, with male and female gametangia intermixed, the leaves are subquadrate to wider than long, and the leaf margins are undulate-crispate and without or with a weak border of elongate cells. Tubers are lacking. The species is abundant on the northern slope of the Sierra Nevada de Santa Marta and on road banks between Santa Marta and San Lorenzo (S. R. Gradstein, pers. obs., Nov 2015). Winkler (1976) recorded F. lamellata in the Sierra Nevada de Santa Marta instead of F. porphyrorhiza, but the occurrence of F. lamellata in the area could not be confirmed (Gradstein et al. 2016). 7. Fossombronia pusilla (L.) Nees  Fossombronia tenuifolia Spruce (fide Freire 2002)

Fig. 6M–P

On moist soil over volcanic rock, 1100–3450 m. Ecuador: Ch (volcán El Altar; Spruce 1885 as F. tenuifolia), Ga (summit of volcán Alcedo; Gradstein 2009). General distribution: Europe, western North America, northern Andes, Africa, Australasia. Fossombronia pusilla is a widespread temperate species that is very rare in the neotropics. In the northern Andean region, it is known only from two records from Ecuador. The species is closely similar to F. crassifolia but differs by spores with fewer and more widely spaced lamellae, forming less than 20 (15–19) lamellae projections on the spore margin, and by leaves subquadrate and 2–3 cell layers thick at the base (wider than long and 4–5 cell layers thick in F. crassifolia). Material

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from Ecuador (old road Quito to St. Domingo, Gradstein et al. 6717, U), listed as F. pusilla by Freire (2002), belongs to F. fernandeziensis (see above). Similarly, the records of F. pusilla from Colombia in Freire (2002) refer to F. fernandeziensis. Erroneous Record Fossombronia paranapanema Schiffn. – Colombia: Sierra Nevada de Santa Marta (Winkler 1976). The material belongs to Noteroclada confluens.

Pallaviciniaceae Plants thalloid, 3–9  mm wide, with a distinct, swollen midrib and a unistratose lamina, prostrate or erect, simple or forked, sometimes with ventral branches. Midrib with 1(−3) central strands. Oil bodies finely granular. Dioicous. Gametoecia on the dorsal surface of the midrib, protected by scales or an involucre; antheridia in a long band on the midrib, hidden under laciniate scales; archegonia in small groups on the midrib, surrounded by a scale-like or closed-tubular involucre or both. Sporophytes surrounded by a deeply laciniate pseudoperianth or a fleshy calyptra. Capsules ellipsoidal to narrow-cylindrical, opening into (1–)2(−4) valves. Spores small, unicellular. Elaters with 2–3 spirals, free, not attached to capsule valves. Vegetative reproduction absent in neotropical taxa. About eight genera. The family is mainly characterized by (1) thallus simple or dichotomously branched, with a distinct, swollen midrib; (2) midrib with 1(−3) central strands; (3) capsule elongate, opening by 2 valves (sometimes by 4 imperfect valves, occasionally by only 1 valve); and (4) elaters free, not attached to the capsule valves. 1. Thallus divided into lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symphyogyna 1. Thallus not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Thallus ascending to erect, arising from a stolon-like rhizome. . . . . . . . . . . . 3 2. Thallus prostrate or ascending, stolon-like rhizome absent. . . . . . . . . . . . . . . 5 3. Thallus simple, unbranched. Thallus margin with stalked slime papillae, the stalk 2–3 cells long. Archegonia covered by a circle of free to slightly connate scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pallavicinia 3. Thallus dichotomously branched. Thallus margin with teeth (teeth rarely absent), stalked slime papillae absent. Archegonia covered by a cup-shaped involucre (= circle of highly connate scales all around the archegonia) or by a laciniate scale on one side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Archegonia covered by a cup-like involucre. After fertilization, a long-tubular pseudoperianth (several times longer than the involucre) is developed from within the involucre, containing the young sporophyte. Plants from above 1500 m, common in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jensenia 4. Archegonia covered by a laciniate scale. Pseudoperianth absent. Plants from 500 m up to the páramo. . . . . . . . . . . . . . . . . . . . . . Symphyogyna podophylla

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5. Thallus margins with slime papillae, each slime papilla on a 2–3-cell-long stalk. Archegonia covered by a circle of ciliate-laciniate scales. After fertilization, a long-tubular pseudoperianth (several times longer than the scales) develops, containing the young sporophyte (like in Jensenia). . . . . . . . Pallavicinia 5. Thallus margins without slime papillae or with slime papillae on a 1-cell-long stalk. Archegonia covered on one side by an entire, bifid, or ciliate-laciniate scale. Pseudoperianth absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symphyogyna Jensenia Lindb. Plants pale green to reddish, to 5 cm long, 3–9 mm wide, dendroid, differentiated into an upright stipe and frond arising from a prostrate, stolon-like base, the frond 2–4 × forked, the branches in one plane, flat or deeply concave, winged. Midrib of the branches very broad and thick, to 1/2 branch width, with one central strand of narrow, thick-walled cells. Branch margins entire or toothed. Archegonia in clusters at the lower furcations of the thallus, each cluster of archegionia surrounded by a cup-like involucre with a fringed mouth. Sporophytes surrounded by a long-tubular pseudoperianth several times longer than the involucre. Spore surface cristate, with numerous irregular ridges. A tropical and southern-temperate genus (7 spp.), three species in Colombia and Ecuador. By its dendroid habit, Jensenia is readily separated from other thalloid liverworts except Symphyogyna podophylla. For differences see the key; when sterile, Jensenia and Symphyogyna podophylla cannot always be distinguished with certainty. Literature: van der Gronde (1980), Schuette and Stotler (2005) 1. Thallus margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. difformis 1. Thallus margins toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Teeth at the thallus margin very large, (3–)4–10 cells long and 1–5 cells wide, teeth sometimes fused. Thallus branches of mature female plants strongly inrolled, with overlapping margins. Thallus wings narrowly unistratose over a width of 1–2(−3) cells. Plants rare, in páramo above 3700 m . . J. florschuetzii 2. Teeth at the thallus margin smaller, 1–4(−6) cells long and 1–2 cells wide, never fused, sometimes absent. Thallus branches plane or slightly inrolled, margins not overlapping in mature female plants. Thallus wings broadly unistratose over a width of 3–11 cells. Plants common, 1500–4000 m . J. spinosa 1. Jensenia difformis (Nees) Grolle On soil, tree bases, and rock, 500–3600  m. Colombia: Boy, Hui, Met, Ris (Gradstein and Uribe 2016a). Ecuador: MS (Río Quimi, Pérez et al. 11613, QCA), ZC (Parolly et  al. 2004). General distribution: northern Andes, southeastern Brazil, southern South America, and the subantarctic islands eastward to Marion Island.

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Jensenia difformis is readily recognized by the dendroid thallus with fully entire margins, without teeth or slime hairs. 2. Jensenia florschuetzii Gronde

Fig. 5I

On moist soil in páramo, 3700–4150 m. Colombia: Boy, Cun, Met. Ecuador: Ca (Gradstein 1999), Pi (Burghardt 2019). General distribution: northern Andes. Jensenia florschuetzii is a robust, northern Andean páramo species close to J. spinosa; for differences see the key. The thallus margins in J. florschuetzii are sometimes reddish-brown pigmented. 3. Jensenia spinosa (Lindenb. & Gottsche) Grolle Jensenia erythropus (Gottsche) Grolle Jensenia erythropus var. nobandae Gronde Jensenia wallisii (J.B. Jack & Steph.) Grolle

Fig. 5E–H

On moist soil and rock, 1500–4000 m. Colombia: Ant, Boy, Cas, Cau, Cun, Hui, Mag, Met, Nar, Qui, Ris, San, Tol. Ecuador: Ca, Co, Lo, Na, Pi, ZC. General distribution: tropical America, Africa. Jensenia spinosa is a common and variable Afro-American montane species that is distinguished by the dendroid thalli with subentire to distinctly toothed margins (teeth 1–4(−6) cells long and 1–2 cells wide) and with unistratose portions of the thallus wings rather broad, 3–11 cells wide. Van der Gronde (1980) described two varieties based on Colombian material: var. erythropus with small teeth (1–4 cells long) and var. nobandae with large teeth (4–7 cells long). Jensenia wallisii, described from Colombia and characterized by broad, unistratose wings and small, scattered teeth on thallus margins, was accepted as a separate species by Schuette and Stotler (2005) but is included here in J. spinosa following van der Gronde (1980). Sterile plants of J. spinosa cannot be distinguished with certainty from Symphyogyna podophylla. Pallavicinia Gray Thallus green, about 2–6 cm long, 3–6 mm wide, prostrate to ascending, usually simple, sometimes with ventral branches, rhizome-like base absent or present, short. Midrib ca. 1/8–1/6 of the thallus width, with one central strand of narrow, thick-walled cells. Thallus wings undivided, unbordered or slightly bordered by somewhat narrower cells, margins with slime papillae on a 2–3-cell-long stalk, otherwise entire. Archegonia in clusters scattered on the midrib, surrounded by a circle of free or slightly connate, ciliate-laciniate scales. Sporophytes surrounded by a long, tubular pseudoperianth several times longer than the scales. Spore surface reticulate or punctate. About 12 species worldwide, two in Colombia and Ecuador. Pallavicinia is recognized by the prostrate to ascending, unlobed, ± simple thalli with entire margins, presence of longly stalked slime hairs on thallus margins and gynoecia covered by free or slightly connate, ciliate-laciniate scales.

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Literature: Gradstein et al. (2001) 1. Thalli arising from a short rhizome-like base, prostrate or ascending. Plants from Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sp. 1. Thalli not arising from a rhizome-like base, prostrate . . . . . . . . . . . . . P. lyellii 1. Pallavicinia lyellii (Hook.) Gray

Fig. 5J–L

On rotten wood, tree base, litter, humus and soil over rock, in ± undisturbed lowland, and montane rainforest, from sea level to 2800 m. Colombia: Ama, Boy, Ces, Cho, Hui, Nar, Ris, San. Ecuador: Lo, MS, Na, Pa, Su, Tu, ZC. General distribution: widespread in the Holarctic region, less common in the tropics. Female plants of P. lyellii are immediately recognized by the group of free or slightly connate, ciliate-laciniate scales surrounding the archegonia. After fertilization, a longly tubular pseudoperianth is developed from within the scales, surrounding the sporophyte. Sterile plants are very similar to unlobed species of Symphyogyna, especially S. brasiliensis and S. trivittata, but differ from Symphyogyna in the presence of longly stalked slime papillae on thallus margins, each with a 2–3-celllong stalk. 2. Pallavicinia sp. On trunk bases and rock in Amazonian rainforest, 250 m. Colombia: Vau (Río Apoporis, vicinity of Soratama, Schultes & Cabrera 12540, 12541, 12851, 13202, 13356, CINC, PC; Cerro de Mitu, Schultes & Cabrera 13858, CINC, PC). This Amazonian taxon resembles Pallavicinia lyellii in the entire thallus margins with scattered slime hairs but differs in the presence of a short basal rhizome, from which several thalli arise. Possibly the plants represent a new species; because pseudoperianths and sporophytes have not been seen, the species is not formally described here. The plants grow prostrate or ascending and are dioicous; both male and immature female thalli are present among the studied collections. The species seems to be rather common in Amazonia and may have been identified previously as P. lyellii. All Amazonian records of P. lyellii need to checked; they may belong to this undescribed species. Symphyogyna Nees & Mont. Thallus green, about 2–6  cm long, 3–6  mm wide, mostly prostrate, occasionally upright and arising from a stoloniferous base and with a stalk-like lower half, simple or forked, sometimes with ventral branches. Midrib to 1/6 of the thallus width, with 1(−3) central strands of narrow, thick-walled cells. Thallus wings undivided or lobed, sometimes bordered by narrow cells, the margins entire or toothed, with or without slime papillae, stalk of the slime papillae 1 cell long. Archegonia in clusters scattered on the midrib, each cluster protected by a tiny scale behind the archegonia (ring of scales surrounding the archegonia lacking), a closed involucre lacking. Sporophytes surrounded by a fleshy calyptra; pseudoperianth lacking. Spore surface variously ornamented.

Pallaviciniaceae

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A tropical and southern-temperate genus (ca. 20 spp.), nine in Colombia and Ecuador. The main features of Symphyogyna are the prostrate or ascending, undivided or lobed thalli with a thick midrib with 1(−3) central strands and the presence of a small scale protecting the archegonial cluster from the basal side and not surrounding the archegonia. In the other two neotropical genera of Pallaviciniaceae (Pallavicinia, Jensenia), the archegonia are fully surrounded, either by a cup-like involucre consisting of a group of connate scales (Jensenia) or by a group of free scales which are not fused (Pallavicinia). A further character of Symphyogyna is the absence of a tubular pseudoperianth; instead, the sporophyte is protected by a fleshy calyptra. Literature: Schuster (1992a), Uribe and Aguirre (1995), Uribe (1999), Gradstein and Costa (2003) 1. Thallus very narrow, less than 1 mm wide . . . . . . . . . . . . . . . . . . . S. rubescens 1. Thallus broader, 3–8 mm wide (thallus base sometimes narrower) . . . . . . . . . 2 2. Thallus ascending to erect, arising from a long and narrow, creeping, stolonlike base; thallus dichotomously branched . . . . . . . . . . . . . . . . . S. podophylla 2. Thallus ± prostrate, without long stolon-like base; thallus unbranched or dichotomously branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Thallus divided into lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Thallus undivided (but margins sometimes undulate-sinuate) . . . . . . . . . . . . . 6 4. All or some thallus lobes with a distinct tooth or acute to mucronate. Thallus shallowly or deeply lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Thallus lobes without teeth, rounded. Thallus mostly shallowly lobed (to 1/3–1/2 of the width of the thallus wing), occasionally more deeply divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aspera 5. Thallus deeply divided into lobes, almost to the midrib . . . . . . . S. brogniartii 5. Thallus shallowly divided into lobes, to 1/3–1/2 of the width of the thallus wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. apiculispina 6. Thallus margins coarsely toothed, the teeth to 5–6 cells long. Thallus margins without border . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. leptothelia 6. Thallus margins entire or with an occasional short, blunt tooth. Thallus margins with or without border. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Thallus bordered by elongate, prosenchymatic cells with narrow tips, the border cells up to more than 200 μm long . . . . . . . . . . . . . . . . . . . . . . S. marginata 7. Thallus bordered by quadrate to rectangular, parenchymatic cells with broad, truncate tips, the border cells less than 200 μm long. . . . . . . . . . . . . . . . . . . . 8 8. Thallus margins with slime papillae. Midrib with 2–3 central strands (with only 1 strand on slender branches) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. trivittata 8. Thallus margins without slime papillae. Midrib with 1 central strand . . . . . . 9 9. Border cells thin-walled or slightly thickened. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. brasiliensis 9. Border cells strongly thick-walled. Caribbean species known from Venezuela, not yet known from Colombia and Ecuador . . . . . . . . . S. digitisquama Steph.

82

1. Symphyogyna apiculispina Steph.

 

Keys and Descriptions

Fig. 7E

On rotten logs, leaf litter, and soil in submontane and montane forest, 500–3350 m. Colombia: Boy, Cun, Met, Qui. Ecuador: Lo (Benitez et al. 2012). General distribution: tropical Andes. Symphyogyna apiculispina is an Andean species that resembles S. aspera in the shallowly lobed thallus but differs in the thallus lobes with a distinct tooth (or being acute to mucronate). 2. Symphyogyna aspera F.A.McCormick On tree bases, rotten wood, and soil in submontane and montane rainforests, 450–3700 m. Colombia: Boy, Cas, Ces, Cho, Cun, Hui, Mag, Met, Nar, Ris, San, Tol, Val. Ecuador: EO, MS, Na, ZC. General distribution: tropical America. Symphyogyna aspera is a rather common neotropical species that is distinguished by the rounded thallus lobes with entire margins. The thallus is usually shallowly lobed, to 1/3–1/2 of the width of the thallus wing, occasionally more deeply lobed. 3. Symphyogyna brasiliensis Nees & Mont.  Symphyogyna canaliculata Steph. Symphyogyna circinata Nees & Mont. (?)

Fig. 7A–C

On moist soil, peat, rotten wood, tree bases, and rock from sea level to 4300 m. Colombia: Ant, Boy, Caq, Cal, Cas, Cau, Ces, Cho, Cun, Hui, Mag, Met, Nar, Put, Qui, Ris, San, Tol, Val, Vau. Ecuador: Ca, Co, Ga, MS, Lo, Na, Tu, ZC. General distribution: widespread in tropical America and Africa. Symphyogyna brasiliensis is a very common Afro-American species with a remarkably large elevational range. It is recognized by the unlobed, prostrate thallus and the entire thallus margins without slime papillae and without teeth or, occasionally, with a few short bluntish teeth. A further character of the species is the violetbrown oil bodies (T. Pócs, pers. comm.). The margin cells are quadrate or rectangular (in the latter case forming a border) and thin-walled or slightly thickened. Conspicuously thick-walled margin cells are characteristic of S. digitisquama Steph., a Caribbean species that has been recorded once from Colombia (Cleef 1981, p. 141). The Colombian specimen belongs to S. brasiliensis (fide R. Grolle, in sched.). Plants with an occasional bluntish tooth on thallus margins have been named S. circinata (Uribe and Aguirre 1995). Gradstein and Cuvertino (2015) considered S. circinata doubtfully distinct from S. brasiliensis and possibly a synonym. 4. Symphyogyna brongniartii Mont. 

Fig. 7F

On moist soil, rotten wood, or rock in montane rainforest, 200–3700  m. Colombia: Ant, Boy, Cal, Cau, Ces, Cho, Cun, Hui, Mag, Met, Qui, Ris, San, Tol, Val. Ecuador: Ch, EO, LR, Lo, MS, Na, Tu, ZC. General distribution: tropical America. Symphyogyna brongniartii is a very common neotropical species that is readily distinguished by the deeply lobed thallus, being lobed up to the midrib. The lobes are rounded, acute, or mucronate; when rounded, each lobe has a small tooth. Symphyogyna brongniartii can be confused with S. aspera, but the thallus in the

Fig. 7 (A–C) Symphyogyna brasiliensis. (A) Habit with gynoecia and young sporophyte, dorsal view. (B) Habit with androecia, dorsal view. (C) Thallus margin. (D) Symphyogyna marginata, thallus margin showing border. (E) Symphyogyna apiculispina, habit. (F) Symphyogyna brogniartii, habit. (G and H) Symphyogyna podophylla. (G) Habit with young gynoecia, dorsal view. (H) Thallus margin with tooth. (I–L) Noteroclada confluens. (I) Habit with gynoecium and antheridia, dorsal view. (J) Portion of stem with leaf. (K) Midleaf cells. (L) Dehisced capsule with elaters. (C–E, G, H redrawn from Uribe and Aguirre 1995; rest from Gradstein et al. 2001)

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Keys and Descriptions

latter species is less deeply divided, to 1/2 of the width of the thallus wing (very rarely divided to the midrib), and the lobes are rounded and entire, without tooth. 5. Symphyogyna leptothelia Taylor On moist soil, 3450–3750 m. Colombia: Cau, Ris. Ecuador: Tu (Spruce 1885). General distribution: tropical America. Symphyogyna leptothelia is a rare neotropical species that is readily recognized by the unlobed thallus with coarsely toothed margins without a border. The teeth are up to 5–6 cells long and 1–3 cells wide. The species may be confused with Jensenia spinosa and S. podophylla, but the latter two species are growing upright or ascending, from a stoloniferous base, and the base of the thallus is longly stalk-like. In S. leptothelia, the thallus is prostrate and a stalk-like base is lacking. 6. Symphyogyna marginata Steph.

Fig. 7D

On rotten wood, rock, and moist soil, sometimes growing associated with Sphagnum, (1600–)2000–3700 m. Colombia: Ant, Boy, Cau, Cho, Cun, Hui, Qui, Ris, Tol. Ecuador: Na (Gradstein 12793, QCA), ZC (Parolly et al. 2004). General distribution: tropical America. Symphyogyna marginata is an uncommon neotropical species that stands out by the unlobed, fleshy thallus with a stoloniform base, a broad, ill-defined midrib, entire margins without slime papillae and without teeth and, especially, the 1–4-cellwide hyaline thallus border made up of elongate, prosenchymatic cells with narrow tips. The border cells are about 2–8× longer than wide and up to over 200 μm long and lack chlorophyll. Symphyogyna marginata resembles S. brasiliensis in the unlobed thalli without teeth and without slime papillae but clearly differs from the latter species by the thallus border made up of prosenchymatic cells with narrow tips. 7. Symphyogyna podophylla (Thunb.) Nees & Mont. Fig. 7G, H Symphyogyna bogotensis Steph. (Symphyogyna hymenophyllum (Hook.) Nees & Mont. var. bogotensis Gottsche, nom. inval.), syn. nov. Type: Colombia, “Bogotá, ad La Peña, altitudine 2900 m, mense augusto 1859...,” Lindig 1711 (ex parte) (lectotype, designated here, PC-0081466!) Symphyogyna hymenophyllum (Hook.) Nees & Mont. var. heterogena Spruce On shaded, moist soil and rock in montane forest areas and páramo, often near rivulets, 500–3900 m. Colombia: Ant, Boy, Cal, Cau, Cun, Hui, Mag, Met, Qui, Ris, San Tol. Ecuador: ZC (Nöske et al. 2003). General distribution: tropical Andes (Colombia to Bolivia), southeastern Brazil, Patagonia, Africa, Australasia. Symphyogyna podophylla is a widespread southern-temperate and tropical-montane species that differs from other Symphyogyna species in the northern Andes by the ascending to erect thallus with a long and narrow, creeping, stolon-like base. The thallus is dichotomous branched, and the margins are toothed, with teeth being 1–6 cells long. Plants from páramo sometimes have strongly undulate thalli (Quindío, Gradstein 12739, HUQ). Symphyogyna podophylla is rare in Ecuador but common in Colombia where it was first reported by Gottsche (1864) based on a collection made by Lindig near Bogotá.

Pelliaceae

85

Gottsche recognized two different taxa in the specimen, S. podophylla and S. hymenophyllum var. bogotense Gottsche (nom. inval., without description); the latter variety was subsequently treated as a separate species, S. bogotensis, by Stephani (e.g., Uribe and Aguirre 1995). I have studied part of the original material of S. bogotensis (PC) and found that it fully fits the variation of S. podophylla, with teeth on the thallus margins being 2–6 cells long. The two taxa are therefore united here. Schaumann et al. (2003), in a study of the genetic diversity in S. podophylla, found evidence for cryptic speciation in S. podophylla, with populations from southern South America and Australasia being genetically different. No morphological differences were detected among the South American and Australasian populations. Surprisingly, a specimen from Colombia (identified as S. bogotensis) proved to be genetically more similar to populations from Australasia than from southern South America. 8. Symphyogyna rubescens Steph. On moist soil in páramo, 3600 m. Colombia: Cun (Uribe and Aguirre 1995). General distribution: southeastern Brazil, Colombia. Symphyogyna rubescens is a rare neotropical species that is readily recognized by the very small thalli, which are less than than 1 mm wide. The thallus is unlobed and dichotomously branched, the margins are entire or weakly toothed, and the base of the thallus is stoloniform. 9. Symphyogyna trivittata Spruce On moist soil on earth walls, 600–3200  m. Colombia: Ant, Cho, Cun, Ris. Ecuador: Pa (Gradstein and Benitez 2017). General distribution: West Indies, northern Andes (Colombia, Ecuador). Symphyogyna trivittata is an uncommon neotropical species that resembles S. brasiliensis but differs from the latter in the presence of 2–3 central strands in the midrib (one strand only on slender branches) and small slime papillae on thallus margins. In S. brasiliensis the midrib has only one central strand and slime papillae are absent. Sterile plants of S. trivittata may be confused with Pallavicinia lyellii, but in the latter species, the midrib has only one central strand and the slime papillae have a 2–3-cell-long stalk.

Pelliaceae A family of two genera, one in Ecuador. Noteroclada Hook.f. & Wilson Plants foliose, green to glaucous, rather large, 0.5–1 cm wide, very flat, prostrate, unbranched or forked. Stems fleshy, dorsally flattened. Leaves succubous, almost longitudinally inserted and attached to the lateral side of the axis (dorsal surface of the stem leaf-free), ovate, plane to deeply concave, margins entire, lower half of the leaves 2–3 cell layers thick, otherwise leaf unistratose. Central strand lacking. Cells

86

 

Keys and Descriptions

large, fully thin-walled, each cell with 5–20 small, finely segmented oil bodies. Rhizoids colorless or pale brown. Monoicous or, occasionally, dioicous. Gametoecia on the dorsal surface of the stem, antheridia scattered in small cavities in the dorsal stem surface, archegonia in clusters. Sporophytes surrounded at the base by a shorttubular pseudoperianth with a wide mouth. Setae very long, to 10 cm long, thick. Capsules spherical, opening by 4 valves. Spores green, very large, to 100  μm in diameter, multicellular, germination endosporous. Elaters with 2–4 spirals, attached to the base of the capsule. Vegetative reproduction unknown. A Latin American genus with one species; for recognition of Noteroclada, see under the species. The genus is sometimes placed in a separate family, Noterocladaceae (Söderström et al. 2016). Literature: Crandall-Stotler et al. (2010) 1. Noteroclada confluens Taylor Noteroclada leucorhiza Spruce

Fig. 7I–L

On wet soil along rivers and on road banks, usually near running water, (1800–) 2800–4300 m. Colombia: Ant, Boy, Cau, Ces, Cun, Mag, Met, Nar, Qui, Put, Ris, Tol. Ecuador: Az, Ca, Ch, Co, Lo, MS, Pi, ZC. General distribution: throughout the mountains of Latin America, from Mexico to Tierra del Fuego. Noteroclada confluens is characteristic of moist banks of rivulets in páramo in the Andes and Central America, above 3000 m; occasionally the species is found at lower elevation. It is readily distinguished by the robust, green to glaucous plants with colorless rhizoids (not violet) and large, ovate, undivided, succubous leaves, which are attached to the lateral side of the stem, leaving the dorsal surface leaffree, and are several layers of cells thick toward the leaf base. Underleaves are absent. A further characteristic feature is the spherical capsule, which contains large, multicellular, green spores and is elevated when mature on a long seta, to 10 cm or more in length. Noteroclada confluens has sometimes been confused with Fossombronia peruviana, but the leaves of F. peruviana are undulate-plicate and usually much broader than long, the rhizoids in F. peruviana are violet, the antheridia are borne naked on the stem (not in cavities in the dorsal stem surface), the mature seta is very short, and the capsules open irregularly, not by 4 valves. Subclass Metzgeriidae (Simple Thalloid Liverworts II) Plants thalloid, growing with a two-sided apical cell, divided into two lateral merophytes, ventral merophyte absent. Thallus usually differentiated into a midrib and thinner thallus wings, midrib and thallus wings sharply separated from each other or not, wings usually undivided. Cells with or without trigones; oil bodies usually present. Rhizoids with smooth walls. Antheridia and archegonia on abbreviated branches at the thallus margin or on the ventral surface of the midrib;

Aneuraceae

87

archegonia not arising from the apical cell of thallus (plants anacrogynous). Sporophytes surrounded by a fleshy calyptra. Setae well developed, 1–3 cm long, thin to very thick. Capsules ellipsoid, opening by 4 valves. Contains about 52 species in Colombia and Ecuador, in four genera and two families. Key to the families of Metzgeriidae of Colombia and Ecuador 1. Thallus margins usually with hairs. Thallus dichotomous (rarely pinnate), with a very narrow midrib and broad, unistratose wings. Gametoecia on the ventral side of the midrib. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metzgeriaceae 1. Thallus margins without hairs. Thallus unbranched or pinnate to palmate, with a broad, indistinct midrib or with a distinct midrib on branches; unistratose wings present or absent. Gametoecia on short lateral branches or on the underside of the thallus margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aneuraceae

Aneuraceae Plants thalloid, green to brown to black, 1–10 cm long, 0.3–10 mm wide, prostrate to erect, unbranched or pinnate to palmate, without midrib or with a distinct midrib on branches; thallus apex and ventral surface with slime papillae. Oil bodies finely to coarsely granular, sometimes absent. Monoicous or dioicous. Gametoecia in cavities on short lateral branches or on small receptacles on the underside of the thallus margin. Sporophytes surrounded by a fleshy calyptra; pseudoperianth lacking. Setae 4–15 cells thick. Capsules ellipsoidal, opening by 4 valves, wall 2-layered. Spores small, unicellular. Elaters with 1(−2) spirals, attached to the tips of the capsule valves. Vegetative reproduction by multicellular gemmae. A family of five genera worldwide. The principal characteristics of Aneuraceae are (1) thallus pinnate or unbranched; (2) midrib usually indistinct; (3) gametoecia at thallus margins or on short lateral branches; (4) sporophyte in a fleshy calyptra; and (5) elaters attached to the tips of the valves. 1. Thallus 2–4-pinnate, main axis 0.3–1.5(−2) mm wide. Male branches with antheridia in 2 rows. Archegonia on short branches. Seta 4 cells thick . . . . . . Riccardia 1. Thallus unbranched or irregularly 1-pinnate, main axis 2–10 mm wide. Male branches with antheridia in 2–6 rows. Archegonia on small receptacles hidden under the thallus margin, not on short branches. Seta 8–15 cells thick. . . . . . 2 2. Thallus irregularly pinnately branched. Male branches with antheridia in 2 rows. Plants of upper montane forest and wet páramo of Ecuador, rare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lobatiriccardia 2. Thallus branches absent or very scarce. Male branches with antheridia in 2–4(−6) rows. Plants of forests and páramo, from sea level to 4000  m, common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aneura

88

 

Keys and Descriptions

Aneura Dumort. Thallus fleshy, bright green to dark green, 1–5 cm long, 3–10 mm wide, prostrate, flat, or convex, sometimes crisped, ± simple, branches lacking or very scarce, thallus thickened to near the margins, without midrib, wings lacking or narrow, margins entire or lobed. Oil bodies numerous per cell (usually more than 10, fewer in thallus margin cells), small, finely granular. Dioicous. Antheridia in 2–4(−6) transverse rows on short or long branches. Archegonia in clusters in incisions of the thallus margin, hidden under small thallus flaps. Setae thick, in cross section 8–16 cells in diameter. Vegetative reproduction lacking. About 20 species worldwide, 2 in Colombia and Ecuador. Aneura is recognized by the rather broad (3–10 mm wide), fleshy, scarcely branched thallus with entire to somewhat crisped margins and without midrib. Literature: Meenks and Pócs (1985) 1. Thallus bright green, 9–20 cells thick in the middle (cross section). Male branches with 2–5 pairs of antheridial chambers along its length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pinguis 1. Thallus usually dark green, 5–8 cells thick in the middle. Male branches longer, with 6–25 antheridial chambers along its length. . . . . . . . . . . . . . A. latissima 1. Aneura latissima Spruce Aneura pseudopinguis Herzog Mostly on rotten logs, also found on rock and soil, in lowland, and lower montane rainforest. Colombia (new): Vau (Río Vaupés, Javareté, May 1953, Schultes & Cabrera 19412, c. andr., CINC, PC). Ecuador: Na, Or (Estación Tiputini, Gradstein, Deleg & Bader s.n., HUTPL), Pi (?). General distribution: tropical South America, Africa. Aneura latissima, originally described from Amazonia but possibly widespread in tropical America and tropical Africa, is distinguished by the rather thin, glossy dark green thallus, (3–)5–8 cells thick, and long and broad antheridial branches, measuring 6–25 antheridial chambers in length and 2–4 antheridial chambers in width. The species resembles A. pinguis, but the latter species has a thicker thallus (9–20 cells thick in the middle of the thallus) and shorter and narrower antheridial branches, to maximally 5 antheridial chambers long and 2(−3) antheridial chambers wide. Moreover, the thallus of A. pinguis is more bright green in color. Aneura latissima has been confused with A. pinguis, and the neotropical distribution of the two species is incompletely known. Some neotropical specimens named A. pinguis may belong to A. latissima. 2. Aneura pinguis (L.) Dumort. Riccardia pinguis (L.) Gray

Fig. 8A–C

On rotten logs and soil in montane rainforest and páramo, 500–4300  m. Colombia: Boy, Ces, Cho, Hui, Put, Qui, Ris, Tol. Ecuador: Na, Tu, ZC. General distribution: subcosmopolitan. For characters of the widespread and variable A. pinguis, see the key and the discussion under A. latissima. Aneura pinguis is genetically highly diverse, and

Aneuraceae

89

Fig. 8 (A–C) Aneura pinguis. (A) Habit with androecia. (B) Habit with gynoecia. (C) Cross secion of thallus. (D) Lobatiriccardia oberwinkleri, habit. (E) Lobatiriccardia verdoornioides, habit with antheridia. (F) Riccardia regnellii, habit with gynoecia and sporophytes. (G and H) Riccardia fucoidea. (G) Habit, dorsal view. (H) Branches, showing broad wings. (I and J) Riccardia wallisii. (I) Habit. (J) Portion of cross section of thallus. (K and L) Riccardia hymenophytoides. (K) Portion of habit, showing tongue-shaped branches and axis with short stolons. (L) Cross section of axis. (A–C, G, H from Gradstein et al. 2001; D, E redrawn from Preußing et al. 2010; I, J redrawn from Meenks and de Jong 1985; rest from Gradstein et al. 2001; Gradstein and Ilkiu-Borges 2009)

90

 

Keys and Descriptions

much cryptic speciation has been reported in the species (Bączkiewicz et al. 2017), but the neotropical populations have not been studied. The collections of A. pinguis from Colombia and Ecuador need revision; some of them may belong to A. latissima. Lobatiriccardia (Mizut. & S.Hatt.) Furuki Thallus fleshy, glistening green to darkish-green to bluish-green, 1–5  cm long, 2–8  mm wide, prostrate, concave, pinnate, without a midrib, margins with a 2–4-cell-wide unistratose wing. Oil bodies 1–8 per cell, brownish, finely granular, sometimes lacking in the epidermis. Dioicous. Antheridia in 2 rows on short branches. Archegonia in clusters on receptacles at incisions of the thallus margin (sometimes hidden under thallus wings), surrounded by multicellular paraphyses and thick-walled unicellular rhizoids. Setae thick, in cross section 8–15 cells in diameter. Vegetative reproduction lacking. A genus of seven species, mainly in Southeast Asia and New Zealand (Preußing et al. 2010), two species in Ecuador. Branching and oil bodies in Lobatiriccardia are like in Riccardia, but in other respects (thallus width, position of archegonia, thickness of seta), the genus is more similar to Aneura. Literature: Preußing et al. (2010) 1. Thallus more than 10 cells thick in the middle (cross section). Female receptacles extending beyond the thallus margin. Plants of wet páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. verdoornioides 1. Thallus thinner, 5–10 cells thick in the middle. Female receptacles not extending beyong the thallus margin. Plants of upper montane forest. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. oberwinkleri 1. Lobatiriccardia oberwinkleri Nebel et al.

Fig. 8D

On wet rock, soil, and rotten wood in upper montane forest environments, 1850–3500 m. Ecuador: Lo, MS (Sangay National Park, Ramsay et al. 168, QCA), ZC (Preußing et al. 2010; Schäfer-Verwimp et al. 2013a). General distribution: only known from Ecuador. Lobatiriccardia oberwinkleri is a rare Ecuadorian endemic that is known from only few collections. For characters of the species, see the key. 2. Lobatiriccardia verdoornioides Nebel et al.

Fig. 8E

On wet soil and rock in páramo, in bogs, and along rivulets or lakes, 3700–4100 m. Ecuador: Az, Pi (Preußing et  al. 2010; Schäfer-Verwimp et  al. 2013a); to be expected in Colombia. General distribution: only known from Ecuador. Lobatiriccardia verdoornioides is a rare Ecuadorian páramo species that is characterized by the fleshy thallus (more than 2  mm wide, 10–20 cells thick in the middle) with a concave surface and undulate margins with short and plump branches; in addition, one or a few long branches may be present. The species resembles Aneura pinguis but differs from the latter by the more branched, concave thallus and thallus cells with only one to eight oil bodies (10–40 per cell in A. pinguis).

Aneuraceae

91

Riccardia Gray by S. Robbert Gradstein and Catherine Reeb Thallus green to brown to black, 1–15 cm long, axis 0.3–2 mm wide, creeping, ascending, or erect, sometimes with short stolons at the base, irregularly to regularly (1–)2–4-pinnate to palmate, the branches suberect (branching palmate) to widely spreading (branching pinnate), sometimes with a distinct midrib. Axis ca. 3–20(−40) cells in diameter, concave-convex, plano-convex, bi-convex, or rounded in diameter, sometimes with a subepidermis of cells with thick-walled, brownish walls, axis and branches with or without unistratose lateral wings, the wings entire or toothed, sometimes crispate (R. gradsteinii). Oil bodies finely to coarsely granular, 1–10(−25) per cell, sometimes scarce or absent in the epidermis. Monoicous or dioicous. Gametoecia in 2 rows on short branches. Archegonia surrounded by unicellular and thick-walled rhizoids or by multicellular paraphyses (in some robust Andean species). Setae thin, 4 cells in diameter. Vegetative reproduction by gemmae from the apex and surface of narrow thallus branches. About 100 species worldwide, 15 in Colombia and Ecuador. Riccardia stands out by the pinnately or palmately branched thalli with a narrow, 0.3–2-mm-wide axis and gametoecia on short lateral branches; the archegonia and antheridia are arranged in 2 straight rows on separate male and female branches. The genus is related to Aneura and Lobatiriccardia, but the latter two groups have a broader axis (usually more than 2  mm wide) and a thicker seta. Moreover, Aneura has many more oil bodies per cell (10–40). In the older literature, Riccardia was considered part of Aneura (e.g., Spruce 1885; Stephani 1898–1924). The species of Riccardia of the tropical Andes have been studied by Meenks (1987) who accepted 29 species in Colombia and Ecuador. However, no descriptions were given and the key was difficult to work with. Gradstein and Reeb (2018b) presented a new revision of the regional species, with descriptions and illustrations, and recognized much fewer species. The present treatment is based on the latter study. Note that the data on the distribution of the species within Colombia and Ecuador given here are in part based on the literature and unreliable and in need of revision as many published records and identifications of herbarium specimens proved to be erroneous (Gradstein and Reeb 2018b). For identification of the species, complete plants consisting of main axis and branches are required. As thallus cells are often collapsed in dried material, rehydration of the thallus with sodium hypochlorite (Rico 2011) is recommended for the study of old herbarium material. Literature: Gradstein and Reeb (2018b) 1. Thallus cells densely covered by lamellate papillae . . . . . . . . . . . . R. aberrans 1. Thallus cells smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Ultimate branches winged (wings 1–6 cells wide, ± unistratose, translucent) . . . . 3 2. Ultimate branches not or scarcely winged [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 3. Main axis winged (sometimes only partly and very narrowly). . . . . . . . . . . . 4

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3. Main axis not winged [when in doubt try both leads]. . . . . . . . . . . . . . . . . . 11 4. Plants prostrate, delicate. Axis 3–7 cells thick, dorsal surface of axis flat. Subepidermis lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Plants ascending to erect, robust. Axis (7–)8–25 cells thick, dorsal surface of axis flat to convex. Subepidermis present or absent . . . . . . . . . . . . . . . . . . . . 6 5. Branch wings as wide as or wider than the midrib. Surface cells on branches in straight rows. Plants autoicous (rarely dioicous). . . . . . . . . . . . . R. leptophylla 5. Branch wings narrower than the midrib. Surface cells on branches often in oblique rows. Plants dioicous or paroicous (= with a small male branch at the base of the female branch). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. regnellii 6. Main axis with strongly crispate wings. Plants from Peru (Pócs 2019b), not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . R. gradsteinii Pócs 6. Main axis not with strongly crispate wings. . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Wings with ciliate teeth, the teeth 1–5 cells long, consisting of elongate cells. Branches obliquely spreading and short and plump . . . . . . . . . . . . R. ciliolata 7. Wings entire or with short teeth, the teeth 0.5–1 cell long, consisting of a subquadrate cell. Branches widely spreading and long and slender or obliquely spreading and short and plump . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Plants with broad, very short and deeply concave gemmiparous branches. Plants found in páramo of Ecuador, ca. 4000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ������������������������������������������������������������������������� gemmiparous phenotype of R. pallida 8. Plants without such gemmiparous branches . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Plants irregularly 1–2(−3)-pinnate. Axis broad, mostly (0.8–)1–2  mm wide, plano-convex, gradually broadened toward the branch bases. Branches suberect to obliquely spreading, not deflexed, short and plump with a broad apex, branch base decurrent. Branch apex often with gemmae . . . . . . . . . . . . . . . R. wallisii 9. Plants ± regularly 2–4-pinnate. Axis less than 1 mm wide, bi-convex to rounded, not gradually broadened toward the branch bases. Branches widely spreading, often deflexed, long and slender, linear to subulate, branch base not decurrent. Gemmae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Wing cells large, 35–50 μm wide. All branches broadly winged. Wings of ultimate branches wider than the midrib. . . . . . . . . . . . . . . . . . . . . . . . R. fucoidea 10. Wing cells smaller, 20–30(−35) μm wide. Primary branches narrowly and interruptedly winged. Wings of ultimate branches narrower than the midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pallida 11. Axis with a subepidermis (sometimes only weakly developed in R. poeppigiana). Plants growing erect or ascending, 2–4-pinnate, primary branches opposite or alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11. Axis without subepidermis. Plants creeping or ascending, 1–3-pinnate, primary branches usually alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 12. Plants very robust (to 10 cm long), with regularly spaced whorls of horizontally spreading fronds. Primary branches without wings. Wings of ultimate branches 3–4× wider than the midrib, wing cells in oblique rows, with trigones. Plants only known from the Cordillera del Cóndor, Ecuador . . . . . . . . . R. verticillata 12. Plants without whorls of horizontally spreading fronds. Primary branches winged or not. Wings of ultimate branches narrower or wider than the midrib,

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wing cells in straight or slightly oblique rows, without trigones. Plants widespread. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Branches usually decurved, narrowed to the tip. Ultimate branches winged but primary branches ± without wing. Wing cells 15–30(−35) μm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. poeppigiana 13. Branches straight, not narrowed to the tip. All branches regularly and broadly winged. Wing cells larger, 35–50 μm wide . . . . . . . . . . . . . . . . . . . R. fucoidea 14. Stoloniform branches (± rounded in cross section) present in the lower half of the thallus. Branching in the upper half of the thallus palmate . . . R. hymenophytoides 14. Stoloniform branches lacking, all branches clearly flattened. Branching mostly pinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15. Axis broad, (0.5–)0.8–2  mm wide. Branches short and plump, suberect to obliquely spreading, branch bases distinctly decurrent. . . . . . . . . . . R. wallisii 15. Axis narrower, 0.3–0.8  mm wide. Branches linear or tongue-shaped, rather widely spreading, branch bases not distinctly decurrent . . . . . . . . . . . . . . . . 16 16. Branch wings 2–5 cells wide, surface cells on branches often arranged in oblique rows. Branches often tongue-shaped. Plants growing in lowland and montane forests up to the páramo, from sea level to 3500 m . . . . . R. regnellii 16. Branch wings 1–2(−3) cells wide (wings lacking in older parts of the thallus), surface cells on branches in straight rows. Branches linear, rarely tongueshaped. Plants growing in upper montane forest and páramo, ca. 2000–4000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. Epidemis cells not or little narrower (1–2×) than medullary cells. Main axis (6–)7–12 cells thick, dorsal surface convex. Gemmae present. Plants common in páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hans-meyeri 17. Epidermis cells distinctly narrower (2–3×) than medullary cells. Main axis thinner, 4–7 cells thick, dorsal surface flat or concave. Gemmae absent. Plants rare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18. Plants semi-aquatic, often growing submerged. Dioicous. Thallus conspicuously elongate, to 5  cm long, remotely pinnate, branches mostly very short or reduced, linear, most branches without wing. Sexual branches winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. cataractarum 18. Plants not semi-aquatic, on soil or rock. Autoicous. Thallus shorter, 1–2 cm long, rather densely branched. Branches well developed, long, linear or tongue-shaped, all branches narrowly winged. Sexual branches without wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. chamedryfolia 19. Axis with a subepidermis. (Dioicous. Branches often decurved, long-linear or lanceolate. Epidermis cells ± as wide as inner cells). . . . . . . . . R. poeppigiana 19. Axis without subepidermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Plants paroicous (= with a small male branch at the base of the female branch), rarely dioicous, common in páramo. Branching in upper part of thallus palmate, branches linear to lanceolate to tongue-shaped. Epidermis cells on ultimate branches often arranged in oblique rows. Main axis stoloniform (1–2× wider than high, bi-convex to rounded). Gemmae absent . . . . . . R. latifrons subsp. parasitans

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20. Plants dioicous, often sterile. Branching not palmate, all branches linear. Epidermis cells on ultimate branches arranged in straight rows. Main axis not stoloniform. Gemmae present or lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21. Epidermis cells not or scarcely narrower than medullary cells. Plants regularly or irregularly 2-pinnate, common in páramo. Gemmae frequently present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hans-meyeri 21. Epidermis cells distinctly narrower (2–3×) than medullary cells. Plants irregularly 1–2-pinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Gemmae absent. Axis thin, 4–6(−7) cells thick, upper surface flat (axis sometimes thicker at the base, bi-convex). Sexual branches winged. Plants growing submerged in lakes and marshes in páramo . . . . . . . . . . . . . . . R. cataractarum 22. Gemmae present. Axis thicker, 6–12 cells thick, upper surface convex. Sexual branches not winged. Plants of lowland or montane forest areas, not growing submerged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. digitiloba 1. Riccardia aberrans (Steph.) Gradst.

Fig. 9A, B

On very humid soil and among Sphagnum in upper montane scrub and páramo, ca. 2700–3700  m. Colombia: Ant (type), Qui (Gradstein and Reeb 2018b). Ecuador: Na (Parque Nac. Cayambe-Coca, Gradstein 12792, QCA), ZC (Gradstein and Reeb 2018b). General distribution: northern Andes (Colombia, Ecuador). Riccardia aberrans is a rather robust northern Andean species that is readily distinguished by the numerous lamellate papillae on the thallus surface; the papillae are particularly conspicuous on the hyaline wing cells of the branches and are protuberant on the margins. Further characters of R. aberrans are (1) plants monoicous, 2–4  cm long, light green to brownish-green, 2–3-pinnate, branches alternate or opposite, obliquely spreading, margins of axis and branches with very irregularly developed wings, appearing eroded; (2) axis ca. 1 mm wide, plano-convex to slightly bi-convex, 12–13 cells thick, axial cells large, medullary cells thin-walled, hyaline, walls becoming slightly thickened and brownish toward the epidermis, a weak subepidermis of 1–2 cell layers present, axis rudimentarily winged (wing to 2 cells wide), wing becoming wider and more conspicuous toward the branch bases; (3) branches obliquely spreading, short or long, narrowly rectangular, margin irregularly winged, the wings lobed-undulate, to 5 cells wide, margins crenate and with a few blunt teeth, cells thin-walled and very transparent, ca. 2× larger than median surface cells; (4) oil bodies lacking in the epidermis, present in the inner cells of the thallus, few per cell, ovate, pale-brownish; and (5) gemmae not observed. By the papillose thallus surface, R. aberrans differs from all other neotropical Riccardia species and is similar to R. crassa (Schwägr.) Nees from southern South America and Australasia. Meenks (1987) apparently overlooked the papillae and confused the species with R. wallisii. All recent records of R. aberrans from Colombia and Ecuador must be verified; the material may be misidentified. Gradstein and Reeb (2018b) suggested that oil bodies are lacking in R. aberrans, but observation of fresh materal from Ecuador (Napo, Gradstein 12791, QCA) showed their presence in the inner cells of the thallus.

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Fig. 9 (A and B) Riccardia aberrans. (A) Habit. (B) Portion of thallus surface, showing striatepapillose cells. (C and D) Riccardia cataractarum. (C) Habit. (D) Cross section of the main axis. (E and F) Riccardia digitiloba. (E) Habit. (F) Cross section of the main axis. (G and H) Riccardia poeppigiana, habit with sexual branches and sporophytes. (I and J) Riccardia pallida. (I) Habit. (J) Cross section of main axis. (K and L). Riccardia latifrons subsp. parasitans. (K) Habit. (L) Cross section of the main axis. (A, B from Holz 536, QCA; C, D from the isotype of R. paramorum, G; E, F from Gradstein & Varón 11072, GOET; G, H from Gradstein et al. 12709, PC; I from SchäferVerwimp & Nebel 32071, QCA; J from Gradstein et al. 12661, QCA; K, L from Schäfer-Verwimp & Nebel 32192, QCA). Scale bars: A, G–I, K = 1 mm; C, E, J, L = 0.5 mm; D, F = 0.25 mm

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2. Riccardia cataractarum (Spruce) Schiffn. Riccardia paramorum Meenks

 

Keys and Descriptions

Fig. 9C, D

On wet soil and submerged in lakes, in páramo, 3400–4000 m; elsewhere in the neotropics found at lower elevations (Meenks 1987). Colombia: Boy, Cun, Met. Ecuador: Na (Gradstein and Reeb 2018b). General distribution: Dominican Republic, tropical Andes (Colombia to Bolivia), Brazil, Paraguay. Riccardia cataractarum is a semi-aquatic neotropical species found a few times in páramo in the northern Andes where it was called R. paramorum. It is recognized by (1) plants dioicous, pale green to dark brown, 1–5 cm long, sparsely to densely 2-pinnate, branches often short or rudimentarily developed, wingless or with very narrow wings; (2) main axis usually bi-convex at the base (10–15 cells thick) but soon becoming thin, plano-convex to slightly ellipsoid, 4–6(−7) cells thick, cells walls thin or slightly thickened, epidermis cells ca. 2–5× longer than wide, flat (not mammillose), walls colorless to brownish, much smaller (2–3×) than the medullary cells in cross section, medullary walls ± colorless; (3) branches alternate to subopposite, linear, varying in length, often very short or rudimentary, wingless or with very narrow, 1–2-cell-wide wings, margins entire, apex rounded or retuse, papillae scattered on the ventral surface of the branches; (4) male branches and female branches usually winged; and (5) gemmae not observed. The elongate (to 5 cm long) thallus with a rather flat and thin axis, ± wingless branches, small-celled epidermis, winged sexual branches, and absence of gemmae are especially characteristic of R. cataractarum. Sometimes the basal portion of the axis (or primary branch) is swollen and bi-convex. Plants growing permanently submerged are more elongate and less branched. 3. Riccardia chamedryfolia (With.) Grolle Riccardia pinnatifida (F.Weber) Trevis. Riccardia sinuata (Hook.) Trevis. On moist soil or rock in upper montane forest and páramo, 2800–4000  m. Colombia: Cun (Gottsche 1864). Ecuador: Ca, Lo, Na, Pi. General distribution: common and widespread in the temperate regions of the sourthern Hemisphere, scattered in the tropics, and in the neotropics recorded from the West Indies, the northern Andes, and Brazil. Riccardia chamedryfolia is a common Holarctic species that has been collected a few times in the northern Andes. The species is distinguished by (1) plants monoicous, rather small, 1–2  cm long, 0.3–0.6  mm wide, prostrate, irregularly 1(−2)-pinnate, without stolons; (2) main axis without wing, 4–7 cells thick, planoconvex with flat (to slightly concave) dorsal side and convex ventral side, all cells thin-walled, epidermis cells scarcely smaller than medullary cells; (3) branches linear or tongue-shaped, with rounded to retuse tips, ultimate branches narrowly winged (wings lacking in older parts of the thallus), wings 1–2(−3) cells wide, margins entire, margin cells not enlarged; (4) oil bodies present in all epidermis cells, finely granular; and (5) gemmae not observed.

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Riccardia chamedryfolia has been overlooked in the northern Andes and is probably not rare in páramo. The occurrence of R. chamedryfolia in the region was confirmed by Schäfer-Verwimp et al. (2013a) using DNA sequences. The species may be confused with R. hans-meyeri and R. cataractarum, but the latter two species are dioicous. Riccardia hans-meyeri furthermore differs by the thicker axis with epidermis cells not or scarcely smaller than inner cells and more or less mammillose as well as the frequent presence of gemmae, and R. cataractarum by semiaquatic growth, more elongate and much less branched thalli, and winged sexual branches. 4. Riccardia ciliolata (Spruce) Horik. On bark, rotten wood, and humus, 2650–3750  m. Colombia: Ant, Ara, Boy, Cau, Cho, Cun, Met, Qui, Ris. Ecuador: Lo, Tu. General distribution: tropical Andes. Riccardia ciliolata is a characteristic high-Andean species that is recognized by (1) plants dioicous, robust, 3–10 cm long, 2-pinnate, primary branches obliquely spreading, short and plump; (2) axis robust, irregularly and narrowly winged, biconvex, 10–15 cells thick, with a subepidermis, dorsal epidermis cells ± larger than subepidermis cells; (3) branches with a broad midrib and narrow, irregular wings, the wings on ultimate branches to 5 cells broad, wing cells 25–40 μm wide, irregularly elongate-hexagonal, cells becoming conspicuously smaller to the margin, the margins sharply toothed by 1–5 cells long, linear teeth made up of elongate cells, apex of branches rounded to emarginate; (4) oil bodies present in most epidermal cells but rare or absent in wing cells (Meenks and de Jong 1985); and (5) gemmae frequently present. The robust thalli with obliquely spreading, short and plump, winged branches with ciliate margins, the cilia being 1–5 cells long and made up of elongate cells, are the distinctive characters of R. ciliolata. 5. Riccardia digitiloba (Spruce) Pagán Riccardia calcarea (Steph.) Meenks Riccardia metzgeriiformis (Steph.) R.M.Schust.

Fig. 9E, F

On soil, rock, rotten wood, and tree bases in shaded and humid sites in lowland and montane forest areas, from sea level to ca. 3200 m. Colombia: Cau, Cun, Hui, Mag, Qui. Ecuador: EO, Lo, Na, Pa, Pi, ZC.  General distribution: tropical America. Riccardia digitiloba (= R. metzgeriiformis) is a widespread neotropical species that is distinguished by (1) plants dioicous, delicate, to 1 cm long, pale green, prostrate, irregularly 2-pinnate to dichotomous, primary branches different in length, often very long, finger-like, producing gemmae at the tip, the gemmiparous tips sometimes attenuate (narrowed to the apex), branch bases sometimes stolon-like; (2) axis without wing, ellipsoid, ca. 6–12 cells thick, medullary cells large, thinwalled, epidermis cells 3–4× smaller than medullary cells, thin-walled or somewhat

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thick-walled, sometimes a few large, swollen epidermis cells occurring mixed with the small epidermis cells; (3) branches linear, without wing or, occasionally, with a very narrow, 1(−2)-cell-wide wing, margins ± straight, apex rounded; (4) calyptra surface almost smooth; and (5) gemmae usually abundant, produced on decurved branch tips. The small, pinnate to dichotomous thalli with numerous linear, wingless, or rudimentarily winged branches producing gemmae at the tips, and the small epidermis cells, 2–3× narrower than medullary cells, are the principal characters of R. digitiloba. The large hyaline medullary cells are readily visible below the small-celled epidermis, without cross sectioning. The gemmiparous branches are sometimes tapering to narrow tips. 6. Riccardia fucoidea (Sw.) C.Massal. Riccardia andina (Spruce) Herzog Riccardia bogotensis (Gottsche) Pagán Riccardia crassicaulis (Steph.) Meenks & C.De Jong Riccardia lepidomitra (Spruce) Gradst. Riccardia papillata (Gottsche) Hässel Riccardia plumiformis (Spruce) Meenks [“plumaeformis”]

Fig. 8G

On moist soil, bark, rotten wood, and rock in submontane and montane rainforests and cloud forests, (300–)1000–3700 m. Colombia: Ant, Boy, Cau, Cho, Hui, Mag, Met, Nar, Put, Qui, Ris, San, Tol. Ecuador: Ca, Lo, LR, MS, Na, Pa, Pi, Tu, ZC. General distribution: tropical America. Riccardia fucoidea is a common neotropical-montane species that is recognized by (1) plants dioicous, rarely monoicous, growing upright, rather robust (but varying greatly in length), 2–10 cm long, (2–)3–4-pinnate, primary branches subopposite, spreading at straight or oblique angles (becoming more oblique toward the apex of the thallus), branches straight (not deflexed), stolons present at the base; (2) axis robust, bi-convex to rounded, 10–25 cells thick, with a subepidermis, without wing or slightly winged at branch bases, rarely narrowly and interruptedly winged, margins entire, epidermis cells usually smaller than inner cells; (3) branches linear, straight, with broad transparent wings (wing ca. 4–7 cells wide), the wings of ultimate branches wider than the midrib, wing cells large, 35–50 μm wide and 40–75 μm long, hexagonal, thin-walled or slightly thickened but without trigones, arranged in straight or slightly oblique rows, margins entire to crenate to bluntly toothed; (4) male branches winged, margins crenate; (5) calyptra irregularly hairy; and (6) gemmae absent. Upright growth, regularly 2–4-pinnate branching, broad branch wings made up of large hexagonal cells (35–50 μm wide), a thick axis with a subepidermis and with short stolons, and absence or rudimentary presence of an axial wing are the main characters of R. fucoidea. The branch margins are entire or bluntly toothed. Phenotypes of R. fucoidea with a rudimentarily winged main axis can be confused with R. pallida, but the latter species has smaller wing cells (20–30 μm wide) and narrower branch wings not wider than the midrib and only weakly developed on the primary branches.

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7. Riccardia hans-meyeri (Steph.) Meenks & C.De Jong Riccardia columbica (Steph.) Gradst. & Hekking Riccardia hans-meyeri var. dentata Meenks On humic soil, peat, rotten wood, and rock in upper montane forests and páramo, occasionally in running water, 2700–4500 m. Colombia: Boy, Cau, Cho, Hui, Met, Qui, Ris. Ecuador: Az, Ca, Ch (type), Co, Lo, MS, Na, Pi, ZC. General distribution: northern Andes (Venezuela to Peru); new to Peru: La Convención, S. Teresa, lower Phaccaq valley, 4236 m, Sylvester s.n. (PC). Riccardia hans-meyeri (= R. columbica) is very characteristic of northern Andean páramo and is distinguished by (1) plants dioicous, small but variable in size, 0.5–1.5 mm long, axis ca. 0.2–0.6 mm wide, prostrate or growing upright in dense mats, regularly or irregularly 2-pinnate, branches obliquely to widely spreading, little narrower than the axis, cells of the thallus surface smooth or mammillose, sometimes with projecting tips (especially on the ventral thallus surface), walls thin or slightly thickened, especially the transverse walls; (2) main axis ± without wing, bi-convex, (6–)7–12(−17) cells thick, without subepidermis, dorsal and ventral epidermis cells similar in size and scarcely smaller than medullary cells, with brownish walls in older portions of the thallus, medullary cells with colorless or yellowish walls, all cell walls thin or thickened; (3) branches varying in length, short-rectangular to linear, plano-convex, margins entire to crenate, apex rounded or retuse, wings absent or present and narrow, 0–2(−3) cells wide, marginal wing cells sometimes conspicuously larger than medullary cells, quadrate to subrectangular, thinwalled or walls thickened, radial walls straight or oblique; (4) papillae sometimes present on dorsal surface of branches; and (5) gemmae frequently produced at the retuse thallus tips, 2–celled, thin-walled. The delicate, creeping, regularly or irregularly 2-pinnate, dioicous thalli with wingless or very narrowly winged branches, frequent presence of gemmae at branch tips and a bi-convex main axis (branches plano-convex) with epidermis cells not or  scarcely smaller than inner cells are the main characters of R. hans-meyeri. The epidermis cells are sometimes mammillose, and the walls of the epidermis cells can be colorless or browish pigmented and thin or somewhat thickened. Riccardia hans-meyeri may be confused with R. latifrons subsp. parasitans, which is also common in páramo, but the latter taxon is usually paroicous and is ± palmately branched toward the thallus apex (branches not widely spreading near apex) and lacks gemmae. Moreover, the main axis in R. latifrons subsp. parasitans is ± rounded in cross section (stoloniform), and the epidermis cells are conspicuously narrower than the inner cells and often orientated obliquely on ultimate branches. The specimen from Quindio (Gradstein 12732) cited by Gradstein and Reeb (2018b) as R. hans-meyeri belongs to R. latifrons subsp. parasitans. 8. Riccardia hymenophytoides (Spruce) Meenks Aneura pectinata Spruce, nom. illeg. Riccardia sprucei (Steph.) Meenks & C.De Jong

Fig. 8K, L

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On rotten wood, shaded soil, and rock in montane rainforest, ca. 200–3000 m. Colombia: Cho, Mag, Met, Qui, Ris, San. Ecuador: EO (Reserva Buenaventura, 300–400 m, Benitez et al. 1320, HUTPL), Pi (Gradstein and Reeb 2018b), Tu (slope of Mt Tungurahua, 3000  m, Thiers 4519, QCA). General distribution: West Indies, tropical Andes (Venezuela to Bolivia), Guianas, Brazil. Riccardia hymenophytoides is a widespread neotropical species that is distinguished by (1) plants monoicous, more rarely dioicous, pale green, small to rather robust, ca. 1–2  cm long, ascending to erect from a creeping base, 2-pinnate and usually palmate (branches suberect) in the upper part, in the lower half with short or long stoloniform branches; (2) main axis stolon-like, not winged, broadly ellipsoid to rounded, in cross section, 7–15 cells thick, without subepidermis, all cells thinwalled, epidermis cells much smaller than medullary cells; (3) normal branches flat, linear or tongue-shaped, mostly suberect, with broad transparent wings, the wings 4–8 cells wide, broader that the midrib; (4) calyptra covered by thick-walled papillae; (5) and gemmae not observed. Riccardia hymenophytoides is close to R. fucoidea but differs from the latter in the creeping main axis without subepidermis and palmate branching. The species is normally monoicous, but unisexual populations may occur (e.g., the type of R. hymenophytoides). Plants from higher elevation are more robust. 9. Riccardia latifrons (Lindb.) Lindb. subsp. parasitans (Steph.) Gradst. & Reeb Riccardia parasitans (Steph.) Meenks & C.De Jong Fig. 9K, L On humic soil, rock, and rotten wood in upper montane cloud forest and páramo, 2600–4500 m. Colombia: Ant, Ara, Boy, Cho, Cun, Hui, Met, Qui, Ris. Ecuador: Ca, Co, Lo, Na, Pi. General distribution: tropical Andes, usually at high elevation. Riccardia latifrons subsp. parasitans is characteristic of páramo and recognized by (1) plants paroicous or autoicous, small, to 1 cm long, with a prostrate main axis and numerous small ascending branches, irregularly 2(−3)-pinnate to palmate, primary branches widely to obliquely spreading, ultimate branches suberect forming a palmate branching pattern; (2) axis without wing, swollen, stoloniform, bi-convex to rounded, 10–15 cells thick, without or with a subepidermis, axial epidermis cells subquadrate, usually narrower than medullary cells, thin-walled or slightly thickwalled, flat to slightly mammillose, trigones absent; (3) branches ligulate to lanceolate, usually wingless, occasionally with a rudimentary, 1-cell-wide wing, margins entire, apex rounded, epidermis cells on ultimate branches often orientated obliquely toward the margin; (4) male branches usually developed on the side of the female branch (paroicous), more rarely separate (autoicous); and (5) gemmae not observed (Gradstein and Reeb 2018a, b). The paroicous sexuality, palmate branching (in the upper part of the thallus), wingless branches, stoloniform axis, small-celled epidermis cells, and absence of gemmae are the principal characters of R. latifrons subsp. parasitans. The epidermis cells of the ultimate branches are often orientated obliquely toward the thallus margin. The subspecies is closely similar to subsp. latifrons, a widespread Holarctic taxon, but differs from the latter in the thicker axis and, possibly, in the distribution of the oil bodies which are absent in the epidermis of subsp. latifrons but were

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observed in epidermis cells of subsp. parasitans by Meenks and de Jong (1985). The close relationship of the two subspecies was supported by molecular analysis (Schäfer-Verwimp et al. 2013a). 10. Riccardia leptophylla (Spruce) Herzog Aneura allionii Steph. Riccardia tenuicula (Spruce) Meenks (?) On rotten wood and trunk bases in moist lowland and montane forests up to the lower páramo, 100–3350 m. Colombia: Ant, Boy, Cho, Cun, Hui, Ris, San, Vau (Schultes & Cabrera 13952, CINC, PC). Ecuador: Co, LR, MS, Or, ZC. General distribution: tropical America. Riccardia leptophylla is one of the smallest Riccardia species and characteristic of rotten wood. The species is distinguished by (1) plants monoicous, occasionally dioicous, very delicate, less than 1 cm long, translucent, prostrate, 2-pinnate to palmate; (2) axis only 3–6 cells thick, plano-convex to ellipsoid, epidermis cells smaller than medullary cells, margins winged, wings 1–3 cells wide; (3) branches mostly linear, sometimes tongue-shaped, broadly winged, wings usually wider than the midrib, the midrib vanishing below apex; (4) epidermis cells on branches usually in straight rows; and (5) gemmae present. Riccardia leptophylla is rather similar to R. regnellii, but the branches are mostly linear and the wings in R. leptophylla are wider and are usually broader than the midrib. Riccardia tenuicula, described from Amazonia, is probably a synonym of R. leptophylla. 11. Riccardia pallida (Spruce) Meenks & C.De Jong Fig. 9I, J Riccardia boliviensis (Steph.) Meenks Riccardia capillacea (Steph.) Meenks & C.De Jong Riccardia capillacea var. dentata Meenks Riccardia foliacea Meenks & C.De Jong, syn. nov. Type: Ecuador, Napo, páramo de Guamani, 4000 m, Apr 1982, Meenks & De Vries 214, very sparse, c. gyn. (holotype U-0033759!; isotype QCA-203238!) Riccardia herzogiana (Steph.) Meenks & C.De Jong Riccardia trichomanoides (Spruce) Meenks On rough bark, rotten wood, and humic soil, in humid montane forests and páramo, and also in Sphagnum bogs, 1000–4000 m. Colombia: Ant, Ara, Boy, Cas, Cau, Cho, Cun, Hui, Mag, Met, Qui, Ris, San. Tol. Ecuador: Az, Ca, Ch, Co, Na, Pi, Tu, ZC. General distribution: tropical Andes. Riccardia pallida is a common Andean species that is distinguished by (1) plants dioicous, very slender, to 6 cm long, regularly to irregularly 2–4-pinnate, growing upright or pendent, primary branches often deflexed, opposite or alternate, stolons lacking; (2) axis 0.5–0.75 mm wide, usually bi-convex, (7–)8–12 cells thick, with a subepidermis (sometimes only weakly developed) and with a narrow, interrupted, 1–2(−3)-cell-wide wing (part of the axis without wing), axial wing cells quadrate to narrowly rectangular, 15–30(−35) μm wide and 25–80 μm long, thin-walled, without trigones; (3) primary branches winged like the axis, ultimate branches very thin, linear to subulate, straight or deflexed, with a conspicuous, 2–4-cell-wide wing, the

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wings as wide as or narrower than the midrib, wing cells elongate-hexagonal, ca. 20–30(−35) μm wide and 30–50 μm long, in straight or oblique rows, thin-walled with small trigones, ultimate branch margins entire to crenate to remotely toothed, teeth 0.5–1(−2) cells long; (4) oil bodies finely granular (Jungermannia-type), 1–4 per cell, present in 60–80 percent of the epidermis cells; (5) antheridial branches winged, the wing toothed or subentire; (6) gynoecial branches subtended by 1–2 innovating sterile branches, margins with scale-like paraphyses; and (7) gemmae very rare, produced on ± rounded, deeply concave, leaf-like gemmiparous branches. Riccardia pallida may be confused with R. fucoidea; for differences see under the latter species. Riccardia trichomanoides and R. herzogiana are robust and rather densely branched, 3–4-pinnate phenotypes of R. pallida, while R. capillacea is a delicate, 2-pinnate phenotype with deflexed branches. A phenotype with more or less rounded and deeply concave, leaf-like gemmiparous branches was described from Ecuador as R. foliacea. Gradstein and Reeb (2018b) treated this species as a synonym of R. chamedryfolia based on examination of the scanty isotype in QCA. However, study of a copious collection of R. foliacea from a 5 × 5 m vegetation plot in the páramo of Cotopaxi, Ecuador (Burghardt et al. 140308M3, PC, QCA), showed the presence of a winged bi-convex axis with a subepidermis. Except for the peculiar gemmiparous branches, these plants are fully identical to R. pallida. As the latter species was very common in the plot and mostly lacked gemmiparous branches – of ten collections of R. pallida made in the plot, only one had gemmiparous branches  – R. foliacea is considered a phenotype of R. pallida. Thus far, the gemmiparous phenotype is only known from two páramos of Ecuador (Napo, Cotopaxi). 12. Riccardia poeppigiana (Lehm. & Lindenb.) Meenks & C.De Jong Fig. 9G, H Riccardia algoides (Taylor) Meenks Riccardia cervicornis (Spruce) Gradst. & Hekking On loamy soil, rotten wood, and twigs in montane forests and páramo, also in running water, (1200–)2500–4200  m. Colombia: Ant, Boy, Ces, Cho, Cun, Hui, Met, Qui, Ris, San, Tol, Val. Ecuador: Az, Co, Lo, MS, Na (Gradstein 12795, QCA), Pi, Tu. General distribution: probably widespread in tropical America. Riccardia poeppigiana is the earliest Riccardia species described from tropical America and in the Andes largely restricted to upper montane forest and páramo, above 2500  m. The species is recognized by (1) plants dioicous, medium-sized, ascending to erect, 1.5–3(−4) cm long, the ascending shoots rather densely 2–3-pinnate, branches suberect or spreading, alternate or opposite, usually curved downward; (2) main axis without wing, bi-convex, 8–22 cells thick, with a subepidermis especially in older portions of the axis, epidermis cells as large as inner cells or slightly smaller; (3) primary branches ± without wings, often deflexed, ultimate branches narrowly linear to laneolate and narrowed to the tip, curved, winged or not, the wings 0–3(−4) cells broad, usually narrower than the midrib, the wing cells rather small, 15–35 μm wide, margins entire to irregularly crenate, marginal cells as large as or smaller than adjacent cells and sometimes slightly thick-walled, apex rounded or retuse; (4) oil bodies present in most epidermis cells, finely granular, 1–5(−10) per cell, globose to ellipsoid; (5) male branches with broad, crenulate

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wings (1–4 cells wide), female branches short and broad, with strongly laciniate margins; and (6) gemmae common, produced on decurved branch tips. Riccardia poeppigiana can be confused with R. fucoidea; both have ascending to erect-growing, freely 2–3(−4)-pinnate thalli with an unwinged axis, a subepidermis, and broadly winged androecia. However, R. fucoidea has straight, regularly and broadly winged branches with broad tips and large, 35–50-μm-wide wing cells and lacks gemmae. The development of the branch wings varies considerably; a robust phenotype with rather broadly winged ultimate branches was described as R. algoides, whereas phenotypes with ± unwinged branches were called R. cervicornis. 13. Riccardia regnellii (Ǻngstr.) K.G.Hell Riccardia amazonica (Spruce) Gradst. & Hekking

Fig. 8F

On decaying wood, soil, tree bases, and rock in lowland and montane rainforests up to the páramo, from sea level to 3500 m. Colombia: Ama, Boy, Cau, Cho, Cun, Hui, Mag, Ris, San, Vau. Ecuador: Co, Es, Im, Lo, MS, Na, Pa, Pi, Tu, ZC. General distribution: widespread in tropical South America, tropical Africa (?) (see Reeb and Bardat 2014). Riccardia regnellii is a common tropical South American species that was previously called R. amazonica (Gradstein and Reeb 2018a). The species is distinguished by (1) plants dioicous or paroicous, delicate, 0.5–1.5 cm long, prostrate, irregularly 1–2-pinnate, without stolons; (2) axis 4–7 cells thick, plano-convex, epidermis cells smaller than medullary cells, wing absent or narrow, 1–3 cells wide; (3) branches flat, broadly winged, often tongue-shaped and narrowed to the apex, margins straight or curved, apex rounded or retuse, wing cells thin-walled or slightly thickened, margin cells not conspicuously larger than adjacent cells; (4) surface cells of branches arranged in oblique rows; (5) oil bodies present in almost all epidermis cells, 1–2 per cell; (6) apex of the calyptra umbonate; and (7) gemmae occasionally present at branch tips. Dioicous populations of R. regnellii are sexually dimorphous with male thalli being smaller than female thalli (Reeb and Bardat 2014). The two types of thalli may occur mixed in the same mat. A specimen from Ecuador (Imbabura, on roadside near El Paraiso, 974 m, Brokamp & Mittelbach 2011–17, QCA) stands out by the presence of numerous brownish papillae on the ventral surface of the thallus. 14. Riccardia verticillata Gradst. & Reeb

Fig.: see page v

On shaded, humic soil in evergreen montane forest, 1900–2200 m. Ecuador: MS (Cordillera del Condor, El Quimi Reserve; Gradstein et al. 2019b). General distribution: only known from Ecuador; to be expected in Peru. Riccardia verticillata is a magnificent, robust dendroid species that has been found once in Ecuador, near the border with Peru. The species is readily recognized by the erect axes (to 10  cm long) with regularly spaced whorls of horizontally spreading, flattened, very densely 3–5-pinnate fronds. The axes are fully wingless, ellipsoid or bi-convex, 30–40 cells thick and with a subepidermis, and the fronds are wingless below and broadly winged above. The ultimate branches are linear, broadly winged and with crenate to toothed to incised-lobed margins, and the wings are

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3–4× wider than the midrib and are made up of very large cells, 50–80 × 30–50 μm, with small trigones (Gradstein et al. 2019b). Riccardia verticillata is related to R. fucoidea but is readily recognized from the latter by the characteristic, verticillate habit. Moreover, the axis in R. fucoidea is thinner, all the branches are winged also the primary ones, and the wing cells in R. fucoidea are usually thin-walled without trigones. In its strikingly verticillate habit, R. verticillata is similar only to R. prehensilis (Hook.f. & Taylor) C.Massal. from Tierra del Fuego and R. eriocaula (Hook.) C.Massal. from Australia and New Zealand. In the latter two species, however, the branches are scarcely winged, and the surface of axis and branches is roughened by strongly papilliform epidermis cells. 15. Riccardia wallisii (Steph.) Gradst. Fig. 8I, J Riccardia smaragdina Meenks & C.De Jong Riccardia judithae Meenks & C.De Jong, syn. nov. Type: Bolivia, La Paz, Prov. Yungas, ca. 10 km from Unduavi, 3225 m, Meenks & Ybert 439 (holotype U!), 424 (paratype U!) Riccardia regina Meens & C.De Jong, syn. nov. Type: Bolivia, La Paz, Prov. Yungas, ca. 10 km from Unduavi, 3225 m, Meenks & Ybert 431 (holotype U!), 420 (paratype U!) On humic soil, bark, rotten wood, and rock in upper montane forests and páramo, also among Sphagnum in bogs, 2300–4100 m. Colombia: Ant (type), Cho, Cun, Met, Qui, Ris, San. Ecuador: Co, Lo, Pi, ZC. General distribution: tropical Andes (Colombia to Bolivia). Riccardia wallisii is a rather robust Andean species from upper montane forests and páramo that is characterized as follows: Plants dioicous, prostrate to ascending, 1–6 cm long, irregularly 1–2(−3)-pinnate to palmate, stolons absent. Axis (0.5–) 0.8–2  mm wide, widened toward branch bases, surface sometimes spotted by brownish-colored cells, plano-convex to slightly ellipsoid, 7–14 cells thick in the middle, subepidermis absent or present, 0–2 cells wide, epidermis cells as large as inner cells or smaller; axis irregularly winged, wings 1–6(−8) cells wide, wing cells quadrate to rectangular, varying in size, 20–50 μm wide, margin cells often thickwalled. Branches alternate, suberect to obliquely spreading, gradually arising from the axis and with a decurrent base, short and plump with a broad apex, little narrower than the axis, not deflexed, irregularly winged, wings narrow or broad, 1–7 cells wide, wing margin cells thin-walled or thick-walled, margins entire or crenulate, branch apex truncate to retuse; oil bodies present in almost all epidermis cells, ca. 1–15 per cell, finely segmented. Male and female branches ± winged, female branches plump, short and broad, solitary or in pairs, with lacinia. Male plants sometimes quite small. Gemmae frequently present at branch tips. The rather broad and flat, plano-convex to slightly ellipsoid axis and short and plump, obliquely spreading branches with decurrent bases and broad tips are the main characteristics of R. wallisii. The axis is often widened toward the branch bases, the margin cells are often thick-walled, and the axial surface is sometimes dotted by brownish-colored cells. The wings vary considerably in width and a subepidermis is present or absent. Plants with rudimentary wings have been called R. smaragdina.

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Excluded Records Riccardia multifida (L.) Gray  – Colombia (Kunth 1822; Gottsche 1864). Widespread Holarctic species, in tropical America rare and recorded from Brazil (Gradstein and Costa 2003). The specimens from Colombia were probably misidentified but are lost (Meenks 1987). Riccardia palmata (Hedw.) Carruth. – Colombia (Gottsche 1864). Widespread Holarctic species, not known with certainty from the tropics (records from Mexico and Cuba may belong to R. digitiloba). The specimens from Colombia were presumably misidentified but are lost (Meenks 1987).

Metzgeriaceae A family of two to four genera worldwide, one genus in Ecuador. Metzgeria Raddi Plants thalloid, thin and with a narrow midrib (midrib rarely absent), pale green to yellowish, sometimes blue when dry, 0.5–15 cm long, to 2(−3) mm wide, prostrate to erect or pendent, forked, rarely pinnate, usually with hairs (= rhizoids) on the margins, on the underside of the midrib and, sometimes, on the ventral surface of the thallus. Hairs unicellular, arising singly or in pairs, sometimes in groups of three to five. Thallus surface plane to strongly convex with reflexed margins. Midrib 2–8(−12) cells wide, epidermis cells smooth or papillose, larger than the inner cells; central strand absent. Thallus wings unbordered or with a narrow border of elongate cells. Oil bodies absent or very small, homogeneous. Monoicous or dioicous. Gametoecia on highly abbreviated branchlets scattered along the ventral side of the midrib, the antheridia inside a globose sac (= inrolled male branchlet), the archegonia hidden under a thallus flap (= female branchlet or “involucre”). Sporophytes surrounded by a fleshy calyptra, the calyptra often densely hairy; pseudoperianth lacking. Capsules spherical to ovoid, opening by 4 valves, wall 1–2-layered. Spores small, unicellular. Elaters 1-spiralled, attached to valve apices. Vegetative reproduction by gemmae or caducous branches. About 75 species worldwide, 33  in Colombia and Ecuador. The main generic characters of Metzgeria are (1) thallus dichotomously branched (rarely pinnate), very thin, of 1 layer of cells, usually with a narrow midrib, margins usually with hairs (= rhizoids); (2) gametoecia on the underside of the midrib; (3) sporophyte in a fleshy calyptra, pseudoperianth lacking; and (4) elaters attached to the tips of the capsule valves. The last two characteristics are shared with Aneuraceae. Important characters for identification of Metzgeria species include (1) ramification dichotomous or pinnate; (2) thallus inrolled or not; (3) dry thallus with or without blue coloration; (4) attenuate thallus shoots (producing gemmae near apex)

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present or absent; (5) ventral thallus surface hairy or smooth; (6) hairs single or paired, straight or falcate; (7) number of epidermal cells across the dorsal and ventral surfaces of the midrib; (8) thallus margins entire or crenulate, bordered or not; (9) gemmae produced on the thallus surface or only on the margins and (10) plants monoicous or dioicous. Unless otherwise indicated the number of epidermal cells across the surface of the midrib should be observed on older, mature portions of the thallus, not on young shoots nor at bifurcations. Literature: Costa (2008) 1. Thallus mostly consisting of large, globose sacs. Plants growing on twigs in páramo and upper montane forest in southern Ecuador . . . . . . . . . . . . M. holzii 1. Thallus not consisting of large globose sacs . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Branching pinnate or mixed dichotomous-pinnate. Midrib on main axis 4–13 cells wide on dorsal and ventral surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Branching purely dichotomous or branches lacking. Midrib 2–4(−8) cells wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Marginal hairs 1 per cell, rather scarce, straight. Thallus 1–3-pinnate, strongly convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. filicina 3. Marginal hairs 2(−4) per cell, numerous, falcate. Thallus dichotomous to irregularly 1-pinnate, plane to somewhat convex . . . . . . . . . . . . . . . . . M. claviflora 4. Large parts of the thallus strongly inrolled, ventral thallus surface almost invisible (at least in younger, green portions of the thallus). Dioicous. . . . . . . . . . 5 4. Thallus not or partly inrolled, at least part of ventral thallus surface visible. Monoicous or dioicous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5. Hairs at thallus margins of well-developed plants mostly 1 per cell or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Hairs at thallus margins of well-developed plants mostly 2 per cell. . . . . . . . 7 6. Calyptra with large, laminate appendices. Ventral surface of midrib 2 cells wide. Thallus and gemmae not becoming blue when dry. Plants endemic to the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. grandiflora 6. Calyptra without large, laminate appendices. Ventral surface of midrib 2–4 cells wide. Thallus and gemmae sometimes becoming blue when dry. Plants of upper montane forest and páramo, not on the Galápagos Islands����������� M. dorsipara 7. Thallus strongly swollen and undulate. Hairs at thallus margins ± straight. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. inflata 7. Thallus very slender, not strongly swollen and not undulate. Hairs at thallus margins mostly strongly falcate. Plants very common . . . . . . . . M. leptoneura 8. Thallus margins without hairs. (Thallus plane, ventral surface strongly hairy, ventral surface of midrib 4–7 cells wide] . . . . . . . . . . . . . . . . . . . . M. mexicana 8. Thallus margins with hairs (but hairs sometimes scarce) . . . . . . . . . . . . . . . . 9 9. Some attenuate thallus branches present (branch gradually narrowed to the apex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Attenuate branches absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 10. Thallus and gemmae becoming blue when dry . . . . . . . . . . . . . . . . . . . . . . . 11 10. Thallus and gemmae not becoming blue when dry. . . . . . . . . . . . . . . . . . . . . 13

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11. Ventral and dorsal surfaces of midrib up to 8 cells wide on attenuate branches. Gemmae produced on thallus margin and surface . . . . . . . . . . . . . . . M. violacea 11. Ventral and dorsal surfaces of midrib 2 cells wide on attenuate branches. Gemmae produced on thallus margin only . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Hairs at thallus margins 1 per cell. Gemmae ± plane . . . . . . . .   M. scyphigera 12. Hairs at thallus margins (1–)2 per cell (never only 1). Gemmae strongly concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. agnewiae 13. Dorsal surface of midrib only 2 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13. Dorsal surface of midrib 2–5 cells wide . . . . . . . . . . . . . M. parvi-involucrata 14. Ventral surface of midrib 2 cells wide. Marginal hairs straight or slightly falcate. Gemmae concave or plane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Ventral surface of midrib mostly more than 2 cells wide. Marginal hairs strongly falcate. Gemmae plane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. cleefii 15. Branches of two types: attenuate and non-attenuate. Gemmae concave . . . . . . ����������������������������������������������������������������������������������������������� M. consanguinea 15. All branches attenuate. Gemmae plane . . . . . . . . . . . . M. albinea var. angusta 16. Plants monoicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Plants dioicous, or sterile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Thallus flat, margins in part without hairs, the hairs single (1 per cell) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lindbergii 17. Thallus convex and with ± recurved margins, the margins densely hairy, hairs mostly paired (2 per cell). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. conjugata 18. Ventral surface of midrib mostly 2 cells wide. . . . . . . . . . . . . . . . . . . . . . . . 19 18. Ventral surface of midrib more than 2 cells wide (to be examined in the middle of the thallus branches, away from the bifurcation) . . . . . . . . . . . . . . . . . . . . 27 19. Thallus becoming blue when dry . . . . . . . . . . . . . . . . . . . . . . . . . M. jamesonii 19. Thallus not becoming blue when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Hairs at thallus margins mostly 1 per cell (exceptionally 2). [Plants delicate, less than 1 mm wide]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20. Hairs at thallus margins mostly 2 or more per cell . . . . . . . . . . . . . . . . . . . . 23 21. Ventral thallus surface without hairs. Thallus flat. . . . . . . . . . . . . . . . M. ciliata 21. Ventral thallus surface with numerous hairs. Thallus strongly convex to ± flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Thallus ± plane. Gemmae commonly present, rhizoids on gemmae with hooked tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. uncigera 22. Thallus convex. Gemmae lacking . . . . . . . . . . . . . . . . . . . . . . . . . M. chilensis 23. Marginal hairs varying from straight to flexuose to somewhat falcate��������������������������������������������������������������������������� M. albinea var. albinea 23. Marginal hairs mostly falcate������������������������������������������������������������������������� 24 24. Ventral thallus surface with hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24. Ventral thallus surface without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25. Marginal hairs 3–5 per cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. polytricha 25. Marginal hairs 2 per cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. bischlerae 26. Thallus margins plane or narrowly recurved, with a weak border of elongate cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. procera 26. Thallus margins broadly and strongly recurved, without border of elongate cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. leptoneura

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27. Thallus margins bordered by narrowly elongate cells. Marginal hairs mostly 1 per cell, straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 27. Thallus margins not bordered by narrowly elongate cells. Marginal hairs 1–3 per cell, straight or falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 28. Plants becoming blue when dry. Dorsal surface of midrib 2–5 cells wide. Thallus cells 20–55  μm long. Gemmae abundant, produced on thallus surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lechleri 28. Plants not becoming blue when dry. Dorsal surface of midrib only 2 cells wide. Thallus cells larger, 60–110 μm long. Gemmae absent or scarce, produced on thallus margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. maegdefrauii 29. Marginal hairs only 1 per cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 29. Marginal hairs 1–2(−4) per cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 30. Dorsal surface of midrib 2 cells wide. Gemmae produced on thallus margins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. furcata 30. Dorsal surface of midrib more than 2 cells wide. Gemmae produced on thallus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. crassipilis 31. Dorsal surface of midrib 2 cells wide. Ventral thallus surface ± smooth. Thallus margins ± recurved, abundantly with gemmae . . . . . . . . . . . . . . M. myriopoda 31. Dorsal surface of midrib at least in part more than 2 cells wide. Ventral thallus surface hairy or smooth. Thallus margins plane or recurved, with or without gemmae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 32. Thallus strongly convex (with strongly recurved margins), becoming blue when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pulvinata 32. Thallus plane or convex, not becoming blue when dry . . . . . . . . . . . . . . . . . 33 33. Marginal hairs 1–2 per cell, ± straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33. Marginal hairs densely crowded, 2–4 per cell, straight to strongly falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 34. Gemmae produced on the thallus surface. Thallus margins not crenulate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. liebmanniana 34. Gemmae produced only at the thallus margin, not on the surface. Thallus margins ± crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. rufula 35. Ventral thallus surface densely hairy. Marginal hairs 0.1–0.3  mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. fruticola 35. Ventral thallus surface ± smooth. Marginal hairs less than 0.1  mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. claviflora 1. Metzgeria agnewiae Kuwah. On branches and twigs of shrubs, rotten wood, and living leaves in montane forests and páramo, 1800–4430 m. Colombia: Boy, Cal, Cun, Tol. Ecuador: Az, Ca, Na, Pi, ZC. General distribution: tropical America, Africa. Metzgeria agnewiae is a widespread Afro-American species that is distinguished by the blue coloration of the thallus, the presence of attenuate branches, the 2(−3)-cell-wide surface of the midrib (both dorsally and ventrally), marginal hairs 1–2 per cell, and gemmae deeply concave, produced on apical margins of attenuate branches.

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Metzgeria agnewiae belongs to the group of Metzgerias with attenuate, gemmiparous branches; other species in this group include M. cleefii, M. consanguinea, M. parvi-involucrata, M. scyphigera, and M. violacea. Metzgeria cleefii and M. consanguinea differ from M. agnewiae by the absence of blue coloration, M. scyphigera by plane gemmae and marginal hairs one per cell, M. parvi-involucrata by the 2–5-cell-wide dorsal and ventral surface of the midrib, and M. violacea by the broad, to 8-cell-wide surface of the midrib on attenuate branches (2 cells wide on non-attenuate branches). 2a. Metzgeria albinea Spruce var. albineaFig. 10A, B Metzgeria australis sensu Kuwah. (non typus) Metzgeria thomeensis sensu Kuwah. (non typus) On bark, living leaves, rotten wood, rock, and soil in montane forests, 700–4000 m. Colombia: Boy, Ces, Cho, Cun, Nar, NSa, Qui, Ris. Ecuador: EO, Na, Pa, Pi, ZC, Tu. General distribution: pantropical. Metzgeria albinea is a common and widespread tropical species that is readily recognized by the thallus with the midrib mostly 2 cells wide both dorsally and ventrally (rarely 2–3 cells wide ventrally) and with mostly twinned marginal hairs, (1–)2 per cell, that are straight to flexuose to falcate, but not regularly strongly falcate. The thallus margins are flat or inrolled. Almost all collections of M. albinea from Colombia and Ecuador belong to var. albinea; two records – both from Ecuador – belong to var. angusta (Costa 2008). For differences between the two varieties, see under var. angusta. Metzgeria albinea is morphologically close to M. leptoneura, but in the latter the thallus is more strongly inrolled and the hairs are regularly and strongly falcate. 2b. Metzgeria albinea var. angusta (Steph.) Costa & Gradst. Metzgeria angusta Steph. On bark, 2700 m. Ecuador: Su (Herzog 1942; Arnell 1962). General distribution: tropical and subtropical America. Metzgeria albinea var. angusta differs from the typical vatiety in the thallus being slightly narrowed to the apex and in the larger spores, 30–35 μm in diameter (10–25 μm in var. albinea). Moreover, the ventral thallus surface is always hairy in var. angusta while being smooth or hairy in var. albinea (Costa 2008). 3. Metzgeria bischlerae Kuwah. On rock in páramo, (1800–)3500–3650 m. Colombia: Boy, Cau, Hui. Ecuador: ZC (Parolly et  al. 2004). General distribution: northern Andes (Colombia, Ecuador). Metzgeria bischlerae is a rare páramo species that is distinguished by (1) ventral thallus surface densely hairy: (2) dorsal and ventral surfaces of midrib 2 cells wide; and (3) marginal hairs paired, falcate. The species approaches M. polytricha and M. leptoneura, but M. polytricha differs in having 3–5 marginal hairs per cell and M. leptoneura in the smooth ventral thallus surface.

Fig. 10 (A and B) Metzgeria albinea var. albinea (A) Portion of cross section of thallus. (B) Thallus margin showing twinned hairs (two per cell). (C) Metzgeria conjugata, habit showing androecia, gynoecia, and sporophyte, ventral view. (D) Metzgeria dorsipara, ventral view. (E) Metzgeria claviflora, branching pattern. (F–H) Metzgeria filicina. (F) Branching pattern, 1.5×. (G) Portion of cross section of main axis, 150×. (H) Portion of cross section of ultimate branch, 150×. (I) Metzgeria fruticola, portion of thallus in cross section, showing hairs on ventral thallus surface and margin cell with 4 hairs. (J) Metzgeria myriopoda, cross section of costa. (K) Metzgeria lechleri, thallus margin. (L) Metzgeria leptoneura, portion of cross section of thallus. (M and N) Metzgeria violacea. (M) Habit, showing attenuate branches with a broad costa, 10×. (N) Branch apex with gemmae, dorsal view, 40×. (A, B from Gradstein and Costa 2003; C, L redrawn from Paton 1999; D, E, J, K, M, N redrawn from Costa 2008; F–I redrawn from Kuwahara 1976)

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4. Metzgeria chilensis Steph. On soil and rotten wood in páramo, 3050–4000 m. Colombia: Cun. Ecuador: Na, Pi, ZC. General distribution: northern Andes (Colombia, Venezuela), central Chile (Concepción); the record from New Zealand belongs to M. furcata (So 2002). Metzgeria chilensis is a delicate, rare Andean species that is characterized by the convex thalli with a densely hairy ventral surface, single marginal hairs, and a 2-cell-wide midrib surface (both dorsally and ventrally). The species approaches M. ciliata, but in the latter the thallus is very flat and the ventral leaf surface is smooth. 5. Metzgeria ciliata Raddi Metzgeria furcata (L.) Corda var. decipiens C.Massal. (Metzgeria decipiens [C.Massal.] Schiffn.) Metzgeria tonsa Herzog (?) (fide Costa 2008) On bark and living leaves, occasionally on rotten wood or rock, 200–3000(−4100) m; on the Galápagos Islands between 500 and 750 m. Colombia: Ant, Ces, Cun, Mag, Met, Qui, Ris. Ecuador: Ga, Lo, LR, Na, Or, ZC. General distribution: pantropical. Metzgeria ciliata is a common and widespread tropical species that is characterized by (1) thallus rather delicate, 0.5–1 mm wide, ± flat; (2) marginal hairs scarce, single (one per cell); (3) surface of midrib 2 cells wide, both dorsally and ventrally; and (4) common presence of small, caducous thallus branches, functioning as propagules. Very rarely the midrib is absent (Kuwahara 1978). Metzgeria ciliata is close to M. furcata and has been considered a variety or forma of the latter but differs by the narrower, 2-cell-wide ventral surface of the midrib (3–4 cells wide in M. furcata). The species is also close to M. chilensis and M. uncigera but differs from the latter two in the lack of hairs on the ventral thallus surface. 6. Metzgeria claviflora Spruce Metzgeria gigantea Steph. Metzgeria hygrophila Spruce

Fig. 10E

On bark, rotten wood, and moist soil along rivers and near waterfalls, in montane forest areas, and in páramo, 900–4350  m. Colombia: Ara, Boy, Cun, Met, Ris. Ecuador: Ca, Ch, Co, Na, Pi, ZC. General distribution: Nicaragua to Bolivia, West Indies, Guianas. Metzgeria claviflora is a rather widespread neotropical species that is recognized by the numerous short, falcate hairs at the thallus margin (1–4 hairs per cell) and the broad, 4–5-cell-wide ventral and dorsal surface of the midrib. The ramification varies from dichotomous to irregularly pinnate. Metzgeria claviflora may be confused with M. filicina and M. fruticola; for differences see the key. Spruce (1885) distinguished three varieties in the species, all of which are synonyms (Costa 2008). Based on the detailed descriptions and illustrations in Costa (2008), Metgeria spindleri Steph. described from Bolivia and recorded once from Ecuador might be conspecific with M. claviflora.

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7. Metzgeria cleefii Kuwah. On tree trunk in dwarf forest near the forest line, ca. 3300 m. Colombia: Cau (Puracé; Costa 2008). General distribution: only known from the type from Colombia. Metzgeria cleefii is a rare Colombian páramo endemic that is distinguished by (1) thallus with attenuate branches; (2) marginal hairs (1–)2(−3) per cell; (3) ventral surface of the midrib 2–6 cells wide; (4) gemmae flat, produced on the apical margins of attenuate branches; and (5) thallus and gemmae not becoming blue when dry. By the presence of attenuate branches and the absence of blue coloration, M. cleefii is similar to M. consanguinea; for differences between these two species, see the key. The record of M. cleefii from Boyacá (Álvaro Alba et al. 2007) needs verification; the characteristic attenuate branches are not shown in the illustrations in the paper. 8. Metzgeria conjugata Lindb. Metzgeria minor (Schiffn.) Kuwah. var. cyanogutta Kuwah.

Fig. 10C

On bark in disturbed upper montane forest, 2300–3820  m. Colombia: Boy (páramo of Belén, Cleef 2038, PC), Cun (Kuwahara 1976). General distribution: common and widely distributed in the temperate regions of the Northern Hemisphere, scattered in the tropics. Metzgeria conjugata is a Holarctic species that is very rare in the northern Andes; a few collections are available from the Eastern Cordillera of Colombia. The species is readily characterized by the convex thallus with recurved margins and frequent ventral branches, thallus margins with numerous twinned hairs (2 hairs per cell), and monoicous sexuality. Androecia and gynoecia are produced mixed and usually in great abundance on the ventral midrib of the thallus. Gemmae are rare. Metzgeria conjugata may be confused with M. lindbergii, the only other monoicous Metzgeria in tropical America (Gradstein et al. 2016). However, M. lindbergii clearly differs from M. conjugata in the flat thallus with margins not recurved, the scarcity of marginal hairs (some parts of the thallus may be without hairs), and hairs only one per cell. 9. Metzgeria consanguinea Schiffn. Metzgeria attenuata Steph. Metzgeria neotropica Kuwah. (?) Type: Peru, Cajamarca, 3350 m, Hegewald 6614 (MO n.v.) Metzgeria temperata Kuwah. On branches and twigs of trees and shrubs and on living leaves in rather open, montane environments up to the páramo, 1500–4000 m. Colombia: Ara, Boy, Cas, Cun, Met, Qui. Ecuador: Ca, Co, Im, Na, Pi, ZC. General distribution: pantropical; also in western Europe and eastern North America where the species was known as M. temperata. Metzgeria consanguinea is a very common and widespread, pantropical species that is recognized by (1) thallus green in the shade and yellowish in open sites, not becoming blue when dry, variable in size, broad or very small and flagelliform,

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attenuate branches with marginal gemmae; (2) marginal hairs ± straight, 1–2 per cell; (3) ventral and dorsal surface of the midrib mostly 2 cells wide; (4) ventral thallus surface smooth or hairy; and (5) gemmae concave, rounded or elongate, produced on the apical margins of attenuate branches, not becoming blue when dry. The attenuate branches of M. consanguinea vary considerably in width and are usually very long and narrow in small, yellowish plants from open habitas, while being more plump and broad in green plants from more shaded sites. The midrib may be absent in weakly developed plants (e.g., Venezuela, Mérida, Pócs 9718/AU, EGR). Metzgeria attenuata was treated as synonym of M. consanguinea by So (2004) but as a separate species by Costa (2008). Metzgeria neotropica – recorded from Colombia and Ecuador – is morphologically very similar to M. consanguinea but differs in male thalli being somewhat smaller than female thalli (not smaller in M. consanguinea). Possibly, M. neotropica is a synonym or variety of M. consanguinea. Metzgeria scyphigera is very similar to M. consanguinea but differs from the latter by blue coloration of dried plants. Costa (2008), however, reported the occurrence of occasional blue coloration in M. consanguinea. 10. Metzgeria crassipilis (Lindb.) A.Evans On bark and rock in open montane forests, plantations, and isolated trees, 800–2500  m. Colombia: Cun (Onraedt 72-A52, GOET). Ecuador (new): EO (road El Cisne to Zaruma km 5, Jörgensen 56375, AAU, PC), MS (Gualaquiza, Pérez et al. s.n., QCA). General distribution: widely distributed in tropical and warm-temperate regions but rare in tropical America. Metzgeria crassipilis closely resembles the widespread M. furcata and was originally considered a subspecies of the latter but differs by the broader dorsal surface of the midrib (2–5 cells wide) and the occurrence of gemmae on the thallus surface (on thallus margins in M. furcata). The marginal hairs are one per cell and ± straight. The plants from Ecuador stand out by slight blue coloration of the dry thallus. Metzgeria allionii Steph., described from Ecuador (Gualaquiza, Allioni s.n., Bryotheca Levier 6396, holotype G-00069514), is identical to M. crassipilis but differs by young part of the thallus and gemmae with falcate hairs (Costa 2008). Possibly, M. allionii is a variety or phenotype of M. crasspilis. 11. Metzgeria dorsipara (Herzog) Kuwah. Fig. 10E Metzgeria violacea (Ach.) Dumort. var. dorsipara Herzog Metzgeria cylindra Kuwah., syn. nov. Type: Peru, Chachapoyas, between Balsas and Leimebamba, 3100 m, Hegewald 6956 (isotype F-C0000281!) Metzgeria metaensis Kuwah., syn. nov. Type: Colombia, Meta, Sumapaz, 4120 m, Cleef 8110 (isotype U!) On soil and bark in upper montane forest, scrub, and páramo, 2700–4300  m. Colombia: Boy, Cun, Met. Ecuador: Az, Ca, Lo, Pi, Tu. General distribution: northern Andes, Costa Rica. Metzgeria dorsipara is a characteristic páramo species that is distinguished by (1) thalli dioicous, strongly convex and strongly inrolled at least in younger, green portions of the thallus (older, dead portions of the thallus may be more flattened), margins straight or undulate and somewhat crispate, thallus with or without blue

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coloration when dry; (2) attenuate branches absent; (3) dorsal surface of midrib 2 cells wide, ventral surface 2–4 cells wide; (4) hairs on thallus margins mostly single, straight or flexuose, sometimes very scarce, ventral thallus surface smooth; (5) thallus cells rather thick-walled, cuticle smooth or slightly papillose, cells on dorsal surface somewhat mammillose and with thickened walls; (6) gemmae produced on thallus surface; (7) calyptra covered by numerous thick-walled hairs; and (8) spores variable in size, ca. 25–45 μm in diameter, ca. 25 μm in the type of M. cylindra, 32–37 μm in the type of M. dorsipara (Costa 2008), 34–45 μm in diameter in plants from Loja (Benitez 1153, HUTPL). Metzgeria dorsipara is rather broadly defined here and includes plants with and without blue coloration and with straight or undulate thallus margins. Plants without blue coloration were described as M. cylindra, which is here considered a synonym of M. dorsipara. Metzgeria dorsipara may be confused with M. chilensis, which also has convex thalli and marginal hairs one per cell and also occurs in páramo. The thallus of M. chilensis, however, is less strongly inrolled (ventral thallus surface well visible), and the ventral thallus surface is densely hairy (smooth in M. dorsipara). A collection from superpáramo of Ecuador (páramo de la Virgen, 4250 m, on rock, Gradstein et al. 12613, QCA) is tentatively identified as M. dorsipara. The plants stand out by thin-walled cells and virtual absence of hairs on the thallus margins. 12. Metzgeria filicina Mitt. Metzgeria bialata S.Winkl.

Fig. 10F–H

On bark in upper montane forest, scrub, and páramo, 1700–3700 m. Colombia: Boy, Tol. Ecuador: Az, Lo, Na, Pi, Tu. General distribution: tropical Andes. Metzgeria filicina is a very distinct high-Andean species that is readily separated from all other neotropical Metzgeria species by the 1–3-pinnately branched thallus. Moreover, the surface of the midrib in this species is very wide on the main axis (5–13 cell rows wide), both dorsally and ventrally. The midrib becomes narrower on branches, and on ultimate branchlets it may be only 2 cells wide both dorsally and ventrally. 13. Metzgeria fruticola Spruce Metzgeria uleana Steph.

Fig. 10I

On bark, rock, and soil in montane forest and páramo, 700–4000 m. Colombia: Boy, Cal, Cas, Met, Nar, Ris, Val. Ecuador: Co, Pi, Tu, SZ. General distribution: Mexico to Bolivia, SE Brazil. Metzgeria fruticola is a common Andean species recognized by the numerous, crowded hairs on the thallus margins (2–4 hairs per cell), the densely hairy ventral thallus surface, and the broad, 4–5-cell-wide ventral surface of the midrib. Metzgeria fruticola is close to M. rufula and M. polytricha, but M. rufula differs in less densely crowded marginal hairs (1–2 per cell), sparsely hairy ventral thallus surface, and crenulate thallus margins, and M. polytricha in the 2-cell-wide ventral surface of the midrib.

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14. Metzgeria furcata (L.) Corda On small trunks and on branches of shrubs in montane rainforest areas and páramo, rare, 1800–3300 m. Colombia: Cun (Costa 2008). Ecuador: ZC (SchäferVerwimp et al. 2013a). General distribution: common and widespread in temperate regions of the Northern Hemisphere and in Australasia, scattered in the tropics. Metzgeria furcata is a common Holarctic species that is very rare in the northern Andes. The plants are small and are distinguished by (1) marginal hairs single, one per cell; (2) dorsal surface of midrib 2 cells wide, ventral surface 3–4 cells wide; and (3) gemmae frequent on thallus margins, ellipsoid to elongate. The species is close to M. ciliata and M. crassipilis; for differences see under the latter two species. 15. Metzgeria grandiflora A.Evans Pendent epiphyte on bark in moist, open evergreen woodland and summit scrub, occasionally on rock in open pampa vegetation, 450–800  m. Ecuador: Ga (Floreana, Isabela, Pinta, Santa Cruz; Gradstein 2009). General distribution: only known from the Galápagos Islands. Metzgeria grandiflora is the only Metzgeria species endemic to Galápagos archipelago. It is recognized by (1) thallus strongly convex, inrolled, ventral thallus surface scarcely visible; (2) dorsal and ventral surfaces of midrib 2 cells wide; (3) marginal hairs single; and (4) calyptra with large, irregular, laminate appendices. Metzgeria grandiflora can be confused with small phenotypes of M. leptoneura with few, single marginal hairs (e.g., Chocó, Gradstein 8899, COL, GOET). Such abnormal phenotypes of M. leptoneura are sterile and occur together with the typical phenotype with paired hairs. Confusion of the two species is unlikely to occur as M. leptoneura is restricted to the continent and does not occur on the Galápagos Islands. 16. Metzgeria holzii Gradst. & A.R.Benitez Metzgeria saccata auct. (non typus) On twigs and branches of shrubs and bamboo in upper montane cloud forest and páramo, in a species-rich twig community with Blepharolejeunea incongrua, Cheilolejeunea fulva, Colura calyptrifolia, Diplasiolejeunea pauckertii, Drepanolejeunea cutervoensis, Frullania subtilissima, Leptoscyphus autoicus, etc., 2900–3600 m. Ecuador: Az, Lo, ZC (Gradstein and Benitez 2018). General distribution: southern Ecuador, southeastern Brazil (São Paulo); in Brazil occurring at much lower elevation in the Serra do Mar (Costa and Dias dos Santos 2016 as M. saccata). Metzgeria holzii is a rare species of southern Ecuador and Brazil and one of the most distinct species in the genus Metzgeria. It is characterized as follows: Thallus 2–3  mm wide, modified into large, globose sacs, the sacs 1–2× wider than the unlobed thallus. Thallus margins with scattered, straight or slightly curved hairs, the hairs one per cell. Ventral surface of thallus wings and midrib hairy. Midrib with 2 dorsal and 2–4 ventral rows of epidermis cells, the epidermis cells usually thickerwalled than the wing cells. Dioicous. Shoot calyptra invested with thick-walled hairs.

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Keys and Descriptions

Metzgeria holzii is readily distinguished by the thallus with large, globose, bladder-like sacs formed by inrolled, inflated thallus lobes. Bladder-like sacs also occur in M. saccata Mitt. from Australasia, and the South American plants were initially identified as M. saccata (Benitez and Gradstein 2011). However, the sacs of M. saccata are smaller and clavate, the ventral surface of the thallus wings is naked (hairy in M. holzii), and the calyptra is covered by lamellae (by hairs in M. holzii). 17. Metzgeria inflata Steph. On bark in montane forest, 1000–3100  m. Colombia: Mag (Winkler 1976). Ecuador (new): ZC (Burghardt 7138a, QCA). General distribution: northern Andes, Panama, Puerto Rico. Metzgeria inflata is very rare in the northern Andes where it has been recorded once in Colombia (Winkler 1976; to be verified) and once in Ecuador (Reserva Biológica San Francisco; new to Ecuador). It is readily recognized by the swollen, undulate thalli with strongy revolute margins. The dorsal and ventral surfaces of the midrib are only 2 cells wide, and the marginal hairs are paired. In its undulate habit, this species resembles M. dorsipara. However, M. dorsipara differs by marginal hairs single, not paired, and ventral surface of the midrib 2–4 cells wide. 18. Metzgeria jamesonii Kuwah. On soil, 2850–4150  m. Ecuador: Pi (Costa 2008). General distribution: Ecuador, Venezuela (new). Metzgeria jamesonii is a very poorly known northern Andean species that was long known only from the type from Ecuador (Pichincha, around 1850, leg. W.  Jameson) and was recently collected in Venezuela (Mérida, Parque Nacional Sierra Nevada, Loma Redonda, Polylepis sericea forest, alt. 4100–4150 m, April 2005, T. Pócs 05030/A, EGR). The species is characterized by the strongly bluecolored, flat or convex thallus (dry) with plane or reflexed margins, 2-cell-wide dorsal and ventral midrib surfaces, and marginal hairs 1–2 per cell. The gemmae, which were unknown in the species and were seen in the Venezuelan material, are discoid, ca 200 μm in diameter when mature, concave, with rhizoids, and blue-colored when dry. They originate on the thallus margins and on the ventral margins of the midrib, near the apex of the thallus. 19. Metzgeria lechleri Steph. Metzgeria vivipara A.Evans

Fig. 10K

On bark, rotten wood, and rock, 1800–4000 m. Colombia: Boy, Cun. Ecuador: Ch, Lo, Tu, ZC.  General distribution: tropical America, Africa (Réunion), Sri Lanka. Metzgeria lechleri is a widespread tropical species distinguished by the blue coloration of the dry thallus, the 2–5-cell-wide dorsal surface of the midrib, paired marginal hairs, and thallus margins bordered by narrow cells. The species is rather similar to M. liebmanniana, but in the latter species, the lamina is unbordered, the marginal hairs are frequently single, and the dry thalli lack blue coloration.

Metzgeriaceae

20. Metzgeria leptoneura Spruce Metzgeria australis Steph. Metzgeria ecuadorensis Kuwah. Metzgeria furcata (L.) Corda var. extensa Nees Metzgeria hamata Lindb. Metzgeria nudicosta Steph. Metzgeria papulosa Steph. Metzgeria pilosa Steph. Metzgeria sandei Schiffn. Metzgeria wallisiana Steph.

117

Fig. 10L

Common on bark, rotten wood, rock, and soil in montane forest and páramo, 500–4200 m. Colombia: Ant, Ara, Boy, Cas, Cua, Ces, Cho, Cun, Hui, Mag, Met, Nar, Qui, Ris, Tol Val. Ecuador: Az, Ca, Ch, Co, Lo, LR, MS, Na, Pa, Su, Tu, ZC. General distribution: widespread in tropical and subtropical regions. Metzgeria leptoneura is a very common and widespread species that is readily distinguished by the long and slender, strongly convex thallus with broadly reflexed, straight margins (not undulate), paired falcate marginal hairs, and dorsal and ventral surfaces of the midrib only 2 cells wide. Small, sterile phenotypes of M. leptoneura may sometimes produce only few, single, straight hairs on the thallus margins. Such atypical phenotypes, which occur together with the typical form, can be confused with M. grandflora. For differences between M. grandiflora and M. leptoneura, see under M. grandiflora. 21. Metzgeria liebmanniana Lindenb. & Gottsche On bark and rock in upper montane forest, scrub, and páramo, 1900–4000 m. Colombia: Boy, Cau, Ces, Cun, Hui, Met, San, Tol. Ecuador: Ca, Na, Pi. General distribution: Mexico to northern Argentina and Chile. Metzgeria liebmanniana is fairly common in the high Andes and one of the most robust Metzgeria species in the region. The species is characterized by (1) thallus 3–5 mm broad, lamina more than 30 cells wide, ventral surface densely hairy, margins plane or narrowly recurved, entire (but appearing crenate where rhizoids are inserted between margin cells); (2) hairs at thallus margin 1–2 per cell, short, straight or slightly curved; (3) dorsal surface of midrib 4–6 cells wide in older portions of the thallus, ventral surface 5–8 cells wide; and (4) gemmae produced on thallus margins and surface, asymmetrical in shape, concave. Metzgeria liebmanniana may be confused with M. fruticola, M. lechleri, and M. rufula; for differences see under the latter species. Although originally spelled “liebmaniana,” the epithet of this species is changed to “liebmanniana” under ICN art. 60.1 (Ex. 7) since it was named for Frederik Michael Liebmann. 22. Metzgeria lindbergii Schiffn. Metzgeria saxbyi Pearson On small tree trunks in secondary montane forest and on rock in páramo, 2130–3800 m. Colombia: Boy (Gradstein 2019b), Mag (Sierra Nevada de Santa Marta; Gradstein et al. 2016). General distribution: pantropical.

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Keys and Descriptions

Metzgeria lindbergii, originally described from Java, is a pantropical species that is widespread in Africa (where it was known as M. saxbyi) and tropical Asia. In the neotropics, however, it is very rare and only known from two localities in Colombia (Gradstein 2019b). The species is readily characterized by (1) monoicous, with numerous gynoecia and androecia, flat, (0.5–)1–1.5(−1.75) mm wide, wings 15–27 cells wide, ventral branches frequently present; (2) thallus margins, ventral thallus surface, and ventral midrib with scattered single hairs, one per cell (exceptionally 2), part of the thallus margins devoid of hairs, thallus margins flat; (3) dorsal surface of midrib 2–3(−4) cells wide, ventral surface of midrib (2–)3–5(−6) cells wide; (4) thallus cells subisodiametrical, 25–50 μm wide, without or without small trigones; (5) gynoecia hairy, androecia naked; and (6) gemmae absent (not observed). The two Colombian collections differ slightly in size and in midrib width; those from montane forest of Sierra Nevada de Santa Marta (2130  m, on bark) are 0.5–1.5  mm wide with a (2–)3–4-cell-wide midrib (ventral surface), while those from páramo in the Sierra Nevada del Cocuy (3550–3800 m, on shaded rock) are 1–1.75 mm wide with a ventrally 3–5(−6)-cell-wide midrib. Metzgeria lindbergii may be confused with M. conjugata, the only other monoicous Metzgeria species in tropical America, but the latter species is readily separated from M. lindbergii by the convex thallus with recurved margins and paired hairs (2 per cell). 23. Metzgeria maegdefrauii Kuwah. On wet rock in waterfall, 2100 m. Colombia: Mag (San Lorenzo; Costa 2008). General distribution: Colombia (type), Bolivia. Metzgeria maegdefrauii is a rare Andean species that is recognized by (1) thallus ± plane, 2–3 mm wide, without blue coloration, bordered by 1–2 rows of narrowly elongate cells; (2) dorsal surface of midrib 2 cells wide, ventral surface 2–5 cells wide; (3) marginal hairs usually single; and (4) thallus cells large, ca. 60–110 × 45–75 μm (smaller at the border). The record from the Chocó (Vasco et al. 2002) needs checking. 24. Metzgeria mexicana Steph. On bark and rock in montane forest areas, 700–2400 m. Colombia: Boy, Cho, Cun. General distribution: scattered in tropical America. Metzgeria mexicana is a rare neotropical species that is readily distinguished by the absence of hairs on the thallus margin (but present on the ventral thallus surface). It is the only Metzgeria species in the northern Andes with hairless thallus margins. The dorsal surface of the midrib is 2 cells wide and the ventral surface 4–6 cells. Discoid gemmae are produced on the dorsal thallus surface. 25. Metzgeria myriopoda Lindb. Metzgeria leptomitra Spruce Metzgeria leptomitra var. angustifrons Spruce Metzgeria leptomitra var. longifurca Spruce

Fig. 10J

On bark in rather dry, open woodlands at rather low elevations, 450–1800(−2800) m. Colombia: Boy, Met. Ecuador: Ga, MS, Su, Tu, ZC. General distribution: tropical and subtropical America.

Metzgeriaceae

119

Metzgeria myriopoda is a common xerotolerant New World species that is readily recognized by the convex thallus with a 2-cell-wide dorsal midrib surface and a 3–7-cell-wide ventral surface, the recurved thallus margins with 1–3 straight or falcate hairs per cell and with numerous small gemmae, and the ventral thallus surface naked or with a few scattered hairs.The species is dioicous and usually sterile. Metzgeria myriopoda resembles M. laciniata Kuwah., a species endemic to Peru, but the latter species differs in blue coloration of the dry thallus apex and gemmae, mostly plane thallus margins with very few hairs and deeply concave gemmae. 26. Metzgeria parvi-involucrata Kuwah. On bark and rock in upper montane forest, scrub, and páramo near the forest line, ca. 3250–3500 m. Colombia (new): Cun (páramo de Chisacá, Jul 1953, Schultes 20217, CINC, PC). Ecuador: Ca (páramo El Ángel; Costa 2008). General distribution: Mexico to Bolivia, at high elevations. Metzgeria parvi-involucrata is very rare in the northern Andes and is reported here new to Colombia. The species is distinguished by (1) occasional blue coloration of parts of the thallus; (2) thallus with attenuate branches; (3) surface of midrib (2–)3–5 cells wide, both dorsally and ventrally; (4) marginal hairs 1(−2) per cell; (5) gemmae deeply concave, produced on apical margins of attenuate branches; and (6) female involucre very small. Metzgeria parvi-involucrata approaches M. agnewiae, but in the latter species, the surface of the midrib is only 2(−3) cells wide, both dorsally and ventrally, and the marginal hairs are 1–2 per cell. Moreover, M. agnewiae is more strongly blue-colored. 27. Metzgeria polytricha Spruce On bark, soil, and rock in montane forests and scrub, exceptionally in wet lowland rainforest along the Pacific coast, (50–)1000–4200 m. Colombia: Boy, Cho, Cun, Qui, Ris. Ecuador: Ca, Im, Lo, MS, Na, Pa, Pi, Tu, ZC. General distribution: tropical Andes, Costa Rica, French Guiana. Metzgeria polytricha is a common species in the Andes that is recognized by numerous, crowded hairs at the thallus margin (2–4 hairs per cell), the hairy ventral thallus surface, and the 2-cell-wide dorsal and ventral surface of the midrib. By the crowded marginal hairs, M. polytricha is similar to M. fruticola, but in the latter species, the ventral surface of the midrib is broader, 4–5 cells wide. 28. Metzgeria procera Mitt. Metzgeria procera var. columbica Herzog On bark, soil, and rock in montane forests, scrub, and páramo, 1300–4000 m. Colombia: Cau, Cun, Hui, Val. Ecuador: Im, LR, Na, Pa, Pi. General distribution: Mexico to Bolivia, West Indies. Metzgeria procera is an uncommon neotropical species that is distinguished by (1) thallus margins plane or narrowly recurved, crenulate, weakly bordered by ± elongate cells; (2) marginal hairs paired, falcate; and (3) dorsal and ventral surfaces of the midrib 2 cells wide. The species is closely related to M. leptoneura; for differences see the key.

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Keys and Descriptions

29. Metzgeria pulvinata Steph. Metzgeria gracilimascula Kuwah. On bark near the forest line, ca. 3500 m. Ecuador: Im, Pi (Costa 2008). General distribution: Mexico to Bolivia. Metzgeria pulvinata is a rare neotropical high-montane species that is recognized by (1) thallus strongly convex and with recurved margins, becoming bluecolored when dry; (2) attenuate branches absent; (3) ventral and dorsal surfaces of midrib very broad, ca. 4–8 cells wide; (4) marginal hairs falcate, usually paired, sometimes 3–4 per cell; and (5) gemmae ± plane, produced on the thallus margins. By the stronly convex, blue-colored thallus with 4–8-cell-wide dorsal and ventral midrib surfaces and usually paired, falcate marginal hairs, Metzgeria pulvinata cannot be confused with any other Metzgeria species in the region. 30. Metzgeria rufula Spruce Metzgeria crenata Steph. Metzgeria crenatiformis Steph. On bark, rotten wood, living leaves, soil and rock in montane forests, scrub, and páramo in rather open habitats, 500–4000 m. Colombia: Ant, Boy, Cho, Cun, Mag, NSa, Ris, San, Tol. Ecuador: Im, Pi, Tu, ZC.  General distribution: pantropical. Metzgeria rufula is a common and widespread tropical species that is distinguished by (1) ventral surface of midrib 4–5 cells wide, dorsal surface 2–5 cells wide; (2) marginal hairs 1–2 per cell (straight to somewhat falcate); (3) ventral thallus surface sparsely to densely hairy; (4) thallus margins crenulate; and (5) gemmae scarce and produced only on the margins of the thallus, not on the thallus surface. The species approaches M. fruticola and M. liebmanniana, but in the latter two species, the thallus margin is not crenulate (but may appear crenate where rhizoids are inserted between margin cells). Metzgeria fruticola furthermore has marginal hairs more crowded, 2–4 per cell, and M. liebmanniana produces gemmae also on the thallus surface, not only at the margin. Metzgeria bracteata Spruce and M. herminieri Schiffn., recorded from Colombia (as M. bracteata var. robusta Herzog) and from Ecuador, are very similar to M. rufula (see Costa 2008) and may be conspecific. 31. Metzgeria scyphigera A.Evans On bark and living leaves in montane rainforest and páramo, 1800–4200  m. Colombia: Boy, Cun, Ris. Ecuador: Az, Co, Lo, Na, Pi, ZC. General distribution: Mexico to Tierra del Fuego, Brazil, East Africa. Metzgeria scyphigera is a widespread Afro-American species recognized by (1) dry thallus becoming blue-colored; 2) thallus with attenuate branches; 3) ventral and dorsal surfaces of the midrib 2 cells wide; 4) hairs at thallus margins single; and 5) gemmae ± plane, produced on the apical margins of attenuate branches. Metzgeria scyphigera may be confused with M. agnewiae and M. violacea; for differences see the key.

Metzgeriaceae

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32. Metzgeria uncigera A.Evans On trees in upper montane cloud forest, 2800–3000 m. Colombia: NSa (between Pamplona and Toledo; Costa 2008). Scattered and highly disjunct in subtropical and tropical America (Florida, Puerto Rico, Colombia, eastern Brazil). Metzgeria uncigera is a rare neotropical species that is characterized by 1) thallus very delicate, ca. 0.5 mm wide; (2) marginal hairs single, straight or with a falcate tip; (3) dorsal and ventral surfaces of the midrib 2 cells wide; and (4) gemmae common on thallus margins, each gemma with numerous short rhizoids with hooked tips (hence the name “uncigera”). Metzgeria uncigera is close to M. decipiens, but in the latter species, the ventral thallus surface is naked and hooked rhizoids are lacking. 33. Metzgeria violacea (Ach.) Dumort. Metzgeria atramentaria Kuwah. Metzgeria fruticulosa (Dicks.) A.Evans, nom. illeg.

Fig. 10M, N

On bark, rotten wood, and soil in upper montane forest and shrubby páramo, 2400–3600 m. Colombia: Boy, Cun, Met. Ecuador: Ca (Costa 2008). General distribution: Mexico to Tierra del Fuego and warm-temperate regions of the Northern Hemisphere. Metzgeria violacea is a widespread species that is distinguished by the blue coloration of the thallus; the presence of attenuate branches; the broad, 4–8-cell-wide surface of the midrib on attenuate branches (but 2 cells wide on non-attenuate branches); marginal hairs 1(−2) per cell; and gemmae deeply concave, produced on apical margins and surface of attenuate branches. In the broad midrib surface on attenuate branches, this species differs from other Metzgeria species in the region with attenuate branches and blue coloration. Further Records Metzgeria aurantiaca Steph.  – Ecuador: Su (Arnell 1962). Brazilian species (Costa 2008); the record from Ecuador needs verification. Metzgeria convoluta Steph. (= M. jackii Steph.) – Ecuador: Na, Pa, ZC (Arnell 1962 as M. jackii). Species from southeastern Brazil and Paraguay (Costa 2008); the record from Ecuador needs verification. Metzgeria heteroramea Steph. – Ecuador: Co (Arnell 1962). Species not treated by Costa (2008). Subclass Pleuroziidae (Foliose Liverworts I) Plants foliose, growing with a two-sided apical cell, divided into two lateral merophytes, ventral merophyte absent. Stems without central strand. Leaves originating from 2 initial cells, 1 layer of cells thick, undivided or bifid. Underleaves lacking. Cells with or without trigones; oil bodies usually present. Antheridia in leaf

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Keys and Descriptions

axils. Archegonia terminal on long or short shoots (acrogynous), surrounded by bracts. Sporophytes surrounded by a perianth and a thin calyptra. Setae well developed, of numerous cells. Capsules ovoid, opening by 4 valves, wall 6–8-layered. Contains only the genus Pleurozia (Pleuroziaceae). The genus has traditionally been classified with the Jungermanniidae, but molecular data (e.g., He-Nygrén et al. 2004) show that it is closer to the simple thalloid liverworts. The genus shares a two-sided apical cell with the simple thalloid liverworts (3-sided in Jungermanniidae). As a result, the plants have only two rows of lateral merophytes; the ventral merophyte is absent.

Pleuroziaceae A family with a single genus. Pleurozia Dumort. Eopleurozia R.M.Schust. Plants robust, olive green to deep wine-red to purplish, 2–10  cm long, 4–7  mm wide, creeping to ascending with leafy shoots sometimes arising from a rhizomelike base, loosely branched or in dense tufts. Stems rigid, usually brown, with a thick-walled cortex. Branches lateral-intercalary. Leaves deeply concave and imbricate, strongly enveloping the stem, succubous, undivided or divided into two segments, the dorsal segment usually smaller and saccate (sometimes explanate), sac opening with a valve, leaf tips acute or short-bifid, margins entire or toothed, sometimes with a white border. Cells quadrate to rectangular, large, 40–100 μm long, with large trigones, cuticle smooth; oil bodies finely granular. Rhizoids scarce, usually restricted to the stem bases, scattered. Gametoecia on short branches. Perianths large, cylindrical, of two types: plicate fertile perianths and terete sterile perianths. Setae of numerous cells. Capsules ovoid, wall 6–8-layered. Vegetative reproduction not observed. A mostly tropical genus of 12 species, 2 in tropical America. Pleurozia is readily distinguished by the large, greenish to reddish or purplish plants with rigid stems and with deeply concave leaves in 2 rows. Underleaves are lacking. A noteworthy feature that has been little mentioned in the literature is the presence of a creeping, rhizome-like primary stem in the two neotropical Pleurozia species. Literature: Thiers (1993) 1. Leaf margins with a whitish border. Leaves undivided. . . . . . . . . P. paradoxa 1. Leaf margins without whitish border. Leaves bifid, divided into a large ventral lobe and a smaller dorsal lobe, the smaller lobe usually turned into a sac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. heterophylla

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1. Pleurozia heterophylla Fulford Growing in tufts on branches of small trees and shrubs in wet upper montane elfin forest, on poor, acid soil over crystalline or sandstone rock, 2300–2650  m. Ecuador: ZC (Reserva Biológica San Francisco; Nöske et  al. 2003). General distribution: Guayana Highland, Ecuador, Honduras. Pleurozia heterophylla is a rare neotropical species that is readily recognized by the large, reddish-purple plants with deeply concave, unevenly bilobed leaves with acute leaf lobes. The smaller (dorsal) lobe is usually modified into a sac. 2. Pleurozia paradoxa (J.B.Jack) Schiffn. Fig. 32A–C Physotium paradoxum J.B.Jack (Eopleurozia paradoxa [J.B.Jack] R.M.Schust.) On poor, acid soil or on trunk bases in upper montane elfin forest and in páramo, in habitats on crystalline or sandstone rock, 2150–4000 m. Colombia: Cau, Hui, Nar. Ecuador: Az (páramo de Patococha), Lo, Na, Tu, ZC.  General distribution: Guayana Highland, northern Andes, southern Chile. Pleurozia paradoxa is a characteristic high-Andean species that is readily distinguished by the robust, reddish-green plants with ovate-oblong, concave leaves with whitish borders. The species looks like a moss (but the leaves are in 2 rows and without a costa). Subclass Jungermanniidae (Foliose Liverworts II) Plants foliose (rarely thallose), growing with a three-sided apical cell, divided into three merophytes (two lateral, one ventral). Stems without central strand. Branches lateral or ventral, with or without collar, originating from a leaf-initial cell or from the stem. Leaves originating from 2 or 3 initial cells, 1 layer of cells thick, divided or undivided, in 2–3 rows; underleaves usually smaller than leaves or lacking. Leaf cells usually with trigones. Antheridia in leaf axils. Archegonia terminal on long or short shoots (acrogynous), surrounded by bracts and often by bracteoles. Sporophytes surrounded by a perianth and a calyptra or by a fleshy perigynium. Capsules globose to cylindrical, opening by 4 valves, wall 2–10-layered. Contains about 725 species in Colombia and Ecuador, in 117 genera and 26 families. The largest subclass of the liverworts, containing two major orders, Jungermanniales and Porellales. Key to the orders of Jungermanniidae of Colombia and Ecuador 1. Leaves with a ventral lobule (= small or large fold or sac on the ventral side of the leaf) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Porellales (p. 349) 1. Leaves without ventral lobule . . . . . . . . . . . . . . . . . . . . . . . . Jungermanniales

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Keys and Descriptions

Order Jungermanniales Plants terrestrial, saxicolous, or epiphytic. Branching lateral and/or ventral. Leaves transverse, succubous or incubous, originating from 2 initial cells; ventral lobule lacking. Setae long. Spores normally small, usually unicellular, germination exosporic. Contains about 300 species in Colombia and Ecuador, in 65 genera and 21 families. Key to the families of Jungermanniales of Colombia and Ecuador 1. Leaves deeply dissected into hair-like segments (= filaments) which are not more than 1–2 cells wide at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaves undivided or divided into segments which are more than 2 cells wide at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Leaves divided to the base into 1–4 filaments, margins entire. Plants very small, alga-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaves divided into (2–)4–9 segments, margins strongly hairy. Plants larger, not alga-like. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Underleaves ± as large as lateral leaves. Cuticle of leaf cells with narrowly elongate papillae (“striate-papillose”). Perianth at stem apex . . . . . . . . Pseudolepicoleaceae 3. Underleaves smaller than lateral leaves or absent. Cuticle of leaf cells smooth or with small, rounded papillae (“verruculose”). Perianth on a short ventral branch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidoziaceae 4. Leaf segments deeply split into numerous ciliate parts. Leaf cells thin-walled or with uniformly thickened walls, trigones lacking. Underleaves smaller than lateral leaves. Plants whitish-green or brown . . . . . . . . . . . . . . . Trichocoleaceae 4. Leaf segments not split into numerous ciliate parts, leaves 4-lobed. Leaf cells with large trigones. Underleaves as large as lateral leaves. Plants gray-green or brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepicoleaceae 5. Leaves with (1–)2 large, sausage-shaped or globose slime papillae at the tips of the segments or on the rounded margins. Plants very small, growing on decaying wood in rainforest, less than 1 mm wide, whitish-green, the leaves almost longitudinally inserted on the stem . . . . . . . . . . . . Lepidoziaceae: Zoopsidella 5. Leaves without large slime papillae at the tips or on the margins. . . . . . . . . . 6 6. Leaves incubous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key A 6. Leaves succubous or transverse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Underleaves present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key B 7. Underleaves absent (or very small) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key C Key A. Leaves incubous 1. Underleaves present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaves absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2. Underleaves and leaves of similar size, bifid or trifid . . . . . . . . . Herbertaceae 2. Underleaves smaller than leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

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3. Leaves with 3 or more lobes or teeth. . . . . . . . . . . . . . . . . . . . . Lepidoziaceae 3. Leaves with 0–2 lobes or teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Vestigial lobule (consisting of a few cells) present at ventral base of leaf. Stems fragile, ventral stem surface only 2–4 cells wide, hyalodermis often present. Ventral branches absent . . . . . . . . . . . . . . . Lejeuneaceae with reduced lobules 4. Lobule lacking. Stems rigid, ventral stem surface more than 4 cells wide, hyalodermis absent. Ventral branches frequently present. . . . . . . . . . . . . . . . . . . . 5 5. Plants densely pinnate, growing upright on twigs near running water, with numerous clustered perianths on short lateral branches. Underleaves minute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeuneaceae: Colura irrorata 5. Plants forked, irregularly branched or unbranched, not growing on twigs near running water, perianths lacking or on ventral side of stem (not clustered on short lateral branches). Underleaves well developed . . . . . . . . . . . . . . . . . . . . 6 6. Stems forked. Flagelliform ventral branches present. Leaf apex bifid or trifid with sharp teeth (rarely entire). . . . . . . . . . . . . . . . . . Lepidoziaceae: Bazzania 6. Stems unbranched or irregularly branched. Flagelliform ventral branches lacking. Leaf apex entire or short-bifid . . . . . . . . . . . . . . . . . . . . . . Calypogeiaceae 7. Plants robust, more than 3 mm wide, turning reddish or purple. Leaves with a long insertion, strongly clasping the stem. Leaf margin with a whitish border. Perianth without keels, without beak. . . . . Pleuroziaceae: Pleurozia paradoxa 7. Plants less than 2 mm wide, green, not turning reddish or purple. Leaves with a very short insertion of a few cells, not strongly clasping the stem. Leaf margin with or without whitish border. Perianth usually keeled, with a beak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeuneaceae: Cololejeunea Key B. Leaves succubous or transverse, underleaves present 1. Leaf margins toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Leaves 4-lobed, underleaves as large as leaves . . . . . . . . . . . . . Lepicoleaceae 2. Leaves not 4-lobed, underleaves smaller than leaves . . . . . . . . . . . . . . . . . . . 3 3. Dorsal leaf base decurrent. Leaves asymmetrical, ventral margin arched, dorsal margin ± straight. Stems brown (or bluish), with thick-walled cortex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plagiochilaceae 3. Dorsal leaf base not decurrent. Leaves ± symmetrical. Thick-walled cortex lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaves ± transversely inserted, deeply concave. Branches originating from the ventral side of the stem . . . . . . . . . . . . . . . . . . . .   Balantiopsidaceae: Isotachis 4. Leaves ± longitudinally inserted, not deeply concave. Branches originating from the lateral and ventral side of the stem . . . . . . . . . . . . . . Lophocoleaceae 5. Leaves undivided or retuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Leaves distinctly divided into lobes [when in doubt try both leads] . . . . . . . 11 6. Leafy plants at the base arising from a leafless, stoloniform shoot . . . . . . . . . 7 6. Leafy plants not arising from a leafless, stoloniform shoot . . . . . . . . . . . . . . . 8 7. Leaves folded, concave and usually keeled, at least above, keel often winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidoziaceae: Micropterygium

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7. Leaves not folded, flat, keel absent . . . . . . . . . . . . . Cephaloziaceae: Alobiella 8. Ventral side of the stem with stolons. Leaf insertion reaching the dorsal midline of stem or not. Rhizoids scattered . . . . . . . . . . . . . . . . . . . . . . Cephaloziaceae 8. Ventral stolons lacking. Leaf insertion reaching the dorsal midline of stem. Rhizoids scattered or in bundles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Underleaves entire, undivided, oblong or narrowly lanceolate. Rhizoids scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Underleaves bifid or toothed, rarely entire. Rhizoids in bundles. . . Lophocoleaceae 10. Leaves alternate, ovate-orbicular, dorsal base short decurrent. Plants on moist soil and rock, often near sulfur springs or fumaroles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenostomataceae: Nardia succulenta 10. Leaves (sub)opposite, asymmetrically ovate-triangular, dorsal base longly decurrent. Plants on humus in montane forest and in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jamesoniellaceae: Syzygiella 11. Leaves divided into 3–5 lobes. Plants small, less than 2 mm wide, leafless stolons present or lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11. Leaves divided into 2(−3) lobes. Plants small or large . . . . . . . . . . . . . . . . . 15 12. Underleaves very different from leaves, much smaller and only 2-lobed. (Leaves strongly concave, 4–5-lobed, sharply piliferous). Plants on soil in superpáramo of the Sierra Nevada de Santa Marta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balantiopsidaceae: Ruizanthus venezuelanus 12. Underleaves similar to leaves (but sometimes smaller) . . . . . . . . . . . . . . . . . 13 13. Leaves succubous. (Leaves palmate, divided to ca. 1/2 into narrow lanceolate, conspicuously diverging lobes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudolepicoleaceae: Chaetocolea palmata 13. Leaves transverse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Plants pale green, in páramo bogs. Leaves shallowly divided (to 1/4 or less) into 3–4 triangular lobes. Stem base stoloniform. Leaf cells thin-walled, cuticle smooth or sligthly papillose. Perianth on a short ventral branch, 3-keeled . . . . . . . . . . . . . . . . . . Lepidoziaceae: Pseudocephalozia quadriloba 14. Plants brownish, on rotten wood in cloud forest. Leaves deeply divided (to 3/4) into 3–4 stiff, subulate lobes. Stem base not stoloniform. Leaf cells thick-walled, cuticle strongly striate-papillose. Perianth on a long shoot. . . . . . . . . . . . . . . . . . . . . . Pseudolepicoleaceae: Temnoma chaetophylla 15. Leaves in 3 ± equal rows, underleaves about as large as leaves . . . . . . . . . . 16 15. Leaves not in 3 equal rows, underleaves smaller than leaves . . . . . . . . . . . . 20 16. Leaves distinctly elongate, (1.5–)2–7× as long as wide. Leaves with a broad vitta of strongly elongate cells. Trigones very large . . . . . . . . Herbertaceae: Herbertus 16. Leaves shorter. Vitta absent. Trigones small or lacking. . . . . . . . . . . . . . . . . 17 17. Plants very small, on soil in páramo, less than 1  mm wide, creeping or ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Plants larger, growing upright . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18. Leaves tightly appressed to the stem and densely imbricate. Leaf margins smooth. Plants often white in color . . . . . . . . . . . . . . . . . . . . . . . Antheliaceae 18. Leaves ± spreading, not densely imbricate. Leaf margins coarsely papillose. Venezuela, not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnomitriaceae: Paramomitrion paradoxum R.M.Schust.

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19. Midleaf cells rectangular, 2–3× as long as wide. Plants reddish, purple-brown or black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balantiopsidaceae: Isotachis 19. Midleaf cells quadrate. Plants light green to brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Herbertaceae: Triandrophyllum 20. Leaves transverse, concave. Sporophyte in a fleshy perigynium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balantiopsidaceae: Isotachis 20. Leaves succubous. Sporophyte in a perianth. . . . . . . . . . . . . . . . . . . . . . . . . 21 21. Leaves and underleaves divided to the middle into 2–3 narrow lobes (some 3-lobed leaves always present). Rhizoids in bundles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidoziaceae: Paracromastigum 21. Leaves and underleaves 2-lobed or undivided, never 3-lobed, less deeply divided. Rhizoids scattered or in bundles . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Leaves asymmetrically ovate-triangular, apex obtuse and sometimes shallowly bilobed, dorsal base longly decurrent. Underleaves narrowly ligulate, 5–6× longer than wide . . . . . . . . . . . . . . . . . . . . . . Jamesoniellaceae: Syzygiella uleana 22. Plants different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23. Leaf insertion line not reaching dorsal midline of stem, dorsal side of stem “leaf-free.” Rhizoids scattered. . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziaceae 23. Leaf insertion line reaching dorsal midline. Rhizoids in bundles . . . . . . . . . 24 24. Plants often with some reddish coloration (especially in the rhioids). Branching purely ventral. Leaf cells narrowly rectangular, cuticle with elongate papillae (“striate-papillose”) . . . . . . . . . . . . . . . . . . . Balantiopsidaceae: Neesioscyphus 24. Plants without any reddish coloration. Branching lateral and ventral. Leaf cells shorter, cuticle smooth or with small rounded papillae. . . . . . Lophocoleaceae Key C. Leaves succubous or transverse, underleaves absent (or very small) 1. Plants densely pinnate, growing upright on twigs near running water, with numerous clustered perianths on short lateral branches, perianth beaked. Underleaves minute . . . . . . . . . . . . . . . . . . . . . Lejeuneaceae: Colura irrorata 1. Plants different. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants small, worm-like, whitish or pale brown to blackish, on soil or rock in páramo. Leaves densely imbricate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Plants not worm-like, small or large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 3. Stems with numerous pale green, linear to lanceolate paraphyllia between the leaves. Ventral stolons present or absent . . . . . . . . . . . . . . . . Stephaniellaceae 3. Stems without paraphyllia. Stolons absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaves undivided, opposite. Leaf cells conspicuously elongate near the ventral margin…. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arnelliaceae 4. Leaves bifid, alternate. Leaf cells not conspicuously elongate near the ventral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaves transverse. Leaf insertion line extending beyond the dorsal midline of the stem (merophytes “interlocking”) . . . . . . . . . . . . . . . . . . Gymnomitriaceae 5. Leaves succubous. Leaf insertion line not extending beyond the dorsal midline of the stem (merophytes not “interlocking”). . . . . . . . . . . . . . . . Scapaniaceae 6. Plants minute, less than 0.6 mm wide. Leaf cells usually small, ca. 8–20(−25) μm, without trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziellaceae

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6. Plants larger. Leaf cells with or without trigones . . . . . . . . . . . . . . . . . . . . . . 7 7. Leaves opposite, dorsal leaf bases united . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Leaves alternate, dorsal leaf bases not united . . . . . . . . . . . . . . . . . . . . . . . . 10 8. Plant strongly attached to soil, usually whitish, rarely green, growing above 2000  m. Leaf cells conspicuously elongate toward the ventral leaf margin. Trigones very small or lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . Arnelliaceae 8. Plant not strongly attached to soil, green, brown, or reddish, not whitish. Leaf cells not conspicuously elongate toward the ventral leaf margin. Trigones usually large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Plants with reddish or purple pigmentation. Perianth inflated over its whole length, with several plicae . . . . . . . . . . . . . . . . . . . . . . . . . . . Jamesoniellaceae 9. Plants lacking reddish or purple pigmentation. Perianth flattened toward the mouth, with 2 keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plagiochilaceae 10. Dorsal margin of leaves ± recurved, dorsal leaf base decurrent. Leaf surface convex along dorsal margin and concave toward ventral margin . . . . . . . . . 11 10. Dorsal margin of leaves plane or incurved, dorsal leaf base not or slightly decurrent. Leaf surface ± plane or concave. . . . . . . . . . . . . . . . . . . . . . . . . . 13 11. Plants with reddish or purple pigmentation. Sporophyte in a plicate perianth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jamesoniellaceae 11. Plants lacking reddish or purple pigmentation. Sporophyte in a 2-keeled perianth or a marsupium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Plants pale green to yellowish-green when fresh (grayish or yellow brown when dry). Leaf apex obliquely truncate-emarginate. Stems pale-colored. Oil bodies pale brown, numerous per cell. Sporophyte in a marsupium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbaceae: Acrobolbus 12. Plants green or brown. Leaf apex not obliquely truncate-emarginate. Stems brown, darker-colored than leaves. Oil bodies colorless or gray. Sporophyte in a flattened perianth. . . . . . . . . . . . . . . . . . . . . . Plagiochilaceae 13. Leaf base several layers of cells thick . . . . . . . . . . . . . . . . . . . . . . . . . Pelliidae 13. Leaf base only 1 layer of cells thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Leafy plants arising from a leafless, rhizome-like shoot . . . . . . . . . . . . . . . . 15 14. Leafy plants not arising from a leafless, stoloniform shoot . . . . . . . . . . . . . . 19 15. Leaves transverse. (Dorsal margin of leaves ± incurved. Stems rigid, with thickwalled cortex. Gametoecia at stem base on abbreviated branches) . . Adelanthaceae 15. Leaves succubous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Leaf cells small, less than 25  μm, without trigones. Plant less than 1  mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziellaceae 16. Leaf cells larger, with or without trigones. Plants usually wider than 1  mm (rarely less than 1 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. Leaf insertion line not reaching dorsal midline of stem, dorsal side of stem leaf-free. Plants frequently with upright flagelliform branches producing gemmae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziaceae 17. Leaf insertion line reaching dorsal midline of stem, dorsal side of stem not leaffree. Plants without upright flagelliform, gemmiparous branches . . . . . . . . . 18 18. Leaf cells papillose. Plants pale green to yellowish-green (grayish or yellowbrown when dry). Sporophyte in a marsupium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbaceae: Acrobolbus

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18. Leaf cells smooth. Plants green to reddish-brown to black. Sporophyte in a perianth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jamesoniellaceae 19. Leaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 19. Leaves 2(−3)-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 20. Leaf cells very large, ca. 60–80 μm long in midleaf, very thin-walled, without trigones. Leaf tip often with a large, sausage-shaped hyaline papilla. Stems with a hyalodermis. Leaves attached to the lateral side of the stem only, very flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidoziaceae: Zoopsidella 20. Leaf cells smaller, usually with trigones. Sausage-shaped hyaline papilla absent. Stems without hyalodermis. Leaves attached to the lateral and dorsal side of the stem, flat or concave. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21. Leaf cells densely striate-papillose, conspicuously elongate toward the ventral margin. Sporophyte in a marsupium. Plants whitish-green to yellowishgreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbaceae: Lethocolea 21. Leaf cells smooth or papillose, not conspicuously elongate toward the ventral margin. Sporophyte in a perianth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Ventral stolons present. Leaf insertion reaching the dorsal midline of stem or not. Gemmae sometimes present. Gynoecia on a short ventral branch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziaceae: Odontoschisma 22. Ventral stolons absent. Leaf insertion always reaching the dorsal midline of stem. Gemmae absent. Gynoecia terminal on the main stem or on a long branch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23. Plants blackish-green, on rock in rivers above 2500  m, often growing submerged. Leaves longer than wide. Midleaf cells 20–30  μm long, very thinwalled, trigones lacking or very small. Rhizoids few, colorless. Plants without perigynium (bracts not attached to perianth base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jungermanniaceae: Jungermannia ovatotrigona 23. Plants green to brown or reddish, on soil and rock in various habitats. Leaves longer than wide, rounded or wider than long. Midleaf cells (20–)30–60 μm long, trigones present or lacking. Rhzoids often numerous, colorless or reddish. Plants with a perigynium (inner female bracts attached to perianth base). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenostomataceae: Solenostoma 24. Stems with hyalodermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24. Stems without hyalodermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25. Underleaves completely absent. Rhizoids scattered (or absent). Oil bodies lacking. Perianth terminal on a long shoot. Leaf tips without hyaline papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziaceae 25. Underleaves present but very small, consisting of a transverse row of 2–4 large cells. Rhizoids in bundles at the bases of the underleaves (or absent). Oil bodies present. Perianth on a very short ventral branch. Leaf tips with or without hyaline papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidoziaceae: Zoopsidella 26. Leaves on sterile and male shoots with a very narrow base (leaves 2–3× wider in the middle than at the base), often caducous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbaceae: Acrobolbus cuneifolius 26. Leaves not with a very narrow base on sterile and male shoots, not caducous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

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27. Cuticle smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scapaniaceae 27. Cuticle papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28. Plants greenish-brown to reddish or purple. Gemmae present or absent. Oil bodies colorless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scapaniaceae 28. Plants glossy green to light brown (never reddish or purple). Gemmae absent. Oil bodies brown . . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbaceae: Acrobolbus

Acrobolbaceae Plants usually glistening yellowish-green to pale bluish-green, rarely brown, creeping or upright, sometimes with a stoloniferous base. Stems with or without thickwalled cortex. Branches intercalary (lateral or ventral); stolons present or absent. Leaves succubous, undivided or 2-lobed (rarely 3–4-lobed), margins sometimes with long rhizoids. Cells usually with a finely papillose cuticle; oil bodies brown, finely granular. Underleaves ± lacking. Rhizoids scattered. Sporophytes in a fleshy, pendent marsupium; perianth lacking. Setae of numerous cell rows. Capsules with acute tip, wall thick, 4–10-layered. Vegetative reproduction by caducous leaves or lacking. A family of six genera (in 4 subfamilies), mostly in the Southern Hemisphere. The principal features of Acrobolbaceae are (1) plants often yellowish-green; (2) leaves succubous, undivided or bifid, cuticle usually papillose; (3) underleaves absent; (4) oil bodies brown; (5) sporophyte in a pendent marsupium, perianth absent; and (6) capsule tip acute. 1. Leaves bifid to 1/4–1/2. Leaf tips acute to acuminate (rarely obtuse) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbus 1. Leaves undivided or shallowly retuse. Leaf tips rounded to acute . . . . . . . . . 2 2. Leaves symmetrically ovate to lingulate (resembling Solenostoma), apex rounded, margins entire, without long hairs. Leaf cells conspicuously elongate toward the ventral leaf margin. Oil bodies 1–2 per cell. Plants creeping, stolons present or lacking. Stems without brownish cortex. . . . . . . . . . . . . Lethocolea 2. Leaves asymmetrically ovate-oblong (resembling Plagiochila), apex obliquely truncate to retuse, margins toothed or entire, sometimes with long hairs. Leaf cells not conspicuously elongate toward the ventral leaf margin. Oil bodies 10–40 per cell. Plants ascending from a stoloniform base. Stems with brownish cortex (former genus Tylimanthus) . . . . . . . . . . . . . . . . . . . . . . . . . Acrobolbus Acrobolbus Nees Marsupidium Mitt., Tylimanthus Mitt. Plants glossy green, pale green, yellowish-green to light brown, creeping or ascending, with or without stoloniform base. Stems pale-colored, with or without thick-walled cortex in (1–)2–4 cell layers. Leaves succubous, symmetrically or asymmetrically ovate-orbicular to oblong or quadrate, bifid to 1/4–1/2 or obliquely truncate to emarginate to short bifid, tips rounded to acute to acuminate, margins entire or toothed, sometimes with long rhizoids. Cells with trigones, cuticle

Acrobolbaceae

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papillose, rarely smooth; oil bodies 5–40 per cell, granular, brown. Underleaves lacking or rudimentary. Dioicous. Male plants frequently smaller than female plants. Antheridia 1–10 per bract. Archegonia terminal on long shoots or on a short basal branch, surrounded by large bracts which are often strongly spreading, forming a flower-like involucre, margins dentate-ciliate or subentire; female bracteoles lacking. Marsupium short or long. Vegetative reproduction sometimes by caducous leaves. A genus of about 30 species, most of them in the Southern Hemisphere, 4  in Colombia and Ecuador. Acrobolbus is defined here in broad sense following Briscoe et al. (2015, 2017) with Tylimanthus and Marsupidium included as synonyms. The main features of Acrobolbus are the pale green to yellowish-green to pale brown plants with a pale-colored stem, truncate-emarginate to bifid leaves, papillose cuticle, absence of underleaves, and presence of a fleshy marsupium at the stem tip (on a short basal branch in the former genus Marsupidium) instead of a perianth. Literature: Schuster (2001a), Burghardt and Gradstein (2008), Briscoe et  al. (2015, 2017) 1. Leaves undivided to obliquely truncate to retuse, asymmetrical (resembling Plagiochila). Leaf tips rounded to acute . . . . . . . . . . . . . . . . . . . . . . . A. laxus 1. Leaves bifid to 1/4–1/2, symmetrical. Leaf tips acute to acuminate (rarely obtuse) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Trigones large and swollen. Leaf tips ending in a single row of 5–8 cells. Leaf margins with (0–)1–5 teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. setaceus 2. Trigones small, not swollen. Leaf tips ending in a single row of 1–4 cells. Leaf margins entire (except on bracts). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaves on sterile and male shoots strongly narrowed to the base, leaf base ca. 2–3× narrower than leaf middle. Leaves frequently caducous. Female bracts coarsely toothed, with long laciniate or ciliate teeth. Plants growing below 3000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cuneifolius 3. Leaves on sterile and male shoots not or little narrowed to the base. Leaves persistent. Female bracts toothed or subentire. Plants growing above 3000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. wilsonii 1. Acrobolbus cuneifolius (Steph.) Briscoe Fig. 11A–D Tylimanthus cuneifolius Steph. Type: Venezuela, Valencia, Fendler s.n., c. gyn. (lectotype, designated here, G-00120031!; isolectotypes G-00283284!, S-B5768!) Acrobolbus caducifolius R.M.Schust., syn. nov. Type: Colombia, Boyacá, Villa de Leyva, Santuario Iguaque, on tree trunks, 10 Nov 1988, Aguirre 7321, ster. (holotype COL-290566!; isotype F-CO172194!) On bark in upper montane forest, ca. 2000–2750 m. Colombia: Boy (type of A. caducifolius), San (Virolín, Hässel de Menéndez 5898, COL), Tol (Santa Isabel, Aguirre & Gradstein 1486A, c. gyn., COL). Ecuador: ZC (Reserva Biologica San Francisco, Gradstein 10130, c. andr., GOET). General distribution: Costa Rica, northern Andes. Acrobolbus cuneifolius is a rare northern Andean species known from three localities in Colombia and one in Ecuador. The species is distinguished by (1) leaves ovate to ovate-oblong, bifid to 1/3–2/5, leaf margins fully entire, leaf lobes

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Fig. 11  (A–D) Acrobolbus cuneifolius. (A) Habit with gynoecium and sporophyte, dorsal view. (B) Male shoot, 26×. (C) Caducous-leaved shoot, 32×. (D) Midleaf cells, showing papillose cuticle. (E and F) Acrobolbus wilsonii, habit. (G and H) Lethocolea glossophylla. (G) Habit, dorsal view. (H) Midleaf cells, showing papillose cuticle. (I–K) Acrobolbus laxus. (I) Habit, dorsal view. (J) Leaf, 15×. (K) Cross section of stem. (B, C redrawn from Schuster 2001a, as A. caducifolia; E, F redrawn from Paton 1999; I–K redrawn from Burghardt and Gradstein 2008; rest from Gradstein et al. 2001)

Acrobolbaceae

133

triangular, with an obtuse to acute to acuminate tips ending in a row of 1–3(−4) cells; (2) leaves of sterile and male shoots frequently caducous and conspicuously cuneate with a very narrow, cuneate base (about 2–3× narrower than the leaf middle), leaves persistent on female shoots and with a broader base (scarcely narrower than the leaf middle); (3) leaf cells subisodiametrical, ca. 25–30 μm wide in midleaf, trigones small, cuticle finely papillose or smooth; and (4) female bracts large, flower-like spreading, margins coarsely toothed, with long lacinia and cilia. Acrobolbus cuneifolius is readily separated from A. wilsonii by the cuneate, caducous leaves of male and sterile shoots and by the coarsely ciliate-laciniate female bracts. The leaf lobes are usually obtuse to acute to shortly acuminate, but in the female material from Tolima, some leaves end in a long-ligulate tip crowned by a short single row of 2–3 cells. Acrobolbus caducifolius is a sterile phenotype of A. cuneifolius with strongly caducous leaves. Acrobolbus antillanus R.M.Schust., described from the West Indies (Dominica), resembles A. cuneifolius in having cuneate leaves on sterile shoots, (sub)entire leaf margins, and small trigones but differs in the longer leaf tips, terminating in a single row of (4–)5–9 cells. In its long leaf tips, A. antillanus is similar to A. setaceus (see there). Plants from Costa Rica identified as A. antillanus (“Costa Rican phase”; Schuster 2001a, p. 146) belong to A. cuneifolius. The species is rather common on trees and shrubs along the Panamanian Highway in the Cerro de la Muerte, Costa Rica, near Reserva 3 de Julio (Gradstein 9687, GOET). 2. Acrobolbus laxus (Lehm. & Lindenb.) Briscoe Fig. 11I–K Tylimanthus laxus (Lehm. & Lindenb.) Spruce Tylimanthus approximatus (Lindenb.) Besch. Tylimanthus azoricus Grolle & Perss. (Acrobolbus azoricus [Grolle & Perss.] Briscoe) Tylimanthus madeirensis Grolle & Perss. (Acrobolbus madeirensis [Grolle & Perss.] Briscoe) Tylimanthus ruwenzorensis S.W.Arnell (Acrobolbus ruwenzorensis [S.W.Arnell] Briscoe) Tylimanthus setaceociliatus Steph. On soil, humus, damp rock, and tree trunks in montane forest areas and páramo, (500–)1600–4000  m. Colombia: Boy, Cas, Cho, Cun, Met, Nar, Qui, Ris, Tol. Ecuador: Az, Ca, Co, Ga, Lo, Na, Pi, ZC.  General distribution: tropical America, East Africa, Macaronesia. Acrobolbus laxus is a widespread Afro-American species common in the Andes and recognized by the pale green to glossy yellowish-green plants ascending from creeping stoloniform shoots and leaves with an obliquely truncate to shallowly bifid, occasionally rounded, apex. When bifid, the leaf lobes are usually unequal in size with the dorsal lobe being shorter. Leaf shape varies from ovate-oblong to suborbicular, the leaf margins may be entire or irregularly and finely toothed, the leaf tips may be acute or obtuse, the trigones are small or large and nodulose, and the cuticle is usually densely papillose. The frequent presence of long rhizoids on leaf margins is a further characteristic of the species.

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Burghardt and Gradstein (2008) studied the morphological variation of A. laxus based on a broad sampling and found that plants from Macaronesia and East Africa described as Tylimanthus azoricus, T. madeirensis, and T. ruwenzorensis were part of the morphological variation in T. laxus. In a recent molecular study, Briscoe et al. (2017) resolved T. azoricus, T. madeirensis, and T. ruwenzorensis as separate lineages within a broad A. laxus clade, and resurrected them as species. However, no distinctive morphological characters were found to separate the three species from A. laxus, which was underrepresented in the analysis. Since concordant morphological differentiation was lacking, the three species should be interpreted as genetic races, representing a case of cryptic speciation in A. laxus. 3. Acrobolbus setaceus (Steph.) Gradst., comb. nov. Tylimanthus setaceus Steph., Bull. Herb. Boissier sér. 2, 5(12): 1141. 1905 [Sp. Hepat. 3: 13]. Type: Colombia (“Nova Granada”), Antioquía, páramo de Sonsón, on Plagiochila cucullifolia, ca. 3050  m (“10.000  ft.”), 1872, Wallis s.n. (G-00120194!) On bark in upper montane forest and páramo, creeping over other epiphytic bryophytes, ca. 3000–4000 m. Colombia: Ant (type). Ecuador: Co (Burghardt and Gradstein 2008). General distribution: northern Andes (Colombia, Ecuador) Acrobolbus setaceus is a very rare, fragmentarily known northern Andean species; the type material consists of one sterile shoot. The species is characterized as follows: (1) plants pale brown in herbarium, ca. 2.5 mm wide; (2) leaves subhorizontally succubous, plane, ovate-subrectangular, ca. 1.25 × 0.8 mm, bifid to 1/4–1/3 with a wide sinus and narrow-triangular lobes ending in a row of 5–8 somewhat elongate cells; (3) dorsal and ventral leaf margins with (0–)1–5 teeth, the teeth becoming more numerous toward stem apex (female?), 1–3 cells long, made up of elongate cells with an evenly thickened and finely papillose outer wall, sometimes teeth lacking; (4) leaf cells subisodiametric to somewhat elongate, 25–35 × 30–60 μm in midleaf, with large swollen trigones and a finely papillose cuticle; and (5) underleaves absent. Acrobolbus setaceus is readily separated from A. cuneifolius and A. wilsonii by the leaf margins with (0–)1–5 teeth, the longly piliferous leaf tips ending in a row of ca. 5–8 cells (many leaf tips are broken), and the large, swollen trigones. In its longly piliferous leaf tips, A. setaceus is similar to A. antillanus, but in the latter the leaf margins are usually entire (occasionally with a tooth) and the trigones are small, not bulging. Acrobolbus setaceus and A. antillanus are known from only few collections, and little is known about their morphological variation. Further material is needed to determine the status of these two species. 4. Acrobolbus wilsonii Nees Fig. 11E, F Acrobolbus mittenii Steph., syn. nov. Type: Ecuador, Pichincha, near Quito, Jameson s.n. (G-00069022!) Acrobolbus diversifolius R.M.Schust., syn. nov. Type: Venezuela, Mérida (“Tachira”), Sierra de Santo Domingo, in cloud forest on trunk of Polylepis sericea, 3700 m, 1976, Schuster 76-906c (F n.v.); the material cannot be found (L. Briscoe, pers. comm.)

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Acrobolbus wilsonii var. andinus Spruce, syn. nov. Type: Ecuador, Mt. Tungarahua, Spruce s.n. (MANCH?); the material cannot be found (L. Loughtman, pers. comm.) Lophozia debilis J.B.Jack & Steph. (fide J. Váňa, in Gradstein and Uribe 2016a) On bark and wet rock in upper montane cloud forest, Polylepis forest and páramo, ca. 3000–4500 m. Colombia: Boy (Sierra Nevada del Cocuy, Bischler 2827, PC), unknown loc. (Stephani 1898–1924 as Lophozia debilis). Ecuador: Az (páramo de Patococho, Gradstein et  al. 12654, QCA, c. gyn.), Co (páramo de Llanganates, Burghardt 6689, QCA, PC), Na (páramo de Oyacachi, on Polylepis pauta, Benitez & León-Yánez 1300, HUTPL), Pi (old road Quito to Santo Domingo, W of San Juan, Gradstein et al. 6747, GOET, U), Tu (Spruce 1885 as A. wilsonii var. andinus). General distribution: western Europe, Mexico, northern Andes. Acrobolbus wilsonii is disjunct to western Europe and the neotropics, where it is rare and mostly restricted to the northern Andes. The species is separated from A. cuneifolius by the leaves with broad leaf bases on all shoots (never narrow-based, cuneate, as in the latter species), the absence of caducous leaves, and the less coarsely toothed, to subentire, female bracts. Moreover, A. wilsonii occurs at higher elevations, above 3000  m (the records of A. cuneifolius are mostly from below 3000 m). Plants from the superpáramo of the Sierra Nevada del Cocuy stand out by whitish-colored walls of leaf cells. The occurrence of A. wilsonii in the northern Andes has been widely overlooked although it was already collected there in the nineteenth century by William Jameson (type of A. mittenii) and by Richard Spruce. Spruce described the Andean plants as a separate variety (“var. andinus”) characterized by leaves slightly more deeply bifid and leaf apices less finely pointed than in European plants. However, these differences fit the variation in the European A. wilsonii as described by Paton (1999) and others. Acrobolbus diversifolius described from the Venezuelan Andes (type lost) is identical to A. wilsonii judging from the original description and illustration (Schuster 2001a) and a further synonym. Lethocolea Mitt. Neoprasanthus S.Winkl. Plants yellowish-green to bluish-green, creeping, often in dense mats, frequently with a few ventral stolons. Stems rather fragile, lacking a thick-walled cortex. Leaves succubous, ovate to lingulate, with broadly rounded apex and entire margins. Cells conspicuously elongate toward the ventral margin of the leaf, especially in the lower half of the leaf, with distinct trigones, cuticle densely papillose; oil bodies large, brown, 1–2 per cell. Underleaves lacking. Marsupium long, narrowcylindrical, developing at the tip of the stem and penetrating deeply into the substrate. A southern-temperate genus (ca. 5–6 spp.), one species in Colombia and Ecuador. Lethocolea is morphologically very different from Acrobolbus (see the key) and is traditionally placed in a separate subfamily, Acrobolbaceae subfam. Lethocoleaceae (S.W.Arnell) Grolle. This classification is supported by molecular evidence (Briscoe

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et al. 2015). Lethocolea may be confused with Solenostoma (especially S. amplexifolia) but differs from the latter genus in its bluish-green to yellowish-green color, densely striate-papillose cuticle, large brown oil bodies, and the presence of a marsupium. Lethocolea also resembles Gongylanthus, but the leaves in the latter genus are opposite, stolons are absent, and the cuticle is smooth in neotropical species. Literature: Gradstein et al. (2001) 1. Lethocolea glossophylla (Spruce) Grolle Symphyomitra glossophylla Spruce Calypogeia euthemona Spruce Jamesoniella boliviana Steph. Neoprasanthus granatensis S.Winkl.

Fig. 11G, H

On moist soil in upper montane forest and páramo, (1800–)3000–4000  m. Colombia: Ant, Boy, Cal, Cau, Cho, Cun, Mag, Qui, Ris, San, Tol. Ecuador: Ca, Co, Pi, Tu, ZC. General distribution: Mexico to Bolivia, southeastern Brazil, Dominican Republic. Lethocolea glossophylla is a characteristic high-montane neotropical species that is distinguished by the yellowish-green to glaucous green (rarely reddish), creeping plants with rather flat, succubous, ovate to tongue-shaped leaves, large leaf cells (ca. 40–70 × 30–45 μm in midleaf) becoming elongate toward the ventral leaf margin (especially in the lower half of the leaf), large brown oil bodies (1–2 per cell), a densely papillose cuticle, and lack of underleaves. The species resembles Solenostoma, especially S. amplexifolia (= Jungermannia linguifolia), but differs from the latter species in the characteristic yellowish-green to glaucous plant color, brown oil bodies, and densely striate-papillose cuticle. Fertile material is immediately separated by the long, pendent marsupium. Lethocolea glossophylla is characteristic of páramo and usually occurs above 3000  m, rarely at lower elevation. Records from below 1800 m are probably erroneous and should be verified. Excluded Record Lethocolea repens S.Winkl.  – Colombia: Mag (Sierra Nevada de Santa Marta; Winkler 1969). The species is probably a synonym of Gongylanthus liebmannianus (see there).

Adelanthaceae Plants dark green to brown, growing upright from a stoloniferous base. Stems rigid, usually brown, with a thick-walled cortex. Branches exclusively ventral-intercalary. Leaves transversely inserted with conspicuously decurrent dorsal bases, undivided or shallowly bilobed with acute tips, margins entire or toothed, sometimes bordered; dorsal leaf margin plane or incurved (toward the stem). Cells ± quadrate, 12–35 μm, uniformly thick-walled or with trigones, cells along the leaf margin often

Adelanthaceae

137

thicker-walled and forming a border, toward the leaf base sometimes elongate and forming a weak vitta, cuticle smooth; oil bodies finely granular. Underleaves lacking. Rhizoids scarce, usually restricted to the stolons. Gametoecia on very short branches arising from the leafless stem bases or from the stolons. Sporophytes in a perianth or a fleshy calyptra. Setae of numerous cells (cross section). Capsules elongate, wall 4–6-layered. Vegetative reproduction rare, by caducous leaves or gemmae from leaf margins. Three genera, mostly in tropical mountains and the Southern Hemisphere. Adelanthaceae are sister to Jamesoniellaceae, and recent studies indicate the two groups should be treated as subfamilies of a single family, Adelanthaceae (Feldberg et al. 2010a, b; Söderström et al. 2016). Adelanthaceae differ from Jamesoniellaceae by transverse leaves with frequently upcurved to inflexed margins (leaves succubous and margins plane or recurved in Jamesoniellaceae) and by basal gametoecia (terminal in Jamesoniellaceae). Further characteristics of Adelanthaceae are the leafy stems arising from a stoloniform base and the absence of terminal branching. 1. Cells in the upper part of the leaf ± evenly thick-walled, without distinct trigones. Leaf base with a vitta-like area. Stem cortex well developed, brown, of 2–3 cell layers (cross section) . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelanthus 1. All leaf cells with distinct trigones, cells never evenly thick-walled. Vitta lacking. Stem cortex poorly developed, pale-colored, of 1 layer of cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudomarsupidium Adelanthus Mitt. Calyptrocolea R.M.Schust. Plants dark green to brown, arising from a long stoloniferous base. Stems with a brown cortex of thick-walled cells in 2–3 layers. Branches ventral-intercalary only and often stoloniform. Leaves spreading or secund, apex rounded to short bifid, margins bordered or unbordered, entire to strongly toothed, dorsal margin rolled inward or almost plane. Leaf cells ± uniformly thick-walled, without distinct trigones, quadrate, ± elongate toward leaf base, forming a vitta-like area. Sporophytes in a perianth or a shoot calyptra. Gemmae present (but mostly absent in the species from Colombia and Ecuador). A genus widespread in the Southern Hemispere and the tropics (ca. 10 spp.), four species in Colombia and Ecuador. Adelanthus may be confused with Plagiochila but differs in the ± incurved dorsal leaf margin (recurved in Plagiochila), the transverse leaf insertion (oblique in Plagiochila), and ventral-intercalary branching (lateral-intercalary in Plagiochila). The occurrence of gametoecia on very short, leafless branches near the stem bases is another characteristic feature of Adelanthus. In Plagiochila, gametoecia arise on long leafy shoots. As indicated by Feldberg et al. (2010b), based on the results of molecular analysis, some species currently recognized in Adelanthus may be synonyms. Literature: Grolle (1972), Schuster (2002 as Calyptrocolea), Feldberg et al. (2010b)

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Keys and Descriptions

1. Leaf margin clearly toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf margins entire or with 1–2 small teeth . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Leaves conspicuously bordered by yellowish to orange-colored thick-walled cells in 2–3 rows. Leaf margins ± plane . . . . . . . . . . . . . . A. aureomarginatus 2. Leaves not bordered by yellowish to orange-colored thick-walled cells. Dorsal leaf margin rolled inward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Teeth on leaf margin 2–5(−10) cells long, teeth strongly variable in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. carabayensis 3. Teeth smaller, 1–2 cells long, not strongly variable in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lindenbergianus 4. Dorsal leaf margin distinctly incurved in the lower half. Leaves rounded or slightly longer than wide. Leaf apex undivided . . . . . . . . . . A. lindenbergianus 4. Dorsal leaf margin not distinctly incurved, mostly plane. Leaf apex 1.5–2× longer than wide. Leaf apex truncate to emarginate to 2-toothed . . . . . . A. pittieri 1. Adelanthus aureomarginatus R.M.Schust.  Calyptrocolea aureomarginata (R.M.Schust.) R.M.Schust.

Fig. 12D, E

On moist, peaty soil in páramo, ca. 3500  m. Colombia: Cun (El Tablazo, R.  M. Schuster, material not seen). General distribution: northern Andes (Colombia, Venezuela). Adelanthus aureomarginatus is a rare species of moist páramo in the northern Andes and is only known from the type from Venezuela and a further collection from El Tablazo near Bogotá, both gathered by R. M. Schuster. The species is readily distinguished by the rounded leaves with sharply toothed margins all around and with a prominent, yellowish to orange-colored border of strongly thick-walled cells in 2–3 rows (Schuster 2002). The leaves are not secund and the margins are almost plane. Toothed leaf margins are also present in A. lindenbergianus and A. carabayensis, but in the latter species, the leaves are secund, without border, and the dorsal leaf margin is rolled inward. A leaf border is also seen in Pseudomarsupidium aureocinctum, but in the latter species, the border is only 1 cell row wide and teeth on leaf margins are few and restricted to the apex and the upper half of the ventral margin. 2. Adelanthus carabayensis (Mont.) Grolle Calyptrocolea carabayensis (Mont.) R.M.Schust. Adelanthus decurvus Mitt. On bark and rotten wood in montane rainforest, 1000–3000 m. Colombia: Cho, Tol. Ecuador: ZC (Schäfer-Verwimp et al. 2013a). General distribution: scattered in tropical America. Adelanthus carabayensis is rather similar to A. lindenbergianus, but the teeth on the ventral leaf margin are longer in A. carabayensis, 2–10 cells long, and quite variable in length. The species is much rarer than A. lindenbergianus and seems to be restricted to montane forest areas, while A. lindenbergianus occurs mainly in páramo. A preliminary molecular study, based on limited sampling, however,

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indicated that the two species might be conspecific (Feldberg et  al. 2010b). This needs further work. 3. Adelanthus lindenbergianus (Lehm.) Mitt.  Calyptrocolea lindenbergiana (Lehm.) R.M.Schust. Plagiochila magellanica Lindenb.

Fig. 12A–C

On soil and rock on road banks and in páramo, also epiphytic on trees in upper monate cloud forest, 1800–4200 m. Colombia: Ara, Boy, Cas, Cun, Hui, Mag, Met, Nar, Ris, San, Tol. Ecuador: Lo, Pi, Tu, ZC. General distribution: temperate regions of the Southern Hemisphere, tropical Andes, western Europe. Adelanthus lindenbergianus is a common páramo species that is characterized by brownish plants with laterally appressed and somewhat secund leaves with an incurved, entire dorsal margin and plane, toothed apical and ventral margins with small, 1–2-celled teeth. A weakly developed vitta is present at the leaf base. Adelanthus lindenbergianus may be confused with Plagiochila bifaria, but in the latter species, the dorsal leaf margin is curved outward, not inward, and the teeth in P. bifaria are much longer; both species have a short vitta-like area. Adelanthus lindenbergianus may also be confused with A. carabayensis; for differences see under the latter species. An almost entire-leaved phenotype of A. lindenbergianus, with a few rudimentary teeth present on upper stem leaves and about 90 percent of the leaves fully entire, was found in the páramo of Saraguro and Yacuambi in southern Ecuador (Nov 2018, Benitez et al. 1306, HUTPL). These plants are very similar to Adelanthus integerrimus Grolle from Patagonia; in the latter species, all leaves are entire (Grolle 1972). Possibly, A. integerrimus is just a variety or synonym of A. lindenbergianus; the results of a preliminary molecular study based on limited sampling support this (Feldberg et al. 2010b). 4. Adelanthus pittieri (Steph.) Grolle Calyptrocolea pittieri (Steph.) R.M.Schust. Adelanthus corticola S.Winkl. [“corticolus”]

Fig. 12F, G

On bark and rotten wood in upper montane cloud forest, 2500–3700  m. Colombia: Boy, Cau, Cun, Hui, Mag, Met, Nar, NSa, Ris, San, Tol. Ecuador: Lo, Na. General distribution: Central America, tropical Andes, Jamaica. Adelanthus pittieri is rather common in the Andes and recognized by the somewhat elongate (1.4–2× longer than wide), slightly concave, ventrad leaves with entire and almost plane margins, small thick-walled. Occasionally, the leaves are caducous (seen in Ecuador, Napo, Guagra Urcu, Holm-Nielsen et al. 27484, AAU, PC). The leaf apex is usually emarginate, sometimes entire or 2-toothed with 2 small, unequal teeth, and the leaf cells are clearly elongate in the lower half of the leaf, forming a vitta-like area. The species usually grows on shaded tree trunks in cloud forests and may form large, dark green to brownish mats.

Fig. 12 (A–C) Adelanthus lindenbergianus. (A) Habit, lateral view, 16×. (B) Leaf, 20×. (C) Leaf margin, 250×. (D and E) Adelanthus aureomarginatus. (D) Leaf, 25×. (E) Apical portion of leaf showing border and toothed margin. (F and G) Adelanthus pittieri. (F) Habit, lateral view. (G) Leaves, 25×. (H–K) Pseudomarsupidium decipiens. (H) Habit, dorsal view. (I and J) Leaves, 15×. (K) Gynoecium. (L–N) Pseudomarsupidium aureocinctum. (L) Leaf, 15×. (M) Leaf margin, 100×. (N) Cross section of portion of leaf lamina, showing thickened border, 170×. (O and P) Anthelia juratzkana. (O) Habit with androecia, gynoecium, and sporophyte, lateral view. (P) Leaf lobe. (Q–S) Gongylanthus liebmannianus. (Q) Habit, lateral view. (R) Opposite, connate leaves (flattened). (S) Cells near the ventral leaf margin. (T) Gongylanthus limbatus, bordered leaf margin, 100×. (A–C redrawn from Grolle 1972; D, E, I, J, L–N redrawn from Schuster 2002; T redrawn from Herzog 1942; rest from Gradstein et al. 2001)

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Pseudomarsupidium Herzog Adelanthus p.p. Plants dark green to brown, ascending from a stoloniferous base. Stems with a weak, pale-colored, 1-layered cortex of thick-walled cells (= epidermis). Branching ventral-intercalary only. Leaves spreading, apex rounded to truncate, margins bordered or unbordered, plane or slightly incurved, entire or with a few teeth near the apex, dorsal leaf base longly decurrent, ventral base not decurrent. Leaf cells with distinct trigones, not elongate toward the leaf base, vitta lacking. Sporophytes in a perianth or a shoot calyptra. Gemmae lacking. A genus of four species in southern South America, tropical America, tropical Africa, Borneo, and western Europe, two species in Colombia and Ecuador. Pseudomarsupidium was previously considered a subgenus of Adelanthus (e.g., Grolle 1972), but molecular studies (Feldberg et al. 2010b) indicate that the group merits recognition as a separate genus. For differences with Adelanthus, see the key. Literature: Grolle (1972 as Adelanthus), Schuster (2002) 1. Leaves with a conspicuous, yellowish-brown border of thick-walled cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aureocinctum 1. Leaves without yellowish border of thick-walled cells . . . . . . . . . P. decipiens 1. Pseudomarsupidium aureocinctum (R.M.Schust.) J.J.Engel  Fig. 12L–N Adelanthus aureocinctus (R.M.Schust.) R.M.Schust. (Adelanthus decipiens [Hook.] Mitt. subsp. aureocinctus R.M.Schust.) On bark, rock, and moist soil in montane forest, scrub, and páramo, 2200–4050 m. Ecuador: Ca (Gradstein 3428, GOET, U), Lo (Schäfer-Verwimp et al. 2013a), Pi, ZC (Gradstein 10124, GOET). General distribution: northern Andes, southeastern Brazil. Pseudomarsupidium aureocinctum is similar to P. decipiens except for the presence of a yellowish-brown border of thick-walled cells (in 1–2 rows) along the leaf margins. Like in P. decipiens, leaf dentation varies and plants from páramo may have fully entire leaves. Since the two species are sympatric and differ in one morphological feature (presence/absence of a leaf border), P. aureocinctum may be just a variety of P. decipiens. 2. Pseudomarsupidium decipiens (Hook.) Grolle Adelanthus decipiens (Hook.) Mitt. Adelanthus crossii Spruce Tylimanthus bifidus Steph.

Fig. 12H–K

On old, ± rotten trunks and on rock in montane forests and páramo, in rather open situations, (500–)1200–4300 m; on the Galápagos Islands on bark and rock in moist evergreen woodland at 500–700  m. Colombia: Boy, Cal (4300  m, Cleef 2541, COL), Cun, Met, Qui, Ris, Tol. Ecuador: Az, Ca, Ga, Lo, Na, Pa, Tu, ZC.  General distribution: widely distributed in the mountains of tropical America and Africa, also in western Europe.

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Keys and Descriptions

Pseudomarsupidium decipiens is a common and widespread species that is distinguished by the rounded leaves with a rounded to truncate apex and usual presence of 2 small, widely spaced teeth at the apex. Occasionally a few additional small teeth may be present along the upper part of the ventral margin. Plants from páramo are often robust and lack teeth on the leaves; such entire-leaved plants were described as Adelanthus crossii.

Antheliaceae A family with a single genus. Anthelia (Dumort.) Dumort. Plants very small (usually less than 1 cm long), dioicous or paroicous, pale green to brown, becoming white due to the secretion of a whitish crystalline substance, creeping or erect, simple or forked. Stems fragile, with relatively large, thin-walled epidermis cells (cross section). Branches terminal, Frullania-type, stem base stoloniform. Leaves transverse, deeply bifid, tips acute (or obtuse in male bracts), margins entire. Cells uniformly thick-walled, cuticle smooth; oil bodies lacking. Underleaves ± similar to the leaves. Rhizoids scattered. Male bracts strongly convex, with 1 antheridium, male bracteoles flat, without antheridium. Perianths large, conical, deeply plicate. Setae of numerous cells (cross section). Capsules spherical. Vegetative reproduction unknown. A widespread, bipolar genus (2 spp.), occurring scattered in the tropics at high elevation, above 3000  m. For main characteristics of the genus, see under A. juratzkana. Literature: Schuster (2000a), Gradstein et al. (2001) 1. Anthelia juratzkana (Limpr.) Trevis. Fig. 12O, P Anthelia julacea (L.) Dumort. subsp. juratzkana (Limpr.) Meylan On bare soil and rock in páramo, 3100–4300  m. Ecuador: Lo (páramo of Saraguro, abundant, 3100–3200 m, Nov 2018, Benitez & Gradstein s.n., HUTPL), Na (Benitez et al. 2012); not known from Colombia but expected there. General distribution: bipolar, widespread in arctic and alpine regions of the Northern Hemisphere and furthermore scattered in high tropical mountains, New Zealand, southern South America, and the Antarctic Peninsula (Schuster 2000a; BednarekOchyra et al. 2000). In tropical America known from Mexico and the high Andes of Venezuela, Ecuador, and Bolivia. Characteristics of Anthelia juratzkana are the minute, reddish-brown or white, creeping or ascending plants with 3 equal rows of imbricate, transverse, bifid leaves and underleaves with acute tips and with uniformly thick-walled cells, without trigones and without oil bodies. The cuticle of the leaf cells is smooth or finely striate-papillose. Anthelia juratzkana is paroicous, and the antheridia are produced

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in the axils of swollen, densely imbricate bracts on elongate shoots terminating in a gynoecium. The young fertile shoots are strongly swollen and bud-like. The species is close to A. julacea (L.) Dumort. and is sometimes considered a subspecies of the latter, but A. julacea is dioicous and has smaller spores (Paton 1999). Anthelia julacea is restricted to temperate regions of the Northern Hemisphere and is not known from the tropics. The new record of A. juratzkana from the páramo de Saraguro in southern Ecuador, where the species is locally abundant, represents the lowermost locality of the species in tropical America. The area is part of the “Amotape-Huancabamba depression” of northern Peru and southern Ecuador, a rather low-lying area linking the Northern and Central Andes and characterized by high species diversity and endemism (e.g., Weigend 2002; Richter et  al. 2009; Atwood et  al. 2018). Many high-Andean species descend here to relatively low elevation. The occurrence of Anthelia juratzkana at 3100–3200 m in the páramo of Saraguro reflects this.

Arnelliaceae (Southbyaceae) A small family of three genera worldwide, in Colombia and Ecuador only the genus Gongylanthus. The main characters of Arnelliaceae are (1) small, creeping, terrestrial plants with densely imbricate, opposite leaves; (2) leaf cells very large, transparent, with little or no chlorophyll, frequently elongate along the ventral leaf margin; (3) underleaves present (Arnellia) or absent (Gongylanthus, Southbya); (4) rhizoids scattered; and (5) sporophyte elongate, in a marsupium (Gongylanthus) or perianth (Southbya, Arnellia). Váňa et al. (2012a) placed Gongylanthus together with Southbya in a separate family, Southbyaceae. Since morphological differences with Arnelliaceae were very slight and molecular support for family status of Southbyaceae was lacking, the latter family is not accepted here. Gongylanthus Nees Plants glossy, whitish to green or purplish, creeping, attached to the substrate by numerous, long, pale rhizoids; stolons lacking. Merophytes dorsally not interlocking. Leaves succubous, opposite with the dorsal bases united, orbicular to oblong, strongly concave to almost plane, apex rounded, margins entire or crenulate. Cells thin-walled, without or with small trigones, strongly elongate toward the ventral leaf margin, to 150 μm long and about 4–10× as long as wide, cuticle smooth in tropical America species; oil bodies finely granular, 3–8(−12) per cell. Underleaves lacking or very small. Rhizoids scattered. Sporophytes in a fleshy, ventral marsupium; perianth lacking.

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Keys and Descriptions

A genus of seven to eight species, four in Colombia and Ecuador. Gongylanthus is recognized by the small, terrestrial, creeping, often whitish plants from high elevation (usually above 2000 m) with densely imbricate, opposite leaves, large leaf cells, and sporophytes in a marsupium. Opposite leaves are also found in Syzygiella (Jamesoniellaceae), but the latter genus has smaller leaf cells and a perianth instead of a marsupium. Literature: A modern taxonomic treatment of neotropical Gongylanthus is lacking. 1. Leaves flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaves concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Plants large, more than 4 mm wide. Leaves longer than wide, loosely imbricate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. granatensis 2. Plants smaller, less than 3  mm wide. Leaves about as long as wide, densely imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. muelleri 3. Leaf apex bordered by narrowly rectangular cells in 1–2 rows. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. limbatus 3. Leaf apex not bordered. Leaf margins crenulate or entire . . G. liebmannianus 1. Gongylanthus granatensis (Gottsche) Steph. Lindigina granatensis Gottsche (Calypogeia granatensis [Gottsche] Steph.) Haplozia carnosa Herzog Gongylanthus oniscoides (Spruce) Steph. (?) On soil in open or shaded high-montane environments, 1800–4300 m. Colombia: Boy, Cal, Cun, Hui, Ris, Tol. Ecuador: Az, Ca, Lo, Na, Pi, ZC. General distribution: tropical Andes, Costa Rica. Gongylanthus granatensis is a characteristic high-Andean species that is distinguished by the relatively large, often whitish plants (to more than 4 mm wide) and wide-spreading, flat, loosely imbricate, tongue-shaped leaves. Plants growing on open, exposed soil in páramo are whitish in color; those growing on shaded soil in montane forest (e.g., on steep earth banks) may be green. Gongylanthus oniscoides, described from Ecuador, is probably a modification of G. granatensis (Váňa et al. 2012a). 2. Gongylanthus liebmannianus (Lindenb. & Gottsche) Steph. Haplozia cucullata Herzog Lethocolea repens S.Winkl. (?)

Fig. 12Q–S

On soil and rock in páramo and along roads, 2700–4300  m. Colombia: Ara, Boy, Cun, Mag, Qui, Ris. Ecuador: Az, Ch, Lo, Na, Pi, ZC. General distribution: high mountains of Mexico to Bolivia and southeastern Brazil. Gongylanthus liebmannianus is a common high-montane neotropical species that is recognized by the whitish plants with concave, strongly imbricate, broader than long leaves. The species resembles G. limbatus, but the leaves in the latter species are bordered by narrowly rectangular cells (border lacking in G. liebmannianus) and are less strongly concave. Gongylanthus liebmannianus may be confused

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with Stephaniella paraphyllina with which it may often grow mixed, but the latter species has long ventral stolons (absent in Gongylanthus), asymmetrically falcate, pointed, alternate leaves (leaves symmetrical, opposite, with rounded apex in G. liebmannianus) and paraphyllia. Moreover, Stephaniella lacks a marsupium. Lethocolea repens, described from the Sierra Nevada de Santa Marta (Winkler 1969), is a possible synonym of G. liebmannianus. Although the type of L. repens (Winkler C254a) is lost, a specimen in the Winkler herbarium identified by S. Winkler as L. repens (Sierra Nevada de Santa Marta, Winkler 315, ULM) consists of Stephaniella paraphyllina and a few degenerate shoots of G. cf. liebmannianus. Winkler (1969) described the leaves of L. repens as “alternate,” but they look opposite in the original illustration and similar to those of Gongylanthus. Lethocolea repens also fits Gongylanthus in the smooth cuticle (papillose in Lethocolea). Although originally spelled “liebmaniana,” the epithet of this species is changed to “liebmanniana” under ICN art. 60.1 (Ex. 7) since it was named for Frederik Michael Liebmann. 3. Gongylanthus limbatus (Herzog) Grolle & VáňaFig. 12T Haplozia limbata Herzog Gongylanthus innovans S.Winkl. On soil and wet rock in upper montane forest environments and páramo, 3100–4450 m. Colombia: Boy, Cun, Mag, Met. Ecuador: Co, Pi. General distribution: tropical Andes, southern Central America (Panama, Costa Rica). Gongylanthus limbatus is an uncommon Andean species resembling G. liebmannianus but differing in the apical leaf margin bordered by narrowly rectangular cells. 4. Gongylanthus muelleri (Gottsche) Steph. On soil in high-montane environments, ca. 2000–4000 m. Colombia: Boy, Cun, Met (A. M. Cleef, U), Qui (Gradstein 12732 p.p., HUQ). Ecuador: Pi (SchäferVerwimp et al. 2006). General distribution: northern Andes, Central America, Mexico, Dominican Republic. Gongylanthus muelleri closely resembles G. granatensis but is smaller in size, less than 3 mm wide, more darkish in color, and the flat leaves are more densely imbricate and usually obliquely spreading. The species is newly recorded here from Colombia.

Balantiopsidaceae Plants green, red, purple, brown, or black (green in shade), creeping or erect. Stems usually with a thick-walled cortex. Branches ventral-intercalary, occasionally terminal, Frullania-type; stolons mostly lacking. Leaves transverse to succubous or weakly incubous, usually bifid, margins entire or toothed. Cells ± rectangular, thin-walled or slightly thickened, with or without trigones, cuticle smooth or striate-papillose; oil bodies finely granular, colorless, 2–3(−5) per cell. Underleaves well developed, similar

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Keys and Descriptions

to the lateral leaves but smaller. Rhizoids in bundles from underleaf bases. Gametoecia on long shoots. Sporophytes surrounded by a perigynium or a fleshy perianth. Setae of numerous cells (cross section). Capsules elongate, wall 3–4-layered, valves linear and spirally twisted (except Ruizanthus). Vegetative reproduction unknown. Seven or eight genera, mostly in the Southern Hemisphere. The most important family characteristics are (1) transverse or succubous, bifid leaves; (2) well-developed underleaves; (3) tufted rhizoids; (4) sporophyte surrounded by rather fleshy structures (fleshy perianth, perigynium, marsupium); and (5) capsule valves spirally twisted. The spirally twisted capsule valves are the main feature of this family that is otherwise only seen in Calypogeiaceae. Literature: Campos and Uribe (2012) 1. Leaves 4–5-lobed, with longly piliferous tips. Plants growing in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ruizanthus 1. Leaves 2(−3)-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves ± transverse, strongly concave. Underleaves usually half or more the length of the leaves, occasionally smaller. Plants creeping to erect. Sporophyte produced in a fleshy perigynium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isotachis 2. Leaves clearly succubous, flat to concave. Underleaves less than half the length of the leaves. Plants creeping. Sporophyte produced in a perianth with a fleshy base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neesioscyphus Isotachis Mitt. Hypoisotachis (R.M.Schust.) J.J.Engel & G.L.Merr. Plants green, red, brown, or black, usually erect, sometimes creeping. Branches ventral-intercalary. Leaves transverse (sometimes slightly succubous or incubous), bifid, about as long as wide, strongly concave, leaf tips rounded, acute or mucronate, margins entire or toothed, sometimes bordered. Cells as in the family. Underleaves similar to the leaves but smaller, about half the size (sometimes larger or smaller) and flatter. Sporophytes in a conical, fleshy perigynium at the tip of long shoots. A Southern Hemispheric genus (ca. 15 spp.), four species in Colombia and Ecuador. The transverse, bifid leaves, the large underleaves, tufted rhizoids, and fleshy, terminal perigynium are the main characteristics of Isotachis. Literature: Campos and Uribe (2012) 1. Plants creeping to ascending, 0.5–1(−2) cm long. Rhizoids usually abundant (except at stem apex), in long tufts at underleaf bases . . . . . . . . . . . . . . . . . . . 2 1. Plants growing erect, (2–)3–9 cm long. Rhizoids ± absent. . . . . . . . . . . . . . . 3 2. Leaves with a border of larger, quadrate to rectangular cells. Leaf apex sharply mucronate to cuspidate. Plant glossy light green, fleshy, growing in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. lopezii

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2. Leaf margins not bordered by elongate cells. Leaf apex obtuse to subacute. Plant dull green to blackish-brown, in the montane belt and páramo . . . . I. multiceps 3. Margins of leaves and underleaves toothed (rarely subentire). Leaves asymmetrical, bifid to 1/4–1/2, apices acute to obtuse. Midleaf cells 2–3× longer than wide. Plants reddish or brown, common throughout the montane belt and in páramo, on moist soil and rock . . . . . . . . . . . . . . . . . . . . . . . . . . I. serrulata 3. Margins of leaves and underleaves entire. Leaves symmetrical, bifid to 1/8–1/5(−1/4), apices rounded. Midleaf cells 3–5× longer than wide. Plants brown to blackish, rarely reddish-brown, in stagnant or running water in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. obtusa 1. Isotachis lopezii (R.M.Schust.) Gradst. Ruizanthus lopezii R.M.Schust. On moist soil in páramo, growing prostrate in ditches, and on steep earth walls in deep shade, 2800–4300  m. Colombia: Boy, Cun, Nar, Ris, Tol. Ecuador: Az (páramo de Cajas), Ca, Co, Na, ZC. General distribution: tropical Andes, Costa Rica, Jamaica. Isotachis lopezii is a characteristic páramo species that is readily distinguished by its fleshy, glossy light green appearance, its bifid leaves with sharply pointed lobes and almost entire margins, and rather small underleaves with sharply acuminate lobes and teeth. The most outstanding character of the species is the leaf border of large, quadrate to rectangular cells. Isotachis lopezii was originally described as a member of Ruizanthus but differs from the latter genus by bifid leaves (quadrifid in Ruizanthus) and spiralled capsule valves (straight in Ruizanthus). Vasco et al. (2002) reported the species from 1900 m in the Chocó; this record is dubious and should be checked. 2. Isotachis multiceps (Lindenb. & Gottsche) Gottsche Fig. 13A–D Hypoisotachis multiceps (Lindenb. & Gottsche) J.J.Engel & G.L.Merr. Cephalozia hians Steph., syn. nov. Type: Ecuador, Morona Santiago, Gualaquiza, Bomboiza, “ad truncum putrescentem et jacentem in silva supra Missionariorum domum,” 980 m, 8 Sept 1910, Allioni 532 (holotype G-00061125!); the new synonymy was already noted by J. Váňa (in sched.) Isotachis parva Steph. Isotachis tenax Steph. Common on moist soil and rock, often on cut road banks, (500–)2200–4350 m. Colombia: Boy, Cas, Cau, Cho, Cun, Hui, Mag, Met, Put, Ris, San. Ecuador: Pi, ZC. General distribution: tropical America. Isotachis multiceps is a widespread neotropical species that differs from other species of the genus by the creeping habit, presence of stolons near the stem base, gynoecia on short ventral branches (not terminal on main stems), and bifid leaves with broad, obtuse to subacute lobes. The underleaves in this species are maximally half the length of the leaves, deeply bifid and with entire or toothed margins, and the leaf margins are entire and unbordered.

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Keys and Descriptions

Fig. 13 (A–C) Isotachis lopezii. (A) Leaf. (B) Underleaf. (C) Apex and apical margin of leaf lobe. (D and E) Isotachis multiceps. (D) Female branch with perigynium, lateral view. (E) Leaf. (F and G) Isotachis obtusa. (F) Habit, ventral view. (G) Median leaf cells. (H and I) Isotachis serrulata. (H) Leaf. (I) Habit, ventral view. (J and K) Neesioscyphus argillaceus. (J) Habit, lateral view. (K) Leaf. (L and M) Ruizanthus venezuelanus. (L) Habit with gynoecium and perianth. (M) Apex of leaf lobe. (A–C, F, G, J–M redrawn from Campos and Uribe 2012; rest from Gradstein et al. 2001). Scale bars: A, B, F = 1500 μm; C, G = 90 μm; D, E, K = 500 μm; I, J, L = 1000 μm; M = 50 μm

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3. Isotachis obtusa Steph. Type: Ecuador, Azuay, “in monte Matanga, submers, in rivulo, 3200 m,” Oct 1909, Allioni s.n., Bryotheca Levier 6568 (G-G00053945!) Isotachis lacustris Herzog, syn. nov. Type: Bolivia, “Cordillera Real, Seen am Illampu,” ca. 4700 m, Troll 56 (JE!) Triandrophyllum maegdefrauii S.Winkl. On wet soil or rock in running water or submerged in shallow lakes or pools in páramo, 3200–4150 m. Colombia: Boy, Cau, Cun, Mag, Met, Tol. Ecuador: Az/ MS, Ca, Co, Lo, Na/Pi (páramo de la Virgen; Benitez et al. 2012), ZC (páramo of Saraguro and Yacuambi, Benitez & Gradstein s.n., HUTPL). General distribution: tropical Andes (Colombia, Ecuador, Bolivia). Isotachis obtusa is characteristic of aquatic habitats in the high Andes (shallow lakes, pools, streamlets) and is distinguished by (1) plants usually dark-brown to blackish in color, rarely reddish, growing upright, unbranched, 2.5–3 mm wide; (2) leaves with entire margins, symmetrical, ± as long as wide or wider than long, shallowly bifid to 1/8–1/5(−1/4), incision narrow or wide, depth of incision sometimes variable on a single stem, leaf apices narrowly rounded; (3) underleaves similar to leaves but slightly smaller, as long as wide to slightly longer than wide, margins entire, apices narrowly rounded to obtuse, incision narrow (in the type) or rather wide; and (4) midleaf cells 3–5× longer than wide. Isotachis obtusa has sometimes been confused with I. serrulata, which may grow in the same habitat and may occasionally have subentire leaf margins. The latter plants are distinguished from I. obtusa by the shorter leaf cells (2–3× as long as wide in midleaf), somewhat asymmetrical leaves (dorsal margin more strongly arched than ventral margin) and narrower leaf apices (obtuse to acute). Isotachis lacustris, described from high elevation in Bolivia (4700 m), is a slender phenotype of I. obtusa with rather distant leaves that are as wide as long to slightly longer than wide. Gynoecia and sporophytes of I. obtusa, which have not yet been described in this species, are present in a collection from Ecuador (páramo El Ángel, Gradstein et al. 3458, GOET, U). 4. Isotachis serrulata (Sw.) Gottsche Isotachis ecuadorensis Steph. Isotachis haematodes (Lehm. & Lindenb.) Gottsche Isotachis lindigiana Gottsche Isotachis madida (Hook.f. & Taylor) Mitt. Isotachis mascula Gottsche Isotachis piliflora Herzog Isotachis woronowii Herzog

Fig. 13E, F

On moist soil and rock, often in or near running water, sometimes submerged, 800–4500 m. Colombia: Ant, Ara, Boy, Caq, Cas, Cau, Cho, Cun, Hui, Mag, Met, Nar, Put, Ris, San, Tol. Ecuador: Az, Es, Lo, MS, Na, Pi, ZC. General distribution: widespread in tropical and southern-temperate America, extending southward to Tierra del Fuego but rare in Brazil where the species is replaced by I. aubertii (Schwägr.) Mitt.

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Keys and Descriptions

This common and variable Latin American species is readily recognized by the rather large, (2–)3–9-cm-long, upright reddish plants with deeply bifid, toothed leaves and underleaves (rarely subentire). The leaves are almost transversely inserted, ± asymmetrical with the dorsal margin more strongly arched than the ventral margin, and the underleaves are large, about 1/2–4/5 the length of the leaves. A phenotype of I. serrulata with subentire leaf margins occurs in aquatic habitats in páramo and may be confused with I. obtusa; for differences see under the latter species. Isotachis serrulata is widespread in temperate regions of Chile and Argentina where it was described as I. madida. The latter species is morphologically inseparable from I. serrulata and considered a synonym (e.g., Gradstein et  al. 1977; Dauphin 2005; Campos and Uribe 2012; Söderström et al. 2016). Fulford (1963), however, accepted I. madida as a separate species. Neesioscyphus Grolle Plants pale green to reddish, creeping. Branches ventral-intercalary. Leaves distinctly succubous, bifid with the segments equal or unequal in size, about as long as wide, flat or weakly concave, leaf tips rounded to acute to acuminate, margins entire or somewhat toothed. Cells large, rectangular, densely striate-papillose. Underleaves small, to 1/3 the size of the leaves or less, deeply bifid, bases free, not attached to leaves. Rhizoids in bundles from underleaf bases. Sporophytes produced in a perianth which is rather fleshy (2–3-stratose) in the lower half. A neotropical genus of five species, three in Colombia and Ecuador. Neesioscyphus may be confused with Lophocolea but differs from the latter genus in the elongate, rectangular leaf cells with a densely striate-papillose cuticle and purely ventral branching. Moreover, Neesioscyphus species sometimes have some reddish coloration (never reddish coloration in Lophocolea). Literature: Campos and Uribe (2012) 1. Dorsal leaf lobe larger than ventral lobe. Underleaves relatively large, ca. 2× wider than the stem. Leaf cells without trigones. . . . . . . . . . . . . . . . N. allionii 1. Dorsal leaf lobe as large as ventral lobe or smaller. Underleaves smaller, little wider than the stem. Leaf cells with or without trigones . . . . . . . . . . . . . . . . . 2 2. Leaf cells thick-walled, trigones well developed. Plants reddishbrown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. carneus 2. Leaf cells thin-walled, trigones lacking or very small. Plants pale green to brown to pink. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. argillaceus 1. Neesioscyphus allionii (Steph.) Grolle Isotachis allionii Steph. On soil and rock, 1300–2000  m. Colombia: Ris (Campos and Uribe 2012). Ecuador: MS (type), Pi, ZC. General distribution: northern Andes (Colombia, Ecuador). Neesioscyphus allionii is a rare northern Andes species that resembles N. argillaceus, but the dorsal leaf lobe in N. allionii is larger than the ventral lobe (dorsal

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lobe usually smaller or as large as ventral lobe in N. argillaceus) and the underleaves are larger than in N. argillaceus, about twice as wide as the stem. Both species have very thin-walled leaf cells, without or with minute trigones. 2. Neesioscyphus argillaceus (Nees) Grolle Jungermannia heteracria Spruce On steep, moist earth banks, cliffs, and river banks at rather low elevation, from sea level to 1600 m. Colombia: Boy, Cas, Cho (Gradstein 9039 p.p., GOET), Mag, Qui (Gradstein s.n., HUQ), Ris (Gradstein 8374, 8438, COL, GOET), Val. Ecuador: EO (Reserva Buenaventura, Benitez et  al. 1315, HUTPL), Ga (San Cristóbal), Pa (Holm-Nielsen 461 p.p., AAU, PC), ZC (Gradstein and Benitez 2014a). General distribution: Costa Rica, tropical Andes, southeastern Brazil. Neesioscyphus argillaceus is the most common and widely distributed species in the genus. The species is distinguished by thin-walled leaf cells and by the dorsal leaf lobe being frequently smaller than the ventral lobe. The latter character is not constant, however, and sometimes the two lobes are equal in size. The leaves are emarginate or short bifid (to maximally 1/3), and the apices vary from rounded to acute to acuminate. 3. Neesioscyphus carneus (Nees) Grolle Notoscyphus carneus (Nees) Steph. On moist soil along roads, 950–3700 m. Ecuador: Ca (páramo El Angel, Ramsay 2011-132, QCA), MS, Pi (Schäfer-Verwimp et  al. 2013a); to be expected in Colombia. General distribution: mainly distributed in southeastern Brazil, also found in Ecuador and Panama. Neesioscyphus carneus is a rare species characterized by leaf cells with conspicuous trigones (leaf cells thin-walled in other Neesioscyphus species). Ruizanthus R.M.Schust. Plants greenish to somewhat rose-tinged, creeping to ascending, forming small rosettes or mats. Branches ventral-intercalary. Leaves transverse, 4–5-lobed, strongly concave, leaf tips longly piliferous, margins entire or with a few linear teeth. Cells as in the family. Underleaves small, about 1/3 the size of the leaves, bifid. Sporophytes produced in a perianth which is rather fleshy (2–4-stratose) in the lower half. Capsules ovoid, valves straight. A neotropical genus with a single species, differing from other neotropical members of Balantiopsidaceae by straight capsule valves and 4–5-lobed leaves. Literature: Campos and Uribe (2012) 1. Ruizanthus venezuelanus R.M.Schust. On soil in páramo, ca. 3500 m. Colombia: Mag (Sierra Nevada de Santa Marta; van Reenen et  al. 1984). General distribution: northern Andes (Venezuela, Colombia), Costa Rica.

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Keys and Descriptions

This rare neotropical páramo species grows in small, salad-like rosettes on bare soil and is readily distinguished by the transverse, strongly concave, 4–5-lobed leaves with longly piliferous tips and leaf margins bordered by elongate cells.

Calypogeiaceae Plants translucent whitish-green to deep-green to brownish, creeping. Branches ventral-intercalary, usually sparse; stolons lacking. Leaves incubous, undivided to short-bifid, margins entire or crenulate, sometimes bordered. Cells thin-walled, cuticle smooth or finely papillose; oil bodies finely or coarsely granular, colorless, blue, brown, or violet (C. peruviana). Underleaves small, undivided or (bis)bifid. Rhizoids in bundles from underleaf bases. Gametoecia on very short ventral branches. Sporophytes in a fleshy, subterranean marsupium. Setae of numerous cells (cross section). Capsules cylindrical, wall 2-layered, valves spirally twisted. Vegetative reproduction by gemmae or caducous leaves. Four genera worldwide. The main features of the family are (1) incubous leaves with entire to very short-bifid apex and very thin-walled cells; (2) distinct underleaves; (3) rhizoids in tufts from underleaves; (4) gametoecia on very short ventral branches; and (5) sporophyte in a marsupium. The species usually grow on soil or rotten wood in shaded environments. 1. Underleaves entire or short-bifid. Leaf apex entire. Cuticle of leaf cells usually coarsely papillose. Underleaf cells usually different from leaf cells (except in M. nephrostipum), more elongate in shape, without chlorophyll and fully hyaline, cuticle smooth. Plants deep green to brown. Oil bodies brownish, finely granular. Vegetative reproduction by caducous leaves. . . . . . . . . . . . Mnioloma 1. Underleaves clearly bifid. Leaf apex usually bifid. Cuticle of leaf cells smooth or very finely punctate-papillose. Underleaf cells similar to leaf cells, with chlorophyll, not hyaline. Plants pale green to bluish-green. Oil bodies colorless or sepia, coarsely granular. Vegetative reproduction by gemmae produced on upright, flagelliform branches, not by caducous leaves. . . . . . . . . . Calypogeia Calypogeia Raddi Kantia Gray Plants pale green to bluish-green. Leaf apex entire or shallowly bilobed. Cells thin-walled, trigones small- to medium-sized, cuticle smooth or finely papillose. Oil bodies colorless, sepia or bluish, coarsely granular (resembling a small grape cluster). Underleaves variable in size and shape, bilobed to at least 1/4 of their length, not bordered; underleaf cells chlorophyllose, similar to leaf cells. Vegetative reproduction by gemmae. About 30 species worldwide, 12  in Colombia and Ecuador. The usually pale green plant color, incubous leaves, bifid underleaves, very thin-walled cells with

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small trigones, coarsely granular oil bodies, and vegetative reproduction by gemmae produced in clusters at the tips of upright flagelliform branches are very characteristic of Calypogeia. The plants typically grow on bare, moist soil in shaded sites. Literature: Bischler (1962b, c), Fulford (1968), Schuster (1995a) 1. Ventral leaf margin distinctly longer than dorsal margin, leaves asymmetrical. Underleaves usually very small, scarcely wider than the stem . . . . . . . . . . . . 2 1. Ventral margin not distinctly longer than dorsal margin, leaves symmetrical or asymmetrical. Underleaves small or large. . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Leaves bordered by elongate margin cells, at least in the lower half . . . . . . . . 3 2. Leaves unbordered. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rhombifolia 3. Leaves bordered by narrowly elongate cells with oblique, overlapping tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lechleri 3. Leaves bordered by quadrate to rectangular cells with truncate, not overlapping tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Underleaves very small, scarcely wider than the stem. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laxa 4. Underleaves larger, about 2× wider than the stem. Plants very rare, only known from Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. densifolia 5. Leaves wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. oblata 5. Leaves longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Stem leaves more than 1.5× longer than wide. Plants of lowland rainforest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Stem leaves 1–1.5× longer than wide. Plants montane . . . . . . . . . . . . . . . . . . 8 7. Underleaves wider than the stem, bifid to near the base with large divergent lobes, outer margins entire. Plants on soil and rotten wood . . . . . . . . C. tenax 7. Underleaves not wider than the stem, bifid to the middle, lobes upright, outer margins with a tooth. Plants on loamy soil . . . . . . . . . . . . . . . . . . . C. miquelii 8. Underleaves 2–4× wider than the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Underleaves smaller, 1–2× wider than the stem . . . . . . . . . . . . . . . . . . . . . . 10 9. Underleaves 2–3× stem width, distant, with a blunt tooth on the outer margin. Leaf apex bifid. Oil bodies brown-violet. Plants common . . . . . . C. peruviana 9. Underleaves larger, 3–4× stem width, (sub)imbricate, mostly without tooth on the outer margin. Leaf apex mostly entire. Oil bodies colorless. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grandistipula 10. Leaf apex uniformly bifid. Underleaf insertion line straight. . . . C. uncinulata 10. Leaf apex mostly entire (a few leaves may have bifid apices). Underleaf insertion line arched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Leaf apex apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. andicola 11. Leaf apex rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. subintegra 1. Calypogeia andicola Bischl. Calypogeia colombica S.W.Arnell, nom. inval.

Fig. 14A

On moist soil and rotten wood in montane forest and páramo, 1000–4100  m. Colombia: Boy, Cas, Cau, Ces, Cun, Tol. Ecuador: Az (páramo de Cajas, QCA), Pi. General distribution: tropical Andes.

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Fig. 14 (A) Calypogeia andicola, habit in ventral view, 20×. (B and C) Calypogeia lechleri. (B) Habit, ventral view, 25×. (C) Leaf margin, 200×. (D) Calypogeia laxa, leaf margin, 200×. (E) Calypogeia grandistipula, habit in ventral view, 20×. (F) Calypogeia miquelii, habit in ventral view. (G and H) Calypogeia peruviana. (G) Habit, 20×. (H) Underleaf, 40×. (I) Calypogeia rhombifolia, habit in ventral view, 20×. (J) Calypogeia uncinulata, habit in ventral view, 20×. (K) Calypogeia tenax, habit in ventral view, 37×. (L) Calypogeia oblata, habit in ventral view, 15×. (M–O) Mnioloma caespitosum. (M) Habit, ventral view. (N) Midleaf cells. (O) Underleaf margin cells. (P and Q) Mnioloma cellulosum. (P) Habit, ventral view, 37×. (Q) Cells near leaf apex, 200×. (a–e, g–l, p, q from Fulford 1968; rest from Gradstein et al. 2001)

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Calypogeia andicola is a characteristic Andean species recognized by symmetrical, apiculate leaves and very small, bifid underleaves with entire margins. 2. Calypogeia densifolia (Steph.) Steph. Calypogeia lechleri (Steph.) Steph. var. densifolia (Steph.) Bischl. On moist soil and trunk bases in Amazonian rainforest, 200–250 m. Colombia: Ama (Fulford 1968). General distribution: northwestern Amazonia (Colombia, Brazil). This rare Amazonian species is near to C. laxa but differs by larger underleaves, about 2× wider than the stem. In C. laxa the underleaves are very small, scarcely wider than the stem. Calypogeia densifolia has sometimes been considered a variety of C. lechleri, but the latter species has much smaller underleaves, like C. laxa, and, moreover, a leaf border of long, narrow cells with acute tips (leaf border of quadrate to rectangular cells with broad, truncate tips in C. densifolia). The present treatment of C. densifolia follows Fulford (1968); in the World Checklist (Söderström et al. 2016), the species is listed as a variety of C. lechleri. 3. Calypogeia grandistipula (Steph.) Steph. Calypogeia puiggarii Steph.

Fig. 14E

On moist soil and rotten wood in montane rainforest, 1800–2000 m. Colombia: Mag (Winkler 1976), Qui (Circasia, Gradstein 12731, HUQ). Ecuador: ZC (Parolly et al. 2004). General distribution: northern Andes, southeastern Brazil. This is a rare neotropical species recognized by the entire or short-bifid leaf apex and the large underleaves, 3–4× stem width and mostly without tooth on the outer margin. Calypogeia grandistipula is close to C. peruviana, but the leaf apex in the latter species is constantly bifid and the underleaves are smaller (2–3× stem width) and have a blunt tooth on the outer margin. Moreover, the oil bodies are brownishpurple in C. peruviana, whereas they are colorless in C. grandistipula (oil bodies seen in material from Quindio). By the entire leaf apices and bifid underleaves without lateral tooth, C. grandistipula is similar to C. subintegra, but the underleaves in the latter species are much smaller, maximally 2× stem width, and highly distant. The plants from Quindio are tentatively placed in C. grandistipula because the underleaves are contiguous to subimbricate, not densely imbricate as in typical C. grandistipula, and the leaf cells are very thin-walled. Interestingly, the Quindio plants are monoicous and copiously fertile, with numerous long marsupia. Thus far, reproductive organs had not been recorded in this species. 4. Calypogeia laxa Gottsche & Lindenb.

Fig. 14D

On rotten wood, tree bases, soil, and rock in lowland and montane rainforests, from sea level to 3000 m. Colombia: Ama, Ant, Caq, Cun, Mag, Put, Vau (Campos et  al. 2015). Ecuador: ZC (Schäfer-Verwimp et  al. 2013a). General distribution: widespread in tropical America. Calypogeia laxa is the secondmost common species of the genus in tropical America (after C. peruviana) and is readily distinguished by the asymmetrical, bordered leaves and the very small, bifid underleaves with a distinct tooth on the

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margins. The species may be confused with C. lechleri but differs from the latter by the quadrate to rectangular leaf border cells with broadly truncate ends (border cells narrowly elongate with acute ends in C. lechleri). 5. Calypogeia lechleri (Steph.) Steph.

Fig. 14B, C

On soil, tree bases, and rock in lowland and montane rainforest and scrub, from sea level to 2600 m. Colombia: Ama, Cho, Nsa, Ris. Ecuador: Na, ZC. General distribution: tropical America. Calypogeia lechleri is close to C. laxa; for differences see under the latter species. 6. Calypogeia miquelii Gottsche, Lindenb. & Nees Calypogeia amazonica (Spruce) Steph.

Fig. 14F

On moist loamy soil in lowland rainforest, 200–500(−800) m. Colombia: Ama, Cho, Mag, Vau; to be expected in Ecuador. General distribution: Central America and northern South America, also in the West Indies. Calypogeia miquelii is a common neotropical lowland rainforest species, characterized by oblong, symmetrical leaves, which are more than 1.5× longer than wide, and very small bifid underleaves with a distinct tooth on the margins. The leaves are often shorter on one side of the stem than on the other side. The species has been recorded a few times from above 1000 m, but these records may be misidentifications. The record from the Galápagos Islands (Clark 1953) is C. peruviana. Calypogeia amazonica has long been considered a synonym of C. miquelii (e.g., Gradstein and Hekking 1989) but is resurrected in the World Checklist (Söderström et al. 2016) without justification. 7. Calypogeia oblata Herzog On moist soil in upper montane forest and scrub, 2150–3500 m. Ecuador: Tu, ZC; to be expected in Colombia. General distribution: northern Andes (Venezuela, Ecuador, Peru). This rare northern Andean species is recognized by very broad leaves that are wider than long. The leaf apex is short bifid or undivided, and the underleaves are 2–3× as broad as the stem, with an emarginate to short-bifid apex and ± entire outer margins. 8. Calypogeia peruviana Nees & Mont. Calypogeia afrocoerulea E.W.Jones Calypogeia biapiculata (Spruce) Steph. Calypogeia heterophylla (Steph.) Steph. Calypogeia muscicola Steph.

Fig. 14G, H

On moist soil, humus, rock, tree bases, and rotten wood in montane forests and lower páramo, (450–)1000–4000 m; on the Galápagos Islands ocurring on thin soil over rock in deep shade in river gorges and in deep lava cracks with temporary streamlets together with Telaranea nematodes, 450–650 m. Colombia: Ama, Boy, Ces, Cho, Cun, Hui, Mag, Met, NSA, Put, Ris, San. Ecuador: Ga, Lo, Na, ZC. General distribution: widespread in tropical America; also in East Africa (Gradstein 2013b).

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Calypogeia peruviana is the most common member of the genus in tropical America. The species is readily distinguished by the symmetrically ovate leaves with a short-bifid apex and underleaves about 2–3× wider than the stem and usually with 1(−2) blunt teeth on the outer margins. Fresh material is readily recognized by the brownish-purple (sepia) or violet oil bodies; the color of the oil bodies was erroneously described as “blue” by Buczkowska et al. (2018). 9. Calypogeia rhombifolia (Spruce) Steph. Fig. 14I Calypogeia rhombifolia var. colombiana Bischl., syn. nov. Type: Colombia, Valle, road Cali to Buenaventura km 43, on rock, 3 Aug. 1958, Bischler 477 (syntype G-00114758!) On moist soil, rock, and tree bases in lowland and lower montane rainforests, from sea level to 1700  m. Colombia: Ant, Cho, Cun, Hui, Met, Ris, San, Val. Ecuador: Tu, ZC. General distribution: tropical America. Calypogeia rhombifolia is a rather widespread neotropical species that is readily distinguished by the asymmetric, unbordered leaves and tiny, bifid underleaves. The species is related to C. laxa and C. lechleri, but the latter two species have bordered leaves. In the field C. rhombifolia is recognized by its bluish-green color. Calypogeia rhombifolia var. colombiana is a robust phenotype with underleaf margins bulging outward (but not developing into a tooth). 10. Calypogeia subintegra (Gottsche, Lindenb. & Nees) Bischl. On humic soil, litter, rock, and roots in upper montane rainforest, in deep shade, 2800–3300  m. Colombia: Boy (Álvaro Alba et  al. 2007), Cun. Ecuador: ZC (Schäfer-Verwimp et  al. 2013a). General distribution: scattered in tropical America. Calypogeia subintegra is very similar to C. peruviana, but the leaf apex in C. subintegra is mostly entire (bifid in C. peruviana) and the underleaves are smaller, usually less than 2× stem width, lacking a blunt tooth on the outer margin. Oil bodies in C. subintegra are unknown and should be studied; in C. peruviana they are characteristically brownish-purple. 11. Calypogeia tenax (Spruce) Steph.

Fig. 14K

On rotten wood and tree bases in lowland rainforest, 100–350  m. Colombia: Ama, Cau, Put, Vau (Campos et  al. 2015). General distribution: northern Amazonia (Colombia, Venezuela, Brazil). A characteristic Amazonian species distinguished by oblong to subrectangular leaves with a broad, rounded apex and deeply bifid underleaves with long, diverging lobes. 12. Calypogeia uncinulatula Herzog

Fig. 14J

On damp soil in montane forests up to the páramo. Colombia: Tol (Gradstein and Uribe 2016a). Ecuador (new): EO (Reserva Buenavista, ca. 350 m, Gradstein et  al. s.n., HUTPL, QCA). General distribution: Guatemala, tropical Andes, southeastern Brazil.

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Keys and Descriptions

This rare neotropical-montane species is recognized by symmetric, ovate leaves with a bifid apex and by small, bifid underleaves with a straight insertion and with or without tooth on the outer margins. Mnioloma Herzog Calypogeia subg. Mnioloma (Herzog) Bischl. Plants deep green to brown. Leaves ± rectangular. Leaf apex undivided, rounded to apiculate. Leaf margins usually bordered by transversely rectangular or obliquely elongate cells, entire or crenulate. Cells thin-walled, trigones small, cuticle usually coarsely papillose. Oil bodies brownish, finely granular. Underleaves undivided or narrowly divided to 1/5 of their length, lobes obtuse, margin bordered; underleaf cells often elongate and hyaline, without chlorophyll and with a smooth cuticle. Vegetative reproduction by caducous leaves. A pantropical (but mostly neotropical) genus of about ten species, five in Colombia and Ecuador. Mnioloma was included in Calypogeia by Spruce, Fulford, and others but clearly differs from Calypogeia by the characters given in the key (see also Gradstein et al. 2001). Literature: Bischler (1962a), Fulford (1968), Schuster (1995a, 2000a, b) 1. Underleaf cells different from leaf cells, without chlorophyll, with a smooth cuticle and usually more elongate in shape. Underleaves ovate, entire or shortbifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaf cells similar to leaf cells, with chlorophyll. Underleaves ovate to reniform, short-bifid. Plants rare, of Amazonia and the Chocó. . . . . M. nephrostipum 2. Leaf margins conspicuously crenulate . . . . . . . . . . . . . . . . . . . . . M. cellulosum 2. Leaf margins ± entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Underleaves short-bifid, apical margin with a few sharp teeth. Leaf cells with trigones. Leaves about 2× longer than wide. Plants of lowland rainforest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. parallelogrammum 3. Underleaves undivided or short-bifid, apical margin entire. Leaf cells without or with minute trigones. Leaves 1–2× longer than wide. Plants of submontane and montane rainforests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaves at least in part bordered by quadrate to narrowly rectangular cells set at right angles to the margin. Leaves rectangular, 1.4–2× longer than wide. Plants growing below 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. caespitosum 4. Leaves not bordered by quadrate to narrowly rectangular cells. Leaves shorter, subrectangular, 1–1.3× longer than wide. Plants growing mostly above 2000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. cyclostipum 1. Mnioloma caespitosum (Spruce) R.M.Schust. Calypogeia caespitosa (Spruce) Steph.

Fig. 14M–O

On tree trunks, humus, and rock or hanging from twigs in submontane and lower montane rainforests, 400–1900 m. Colombia: Cau, Ces, Cho, Mag, Met, Nsa, Ris.

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159

Ecuador: MS (Río Quimi, Pérez et  al. 11608, 11609, QCA), Pa, ZC.  General distribution: northern Andes, Guayana Highland, Amazonia, East Africa (Mt. Kilimanjaro). Mnioloma caespitosum is a common species of northern South America characterized by rectangular leaves (ca. 1.4–2× longer than wide) with entire margins and with a border of quadrate to narrowly rectangular cells set at right angles to the margin. The leaf cells are thin-walled, without or with minute trigones, the cuticle is papillose, and the underleaves are ovate, undivided or retuse and made up of elongate, hyaline cells. 2. Mnioloma cellulosum (Spreng.) R.M.Schust. Fig. 14P, Q Calypogeia crenulata Bischl. (Mnioloma crenulatum [Bischl.] R.M.Schust.), syn. nov. Type: St. Kitts (“St. Christopher”), Breutel s.n. (isotype G-00064247!, 2 shoots) Mnioloma venezuelanum (Fulford) R.M.Schust. (?) On tree bases, rock, and rotten wood in montane rainforests, 800–2550  m. Colombia: Ant, Cho, Cun, Mag, San. Ecuador: Tu, ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical America. Mnioloma cellulosum is a rather widespread neotropical species distinguished by crenulate to crenate leaf margins bordered by narrowly rectangular cells set at right angles to the margin. Calypogeia crenulata was described as more robust, with plants more than 2 mm wide and large underleaves, but the type of C. crenulata is ca. 1.5 mm wide, with rather small underleaves, and is identical to M. cellulosum. Mnioloma venezuelanum, recorded once from Nariño (Ramírez et al. 2008), may be a phenotype of M. cellulosum with weakly crenulate leaf margins. This needs further study. 3. Mnioloma cyclostipum (Spruce) R.M.Schust. Calypogeia cyclostipa (Spruce) Steph.

Fig. 14A

On shaded soil, roots, and bark in montane rainforests and cloud forests, 1700–3600 m. Colombia: Ant, Boy, Cau, Ces, Cho, Cun, Ris, Tol. Ecuador: Lo, Tu, ZC.  General distribution: tropical Andes, Central America, Guayana Highland, southeastern Brazil. Similar to M. caespitosum but occurring at higher elevation, above 1700 m, and with shorter, subrectangular leaves without a conspicous border of quadrate to narrowly rectangular cells. 4. Mnioloma nephrostipum (Spruce) R.M.Schust. Calypogeia nephrostipa (Spruce) Steph. Calypogeia mastigophora (Spruce) Steph. On rotten wood, tree bases, rock, and soil in lowland rainforest, 30–200  m. Colombia: Cho, Mag. General distribution: scattered throughout Amazonia, Chocó. Mnioloma nephrostipum is a rare Amazonian species (also in the Chocó) that has been recorded twice from Colombia. The species stands out by small, bifid, usually

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Keys and Descriptions

reniform underleaves made up of cells similar to those of the leaves and by crenulate leaf margins. Mnioloma nephrostipum may be confused with M. cellulosum, but the latter species has larger, ovate underleaves made up of elongate, hyaline cells devoid of chlorophyll and with a smooth cuticle. 5. Mnioloma parallelogrammum (Spruce) R.M.Schust. Calypogeia parallelogramma (Spruce) Steph.

Fig. 14B, C

On moist soil, rock, rotten wood, and tree bases in Amazonian rainforest, from sea level to 300 m. Colombia: Ama, Cau, Met, Vau (Campos et al. 2015); records from higher elevation need verification. General distribution: West Indies, northern Amazonia, southeastern Brazil. Mnioloma parallelogrammum is a rare Amazonian species that is characterized by conspicuously elongate, rectangular leaves (ca. 2× longer than wide), leaf cells with distinct trigones, and short-bifid underleaves with acute, toothed apices. The leaf cells are densely papillose (not “punctate”; Fulford 1968). The species name is sometimes spelled “parallelogramum” (Söderström et  al. 2016), but the spelling “parallelogrammum” is correct.

Cephaloziaceae Plants pale green to brown, red, or purplish, creeping to ascending. Stems with or without hyalodermis. Branches ventral-intercalary or lateral-terminal, Frullaniatype; ventral stolons sometimes present. Leaves succubous, rarely transverse, bifid or undivided, inserted laterally on the stem, dorsal side of the stem usually leaf-free (except in some species of Odontoschisma), leaf margins usually entire. Cells often large and pellucid, thin-walled to thickened, cuticle smooth or papillose; oil bodies present or absent. Underleaves lacking or very small (rarely large). Rhizoids scattered. Gametoecia on leading shoots or on short ventral branches. Sporophytes surrounded by a perianth. Setae of 12 rows of cells: 8 outer rows and 4 inner rows. Capsules elongate, wall 2-layered. Vegetative reproduction by 1–2-celled gemmae produced on ascending flagelliform shoots. About 15 genera worldwide. The main features of Cephaloziaceae are (1) succubous leaves which are inserted on the lateral side of the stem (insertion of leaves usually not reaching the dorsal stem midline); (2) usual absence of underleaves (rarely present); (3) scattered rhizoids; (4) seta thin, of only 12 rows of cells; and (5) capsule elongate with a 2-layered wall, produced in a perianth. 1. Underleaves large, to ca. 1/3 of leaf length. Leaves narrowly ovate-lanceolate or oblong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaves lacking or very small. Leaves orbicular to ovate to oblong . . . . 3 2. Leaves entire or shallowly retuse, narrowly ovate-lanceolate . . . . . . Alobiella 2. Leaves bifid to 1/3–1/2 of leaf length, ovate-oblong. Plants from Costa Rica and Venezuela; not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontoschisma sect. Iwatsukia (N.Kitag.) Gradst. et al.

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3. Leaves undivided or retuse. Stems without hyalodermis. Ventral stolons present or lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Leaves bifid. Stems with hyalodermis. Ventral stolons lacking . . . . . . . . . . . . 6 4. Cells in midleaf 15–30(−40) μm long, trigones usually present (rarely lacking). Plants (0.5–)1–2  mm wide, growing on rotten wood, humus, or rock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontoschisma 4. Cells in midleaf larger, 40–100  μm long, cell walls without or with small trigones. Plants less than 1 mm wide, growing on bare soil . . . . . . . . . . . . . . 5 5. Leaves almost longitudinally inserted, leaf apex retuse. Leaf cells with small trigones. Stolons and underleaves lacking. Gynoecia on short ventral branches. Plants only known from elfin forest of southern Ecuador, above 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . Fuscocephaloziopsis subintegra 5. Leaves obliquely inserted, leaf apex mostly entire. Leaf cells without trigones. Stolons and very small underleaves usually present. Gynoecia on elongate shoots. Plants growing below 2000 m . . . . . . . . . . . . . . . . . . . . . Alobiellopsis 6. Leaves deeply concave and with longly piliferous tips, leaf base forming an inflated sac. Plants green to reddish, growing on decaying wood at high elevation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nowellia 6. Leaves not deeply concave and with piliferous tips, leaf base without sac. . . 7 7. Leaves nearly longitudinally inserted, bifid to 1/6–1/3 of leaf length, dorsal leaf base distinctly decurrent. . . . . . . . . . . . . . . . . . . . . . . . .  Fuscocephaloziopsis 7. Leaves obliquely inserted, bifid to 1/2 of leaf length, dorsal leaf base ± not decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalozia Alobiella (Spruce) Schiffn. Plants 1–2 mm wide, pale green, leafy shoots arising from a stoloniform creeping stem. Stems fragile, with hyalodermis, dorsal epidermis cells much larger than other stem cells. Branches ventral-intercalary. Leaves succubous, wide-spreading, very flat, inserted almost longitudinally along the lateral side of the stem, narrowly ovatelanceolate, apex entire and acute or shallowly retuse. Leaf cells long and narrow (40–100 μm long), often becoming narrower toward the leaf margin, cuticle smooth to slightly papillose; oil bodies finely granular, 1–2 per cell. Underleaves large, to 1/3 of leaf length, lanceolate, about as wide as the stem, shallowly to deeply bifid, lobes acuminate. Gemmae not observed. A neotropical genus with a single species. For main features of Alobiella, see under A. husnotii. Literature: Gradstein et al. (2001), Schuster (2002) 1. Alobiella husnotii (Spruce) Schiffn. Alobiella campanensis Steph.

Fig. 15D, E

On soil in open, shrubby vegetation, in Ecuador near the forest line at 2750–2900 m, elsewhere at lower elevation (500–1500 m). Ecuador: ZC (SchäferVerwimp et al. 2013a). General distribution: Lesser Antilles, Guyana, tropical Andes (Venezuela, Ecuador, Peru), southeastern Brazil.

Fig. 15 (A) Mnioloma cyclostipum, habit in ventral view, 20×. (B and C) Mnioloma parallelogrammum. (B) Habit, ventral view, 37×. (C) Underleaf, 70×. (D and E) Alobiella husnotii. (D) Habit, dorsal view. (E) Portion of stem, ventral view. (F and G) Alobiellopsis dominicensis. (F) Habit with gynoecium and perianth, dorsal view, 50×. (G) Midleaf cells, 200×. (H and I) Cephalozia crossii. (H) Habit, ventral view, 50×. (I) Leaf lobe, 200×. (J) Fuscocephaloziopsis pleniceps, habit in dorsal view, 50×. (K and L) Fuscocephaloziopsis crassifolia. (K) Habit, dorsal view. (L) Leaf. (M) Fuscocephaloziopsis pulvinata, habit in ventral view with gynoecium and perianth. (N and O) Nowellia evansii. (N) Habit, ventral view. (O) Leaf. (A–C, H–J from Fulford 1968; F redrawn from Schuster 2002; rest from Gradstein et al. 2001)

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163

Alobiella husnotii is a rare neotropical-montane species that is distinguished by (1) plants small, 1(−2) mm wide, very pale green, creeping on soil; (2) leafy stems arising from a creeping stoloniform shoots; (3) leaves narrowly ovate-lanceolate, very flat, inserted laterally on the stem, apex acute, entire or slightly bifid; (4) leaf cells elongate rectangular, walls uniformly thickened, cuticle smooth or finely papillose; and (5) underleaves well developed, lanceolate, shallowly to deeply bifid. The plants from Ecuador are from unusually high elevation. Alobiellopsis R.M.Schust. Plants very small, less than 1 mm wide, whitish-green to reddish, creeping, with upright gemmiparous shoots. Stems without hyalodermis, fragile, all stem cells ± similar in size, walls thin to uniformly thickened. Branches purely ventral-intercalary, frequently stoloniform. Leaves succubous, inserted laterally on the stem, suborbicular to elongate, apex rounded, truncate or retuse. Cells relatively large, elongate (ca. 35–100  μm long), larger toward leaf base, walls thin or uniformly thickened, without trigones, cuticle smooth; oil bodies large, finely granular, 1–4 per cell. Underleaves minute, of only a few cells, hidden between the long, colorless rhizoids; underleaves on gemmiparous and sexual shoots large, lanceolate, with narrow undivided apex. Gemmae frequently produced at the tips of upright flagelliform shoots. Gametoecia on elongate shoots. A small pantropical genus (3 spp.), one species in Colombia and Ecuador. For main features of the genus, see under A. dominicensis. Literature: Gradstein et al. (2001), Schuster (2002) 1. Alobiellopsis dominicensis (Spruce) Fulford

Fig. 15D, E

On bare, loamy soil, often on cut road banks, 250–2000 m. Colombia: Cau, Ris, Val. Ecuador: Ga, ZC.  General distribution: West Indies, northern Andes, Galápagos Islands. Alobiellopsis dominicensis is a common species of road banks in the West Indies; it has been found a few times in the northern Andes. The species is distinguished by (1) plants very small, 0.7–0.9 mm wide; (2) leaves inserted laterally on the stem (leaving the dorsal side of the stem exposed and leaf-free), ovate, apex narrowly rounded to obtuse, margins entire; (3) leaf cells large, 35–100 μm long, walls thin to uniformly thickened; and (4) underleaves very small (of a few cells). Alobiellopsis dominicensis somewhat resembles Odontoschisma but differs by smaller size, larger leaf cells with uniformly thickened cell walls, gynoecia on elongate shoots, and occurrence on bare, loamy soil. Alobiellopsis also resembles Fuscocephaloziopsis, especially F. subintegra; for differences see the generic key. Alobiellopsis acroscypha (Spruce) R.M.Schust., only known from the type from northern Peru, is doubtfully distinct from A. dominicensis; this needs study. If the two prove to be conspecific, the name A. acroscypha has priority.

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Keys and Descriptions

Cephalozia (Dumort.) Dumort. Plants small- to medium-sized, 0.5–2 mm wide, whitish-green to reddish or brown, creeping. Stems fragile, with a hyalodermis. Branches ventral-intercalary or terminal, Frullania-type; stolons or flagella sometimes present. Leaves succubous, concave, obliquely inserted, insertion line not reaching dorsal midline of stem, orbicular to ovate, deeply bifid, tips acute to narrow-acuminate, margins entire, bases not decurrent. Cells thin-walled, 25–50(−60) μm, cuticle smooth; oil bodies lacking. Underleaves lacking. Gametoecia on leading shoots or on short ventral branches. About 25 species worldwide, 2 in Colombia and Ecuador. Cephalozia is recognized by the small, whitish-green (sometimes reddish or brown) plants with large, translucent epidermal cells (hyalodermis present) and deeply bifid, concave, ovateorbicular leaves, which are substransversely inserted and not decurrent. Cephalozia is close to Fuscocephaloziopsis; the latter genus has often been included in Cephalozia but is separated on molecular grounds (Váňa et al. 2013a). For differences between the two genera, see under Fuscocephaloziopsis. Literature: Fulford (1968), Váňa (1984a) 1. Leaf lobes 4–8 cells wide at base, apex ending in a row of 2 cells. Leaf cells 30–40 μm in midleaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bicuspidata 1. Leaf lobes 3–4 cells wide at base, apex ending in a row of 2–5 cells. Leaf cells 40–60 μm in midleaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. crossii 1. Cephalozia bicuspidata (L.) Dumort. Cephalozia colombica Fulford, nom. inval. Cephalozia grandifolia Steph. On rotten wood and on soil along paths in páramo, 3700–3800 m. Colombia: Boy, Cho, Cun, Mag. General distribution: widespread and common in temperate regions of the Northern and Southern Hemisphere and in Africa; in tropical America rare and restricted to high elevations. This common Holarctic species has been recorded a few times from páramo of Colombia. It may be confused with C. crossii, which is the common Cephalozia species in the northern Andes. For differences see the key. 2. Cephalozia crossii Spruce Cephalozia bischlerae Fulford Cephalozia dussii Fulford Cephalozia patentiloba Steph. Cephalozia venezuelana R.M.Schust.

Fig. 15H, I

On shaded humus and soil over rock and in bogs at high elevations, 1700–4100 m. Colombia: Boy, Cau (type), Cun, Mag, Met, Ris, Tol. Ecuador: MS, ZC. General distribution: widespread in the mountains of tropical America. Cephalozia crossii is a rather common neotropical species and the only member of the genus that is restricted to the neotropics. It is closely similar to C. bicuspidata but differs by narrower leaf lobes with longer tips and larger leaf cells (see key).

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Fuscocephaloziopsis Fulford Cephalozia subg. Macrocephalozia R.M.Schust. Plants small- to medium-sized, 0.5–1.5 mm wide, whitish-green to brown, creeping. Stems with a distinct hyalodermis. Branches ventral-intercalary or terminal, Frullania-type. Leaves almost longitudinally inserted, insertion line not reaching dorsal midline of stem, suborbicular to subquadrate, retuse to bifid to 1/3, tips obtuse to acute, margins entire, dorsal base distinctly decurrent. Cells thin-walled, very large, 35–100 μm long, cuticle smooth; oil bodies lacking. Underleaves lacking. Gametoecia as in Cephalozia. A genus of about 17 species, 4 in Colombia and Ecuador. Fuscocephaloziopsis is close to Cephalozia but differs by leaves nearly longitudinally inserted, dorsal leaf base distinctly decurrrent, and leaf apex less deeply bifid (to max. 1/3 of leaf length). The leaf cells are large, 35–100  μm long. Fuscocephaloziopsis has been separated from Cephalozia on molecular grounds (Váňa et al. 2013a), being more distant to Cephalozia in the molecular tree than Nowellia. If Fuscocephaloziopsis is included in Cephalozia, Nowellia should also be transferred to Cephalozia. Literature: Fulford (1968), Schuster (2002), Váňa et al. (2013a) 1. Leaf lobes 1 cell wide at base. Leaf cells very large, 50–100 μm long. . . . . . 2 1. Leaf lobes more than 1 cell wide at base. Leaf cells smaller, 30–65  μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaf apex bifid to 1/10 of leaf length. Leaf lobes mostly made up of only 1 large cell. Plants of upper montane elfin forest of southern Ecuador, rare . . . . . F. subintegra 2. Leaves more deeply bifid. Leaf lobes made up of 2–3 cells. Plants of wet rainforests along the Pacific coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. pulvinata 3. Leaf lobes 4–6 cells wide at the base, ending in a row of 1–2 cells, the tip cell about as long as wide. Plants of páramo bogs, very rare . . . . . . . . F. pleniceps 3. Leaf lobes 2–3 cells wide at the base, ending in a row of 2–3 cells, the tip cell longer than wide. Plants of montane forests . . . . . . . . . . . . . . . . F. crassifolia 1. Fuscocephaloziopsis crassifolia (Lindenb. & Gottsche) Váňa & L.Söderstr. Fig. 15K, L Cephalozia crassifolia (Lindenb. & Gottsche) Fulford Cephalozia caribbeania Fulford Cephalozia subforficata Herzog On rotten wood, bark, rock, and humic soil in montane forests up to the páramo, 400–3300 m. Colombia: Ant, Boy, Cas, Ces, Cho, Cun, Hui, Mag, Met, NSa, Ris, San, Tol. Ecuador: ZC. General distribution: tropical America. Fuscocephaloziopsis crassifolia is a common neotropical species that is distinguished by the nearly longitudinally inserted, suborbicular, bifid leaves (to ca. 1/3) with narrowly acuminate lobes. The lobes are 2–3 cells wide at base and are ending in a row of 2–3 elongate cells. The plants are dioicous. 2. Fuscocephaloziopsis pleniceps (Austin) Váňa & L.Söderstr. Cephalozia pleniceps (Austin) Lindb. Cephalozia patagonica Fulford

Fig. 15J

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Keys and Descriptions

In páramo bogs, 3500–3700 m. Colombia: Boy, Cun, Tol. General distribution: widespread in temperate regions of the Northern Hemisphere, rare in the northern Andes; also in Tierra del Fuego. This Holarctic species has been found a few times in Colombia. It is near to F. crassifolia but differs in the leaves with wider lobes and shorter tips, made up of subquadrate cells. Moreover, F. pleniceps is much rarer in the Andes than F. crassifolia and is only found in páramo bogs. 3. Fuscocephaloziopsis pulvinata (Steph.) Fulford Fig. 15M Type: Dominica, without locality, Elliott 2000 (lectotype, designated by Fulford (1968), G-00066629!) Alobiella biloba Herzog (Fuscocephaloziopsis biloba [Herzog] Fulford), syn. nov. Type: Colombia, Valle, Córdoba, Dagua valley, Killip 5248 (JE n.v.) Cephalozia infuscata R.M.Schust. Common on bark of trees in wet lowland and submontane rainforest along the Pacific coast, 80–700  m. Colombia: Cho, Val (Rangel and Gradstein 2004). General distribution: Colombia, Trinidad. Fuscocephaloziopsis pulvinata is a characteristic Chocó element that is recognized by shallowly bifid leaves (to 1/4) with a broad, lunulate to truncate sinus and large cells (more than 50  μm long). The species was previously recorded from Colombia as F. biloba, which appears to be a synonym. Examination of a large series of specimens from the Chocó (Rangel and Gradstein 2004) showed full overlap of the characters of the two species. Already Schuster (2002) doubted the separation of F. biloba from F. pulvinata. 4. Fuscocephaloziopsis subintegra Gradst. & Váňa On decaying wood under dense vegetation in deep shade, in wet elfin forest on very poor acid soil, 2400–2600 m. Ecuador: ZC (Parolly et al. 2004). General distribution: only known from the type locality in Ecuador. Fuscocephaloziopsis subintegra has only been found in Purdiaea nutans elfin forest in San Francisco Biological Reserve. Due to its very small size and sheltered growth, the species is easily overlooked. It is recognized by the very shallowly bifid leaves (to 1/10) with connivent lobes consisting of only 1 triangular cell and large cells (60–100 × 50–85 μm in midleaf). The stem lacks a distinct hyalodermis. In the absence of a hyalodermis and the subentire leaf apex, F. subintegra is similar to Alobiellopsis. However, Alobiellopsis has stolons, uniformly thick-walled cells without trigones, small underleaves, and gynoecia on elongate shoots. The species has only been briefly diagnosed (Parolly et al. 2004), and a more complete description is therefore provided here: Plants green, prostrate, creeping among bryophytes. Stems 1–2  cm long, to 1  mm wide, sparsely and irregularly branched, branches ventral-intercalary, elongate and sterile or male or very short and female; stolons lacking. Stems in cross section slightly flattened, ellipsoid, 5–6 cells high, composed of thin-walled cells; epidermis cells 45–65 × 30–50 μm, not forming a distinct hyalodermis, medullar cells slightly smaller, subisodiametric, 40–55(−60) μm in diameter; ventral merophyte broad, 2–4 cells wide (but underleaves absent). Rhizoids very scarce, scattered on the ventral size of the stem.

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Leaves almost longitudinally inserted, widely spreading, obliquely subquadrate, slightly wider than long, 0.7–0.8 mm long and 0.7–1.0 mm wide (10–14 cells long and 10–18 cells wide), nearly flat, in situ somewhat convex, dorsal base slightly decurrent, margins entire, apex very slightly bifid, to 1/10 of leaf length or less, lobes connivent and sometimes overlapping, usually consisting of only 1 elongatetriangular cell. Leaf cells subisodiametric, apical 60–90  μm, median ones 60–100  ×  50–85  μm in midleaf, thin-walled or slightly thickened, with small trigones, cuticle smooth. Underleaves absent. Plants dioicous. Androecia (only one observed) terminal on an elongate branch, male bracts somewhat smaller than vegetative leaves, slightly folded and pouched at base, with a tooth at the swollen base, monandrous; bracteoles absent. Gynoecia on short ventral branches. Asexual reproduction not observed. Nowellia Mitt. Plants small, 0.5–1.5 mm wide, whitish-green to red, rose, or purplish, creeping. Stems fragile, with a distinct hyalodermis. Branches ventral-intercalary or terminal, Frullania-type; stolons lacking. Leaves subtransverse, standing upward away from the stem, bifid (rarely undivided), deeply concave and saccate in the lower half due to inrolling of the ventral margin; leaf tips usually piliferous (sometimes rounded), leaf margins entire or toothed. Cells fully thin-walled, without trigones, cuticle smooth; oil bodies lacking. Underleaves absent. Gametoecia on leading shoots or on short ventral branches. Gemmae not observed. A tropical-Holarctic genus (10 spp.), two species in Colombia and Ecuador. Nowellia is readily recognized by the deeply concave, bifid leaves with a saccate base due to inrolling of the ventral margin. The leaf apices in the two species from Colombia and Ecuador are longly piliferous. Literature: Grolle (1968) 1. Leaf lamina from the base to the middle of the sinus (between the two lobes) 4–8 cells high. Sac of the leaf fully inflated . . . . . . . . . . . . . . . . . . . . N. evansii 1. Leaf lamina higher, from the base to the middle of the sinus 10–15 cells high. Sac partially inflated, upper portion of the sac flattened . . . . . . . . N. curvifolia 1. Nowellia curvifolia (Dicks.) Mitt. On rotten wood and moist soil in montane forests and páramo, 900–3200  m. Colombia: Mag, Ris. Ecuador: Ca (Burghardt 2019), Lo, MS (Río Quimi, Pérez et al. 11598, QCA), ZC (Parolly et al. 2004 as N. evansii; Benitez and Gradstein 2011). General distribution: widespread in the temperate regions of the Northern Hemisphere; scattered in tropical America, from Mexico to Bolivia. This widespread Holarctic species is easily distinguished by its reddish to purplish color and leaves with piliferous lobes and with a partially inflated sac, which is flattened in the upper half. The plants typically grow in extensive mats on dead wood or soil. The species closely resembles N. evansii, and the two species have sometimes been confused; for differences see the key.

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Keys and Descriptions

2. Nowellia evansii Grolle Nowellia caribbeania Fulford

Fig. 15N, O

On rotten wood in montane forest, 2200–2700  m. Colombia: Mag, San. General distribution: West Indies, Central America, Guyana, northern Andes. Nowellia evansii is close to N. curvifolia but differs in leaves with a shorter lamina and a fully inflated sac (see key). The species is not yet known from Ecuador; the record from Zamora Chinchipe (Parolly et al. 2004) belongs to N. curvifolia. Odontoschisma (Dumort.) Dumort. Anomoclada Spruce Plants small- to medium-sized, 0.5–2(−4) mm wide, glistening green to yellowish-brown to reddish or deep purplish, little branched, creeping or ascending from a stoloniform base, sometimes sticky by mucilage when fresh (O. portoricense). Stems rigid, of rather uniform thick-walled cells or epidermis cells smaller and thicker-walled, hyalodermis lacking. Branches intercalary, ventral or lateral or dorsal, ventral stolons present. Leaves succubous, undivided or bifid, flat to strongly concave, orbicular to elongate, apex rounded, margins flat or incurved, plane or undulate, sometimes bordered by thick-walled cells, leaf insertion line extending to the dorsal midline of stem or not. Cells ± isodiametric, thin-walled or with large trigones, cell lumen quadrate to rounded to angular to stellate depending on trigone size, cuticle smooth or papillose; oil bodies rather large, 2–5 per cell, finely granular. Underleaves lacking or small, with few or numerous slime cells. Gametoecia on short ventral branches. Vegetative reproduction by whitish-green gemmae produced in clusters at the tips of short, upright shoots with scale-like leaves and underleaves. A tropical-Holarctic genus of 21 species, 7  in Colombia and Ecuador. Three further species, O. bifidum (Fulford) Gradst. & Ilk.-Borg., O. jishibae (Kitag.) L.Söderstr. & Váňa, and O. steyermarkii Gradst. & Ilk.-Borg., have been recorded from Venezuela (see Gradstein and Ilkiu-Borges 2015). The main features of Odontoschisma are the rather rigid stems without hyalodermis and with ventral stolons; the orbicular to ovate and usually undivided leaves (rarely bifid); cells with small or large trigones and with large, finely segmented oil bodies; and the frequent presence of gemmae produced on upright, flagelliform shoots. The genus may be confused with Solenostoma, but the latter genus lacks stolons and gemmae and the leaf insertion in Solenostoma always extends to the dorsal midline of stem (frequently ending before the dorsal midline in Odontoschisma). All Odontoschisma species from Colombia and Ecuador belong to O. sect. Denudatae R.M.Schust., characterized by undivided leaves and frequent presence of gemmae. Literature: Aranda et al. (2014), Gradstein and Ilkiu-Borges (2015) 1. Leaves 1.5–2× longer than wide. Trigones very large, swollen . . . . . . . . . . . . 2 1. Leaves shorter, 1–1.5× longer than wide. Trigones small or large . . . . . . . . . 3 2. Plants very small, less than 1  mm wide, growing on rock in páramo. Stems fragile. Leaves concave, oblong, margins not undulate. . . . . . . . . . . O. engelii

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2. Plants larger, more than 1.5 mm wide, growing in Amazonian lowland forest. Stems rigid. Leaves ± flat or convex in the middle, (sub)rectangular, margins undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. portoricense 3. Plants purely green. Leaf margins plane, leaves flat. Leaf cells with small trigones, cell lumen mostly quadrate, occasionally rounded. . . . . . . . . . . . . . 4 3. Plants green to brownish to reddish or purple. Dorsal leaf margin ± upcurved, leaves concave or almost flat. Leaf cells with rather large trigones, cell lumen rounded to stellate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Leaf margins bordered by thick-walled cells. Leaves ovate-subrectangular, to 1.2× longer than wide. Gemmae usually present. Plants growing on rotten wood and humus in montane forest. . . . . . . . . . . . . . . . . . . . . . . . . . . O. longiflorum 4. Leaf margins not bordered by thick-walled cells. Leaves rectangular, 1.3–1.5× longer than wide. Gemmae lacking. Plants growing in Amazonian lowland forest, very rare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. soratamum 5. Leaf margins bordered by thick-walled cells (trigones in midleaf with much smaller than at the margin). Plants growing in montane forest below 3000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. longiflorum 5. Leaf margins not bordered by thick-walled cells. In forests and páramo . . . . 6 6. Leaf surface with broad colorless papillae, the papillae 4–15 μm in diameter, less than 10 per cell. Gynoecia occurring in the middle or upper portion of the stem. Plants of páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. cleefii 6. Leaf surface smooth or with very small papillae, the papillae mostly less than 3 μm in diameter, usually more than 10 per cell. Gynoecia usually restricted to the base of the stem. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Leaves flat to slightly concave, usually longer than wide, occasionally as long as wide. Dorsal leaf-free strip of stem mostly 2–3 cells wide. Plants common, from sea level to páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. variabile 7. Leaves distinctly concave, rounded, as wide as long or wider than long. Dorsal leaf-free strip of stem absent or 1 cell wide. Plants very rare, in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. denudatum 1. Odontoschisma cleefii Gradst., S.C.Aranda & Vanderp.

Fig. 16A, B

On damp soil in grass páramo, bamboo páramo and in Sphagnum bogs, 2300–4100  m. Colombia: Ara, Boy, Cho, Cun, Hui, Qui (Gradstein et  al. s.n., HUQ); occurring in all three cordilleras of Colombia, to be expected in Ecuador. General distribution: Colombia, Peru (Graham et al. 2016). Odontoschisma cleefii is a characteristic Colombian páramo species that is recognized by the large, colorless papillae (4–15 μm in diameter, less than 10 per cell) on the leaf surface and at the leaf margin. The papillae vary in height and can sometimes be rather low and somewhat inconspicuous. The plants are 0.8–1.5 mm wide, usually dark purplish or reddish-green in color (rarely green; Graham et al. 2016), and the leaves are somewhat concave and inserted up to the dorsal midline of the stem; a dorsal leaf-free strip is lacking. A further character of O. cleefii is the frequent occurrence of gynoecia in the middle or upper portion of the stem. In other species the gynoecia are normally restricted to the base of the stem. Odontoschisma cleefii was previously identified as O. atropurpureum, O. denudatum, or O.

Fig. 16  Odontoschisma. (A and B) O. cleefii. (A) Habit, lateral view. (B) Midleaf cells. (C and D) O. denudatum. (C) Habit, dorsal view. (D) Leaf. (E and F) O. longiflorum. (E) Cells at leaf margin. (F) Habit with gemmiparous shoot, dorsal view. (G and H) O. engelii. (G) Gemmiparous shoot, ventral view. (H) Habit, dorsal and lateral view. (I–L) O. portoricense. (I) Midleaf cells. (J) Habit (flattened), ventral view. (K) Habit (flattened), dorsal view. (L) Leaf. (M and N) O. soratamum. (M) Cells at leaf margin. (N) Dorsal leaf-free strip. (O–Q) O. variabile. (O) Habit, dorsal view. (P) Habit with gemmiparous shoot, dorsal view. (Q) Habit, dorsal view. (All from Gradstein and Ilkiu-Borges 2015). Scale bars: A, D, H = 0.5 mm; B, E, I, M = 25 μm; C, F, J, K, O–Q = 1 mm; G, L = 250 μm; N = 200 μm

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falcifolium, but the large papillae and molecular evidence separate O. cleefii from other neotropical species (Aranda et al. 2014). 2. Odontoschisma denudatum (Nees) Dumort.  Odontoschisma cordifolium Steph.

Fig. 16C, D

On damp soil in páramo, ca. 3500–4000 m. Colombia: Ara, Hui/Cau. Ecuador: Ca, Lo, ZC (páramo of Saraguro and Yacuambi, Benitez et  al. 1309, HUTPL). General distribution: common and widespread in the temperate regions of the Northern Hemisphere; very rare in tropical America (Mexico, Cuba, northern Andes); also in Madagascar. Odontoschisma denudatum is a widespread Holarctic species that is very rare in the tropics. Most neotropical specimens identified as O. denudatum belong to O. cleefii, O. longiflorum, or O. variabile (Gradstein and Ilkiu-Borges 2015). The species is currently known in the Andes from only three collections, all from páramo. Odontoschisma denudatum differs from O. variabile in having concave, rounded leaves (rather flat and ± elongate in O. variabile), from O. longiflorum in larger trigones and lack of a leaf border, and from O. cleefii in the finely papillose cuticle (coarsely papillose in O. cleefii). 3. Odontoschisma engelii Gradst. & Burghardt

Fig. 16G, H

On rock in páramo, ca. 3000–3150  m. Ecuador: Lo, ZC (Gradstein and Burghardt 2008); to be expected in Colombia. General distribution: northern Andes (Ecuador, Venezuela). This rare northern Andean páramo species is readily separated from other members of the genus Odontoschisma by the distinctly elongate leaves, 1.5–2× longer than wide. Further notable features of O. engelii include the very small size of the plants, the stems without dorsal leaf-free strip (dorsal leaf insertion extending to the midline of the stem), the smooth cuticle, the wine-red gemmae, and the long, microphyllous branches with densely imbricate leaves. Odontoschisma engelii has not been recorded from Colombia but is expected there. 4. Odontoschisma longiflorum (Taylor) Trevis.

Fig. 16E, F

On rotten wood and humus in montane rainforests, 500–2700  m. Colombia: Boy, Hui. Ecuador: Ga, ZC. General distribution: throughout tropical America. Odontoschisma longiflorum is a rather common neotropical species distinguished by the green plants with rather flat leaves bordered by thick-walled cells and by leaf cells with rather small trigones and mostly quadrate lumina (rarely rounded). Gemmae are frequently present. The species is restricted to montane forests, below 3000 m. 5. Odontoschisma portoricense (Hampe & Gottsche) Steph. Anomoclada portoricensis (Hampe & Gottsche) Váňa Anomoclada mucosa Spruce

Fig. 16I–L

On rotten logs, roots, and tree bases in Amazonian rainforest, 100–350  m. Colombia: Ama, Vau; to be expected in Ecuador. General distribution: northern tropical America.

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Keys and Descriptions

Odontoschisma portoricense is common on logs and rotten wood in Amazonia and the Guianas. The species is recognized by the rather large, glistening-green plants (1.5–5  mm wide) with rigid stems, subrectangular leaves with somewhat undulate margins, leaf cells with very large trigones and stellate lumina, and very short and broad underleaves with numerous slime papillae. Fresh plants are sticky due to the abundant secretion of mucilage. 6. Odontoschisma soratamum Fulford

Fig. 16M, N

On rotten log in Amazonian rainforest, 250 m. Colombia: Ama (Río Apoporis: Soratama; Fulford 1968). General distribution: only known from the type from Colombia. Odontoschisma soratamum is a rare Colombian endemic distinguished by the pale green plants with very flat, subrectangular leaves made up of rather thin-walled cells, with small trigones. A leaf border is lacking and gemmae have not been observed. The species is known from one locality in Amazonian rainforest and should be more widespread. 7. Odontoschisma variabile (Lindenb. & Gottsche) Trevis. Fig. 16O–Q Odontoschisma atropurpureum Steph. Odontoschisma denudatum (Nees) Dumort. var. andinum Spruce Odontoschisma falcifolium Steph. Odontoschisma sphagni (Dicks.) Dumort. var. amazonicum Spruce On rotten wood, tree bases, rock, humus, and soil, from sea level to 4100  m. Colombia: Ama, Ara, Boy, Caq, Cho, Cun, Gui (Churchill et al. 17739, GOET), Hui, Met, Put, Ris, Tol, Vau. Ecuador: Ca, Ga, Lo, MS, Na, ZC. General distribution: tropical America, tropical Africa. Odontoschisma variabile is the most common species of the genus Odontoschisma in tropical America, with a remarkably broad elevational range. The species has often been confused with O. denudatum but differs from the latter in the more flattened and often somewhat elongate leaves (1–1.5× longer than wide) and the broader leaf-free strip on the dorsal side of the stem (ca. 2–3 cells wide). The plants are green to brown to reddish or purplish in color, the leaves are not bordered and not undulate, the dorsal leaf margin is narrowly upcurved (but lamina otherwise flat), the trigones are usually large and swollen, the cuticle is finely papillose to almost smooth, underleaves are lacking or very small and with few slime papillae, and gemmae are frequently present.

Cephaloziellaceae (Phycolepidoziaceae) Plants very small, thread-like, usually less than 1 mm wide, creeping or ascending, with or without stoloniform base. Stems without hyalodermis. Branches variable, ventral- or lateral-intercalary, and terminal, Frullania-type. Leaves transverse or

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succubous, bifid or entire, insertion line extending to dorsal midline or not. Cells small, usually less than 25  μm, with thin or uniformly thickened walls, trigones absent, cuticle smooth or papillose; oil bodies finely granular. Underleaves very small or lacking. Rhizoids scattered. Gametoecia on leading shoots or very short branches. Sporophytes surrounded by a long-cylindrical perianth, mouth of the perianth often crenate and bordered by conspicuously elongate, thick-walled cells. Setae very thin, usually of only 8 rows of cells: 4 large outer rows and 4 small inner rows (12 rows of cells in Gymnocoleopsis). Capsules elongate, wall 2-layered. Vegetative reproduction by gemmae produced on upright flagelliform shoots. About seven to eight genera worldwide. The most important features of Cephaloziellaceae are (1) very small plants (less than 1 mm wide); (2) usually bifid leaves; (3) small, uniformly thick-walled cells without trigones; (4) absence of a hyalodermis, (5) underleaves absent or very small; (6) seta very thin, usually made up of only 8 rows of cells: 4 very large outer cells and 4 tiny inner cells. The present family definition follows Gradstein et al. (2001, 2014) and Crandall-Stotler et al. (2009). Váňa et al. (2013b), however, modified the circumscription and transferred several genera of Scapaniaceae to Cephaloziellaceae. 1. Leaves undivided or short-bifid (to 1/5). Leafy stems ascending to erect, arising from a creeping stolon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kymatocalyx 1. Leaves more deeply bifid, to 1/3 or more. Leafy stems creeping, stolon lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves obcuneate, widest in the upper half, apices rounded. Plants light green, minute, ca. 0.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziopsis 2. Leaves not obcuneate, apices acute or narrowly obtuse. Plants green to brown or reddish, 0.3–1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Plants ca. 1 mm wide. Leaves distinctly concave. Perianth terete. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocoleopsis 3. Plants smaller, 0.3–0.6 mm wide. Leaves not concave. Perianth plicate. . . . . 4 4. Leaves subtransverse, insertion reaching the dorsal midline of the stem. Cells in midleaf very small, 8–15  μm, uniformly thick-walled (rarely thinwalled) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloziella 4. Leaves distinctly succubous, insertion not reaching the dorsal midline of the stem. Cells in midleaf larger, 15–30 μm, thin-walled . . . . . . . . . . Cylindrocola Cephaloziella (Spruce) Schiffn. Plants dull green to brown or red, very small, usually less than 0.5 mm wide; stolons lacking. Stems rigid, without hyalodermis. Leaves suberect to spreading, subtransverse, inserted to the dorsal midline of the stem, bifid, tips acute, margins entire or toothed. Cells very small (ca. 8–15 μm), usually uniformly thick-walled, without trigones, cuticle papillose or smooth; oil bodies as in the family. Underleaves lacking or very small. Gemmae sometimes produced on upright flagelliform shoots, green to red. About 40 species worldwide, 2 in Colombia and Ecuador. Cephaloziella is readily distinguished from Cephalozia by the small stem epidermal cells (hyalodermis absent) and the very small, often thick-walled leaf cells without trigones and often with a papillose cuticle. Moreover, the leaves in Cephaloziella are smaller, little

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Keys and Descriptions

wider than the stem, and more transverse, and the cells have oil bodies (absent in Cephalozia). Due to the small cells with thick walls, Cephaloziella species are less transparent than Cephalozia. Cephaloziella may be confused with Lophocolea fragmentissima and Spenolobus austroamericanus, both of which are very small plants with transverse, bifid leaves. However, L. fragmentissima has strongly caducous leaves and grows on bark (Cephaloziella grows on soil or rock), whereas S. austroamericanus is separated from Cephaloziella by reddish-brown plant color, broader leaf lobes (8–10 cells wide at base), obtuse leaf apex, and more irregular leaf areolation. Small, juvenile plants of Plagiochila spp. may also resemble Cephaloziella but are separated by larger leaf cells (usually more than 20 μm in midleaf) and cell walls thin or with trigones (uniformly thickened in Cephaloziella). Literature: Fulford (1968) 1. Gemmae smooth, ellipsoid. Underleaves present on sterile stems (but small and sometimes scarce). Leaf margins entire or toothed. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. divaricata 1. Gemmae mostly (2–)4-angled, each angle with a papilla. Underleaves absent on sterile stems. Leaf margins entire (rarely with a tooth at the base). Plants autoicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. fragillima 1. Cephaloziella divaricata (Sm.) Schiffn. Cephaloziella andina Herzog On humic soil, rotten wood, and thin soil over rock in shaded and exposed sites in scrub, along trails, in pampa and páramo, (400–)3000–3500 m; on the Galápagos Islands widespread on soil and moist lava rock at 400–1200  m Colombia: Boy, Cun. Ecuador: Ca (?), Ga. General distribution: common and widespread in temperate regions of the Northern Hemisphere, scattered in the tropics and the Southern Hemisphere. Cephaloziella divaricata is a very widespread species that is distinguished by (1) plants dioicous, leaves with entire or slightly toothed margins; (2) leaf cells 10–20  μm long, with ± evenly thickened walls and a smooth or finely papillose cuticle; (3) underleaves present on sterile and fertile shoots, small, subulate to bifid; and (4) gemmae ellipsoid, with smooth walls. The species is readily separated from C. fragillima, the other Cephaloziella species in the region, by the presence of underleaves on sterile shoots; the smooth-walled, ellipsoid gemmae; and dioicy. On the Galápagos Islands, C. divaricata occurs at much lower elevation than on the continent, and in some plants underleaves are developed only sparingly (e.g., San Cristóbal, Gradstein H317, GOET, U). Such specimens were previously named C. cf. antillana (Gradstein 2009). 2. Cephaloziella fragillima (Spruce) Fulford Fig. 17A–E Type: Bolivia, Mapiri, 3000 ft., Rusby 3076 (MANCH n.v.) Cephaloziella granatensis (Steph.) Fulford, syn. nov. (Cephalozia granatensis Steph.). Type: Colombia, Antioquia, páramo de San Isabel, 1874, Wallis s.n., ex herb. Jack (lectotype, designated by Váňa et al. (2013b), G-00067742!) Cephaloziella pygmaea (Spruce) Váňa (?), fide Váňa et al. (2014b)

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Fig. 17 (A–E) Cephaloziella fragillima. (A) Gemmiferous shoot, lateral view. (B) Habit, dorsal view. (C) Leaves. (D) Leaf lobe. (E) Gemmae. (F) Cephaloziopsis intertexta, habit in dorsal view. (G and H) Cylindrocolea rhizantha. (G) Habit with gynoecium, dorsal view. (H) Midleaf cells. (I–K) Gymnocoleopsis cylindriformis. (I) Habit with gynoecium, lateral view. (J) Habit, lateral view. (K) Midleaf cells. (L–N) Kymatocalyx dominicensis. (L) Upright leafy shoot with gynoecium and stoloniform base, lateral view. (M) Portion of shoot, dorsal view. (N) Midleaf cells. (O and P) Kymatocalyx rhizomaticus. (O) Habit, ventral view. (P) Habit, dorsal view. (All from Gradstein et al. 2001)

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Keys and Descriptions

On soil in exposed montane and alpine environments, (600–)1300–4000 m; on the Galápagos Islands on thin soil over steep rock and cliffs in moist pampa of volcan Cerro Azul (Isabela) at 600–750  m, together with Cephaloziella divaricata. Colombia: Boy, Cal, Cas, Cau, Ces, Cun, San, Tol. Ecuador: Az (páramo de Cajas, Burghardt s.n., QCA), Ga (Gradstein 2009), Pi. General distribution: tropical America, Madeira. Cephaloziella fragillima is the most common Cephaloziella in the neotropics and is recognized by (1) plants autoicous, usually fertile, without underleaves on sterile shoots; (2) leaves broadly to narrowly triangular, margins entire, apex 1–2-celled; (3) leaf cells 10–13.5 × 7.5–10 μm at leaf-lobe base, thin-walled to uniformly thick-walled, cuticle finely papillose; and (4) gemmae usually present, reddish-brown, 9–15  μm long, ovate-subrectangular, 2-celled, usually (2–)4-angled, each angle with a projecting papilla. Phenotypes of Cephaloziella fragillima with copious gemmae have been called C. granatensis, which is a synonym. The width of the leaf lobes, used by Fulford (1976) to separate C. granatensis from C. fragillima, clearly overlaps. Cephaloziella fragillima is morphologically also close to C. hampeana (Nees) Schiffn., a species widely distributed in the temperate region of the Northern Hemisphere and in addition occurring in Mexico, Guatemala, and northern Argentina (Flores et al. 2017). The latter species is separated from C. fragillima by the larger, smooth gemmae, which are 16–28 μm long and ellipsoid in shape (Fulford 1976; Schuster 1980a; Paton 1999). Flores et al. (2017) listed several further characters separating the two species, including leaf cell dimensions, cell wall thickess, male bract margins, and plant color, but all these characters overlap. Excluded Record Cephaloziella antillana (Spruce) Fulford – Ecuador: Ga (Gradstein and Weber 1982 “cf.”). The Galápagos material belongs to C. divaricata. Cephaloziopsis (Spruce) Schiffn. Plants light green, lacking secondary pigmentation; stolons lacking. Branches sparse, predominantly ventral-intercalary. Leaves almost transverse, inserted to the dorsal midline of the stem, bifid to 1/3–1/2, obovate to obcuneate, widest in the upper half, tips rounded to broadly obtuse, margins entire. Cells relatively large, ca. 20–35 μm, very thin-walled, without trigones, cuticle smooth. Underleaves lacking or very small. Gemmae lacking. A neotropical genus with a single species, characterized by obovate to obcuneate, bifid leaves with rounded to broadly obtuse tips; relatively large, thin-walled cells (20–35 μm); and absence of underleaves. Literature: Gradstein et al. (2001), Schuster (2002) 1. Cephaloziopsis intertexta (Gottsche) R.M.Schust. Cephaloziopsis schistophila (Spruce) R.M.Schust.

Fig. 17F

Cephaloziellaceae

177

On shaded moist soil banks and on rock near rivers, occasionally on bark, 450–2100 m. Colombia: Boy, Mag, Qui (Armenia, Gradstein s.n., HUQ). Ecuador: Ga, MS, ZC. General distribution: scattered in tropical America. The tiny, light green plants with obovate to obcuneate, bifid leaves, and rounded leaf tips are unmistakable. Cylindrocolea R.M.Schust. Plants very small, ca. 0.3–0.6 mm wide (rarely wider), light green to brown or reddish; stolons lacking. Leaves succubous, rather flat, inserted laterally on the stem, insertion not reaching the dorsal midline of the stem, bifid, tips acute or somewhat obtuse, margins entire. Cells 15–25(−30) μm, thin-walled, cuticle smooth; oil bodies as in the family. Underleaves lacking. Gemmae rarely produced. A pantropical genus (ca. 12 spp.), one species in Colombia and Ecuador. Cylindrocolea is closely related to Cephaloziella but differs by leaves clearly succubous (± transverse in Cephaloziella) and inserted laterally on the stem, leaving a dorsal strip of 2 rows of stem cells “leaf-free” (as in Cephaloziaceae). Furthermore, Cylindrocolea has larger, thin-walled leaf cells. Literature: Fulford (1976), Schuster (1980a, b) 1. Plants 0.3–0.6  mm wide. Cells in leaf middle subquadrate, 15–20(−25) μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rhizantha 1. Plants larger, 0.6–1 mm wide. Cells in leaf middle rectangular, 20–30 μm long. In lowland rainforest of the Guianas and the Mata Atlântica of Brazil; to be expected in Colombia and Ecuador . . . . . . C. planifolia (Steph.) R.M.Schust. 1. Cylindrocolea rhizantha (Mont.) R.M.Schust. Cylindrocolea obtusifolia Fulford

Fig. 17G, H

On soil or rock in open and often rather dry environments, also on bark, 200–1300 m. Colombia: Cas, Cho. Ecuador: Ga (Gradstein 2009). General distribution: rather widespread in tropical and subtropical America. Cylindrocolea rhizantha resembles Cephaloziella, but the leaves are ± laterally attached to the stem, with the insertion line not reaching the dorsal midline of the stem, and the leaf cells are larger (to 25 μm long in midleaf) and quite thin-walled. Moreover, the species prefers dry habitats and is restricted to low elevations (below 1500 m). The leaf tips in C. rhizantha vary from acute to obtuse. Gymnocoleopsis R.M.Schust. Plants ca. 1 mm wide, pale green to dark brown, creeping, without stolons. Branches lateral-intercalary only. Leaves loosely imbricate, succubous, concave, bifid, apices obtuse to acute, margins entire, insertion line reaching the dorsal midline of stem. Cells ca. 16–23 μm in midleaf, with uniformly thickened walls, trigones lacking, cuticle smooth; oil bodies finely granular. Underleaves lacking. Rhizoids sparse. Autoicous or dioicous. Perianths smooth, without plicae. Setae of 8 outer cell rows and 4 inner rows. Capsule walls 2-layered. Gemmae usually lacking.

178

 

Keys and Descriptions

Afro-American genus of two species, one in Colombia. For principal characteristics of Gymnocoleopsis, see under G. cylindriformis. Gymnocoleopsis was long classified in Lophoziaceae or Scapaniaceae, but molecular analysis indicated that the genus is a member of Cephaloziellaceae (De Roo et al. 2007). Literature: Gradstein et al. (2001), Schuster (2002) 1. Gymnocoleopsis cylindriformis (Mitt.) R.M.Schust. Fig. 17I–K Gymnocoleopsis multiflora (Steph.) R.M.Schust. (fide Váňa et al. 2012b) On moist, peaty soil and rock in páramo, often near running water, 3300–4460 m. Colombia: Boy, Cun, Ris. Ecuador: Ch, Co, Pi (Váňa 2011). General distribution: at high elevations in Mexico, the tropical Andes, and East Africa; also on subantarctic islands eastward to Kerguelen (Gradstein et al. 2018a). Characteristic of Gymnocoleopsis cylindriformis are the greenish-brown plants (ca. 1 mm wide) with laxly imbricate-concave, bifid leaves with narrowly obtuse to acute lobes, the ± uniformly thickened cell walls without distinct trigones, and the terete perianth. The species closely resembles Lophonardia laxiflora (Scapaniaceae) which grows in the same habitat. The latter species, however, has leaf cells with small trigones (occasionally fully thin-walled), plicate perianths, and a stoloniform stem base. Kymatocalyx Herzog Stenorrhipis Herzog Plants dark green to brown, with upright leafy shoots arising from creeping stolons, forming small mats. Branching purely ventral-intercalary. Leaves succubous or almost transverse, inserted to the dorsal midline of the stem or not, orbicular to elongate, apex rounded to short bifid. Cells 12–22 μm long, walls slightly thickened, cuticle smooth. Underleaves lacking. Gemmae not observed. A pantropical genus (4 spp.), two species in Colombia and Ecuador. Kymatocalyx is characterized by tiny, upright leafy shoots arising from a creeping stolon and succubous, orbicular to somewhat elongate leaves with a rounded or short-bifid apex. The species in this genus usually grow on soil or rock near running water. Literature: Gradstein and Váňa (1999), Schuster (2002) 1. Leaves (sub)orbicular, about as long as wide, apex undivided. Leaf insertion reaching dorsal midline of stem or not, dorsal leaf-free strip of stem 0–2 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. dominicensis 1. Leaves ovate-oblong, 1.3–2× longer than wide, apex truncate to short-bifid. Leaf insertion not reaching dorsal midline of stem, dorsal leaf-free strip of stem broader, 2–6 cells wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. rhizomaticus 1. Kymatocalyx dominicensis (Spruce) VáňaFig. 17L–N Kymatocalyx cubensis (Steph.) Váňa Kymatocalyx stoloniferus Herzog

Gymnomitriaceae

179

On moist rock and soil over rock, frequently in river beds, 900–2000  m. Colombia: Ris, San. Ecuador: ZC (Parolly et al. 2004). General distribution: tropical America, Madagascar. Kymatocalyx dominicensis is a widespread peri-African species and distinguished by the tiny, dark green plants with erect, leafy shoots arising from a creeping stolon and with (sub)orbicular leaves with undivided apex and an insertion line almost reaching the dorsal midline of stem, leaving a dorsal leaf-free strip of 0–2 stem cells wide. The plants look like a small Solenostoma but are immediately separated from the latter genus by the creeping stolons. 2. Kymatocalyx rhizomaticus (Herzog) Gradst. & VáňaFig. 17O, P On moist earth, rock, and logs, in and along rivulets and along roads, in lowland and submontane rainforest areas, from sea level to 1100 m. Colombia: Cho (road Tutunendo – El Carmen, abundant, Gradstein 8981, 9034, 9040, COL, GOET), Vau (Río Apoporis, Schultes & Cabrera 11971, CINC, PC). Ecuador: Pi (Río Toachi; Gradstein and Benitez 2017). General distribution: pantropical but known from only few localities (Borneo, Madagascar, Panama, Colombia, Ecuador, French Guiana); in the neotropics mainly occurring in the Chocó region. Kymatocalyx rhizomaticus is readily separated from K. dominicensis by the pale green plants with somewhat elongate, truncate to short-bifid leaves (bifid to 1/6–1/5) and very thin-walled cells. Moreover, the leaves in K. rhizomaticus are more laterally attached to the stem, and the leaf-free strip on the dorsal side of stem is broader, 2–6 cells wide (0–2 cells wide in K. dominicensis). The plants are often very tiny and almost flagelliform, with leaves very obliquely spreading and becoming smaller to the apex.

Gymnomitriaceae Plants small and with very densely imbricate leaves, worm-like, 0.2–2 mm wide, green to brownish or purplish, sometimes silver-white, creeping to erect, forming small mats or cushions, little branched, stem bases with stolons. Stems with a thickwalled cortex. Branches lateral-intercalary. Leaves transverse, alternate, densely imbricate, bifid or emarginate, insertion line extending beyond the dorsal midline of the stem (merophytes “interlocking”), leaf margins entire or crenulate. Cells usually small, ca. 12–25 μm in midleaf, with trigones, cuticle smooth or papillose; oil bodies usually 2–3 per cell, finely granular. Underleaves usually lacking (present in the extralimital Paramomitrion). Rhizoids scattered. Gametoecia on leading shoots. Sporophytes surrounded by tubular bracts, perianth lacking or very small. Setae of numerous cells (cross section). Capsules spherical, wall 2-layered. Vegetative reproduction unknown. About ten genera in cold regions of the world. Gymnomitriaceae are characteristic of páramo. The main family characteristics are (1) worm-like plants with basal stolons; (2) leaves densely imbricate, transverse; (3) leaf insertion lines dorsally

180

 

Keys and Descriptions

interlocking; (4) underleaves usually absent; (5) rhizoids scattered; and (6) perianth or perigynium lacking or very small. The leaf cells are sometimes devoid of chlorophyll, giving the plants a grayish-green or whitish appearance. This treatment follows follows Váňa (2003) and Váňa et al. (2010). 1. Plants minute, filiform (flagelliform), 0.1–0.2(−0.35) mm wide, creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants larger, ascending to erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Cells of leaves and stem epidermis strongly papillose (one large, rounded papilla per cell). Plants usually pale-colored. . . . . . . . . . . . . . Nanomarsupella 2. Cells of leaves and stem epidermis smooth or somewhat mammillose, papillae lacking. Plants purplish to blackish. . . . . . . . . . . . . . . Gymnomitrion atrofilum 3. Leaves concave from base to apex, shallowly bifid to 1/8–1/6(−1/5), with a broad, lunulate sinus and short, blunt tips . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnomitrion truncatoapiculatum 3. Leaves concave only in the lower half, flattened toward the apex, more deeply bifid, sinus rounded to acute (not broadly lunulate) . . . . . . . . . . . . . . . . . . . . 4 4. Plants whitish to pale brown. Leaf margin cells without oil bodies. Perianth lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnomitrion 4. Plants green to brown to reddish, not whitish. Leaf margin cells with oil bodies. Perianth present but very small, hidden between the bracts . . . . . . Marsupella Gymnomitrion Corda Plants usually silver-white or grayish, rarely purplish to blackish (G. atrofilum) or pale brown (G. truncatoapiculatum). Stems ascending to erect, forming dense little cushions. Leaves very densely imbricate, transverse, bifid, or emarginate; leaf tips rounded, acute, apiculate, or piliferous; margins or almost the entire leaf often white. Leaf margin cells without oil bodies. Perianth lacking. About 25 species worldwide, 4 in Colombia and Ecuador. The genus is separated from Marsupella by the complete absence of a perianth. In addition, the small, silverwhitish or grayish cushions are very characteristic of some species of Gymnomitrion. Literature: Schuster (2002), Váňa (1976, 2003), Váňa et al. (2010) 1. Plants minute, filiform, less than 0.2 mm wide. Leaves shallowly bifid to 1/5. Leaf margins crenulate, whitish. . . . . . . . . . . . . . . . . . . . . . . . . . . G. atrofilum 1. Plants larger. Leaves shallowly to rather deeply bifid (to 1/8–2/5). Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves concave from base to apex, shallowly bifid to 1/8–1/6(−1/5) of leaf length, with a broad, lunulate sinus. Leaf tips obtuse to apiculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. truncatoapiculatum 2. Leaves concave in the lower half, ± flattened toward apex, bifid to 1/5–2/5, sinus rounded to acute (not broadly lunulate). Leaf tips apiculate to hair-like 3 3. Leaf apex hair-like, made up of a short row of strongly elongate cells. Cells of leaf lobes 2–5× as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . G. setaceum 3. Leaf apex not hair-like, made up of short row of quadrate cells. Cells of leaf lobes 1.5–2× as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . G. bolivianum

Gymnomitriaceae

181

1. Gymnomitrion atrofilum Váňa On exposed soil and rock in páramo, 3400–4400 m. Colombia: Cun, Met (Váňa 2003). Ecuador: MS, Tu (Cerro Hermoso). General distribution: northern Andes (Venezuela, Colombia, Ecuador). Gymnomitrion atrofilum is a minute páramo species that is recognized by (1) plants less than 0.2 mm wide, filiform, creeping, dark reddish to blackish in color (except for the whitish leaf margins); (2) leaves very small, transverse, erect and appressed to the stem, shallowly bifid to 1/5, with acute tips and whitish margins; and (3) leaf cells small, rounded. 2. Gymnomitrion bolivianum (Steph.) VáňaFig. 19A, B Gymnomitrion andinum (Herzog) Herzog Gymnomitrion andinum R.M.Schust., nom. illeg. Marsupella lacerata (Steph.) Váňa Marsupella subhyalina R.M.Schust. Marsupella trollii Herzog On soil and rock in páramo, 3000–4800 m. Colombia: Boy, Cal, Cun, Mag, Met. Ecuador: Az, MS, Na (páramo de la Virgen). General distribution: Mexico to Bolivia, South Africa. Gymnomitrion bolivianum is a rather common páramo plant that is distinguished by (1) plants grayish-green to whitish to almost hyaline; (2) leaves concave, bifid to 1/5–2/5, made up of short rectangular cells (1.5–2× longer than wide); and (3) leaf lobes apiculate, the tips made up of a short row of quadrate cells. 3. Gymnomitrion setaceum Grolle & VáňaFig. 19C–E On exposed soil and rock in páramo, 3200–4300 m. Colombia: Boy, Cun, Mag, Met; to be expected in Ecuador. General distribution: northern Andes (Venezuela, Colombia), Costa Rica. Gymnomitrion setaceum is characteristic of páramo and readily recognized by the leaf lobes made up of strongly elongate, thick-walled cells (2–5× as long as wide) and with hair-like tips of narrowly elongate cells. The plants usually form dense, silvery to whitish mats. 4. Gymnomitrion truncatoapiculatum Herzog Marsupella truncatoapiculata (Herzog) Váňa Marsupella involuta Váňa

Fig. 19F

On open soil in dry and wet páramo, 3500–4300 m. Colombia: Ara, Boy, Cun, Mag, Met. Ecuador: Ch (Mt. Chimborazo; Váňa 2003). General distribution: Mexico to Bolivia. Gymnomitrion truncatoapiculatum is a characteristic neotropic-alpine species, distinguished by the pale yellowish to pale brown plants without any whitish tinge and leaves deeply concave, shallowly bifid to 1/8–1/6(−1/5), with a broadly lunulate sinus. The leaves sometimes cannot be flattened without tearing the leaf. The species has been considered a member of Marsupella due to its pale-brownish color but is maintained in Gymnomitrion because of the lack of a perianth.

182

 

Keys and Descriptions

Marsupella Dumort. Plants usually growing erect, green, brown, black, reddish-purple. Stems erect, forming small mats, stem bases stoloniform. Leaves transverse, loosely to densely imbricate, erect-spreading, bifid or emarginate, leaf tips rounded or acute. All leaf cells (including margin cells) with oil bodies. Perianth very small, hidden between the bracts. About 20 species worldwide, 2 in Colombia and Ecuador. Marsupella is closely similar to Gymnomitrion, but the leaf margins in Marsupella are not whitish, the margin cells have oil bodies, and the sporophyte is surrounded by a small perianth (perianth absent in Gymnomitrion). Literature: Váňa (2003), Váňa et al. (2010) 1. Leaves suborbicular, shallowly bifid to 1/5, leaf lobes wider than long, sinus between the lobes narrowly rounded. . . . . . . . . . . . . . . . . . . . . M. emarginata 1. Leaves ovate, bifid to 1/4–1/3, leaf lobes longer than wide, sinus (sub)acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. miniata 1. Marsupella emarginata (Ehrh.) Dumort. Marsupella andina J.B.Jack & Steph.

Fig. 19I

Small, green mats on wet, shaded soil along rivulets in shrubby subpáramo, sometimes growing submerged, 3100–3200  m. Colombia (Gradstein and Váňa 1987): Ant, Cun (Guasca). General distribution: widely distributed in temperate regions of the Northern Hemisphere; scattered at high elevation in the tropics (Mexico, Colombia, East Africa, Southeast Asia). This common Holarctic species is recognized by the weakly concave, suborbicular leaves, which are shallowly bifid (to 1/5). The leaf lobes are wider than long and the sinus between the lobes is narrowly rounded. 2. Marsupella miniata (Lindenb. & Gottsche) Grolle Gymnomitrion miniatum Lindenb. & Gottsche On soil in páramo, 2700–4350  m. Colombia: Ara, Cun, Mag. Ecuador: Co (Váňa 2003), Pi (Schäfer-Verwimp et al. 2013a). General distribution: Mexico to Bolivia. Marsupella miniata is a high-montane neotropical species that has been confused with M. emarginata but differs in leaves more ovate in shape, more deeply bifid (to 1/4–1/3), and sinus between the leaf lobes acute, not rounded. The plants grow in dense brownish to reddish mats and the leafy stems are stoloniform at the base. The placement of the species in Marsupella follows Váňa et al. (2010). In the World Checklist (Söderström et al. 2016), the species is placed in Gymnomitrion without explanation.

Herbertaceae

183

Nanomarsupella A.Hagborg, L.Söderstr. & von Konrat Plants minute, filiform, less than 0.2 mm wide, pale green to whitish, gynoecia reddish brown. Stems creeping, often stoloniform, stem cells very thick-walled, epidermis cells each with a large, rounded papilla. Leaves transverse, imbricate, erect and appressed to the stem, shallowly bifid to 1/5, leaf tips obtuse, sinus V-shaped, leaf margins crenulate, dorsal leaf surface strongly papillose. Leaf cells small, thickwalled, each cell with a large, rounded, dome-shaped papilla; oil bodies unknown. Underleaves lacking. Gynoecia capitate, made up of 1–2 pairs of large, swollen bracts. Perianths unknown. A neotropical genus with a single species, characterized by minute plants with flagelliform stems, appressed leaves, and coarsely tuberculate leaf cells. The genus was invalidly described by Schuster (1978), and the names of genus and species were validated by Hagborg et al. (2013). Literature: Schuster (1978, 2002) 1. Nanomarsupella xenophylla A.Hagborg, L.Söderstr. & von KonratFig. 19G, H On soil in rather dry superpáramo, 4200–4300  m. Ecuador: Ch (Mt. Chimborazo), Na (volcán Antisana) (Váňa 2003). General distribution: northern Andes (Ecuador, Venezuela). Nanomarsupella xenophylla is a rare, minute páramo plant that resembles Cephaloziella but differs by the suberect leaves and the coarsely papillose leaf cells, each cell having one large papilla (Gradstein and Costa 2003). Due to its very small size and pale color, the species has probably been overlooked.

Herbertaceae Plants medium-sized to robust, green to brown or reddish-purple, ascending to erect or pendent, sometimes with a rhizome-like creeping base, irregularly branched. Stems rigid, usually with a brown cortex of small, thick-walled cells in 1–3 layers. Branches ventral-intercalary or Frullania-type, the ventral branches often flagelliform. Leaves transverse or incubous, 2–3-lobed, usually long and narrow and asymmetrical with the dorsal segment broader, margins entire or toothed. Cells with small or large trigones; vitta present or absent; oil bodies granular or homogeneous. Underleaves similar to the leaves but more symmetrical. Rhizoids absent or very scarce, in bundles from underleaf bases. Dioicous. Gametoecia on elongate shoots. Antheridia in the axils of both the bracts and the bracteoles. Sporophytes surrounded by a strongly plicate perianth and a fleshy calyptra. Setae of numerous cells (cross section). Capsules spherical, wall 4–7-layered. Vegetative reproduction unknown. A family of three genera worldwide, two in Colombia and Ecuador. The most important characteristics of Herbertaceae are (1) leaves and underleaves 2–3-lobed, underleaves as long as the leaves; (2) leaves transverse or incubous; (3) stems rigid,

184

 

Keys and Descriptions

with a thick-walled, brown cortex; (4) sporophyte surrounded by plicate perianth and a fleshy calyptra; and (5) presence of antheridia in axils of bracts and bracteoles (in most other families, antheridia are only found in the axils of bracts and are lacking in bracteoles). 1. Leaves without vitta, bifid or trifid. Leaf cells thin-walled or uniformly thickwalled, trigones ± lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triandrophyllum 1. Leaves with a broad vitta of elongate cells, bifid. Leaf cells with large trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Herbertus Herbertus Gray Schisma Dumort., Sendtnera Endl. Plants usually robust and usually very long, to 10–30 cm, but sometimes small, green to brown to reddish-brown or purple, ascending to erect or pendent from a rhizome-like base. Leaves transversely inserted but often incubous in position, spreading or falcate-secund, 1.5–7× longer than wide, deeply bifid with lanceolate to linear lobes, the dorsal leaf lobe usually broader than the ventral lobe, leaf tips acute to long-piliferous, sometimes ending in a whitish hairpoint, leaf margins entire or toothed. Cells short along the margins, elongate toward the middle and forming a broad vitta from leaf base to apex; cell walls with very large trigones and intermediate thickenings, cuticle papillose; oil bodies faintly granular or homogeneous, often numerous. Underleaves similar to leaves but more symmetrical, with equal lobes. A genus of 15–20 species worldwide, 5 in Colombia and Ecuador. Herbertus is characterized by deeply bifid leaves and underleaves with a vitta and leaf cells with large trigones. The northern Andean species were revised by Fulford (1963), van Reenen (1982), and Feldberg and Heinrichs et al. (2005, 2006). The latter authors reduced three common Andean species, H. acanthelius, H. bivittatus, and H. pensilis, to subspecies rank under H. juniperoideus. However, these three taxa merit to be treated as varieties or species, not as subspecies, because they do not differ in geographic range or habitat. Since they key out well and have several diagnostic characters (see below), the three taxa are maintained here at the species level. Literature: van Reenen (1982), Feldberg and Heinrichs (2005, 2006) 1. Plants aquatic. Stem leaves only 0.4–0.8  mm long. Leaf apex obtuse, mostly 1-celled. [Plants very small, growing submerged in páramo lake] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . dwarf form of H. sendtneri 1. Plants not aquatic. Stem leaves more than 0.8  mm long. Leaf apex acute to piliferous, ending in a row of 1–20 cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Vitta bifurcating very high up in the lamina, at 3/4–4/5 up the lamina length or directly below the sinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Vitta mostly bifurcating lower down, at 1/4–3/4 up the lamina length . . . . . . 5 3. Bases of leaf lobes mostly overlapping. Leaf tips ending in a long row of 3–10(−20) cells, the upper cells usually hyaline (but tips often broken) . . . . . H. acanthelius

Herbertaceae

185

3. Bases of leaf lobes not overlapping. Leaf tips ending in a short row of 1–3 cells, the upper cells not hyaline. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaves very long and slender, undivided lamina 1.5–3× longer than wide, narrowly oblong (to be measured on flattened lamina) . . . . . . . . . . . . . . H. pensilis 4. Leaves shorter, undivided lamina 1–1.5× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. juniperoideus 5. Leaf tips ending in a long row of 3–15(−20) cells, the upper cells usually hyaline (but tips often broken). Vitta convex, bifurcating at 1/2–3/4 up to the lamina length, ending below the leaf tips. Flagelliform branches frequent. Leaves mostly falcate-secund, occasionally wide-spreading. Plants very common in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. sendtneri 5. Leaf tips ending in a short row of 1–3 cells, the upper cells not hyaline. Vitta flat, bifurcating very low in the lamina (at 1/4–1/2 up the lamina length), reaching the leaf tips. Flagelliform branches lacking or scarce. Leaves mostly erectspreading. Plants growing below 3000 m, not in páramo . . . . . . . H. bivittatus 1. Herbertus acanthelius Spruce Fig. 18A, B Herbertus juniperoideus subsp. acanthelius (Spruce) K.Feldberg & Heinrichs Herbertus inaequalis Herzog Herbertus limbatus (Steph.) Herzog Herbertus serratus Spruce Schisma wallisianum Steph. On bark and rock in upper montane cloud forest and humid páramo, 2400–4300 m. Colombia: Ant, Ara, Boy, Cau, Cho, Cun, Mag, Met, Nar, Qui, Ris, San, Tol, Val. Ecuador: Az, Ca, Ch, Co, Lo, MS, Pi, Tu, Lo, ZC. General distribution: tropical Andes, Costa Rica, Dominican Republic. Herbertus acanthelius is a common Andean species that is distinguished by the usually overlapping bases of leaf lobes (in the sinus) and the long, often hair-like leaf tips, ending in a row of 3–10 hyaline, elongate cells (but leaf tips very fragile, often broken). Moreover, the vitta bifurcates high up in the lamina, at 3/4 the leaf length to near the sinus. The leaves are usually falcate-secund, occasionally erect-spreading. Long, hair-like leaf tips occur also in H. sendtneri and these two species may be confused. For differences see under H. sendtneri. 2. Herbertus bivittatus Spruce Fig. 18C Herbertus juniperoideus subsp. bivittatus (Spruce) K.Feldberg & Heinrichs Herbertus angustevittatus (Steph.) Fulford Herbertus divergens (Steph.) Herzog On bark in lowland and montane rainforests, from sea level to 3000 m. Colombia: Ant, Cho, Mag, Ris. Ecuador: Ga, Lo, ZC.  General distribution: tropical America. Herbertus bivittatus is a widespread neotropical-montane species that is recognized by the narrow vitta, which bifurcates near the base of leaves. A rather lowbifurcating vitta is otherwise only seen in H. sendtneri; differences between the two species are given in the key. Herbertus bivittatus is the only species in the genus that descends to elevations below 1000  m on the continent. In the Chocó the species occurs abundantly on canopy branches in everwet coastal rainforests at sea level

186

 

Keys and Descriptions

Fig. 18 (A and B) Herbertus acanthelius. (A) Leaf. (B) Apex of leaf lobe, 350×. (C) Herbertus bivittatus, leaf, 18×. (D and E) Herbertus sendtneri. (D) Leaf. (E) Underleaf (scale as in D). (F) Herbertus pensilis, leaf, 18×. (G–J) Herbertus juniperoides. (G) Leaf, 18×. (H) Apex of leaf lobe, 350×. (I) Cells of vitta. (J) Cells near leaf margin. (K) Triandrophyllum eophyllum, habit in ventral view. (L–O) Triandrophyllum subtrifidum. (L) Habit, dorsal view. (M) Leaf. (N) Underleaf. (O) Midleaf cells. (A, D, E redrawn from van Reenen 1982; B, C, F–H from Fulford 1963; K redrawn from Schuster 2002; rest from Gradstein et al. 2001)

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187

(Gradstein 1995); on the Galápagos Islands, it grows at 500–900 m together with H. pensilis. 3. Herbertus juniperoideus (Sw.) Grolle Herbertus juniperinus (Sw.) Spruce Herbertus subdentatus (Steph.) Fulford (fide Hodgetts 2003)

Fig. 18G–J

On bark, rock, and soil in montane forests and páramo, 1000–3750 m. Colombia: Ant, Ara, Boy, Cau, Cun, Hui, Mag, Nar, Ris, Val. Ecuador: At, Pa, Pi, Tu, ZC. General distribution: tropical America, Africa. Herbertus juniperoideus is a widespread species that is distinguished by the short leaf tips made up of a row of 1–3 short cells (elongate hyaline cells lacking), the rather broad leaf lamina (1–1.5 × longer than wide), the leaf lobes not overlapping at the sinus, and the broad vitta bifurcating very high up the lamina, just below the sinus. The species is close to H. acanthelius but is readily separated from the latter by the short leaf tips and the leaf lobes not overlapping in the sinus. Plants of H. acanthelius with broken leaf tips may be confused with H. juniperoideus. 4. Herbertus pensilis (Taylor) Spruce Fig. 18F Herbertus juniperoideus subsp. pensilis (Spruce) K.Feldberg & Heinrichs Herbertus commutatus (Steph.) Pears. Herbertus grandifolius Steph. On bark in montane forest and shrubby páramo, 1500–4000 m; on the Galápagos Islands at much lower elevation, ca. 750 m. Colombia: Ant, Boy, Cau, Cho, Cun, Hui, Mag, Nar, Qui, Ris, Tol, Val. Ecuador: Ch, Ga, Lo, Na, Pi, ZC.  General distribution: tropical America. Herbertus pensilis is a characteristic neotropical species recognized by the long and narrow leaves with a narrowly oblong lamina, 1.5–3× longer than wide (without the leaf lobes), and a very broad vitta that bifurcates just below the sinus. 5. Herbertus sendtneri (Nees) Lindb. Fig. 18D, E Herbertus colombianus G.Reenen, syn. nov. Type: Colombia, Arauca, Sierra Nevada del Cocuy, Quebrada El Playón, subpáramo, 3255 m, 10 Jun 1973, Cleef 10131 (isotype U!) Herbertus grossispinus (Steph.) Fulford Herbertus oblongifolius (Steph.) Gradst. & Cleef Herbertus subdentatus auct. (non typus) Herbertus subnivalis S.Winkl. Schisma columbiae Steph. Schisma granatense Steph. Schisma striolatum Steph. On bark, soil, and rock in upper montane cloud forest and humid páramo, 2150–4350 m. Colombia: Ant, Boy, Cau, Cho, Cun, Hui, Mag, Met, Ris, Tol, Val. Ecuador: Ca, Co, Lo, Pi, ZC. General distribution: at high elevations in the tropics, central Europe, and Himalayas.

188

 

Keys and Descriptions

Herbertus sendtneri is a very widespread species that is distinguished by (1) frequent presence of microphyllous ventral branches; (2) leaves more or less falcate-secund to circinate (sometimes erect-spreading in very small plants); (3) leaf tips usually hyaline to whitish and hair-like, terminating in a uniseriate row of (2–)5–20 quadrate to narrow-rectangular cells, (occasionally, in very small phenotypes, hair-like tips absent and the leaf apex is only 1–2 cells long); (4) leaf bases usually with one or more ciliate or laciniate appendages crowned by hyaline papillae; and (5) vitta convex, bifurcating at about 1/3–3/4(−5/6) up the lamina and ending before the leaf apex. Fulford (1963) and van Reenen (1982) described the neotropical populations of H. sendtneri as H. subdentatus and H. grossispinus, but based on molecular evidence, Feldberg et  al. (2004) showed that they belong to H. sendtneri, a species originally described from the European Alps. Herbertus sendtneri varies considerably in size and leaf shape and may be confused with H. acanthelius, which also has hair-like, whitish leaf tips. In H. acanthelius, however, the leaf lobes are usually overlapping at the base in the sinus (not overlapping in H. sendtneri); the vitta bifurcates very high up in the lamina, just below the sinus; and microphyllous branches are scarce or absent. Herbertus colombianus, described from the Sierra Nevada del Cocuy, is a phenotype of H. sendtneri with undulate leaf margins. Feldberg and Heinrichs (2006) associated this plant with H. juniperoideus s.l., but the piliferous leaf tips place it in H. sendtneri, which is extremely common and variable in the Cocuy. A dwarf phenotype of H. sendtneri with obtuse, 1-celled leaf tips grows in great abundance in a glacial lake in the Sierra Nevada del Cocuy at 4120 m. The aquatic plants were originally identified as H. oblongifolius (Gradstein et  al. 1977), but molecular analysis showed that they belong to H. sendtneri (Gradstein et al. 2018d). Triandrophyllum Fulford & Hatcher Olgantha R.M.Schust. Plants medium-sized, to 6 cm long, green to yellowish-brown, occasionally reddish, creeping or ascending, sometimes arising from a long rhizome-like base, irregularly branched. Leaves incubous, spreading and somewhat reflexed, asymmetrical, often somewhat falcate-secund, shallowly trifid or bifid (to maximally 1/4–1/2 of leaf length) with unequal to subequal, triangular lobes and obtuse to acute-acuminate tips, leaf margins entire or with a few small teeth. Cells small, quadrate to subrectangular, thin-walled or uniformly thick-walled, ± without trigones, vitta lacking, cuticle smooth or finely striate-papillose; oil bodies finely granular. Underleaves similar to leaves but more symmetrical. A southern-temperate and tropical-montane genus (4 spp.), two species in Colombia and Ecuador. Triandrophyllum is distinguished by the rather shallowly bifid or trifid leaves and underleaves with rather thin-walled cells without trigones and no vitta.

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Literature: Fulford (1963), Schuster (2000a), Bednarek-Ochyra et  al. (2000), Gradstein et al. (2001) 1. Leaves very shallowly bifid (rarely trifid) to ca. 1/10 of leaf length, densely imbricate and deeply concave. Plants found in superpáramo of Ecuador, very rare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. eophyllum 1. Leaves more deeply bifid or trifid, to 1/8–1/4(−1/2) of leaf length, weakly to densely imbricate, concave. Plants common in the high Andes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. subtrifidum 1. Triandrophyllum eophyllum (R.M.Schust.) Gradst.  Olgantha eophylla R.M.Schust.

Fig. 19K

On soil in superpáramo, 4250–4300 m. Ecuador: Na/Pi (summit of páramo de la Virgen; Schuster 1996a; Benitez et  al. 2012). General distribution: only known from the type locality in Ecuador. Triandrophyllum eophyllum is a rare Ecuadorian endemic known from superpáramo east of Quito where it was discovered by R.  M. Schuster in 1993 and described as the new genus Olgantha (named after Schuster’s wife, Olga). Gradstein et al. (2001) subsequently transferred Olgantha to Triandrophyllum. Triandrophyllum eophyllum is very similar in general apparance to T. subtrifidum, which grows together with T. eophyllum in the type locality, but differs from T. subtrifidum in the more swollen and somewhat more robust habit, the more deeply concave leaves which are about as wide as long (usually wider than long in T. subtrifidum) and densely imbricate and impossible to flatten without rupturing the apex, and, especially, the very shallowly bifid leaf apex (bifid to ca. 1/10 of leaf length; rarely trifid). 2. Triandrophyllum subtrifidum (Hook.f. & Taylor) Fulford & Hatcher  Fig. 18L–O Isotachis ripensis Spruce Triandrophyllum subtrifidum var. trifidum (Gottsche) Solari On moist soil and rock along rivers in high-montane environments, 1800–4500 m. Colombia: Boy, Cau, Cal, Ces, Cun, Hui, Mag, Met, Nar, Qui, Ris. Ecuador: Ca, Co, Lo, Pi, ZC. General distribution: widespread in temperate regions of the Southern Hemisphere and on high mountains of tropical America and tropical Asia. Triandrophyllum subtrifidum is a widespread Southern-Hemispheric species that is recognized by the green to greenish-brown to glossy yellowish-brown plants (rarely reddish) with leaves and underleaves in three equal ranks and the incubous, ovate-secund leaves which are bifid or trifid to 1/8–1/4(−1/2) of leaf length and made up of small, (sub)quadrate, thin-walled to uniformly thick-walled cells. The leaf margins are entire or somewhat toothed. Plants growing on naked earth at high elevation sometimes produce long, almost leafless flagelliform shoots giving rise at the tips to short, densely leaved shoots. Triandrophyllum subtrifidum varies considerably in size: robust plants have mostly trifid leaves with sharply acute leaf apices, while small plants have purely bifid leaves with obtuse leaf apices. The small, bifid form seems characteristic of very high elevations, above 3900 m. Two variaties have been recognized based on

190

 

Keys and Descriptions

Fig. 19 (A and B) Gymnomitrion bolivianum. (A) Habit, dorsal view. (B) Leaf. (C–E) Gymnomitrion setaceum. (C) Habit. (D) Leaf. (E) Midleaf cells. (F) Gymnomitrion truncatoapiculatum, habit in dorsal view, 15×. (G and H) Nanomarsupella xenophylla. (G) Habit. (H) Leaf lamina in cross section. (I) Marsupella emarginata, habit in dorsal view. (J and K) Syzygiella anomala. (J) Habit, dorsal view. (K) Midleaf cells. (L and M) Syzygiella autumnalis. (L) Habit with male spike, dorsal view. (M) Habit, ventral view. (N) Syzygiella bilobata, leaf, 10×. (O and P) Syzygiella concreta. (O) Habit, ventral view, 18×. (P) Leaves, 18×. (F redrawn from Váňa 1976; G, H redrawn from Schuster 2002; I from Gradstein and van Melick 1996; L, M redrawn from Paton 1999; N–P redrawn from Inoue 1966: rest from Gradstein et al. 2001)

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this variation, var. subtrifidum with mostly bifid leaves and var. trifidum (Gottsche) Solari with mostly trifid leaves; Fulford (1963) treated the two varieties as different species. The two varieties have received little attention, probably because a key to the two taxa is lacking in Fulford’s manual. A study of the variation in T. subtrifidum in the northern Andes, based on a broad sampling and using molecular and morphological evidence, should be undertaken and would make an excellent subject for a small thesis. Such a study should also include T. eophyllum. Plants with purely bifid leaves may be confused with Isotachis. The latter genus, however, is never yellowish-brown in color, the leaves are transverse or succucous (not incubous), and the leaf cells are clearly elongate, 2–3× as long as wide (subquadrate in Triandrophyllum).

Jamesoniellaceae (Adelanthaceae subfam. Jamesoniellaceae, Jungermanniaceae p.p.) A small family of six or seven genera; contains only the genus Syzygiella in Colombia and Ecuador. The family is sister to Adelanthaceae and is also considered a subfamily of the latter (see under Adelanthaceae). The main features of Jamesoniellaceae are (1) stems usually rigid, with a thick-walled cortex; (2) branching terminal and intercalary; (3) leaves ± undivided, succubous to subtransverse, alternate or opposite, laterally appressed or spreading; (4) dorsal leaf margins plane or recurved; (5) underleaves absent or small; (6) rhizoids usually scattered, scarce; (7) plants dioicous, gynoecia terminal on elongate stems; (8) perianth inflated, plicate; (9) capsule wall 4–6-layered; and (10) gemmae lacking. Syzygiella Spruce Cryptochila R.M.Schust., Jamesoniella (Spruce) Lees Plants small to large, 1–10 cm long and 1–6 mm wide, green, yellowish, reddish, purplish, brown, or black, nearly always with reddish or purplish pigmentation, creeping to erect with decurved shoot apex, little branched, basal stolons usually lacking (present in subg. Cryptochila). Stems rigid. Branches terminal and ventralintercalary. Leaves succubous or nearly transverse, alternate or opposite (leaf bases free or connected), laterally appressed or wide spreading, ± undivided, apex rounded to truncate to shallowly bidentate, margins entire or with a few teeth, dorsal margin plane or recurved, dorsal base decurrent or not. Cells with or without trigones, cuticle smooth or papillose, papillae rounded or elongate; oil bodies finely granular. Underleaves lacking or very small (rarely large), subulate. Rhizoids usually scarce, scattered (occasionally in bundles), only present on creeping shoots. Dioicous. Perianths on elongate stems, inflated, plicate, the folds straight, rarely spirally twisted above (subg. Cryptochila). Capsule walls 4–5-layered. Gemmae lacking. Widespread in tropical mountains and in temperate regions of the Southern Hemisphere (ca. 35 spp.); 18 species in Colombia and Ecuador, in 3 subgenera,

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Keys and Descriptions

subg. Anomalae K.Feldberg et  al., subg. Cryptochila (R.M.Schust.) K.Feldberg et al., and subg. Syzygiella (Feldberg et al. 2010a). Syzygiella is recognized by the rather robust, frequent reddish or purplish plants with opposite or alternate, succubous leaves, usually reduced underleaves (present in S. uleana and, rarely, in S. anomala), scattered rhizoids, gynoecia at the tip of the main stem or on long branches, and an inflated, plicate perianth. In plants with opposite leaves, the ventral and dorsal leaf bases are touching each and sometimes broadly connate (rarely the dorsal leaf bases free). The cuticle may be smooth or papillose; in the latter case, the papillae are elongate (striate-papillose) or, less frequently, rounded. Rounded papillae occur in S. campanulata, S. rubricaulis, and S. setulosa. Syzygiella includes the former genera Cryptochila and Jamesoniella, which were united with Syzygiella based on molecular evidence (Feldberg et al. 2010b). Some species have a recurved and decurrent dorsal leaf margin and resemble Plagiochila, but the usually somewhat reddish or purplish color of these plants and the inflated perianth readily separate them from Plagiochila. Literature: Grolle (1971), Inoue (1966), Feldberg et al. (2010a, b), Váňa et al. (2014a, b) 1. Underleaves present, large (at least 1/2 the length of the leaves), narrowly lanceolate or ligulate, occasionally bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaves absent or very small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Cuticle smooth. Leaves fragile. Underleaves regularly developed. Brazilian species, found once in southern Ecuador . . . . . . . . . . . . . . . . . . . . . . S. uleana 2. Cuticle papillose. Leaves not fragile. Underleaves irregularly developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. anomala 3. Ventral leaf bases alternate. Leaves ± rounded . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Ventral leaf bases opposite [when in doubt try both leads]. Leaves rounded to elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 4. Plants creeping on rotten wood and peaty soil, very rare. Leaves clearly succucous, ± horizontally spreading (at least in part), leaf bases not or scarcely decurrent. Leaf cells thin-walled and with very small but distinct trigones, cuticle smooth. Underleaves present, very small, lanceolat . . . . . . . S. autumnalis 4. Plants ascending to erect, common in páramo and the upper montane belt. Leaves subtransverse, not horizontally spreading, leaf bases decurrent. Leaf cells with large bulging trigones or ± uniformly thickened, cuticle smooth or papillose. Underleaves absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaf cells quadrate, smooth, walls ± evenly thickened or thin, trigones not well defined. Stem bases with stolons. Plants green to dark brown to blackish, often growing in or near water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sonderi 5. Leaf cells ± rounded, papillose, with large bulging trigones. Stolons absent. Plants yellowish-green to orange-brown to reddish-brown, not growing near water. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Dorsal leaf bases longly decurrent and  ±  undulate, completely covering the stem. Plants rare, in páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. undata

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6. Dorsal leaf bases shortly decurrent, not undulate, not completely covering the stem. Plants very common, in páramo and the upper montane belt . . . . . . S. rubricaulis 7. Leaves ovate-triangular, usually tapering to a narrow apex. Dorsal leaf margin distinctly decurrent (like in Plagiochila). Leaf apex entire, toothed or bilobed. Plants usually robust, ca. 3–5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Leaves rounded to ovate-oblong, not tapering to a narrow apex. Dorsal leaf margin not or shortly decurrent. Leaf apex broadly rounded, entire. Plants smaller, 1–3 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 8. Leaf apex with a 2–3(−5) very long, whitish, hair-like teeth made up of 5–10 cells in a row. Cuticle densely striate-papillose. . . . . . . . . . . . . . . . . . S. ciliata 8. Teeth at leaf apex shorter or absent. Cuticle smooth or striate-papillose . . . . 9 9. Cuticle smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Cuticle papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 10. Leaf apex with 2–5 distinct, ciliate teeth. . . . . . . . . . . . . . . . . . . . S. tonduzana 10. Leaf apex entire or with 1–2 small, 1(−2) cell long teeth. . . . . . . . . . . . . . . 11 11. Dorsal leaf bases free. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. trigonifolia 11. Dorsal leaf bases connected. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. manca 12. Leaf apex deeply bilobed, lobes acuminate, sinus between the lobes narrow. Auricle at ventral leaf base very large, squarrose. Plants rare. . . . . S. bilobata 12. Leaf apex entire, shallowly bilobed or with 2 teeth, sinus between the lobes or teeth wide. Auricle at ventral leaf base small, not squarrose. . . . . . . . . . . . . 13 13. Leaves 1.5–2.5× longer than wide, often falcate. Leaf apex with 2 sharp teeth. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pectiniformis 13. Leaves 1–1.5× longer than wide, not falcate. Leaf apex entire or shallowly bilobed with obtuse to subacute lobes. Plants common. . . . . . . . . . S. anomala 14. Leaves 1.3–2× longer than wide, ovate-oblong. . . . . . . . . . . . . . . . . . . . . . . 15 14. Leaves 0.7–1.2× longer than wide, ovate to suborbicular to reniform . . . . . 16 15. Cuticle papillose, papillae rounded. Trigones large, ± bulging and sometimes confluent. Underleaves lacking. Female bracts toothed. . . . . . . . . S. perfoliata 15. Cuticle smooth or slightly striate-papillose, papillae narrowly elongate. Trigones small, not bulging and not confluent. Rudimentary underleaves frequently present, filiform, attached to the leaf base. Female bracts entire . . . . . . . S. concreta 16. Leaves reniform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. burghardtii 16. Leaves orbicular or longer than wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. Cuticle smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. contigua 17. Cuticle papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18. Papillae large, 3–10 papillae over the lumen of each cell in the upper half of the leaf. Female bracts toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. setulosa 18. Papillae small, 10–20 papillae over the lumen of each cell in the upper half of the leaf. Female bracts entire. . . . . . . . . . . . . . . . . . . . . . . . . . . S. campanulata 1. Syzygiella anomala (Lindenb. & Gottsche) Steph.

Fig. 19J, K

On bark, moist rock, and soil in montane forests and páramo, 1500–3600  m. Colombia: Boy, Cal, Cun, Hui, Mag, Nar, Qui, Ris, Tol. Ecuador: Az, Lo, MS, Pi, ZC. General distribution: Central America, tropical Andes, southeastern Brazil.

194

 

Keys and Descriptions

Syzygiella anomala (subg. Anomalae) is a common neotropical species that is distinguished by (1) plants robust, usually violet-colored; (2) leaves ovate-triangular, plagiochiloid, with recurved dorsal leaf margin and decurrent dorsal base, 1–1.5× longer than wide, opposite with opposite leaf bases connected both dorsally and ventrally; (3) leaf apex shallowly and asymmetrically bilobed with obtuse to subacute lobes or undivided; and (4) cuticle striate-papillose. Exeptionally, this species may produce underleaves (seen in a specimen from Colombia: Quindio, páramo El Campanario, 3600 m, O. Ruiz s.n., HUQ). The underleaves are large, about half the length of the leaves, undivided or short bifid, and free or connate with leaves on one side. 2. Syzygiella autumnalis (DC.) K.Feldberg et al.  Jamesoniella autumnalis (DC.) Steph.

Fig. 19L, M

On rotten wood and peat in páramo, (2300–)3300–3700  m; the low elevation record is from azonal páramo bog near Hacienda Merenberg, Huila. Colombia: Boy, Cun, Hui. General distribution: widespread in temperate regions of the Northern Hemisphere, scattered in the mountains of tropical America where it may have been introduced (Gradstein and Váňa 1987). A common Holarctic species (subg. Syzygiella) found a few times at high elevation in the neotropics and characterized by (1) plants small, 1–2 mm wide, ± creeping, yellowish-brown to olive brown, rarely wine-red; (2) leaves alternate, ovate-orbicular, at least in part widely spreading, somewhat convex, bases scarcely decurrent; (3) leaf cells with small trigones and a smooth cuticle; and (4) minute underleaves, filiform to lanceolate, ca. 3–10 cells long. 3. Syzygiella bilobata Inoue

Fig. 19N

On soil in lower montane forest, 1780 m. Ecuador: ZC (Reserva Biologica San Francisco; Schäfer-Verwimp et  al. 2013a). General distribution: Costa Rica, Panama, southern Ecuador, Peru. Syzygiella bilobata (subg. Anomalae) is a rare neotropical species that has been recorded once from southern Ecuador. It is recognized by the plagiochiloid, opposite leaves that are distinctly longer than wide, connected both dorsally and ventrally, and deeply bilobed at the apex, with two acuminate-triangular lobes separated by a narrow sinus. A large, squarrose auricle is present at the ventral base. The leaf cells have conspicuous trigones and the cuticle is striate-papillose. 4. Syzygiella burghardtii Gradst. & A.R.Benitez On shrubs in mossy elfin forest near the forest line, ca. 2900–3200 m. Ecuador: Lo/ZC (pass El Tiro; Gradstein and Benitez 2017), ZC (páramo de Yacuambi, Benitez & Gradstein s.n., HUTPL). General distribution: only known from southern Ecuador. The southern Ecuadorian endemic S. burghardtii (subg. Syzygiella) resembles Syzygiella setulosa in having transverse, laterally appressed, opposite leaves, leaf cells with large trigones, and, especially, perianths with a whitish beak, seen only in these two species (J.  Váňa, pers. comm.). Syzygiella burghardtii differs from S.

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setulosa in the following characters: (1) leaves reniform, 1.4× wider than long (1.0–1.2× wider than long in S. setulosa); (2) leaf cells 30–40 μm long in midleaf (ca. 25 μm in S. setulosa); (3) cuticle very faintly verruculose, almost smooth (cuticle with large, rounded papillae in S. setulosa); and (4) female bracts fully entire, similar to leaves (toothed in S. setulosa). Reniform leaves are unusual in neotropical Syzygiella and are otherwise only seen in S. renifolia Gradst. & D.P.Costa (subg. Cryptochila) from Brazil. 5. Syzygiella campanulata Herzog On tree trunks and twigs in upper montane forest and páramo, 2700–3700 m. Colombia: Mag, Ris, Tol. Ecuador: Lo (Feldberg et al. 2010a). General distribution: northern Andes, Costa Rica, Dominican Republic. Syzygiella campanulata (subg. Syzygiella) resembles S. contigua but differs from the latter by the papillose cuticle, with 10–20(−30) small, rounded papillae on the lumen of each cell. Syzygiella campanulata is also quite similar to S. setulosa, but the papillae in the latter species species are larger and fewer per cell and the female bracts are toothed (see key). Syzygiella riclefii Pócs from the Venezuelan Andes is close to S. campanulata, and Váňa et al. (2014a, b) treated the two as conspecific. However, according to Dr. T.  Pócs (pers. comm.), S. riclefii is clearly different from S. campanulata in its highly fragile leaves and leaf cells with hyaline walls and darkish pigmented lumen (leaf cells with darkish walls and hyaline lumen in S. campanulata; see also Pócs 2005). 6. Syzygiella ciliata Gradst. & A.R.Benitez On moist humus in upper montane forest and scrub in rather open situations, 2000–2600 m. Ecuador: Pi (Holm-Nielsen et al. 24779, AAU), ZC (Cordillera del Cóndor: Gradstein and Benitez 2014b). General distribution: only known from Ecuador. Syzygiella ciliata (subg. Anomalae) is thus far known from two localities in the Ecuadorian Andes. It differs from all other species in the genus by the leaf apex with 2–3 very long, whitish, hair-like teeth made up of a row of 5–10 cells. In addition, a few shorter, ciliate teeth (to 5 cells long) may also be present. The leaves are plagiochiloid, opposite and connected both dorsally and ventrally, the stems are flattened and with a cortex of 2–3 cell layers, and the cuticle is densely striate-papillose. 7. Syzygiella concreta (Gottsche) Spruce Syzygiella grollei Inoue

Fig. 19O, P

On bark, rock, and soil in montane forests and shrubby páramo, 1500–3200 m. Colombia: Cas, Cun. Ecuador: Lo, ZC.  General distribution: scattered in tropical America; tropical Africa. Syzygiella concreta (subg. Syzygiella) is a rare neotropical species distinguished by broadly ovate to ovate-oblong, opposite leaves (connate both dorsally and ventrally) with small trigones and a smooth or slightly striate-papillose cuticle (papillae narrowly elongate). The plants are rather robust, to 5 cm long and 3 mm wide, pale green to yellowish-green in color, and the ventral leaf bases are often broadly

196

 

Keys and Descriptions

connate (across 3–5 cells). Very small underleaves, connected to the ventral leaf bases, are usually present. The female bracts are mostly entire. Syzygiella concreta is very similar to S. perfoliata, and morphologically intermediate populations occur, even though being genetically distinct (Feldberg et  al. 2012); for differences between these two species, see under S. perfoliata. 8. Syzygiella contigua Steph. Syzygiella integerrima Steph.

Fig. 20A, B

On bark, rock, and soil in upper montane forest and shrubby páramo, 2900–3600 m. Colombia: Cun, Hui, Ris, San. Ecuador: Lo, ZC. General distribution: scattered in tropical America Syzygiella contigua (subg. Syzygiella) is an uncommon neotropical species that is recognized by the suborbicular, opposite leaves, leaf cells with large trigones and a smooth cuticle, and entire female bracts. The leaves are connate both dorsally and ventrally. 9. Syzygiella manca (Mont.) Steph. Syzygiella colombiana H.Rob. Syzygiella geminifolia (Mitt.) Steph. Syzygiella plagiochiloides Spruce

Fig. 20C, D

On tree trunks, humic soil, and rock along trails in montane forests, in Polylepis forest, and in shrubby páramo, 1500–4000  m. Colombia: Ris, Qui (páramo El Campanario, Gradstein et al. s.n., HUQ), Val (as S. colombiana). Ecuador: Az, Im, Pi, Tu, ZC. General distribution: tropical Andes (Venezuela to Peru), Guayana Highland, tropical Africa. Syzygiella manca (subg. Anomalae) is a characteristic robust, greenish to violetcolored plant with opposite, plagiochiloid leaves that are connate both dorsally and ventrally, as long as wide or longer than wide and the apex entire or with 1–2 small, 1(−2) cell long teeth. The cuticle is smooth. Syzygiella manca is morphologically close to S. tonduzana, but in the latter species, the leaf apices are more distinctly toothed, with 2–5 teeth. Syzygiella manca also resembles S. anomala, but in the latter species, the cuticle is distinctly papillose. 10. Syzygiella pectiniformis Spruce 

Fig. 20E

On moist, humic soil, and rock, sometimes in dripping water, 500–2300  m. Colombia: Hui (A.M. Cleef s.n., U). Ecuador: MS, Na, Pi, Tu, ZC. General distribution: northern Andes (Colombia, Ecuador, Peru), Costa Rica. Syzygiella pectiniformis (subg. Anomalae) is a robust northern Andean species (to 8 cm long and 5 mm wide) with plagiochiloid, opposite leaves that are conspicuously elongate (ca. 1.5–2.5× longer than wide), often falcate, and with 2 sharp teeth at the apex separated by a wide sinus. The leaves are connate dorsally and ventrally, or dorsally free, and the cuticle is striate-papillose. Syzygiella pectiniformis is morphologically close to S. anomala, but the leaves in the latter species are less elongate (1–1.5× longer than wide), not or little falcate, and the apex is entire or shallowly bilobed with obtuse to subacute lobes.

Jamesoniellaceae

197

Fig. 20 (A and B) Syzygiella contigua. (A) Habit, dorsal view, 16×. (B) Connate female bracts and bracteole, 16×. (C and D) Syzygiella manca. (C) Habit, dorsal view, 10×. (D) Leaf, 10×. (E) Syzygiella pectiniformis, habit in dorsal view, 10×. (F) Syzygiella perfoliata, habit in dorsal view, 16×. (G and H). Syzygiella rubricaulis. (G) Habit with gynoecium, lateral view. (H) Midleaf cells. (I and J) Syzygiella sonderi. (I) Habit with gynoecium, lateral view. (J) Midleaf cells. (K) Syzygiella setulosa, habit with gynoecium in lateral view, 10×. (L) Syzygiella tonduzana, leaf, 10×. (M and N) Syzygiella uleana. (M) Habit, dorsal view, 10×. (N) Underleaf, 40×. (O) Syzygiella undata, habit in lateral view, showing undulate-decurrent dorsal leaf bases, 16×. (A–F, K–N redrawn from Inoue 1966; rest from Gradstein et al. 2001)

198

 

Keys and Descriptions

11. Syzygiella perfoliata (Sw.) Spruce Syzygiella linguifolia Schiffn. On bark, rock, and rotten wood in montane forests, 1500–2750 m. Colombia: Boy, Cun, Mag, Nar, Ris, San. Ecuador: ZC (Nöske et al. 2003). General distribution: widespread in tropical America. Syzygiella perfoliata (subg. Syzygiella) is a common and variable neotropical species that is characterized by (1) green to reddish plants with ovate-oblong to lingulate, opposite leaves that are 1.3–2× longer than wide and connected both dorsally and ventrally; (2) leaf cells with large, bulging trigones (trigones sometimes confluent); (4) cuticle densely papillose, with rounded papillae; (5) underleaves absent; and (6) female bracts coarsely toothed. The paucity of records of the species from Ecuador is surprising. Syzygiella perfoliata closely resembles S. concreta, but the latter can usually be separated from S. perfoliata by smaller trigones that are not strongly bulging and never confluent, a smooth or finely striate-papillose cuticle with narrowly elongate papillae (rounded in S. perfoliata), and the presence of minute, filiform underleaves (Feldberg et al. 2012). 12. Syzygiella rubricaulis (Nees) Steph. Jamesoniella rubricaulis (Nees) Grolle Jamesoniella cuneifolia Steph. Syzygiella oppositifolia Spruce

Fig. 20G, H

Common on exposed rock and soil in humid open environments, especially on road banks and in páramo, also on canopy branches in montane forests, 1500–4500 m; in the Galápagos Islands at much lower elevation (650–800 m) at the summit of San Cristóbal, on a vertical crater wall. Colombia: Ant, Ara, Boy, Cas, Cau, Ces, Cun, Hui, Mag, Met, Nar, Qui, Ris, San, Tol. Ecuador: Az, Ca, Cañ, Co, Ga, Lo, Na, Tu, ZC. General distribution: high mountains of tropical America from Mexico to Bolivia; Azores. Syzygiella rubricaulis (subg. Syzygiella) is a common páramo species that is readily distinguished by (1) plants growing upright, brownish-green to yellowish or orange; (2) leaves subtransverse and usually laterally appressed, orbicular, subopposite, leaf bases free but sometimes approaching each other; (3) leaf cells with huge, confluent trigones and rounded papillae; and (4) female bracts and bracteole toothed. 13. Syzygiella setulosa Steph. Jamesoniella rotundifolia Steph. On shrubs, rock, and soil in shrubby páramo, 2900–3700 m. Colombia: Mag (Winkler 1976 as Jamesoniella rotundifolia). Ecuador: Az, Lo, ZC (SchäferVerwimp et al. 2013a). General distribution: tropical Andes. Syzygiella setulosa (subg. Syzygiella) is a common Bolivian species that has recently been collected a few times in southern Ecuador (Váňa et al. 2014a, b). The species is recognized by (1) plants rather small, to 2 mm wide, prostrate to ascending, pale brown to dark violet; (2) leaves subtransverse, not spreading, suborbicular,

Jamesoniellaceae

199

as wide as long or slightly wider than long, opposite, leaf bases connected dorsally and ventrally; (3) leaf cells with large, swollen trigones and cuticle with large, rounded papillae, 3–10 papillae over the lumen of each cell in the upper half of the leaf; (4) female bracts and bracteole irregularly lobed-toothed; and (5) upper part of the perianth contracted to a whitish beak. In its subtransverse, ± orbicular leaves and leaf cells with huge trigones and large, rounded papillae, S. setulosa is similar to S. rubricaulis, but in the latter species, the leaf bases are free. Syzygiella setulosa also resembles S. campanulata and S. burghardtii, but S. campanulata has smaller papillae and entire female bracts, whereas S. burghardtii has reniform leaves, larger leaf cells, very small papillae, and entire female bracts. 14. Syzygiella sonderi (Gottsche) K.Feldberg et al. Cryptochila grandiflora (Lindenb. & Gottsche) Grolle Jamesoniella allionii Steph.

Fig. 20I, J

On humid rock and on wet soil at high elevation, often near running water or in shallow lakes in páramo, 2000–4300 m. Colombia: Boy, Cas, Ces, Cun, Mag, Nar, Ris, San, Val. Ecuador: Az, MS, Pi, Tu, ZC. General distribution: temperate regions of the Southern Hemisphere and on high tropical mountains, especially in tropical America. Syzygiella sonderi (subg. Cryptochila) was long called Cryptochila grandiflora, but molecular evidence showed that Cryptochila is nested in Syzygiella (Feldberg et al. 2010b). The species is widespread in the Southern Hemisphere and is distinguished by (1) plants ascending, dark green to blackish; (2) stem bases stoloniform; (3) leaves orbicular to slightly elongate, subtransverse, alternate, suberect (not wide-horizontally spreading), with a flat or incurved dorsal margin and with decurrent bases; (4) leaf cells quadrate, smooth (without papillae), walls thin or ± uniformly thickened, cells sometimes becoming thicker-walled toward the margin and forming a broad, ill-defined leaf border; and (5) perianths in the upper part with slightly spirally twisted folds. Syzygiella sonderi may grow together with S. rubricaulis in páramo, and the two species have sometimes been confused. However, S. rubricaulis is readily separated from S. sonderi in its lighter, brownish-green to yellowish or orange color, absence of stolons, and, especially, leaf cells with very large bulging trigones and with large, rounded papillae. 15. Syzygiella tonduzana Steph.

Fig. 20L

On humic soil in montane rainforest and shrubby páramo, 1780–3200  m. Ecuador: Lo, Na (Holm-Nielsen et al. 17702, 27342, AAU, PC), ZC; to be expected in Colombia. General distribution: Costa Rica, Ecuador. Syzygiella tonduzana (subg. Anomalae) is an uncommon northern Andean-Costa Rican species with plagiochiloid, opposite leaves (connate both dorsally and ventrally) that are narrowly triangular and irregularly toothed at the apex, with 2–5 ciliate teeth. The cuticle is smooth. Syzygiella tonduzana is close to S. manca, but the leaf apex in the latter species is entire or with 1–2 small teeth. Possibly, S. tonduzana is just a variety of S. manca.

200

 

Keys and Descriptions

16. Syzygiella trigonifolia (Steph.) Herzog Syzygiella liberata Inoue On bark and rock in upper montane forest and páramo, 1900–3900 m. Colombia: Ant, Boy, Cun, Hui, Met. Ecuador: Az (Gradstein and Benitez 2017), Im, Na, Lo/ ZC (páramo of Saraguro and Yacuambi, Benitez & Gradstein s.n., HUTPL). General distribution: tropical Andes, southeastern Brazil. Syzygiella trigonifolia (subg. Anomalae) is a rather robust Andean species, to 8 cm long and 5 mm wide, with asymmetrically ovate, plagiochiloid leaves, which are usually free on the dorsal side and more or less connate on the ventral side; considerable variation is observed in this character. The usually entire leaf apex (rarely with 2 small teeth), the recurved dorsal leaf margin, and the smooth cuticle are further characteristics of this species (Váňa et al. 2014a, b). 17. Syzygiella uleana Steph.

Fig. 20M, N

On soil along trail in lower montane rainforest, 1780 m. Ecuador: ZC (SchäferVerwimp et  al. 2013a). General distribution: southeastern Brazil, southern Ecuador. Syzygiella uleana (subg. Anomalae?) is an unusual Brazilian species that has been found once in southern Ecuador (Reserva Biologica San Francisco, SchäferVerwimp & Nebel 33058, ST-049014). The species is readily recognized by the large underleaves that are almost as long as the leaves (when well developed), narrowly lanceolate to ligulate, 1.5–2.2  mm long and about 6× longer than wide, becoming smaller toward the stem base and stem apex and sometimes lacking near the apex. The underleaves are connected to leaf bases on one or both sides, and, as noted by Schäfer-Verwimp et  al. (2013a), they may occasionally arise from the ventral base of an adjacent leaf, instead of from the stem. Further characters of S. uleana are (1) plants pale green to brown, to 5 cm long and ca. 2 mm wide; (2) leaves rather fragile, subopposite, decurved, asymmetrically ovate-triangular, tapering to an obtuse to asymmetrically bilobed apex (dorsal lobe smaller) with narrowly rounded to obtuse lobes; (3) dorsal leaf bases longly decurrent and alternate, free, ventral leaf bases connate with underleaves on one or both sides, or among opposite leaves when underleaves are lacking; and (4) leaf cells with small trigones and a smooth cuticle. 18. Syzygiella undata (Mont.) K.Feldberg et al. Jamesoniella undata (Mont.) Steph. Jamesoniella paramicola Herzog On shaded rock and soil in páramo and Polylepis forest, 3400–4300 m. Colombia: Ara, Boy, Cal, Cun. Ecuador: Az (Grolle 1971). General distribution: northern Andes. Syzygiella undata (subg. Syzygiella) is a rare northern Andean páramo species that is closely related to S. rubricaulis but differs by the dorsal leaf bases undulate and longly and broadly decurrent, concealing the dorsal side of stem. The leaves are subopposite and the bases are free.

Jungermanniaceae

201

Jungermanniaceae A small family of five genera, one in the neotropics. Jungermanniaceae have undergone many modifications based on molecular evidence (see Crandall-Stotler et al. 2009; Váňa and Long 2009). The diagnostic features of the family in its current definition are obscure. Jungermannia L. Plants small- to medium-sized, 0.6–4 mm wide, green to blackish-green or brown, creeping or ascending, little branched, stolons lacking. Stems without hyalodermis, cortex weakly differentiated. Branches lateral-intercalary, rarely ventral-intercalary or terminal, Frullania-type. Leaves succubous, undivided, spreading, ovate-elongate, leaf apex rounded, margins entire, not incurved, bases not or short-decurrent, insertion line reaching dorsal midline of stem. Cells with very small trigones, cuticle smooth or papillose; oil bodies finely granular. Underleaves lacking. Rhizoids hyaline or pale brown, scattered, never in fascicles. Dioicous or paroicous. Gametoecia on long shoots. Perianths slightly plicate, long-exserted, mouth without beak, perianth base thin (1–2 cells thick), inner bracts and bracteole not connected to the perianth. Capsule walls 2-layered. Gemmae lacking. A genus of about eight species, one in Colombia and Ecuador. The main characters of Jungermannia are (1) leaves undivided, succubous, inserted to the dorsal midline of stem; (2) underleaves absent; (3) rhizoids scattered; (4) gametoecia on long shoots; and (5) perianth inflated, ± smooth, without beak, perianth base thin, 1–2 cells thick, inner bracts and bracteole not connected to the perianth base. The circumscription of Jungermannia has recently been modified based on molecular evidence (see Váňa and Long 2009), and many species previously placed in Jungermannia have been transferred to Solenostoma (Solenostomataceae). The latter genus differs from Jungermannia by the plicate perianth and the presence of a short perigynium consisting of the thickened lower portion of the perianth (3–5 cells wide) with the inner bracts and bracteole attached to it (perigynium absent in Jungermannia). Literature: Váňa (1973), Váňa and Long (2009) 1. Jungermannia ovatotrigona (Steph.) Grolle Haplozia rorida Herzog Jungermannia cordifolia Hook. var. longifolia Spruce

Fig. 21A, B

On rock in rivers at high elevation, often submerged, 2700–4000 m. Colombia: Boy, Cun, Ris. Ecuador: Na (páramo de Oyacachi, Gradstein & Bader 12790, QCA), Pi, Su. General distribution: tropical Andes. Jungermannia ovatotrigona is an uncommon, semi-aquatic high-Andean species of rock in rivers and rivulets. The species often grows submerged. Characteristics are (1) plants dioicous, rather robust, 2–4  mm wide, glossy green to dark green, branching intercalary or terminal, rhizoids absent or few, colorless; (2) leaves

Fig. 21 (A and B) Jungermannia ovatotrigona. (A) Habit, dorsal view, 18×. (B) Leaves, 18×. (C–E) Lepicolea pruinosa. (C) Habit, ventral view. (D) Leaf. (E) Midleaf cells. (F) Bazzania acanthostipa, leaf and underleaf, 8×. (G and H) Bazzania affinis. (G) Underleaf made up of hyaline cells except at leaf base, 15×. (H) Cells near leaf margin, 150×. (I) Bazzania aurescens, habit in ventral view, 8×. (J) Bazzania bidens, habit in ventral view. (K) Bazzania cuneistipula, habit in ventral view. (L and M) Bazzania falcata. (L) Leaf, 15×. (M) Underleaf, 15×. (N) Bazzania gracilis, habit in ventral view, 8×. (O–Q) Bazzania hookeri var. hookeri. (O) Habit, 8×. (P) Underleaf with small auricles, 8×. (Q) Underleaf with large auricles, 8×. (A, B redrawn from Váňa 1973; F–I, L–Q from Fulford 1963; rest from Gradstein et al. 2001; Gradstein and Ilkiu-Borges 2009)

Lepicoleaceae

203

obliquely to widely spreading, distant, concave to canaliculate, elongate-ovate to lingulate, apex rounded or obtuse, sometimes cucullate, bases not or very little decurrent; (3) leaf cells 20–30 μm long, 12–20 μm wide in midleaf, larger toward the base, fully thin-walled or with very small trigones, cuticle smooth, oil bodies (seen in Gradstein & Bader 12790, QCA) 2–4 per cell, irregularly ovoid to ellipsoid, rather coarsely granular, becoming rapidly homogeneous upon age; (4) underleaves absent; and (5) gynoecia as in the genus.

Lepicoleaceae A small family of one to two genera. Lepicolea Dumort. Plants robust, to 10 cm long, light gray-green or bluish-green to yellowish-brown or dark brown, ascending to erect or pendent, regularly pinnate or 2-pinnate. Branches Frullania-type and often becoming flagelliform toward the tips, occasionally lateral-intercalary. Stems rigid, with a cortex of small, very thick-walled cells in 2–4 layers, the stem surface often with paraphyllia. Leaves transverse, deeply 4-lobed (bisbifid), slightly asymmetrical, segments lanceolate, margins densely ciliatelaciniate. Cells with thick walls and large trigones, cuticle papillose; oil bodies very small, few per cell, faintly and finely segmented to almost homogeneous. Underleaves similar to the leaves but more symmetrical. Rhizoids in bundles on the flagelliform branches. Dioicous. Gametoecia on elongate shoots. Sporophytes surrounded by a fleshy perigynium densely covered by paraphyllia and scale-like leaves; perianth lacking. Setae made up of numerous rows of cells. Capsule walls 5-layered. Vegetative reproduction unknown. A mostly southern-temperate genus of ten species, one in Colombia and Ecuador. The main features of Lepicolea are (1) large bipinnate plants; (2) leaves transverse, deeply 4-lobed, leaf margins strongly ciliate-laciniate; (3) underleaves large, similar to leaves but more symmetrical; (4) stems rigid, with a thick-walled, brown cortex, often with paraphyllia; and (5) sporophyte in a fleshy perigynium, perianth lacking. Literature: Fulford (1963), Schuster (2000a, b) 1. Lepicolea pruinosa (Taylor) Spruce Lepidozia columbica De Not. Sendtnera aequabilis Taylor

Fig. 21C–E

On bark and soil in upper montane forest and páramo, 1800–4000 m. Colombia: Ant, Ara, Boy, Cas, Cau, Cho, Cun, Hui, Mag, Met, Nar, Put, Qui, Ris, San, Tol, Val. Ecuador: Az, Lo, Na, Pa, Pi, Tu, ZC.  General distribution: tropical Andes, Costa Rica.

204

 

Keys and Descriptions

Lepicolea pruinosa is a common Andean-Costa Rican species that is readily recognized by the robust, bipinnate plants with deeply 4-lobed leaves and underleaves with densely ciliate margins and by the presence of numerous small paraphyllia on the stem surface, both on the dorsal and ventral side of the stem.

Lepidoziaceae Plants leafy, rarely thalloid, pale green to brown or reddish-brown, creeping to ascending, usually pinnate or dichotomous, sometimes arising from a stoloniferous base. Branches variable, terminal or intercalary; flagella frequent. Stems with or without hyalodermis. Leaves incubous, succubous, or transverse, alternate, usually divided into several segments or teeth, rarely undivided, leaf insertion extending to dorsal midline of stem or not, leaf margins usually entire. Cells variable; oil bodies present or absent. Underleaves well developed, rarely reduced. Rhizoids in tufts from underleaf bases. Gametoecia on short ventral branches, the androecia occasionally on long shoots. Sporophytes in a 3-keeled perianth. Setae of 8 or 16 outer rows of cells surrounding many (rarely only 4) smaller inner rows. Capsules elongate, wall 2-layered. Vegetative reproduction rare, by caducous leaves. About 28 genera worldwide, 12 in Colombia and Ecuador. The main features of the family are (1) gynoecia on very short ventral branches; (2) rhizoids in tufts from underleaf bases; (3) seta thin, with an epidermis of 8–16 rows of cells; (4) leaves usually divided into lobes; (5) underleaves usually present; (6) sporophyte in a 3-keeled perianth; and (7) gemmae absent. Branching and leaf insertion in Lepidoziaceae are quite variable; leaves may be incubous, succubous, or transverse, and branching is pinnate (Lepidozia, Kurzia, Telaranea), dichotomous (Bazzania), or irregular. 1. Plants consisting of a small thallus with a narrow midrib and unistratose wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pteropsiella 1. Plants not thalloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants consisting of minute shoots (less than 0.2 mm wide), 2 cells wide, with rudimentary leaves, each leaf consisting of 1–2 cells with a slime papilla at the tip. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monodactylopsis 2. Plants larger, leaves not rudimentary, made up of more than 2 cells. . . . . . . . 3 3. Leaves consisting of 1–4 uniseriate filaments, undivided leaf lamina lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telaranea 3. Leaves with a short or long, undivided lamina . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaf cells very large, ca. 60–80  μm long in midleaf, thin-walled, without trigones. Leaf apex undivided to short-bifid, frequently with 1–2 large, sausageshaped or globose slime papillae. Leaves attached to the lateral side of the stem only, longitudinally inserted (see Fig. 25C, D) . . . . . . . . . . . . . . . Zoopsidella 4. Leaf cells mostly smaller, with or without trigones. Leaf apex acute or divided into several teeth or lobes, without sausage-shaped papillae. Leaves different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaves incubous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Lepidoziaceae

205

5. Leaves succubous or transverse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 6. Leaves divided into 4–6 lobes. Branching (bi)pinnate. Ventral flagella usually absent. Plants 0.5–1.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Leaves with 2–3 teeth at apex or undivided. Branching forked or ventral. Ventral flagella common. Plants 1–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . Bazzania 7. Stems with a distinct hyalodermis. Leaf cells thin-walled. Leaf lobes only 1 cell wide except at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neolepidozia 7. Stems without hyalodermis. Leaf cells with thickened walls. Leaf lobes mostly more than 1 cell wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidozia 8. Leaves folded and usually keeled, at least above, the keel often winged . . . . 9 8. Leaves not folded, 2–4-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Leaf apex bifid to 1/4. Underleaves absent . . . . . . . . . . . . . . . . . . . . Mytilopsis 9. Leaf apex undivided or very short-bifid. Underleaves present (sometimes very small) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micropterygium 10. Leaves and underleaves shallowly divided (to 1/4 or less) into 2–4 triangular lobes. Plants growing upright from a stoloniferous base and with ventral stolons. In páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudocephalozia 10. Leaves and underleaves more deeply divided. Plants creeping or ascending, stoloniferous stem base present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Stem leaves 2–3-lobed. Stems with a stoloniferous base . . . . Paracromastigum 11. Stem leaves (3–)4-lobed. Stems without stoloniferous base . . . . . . . . . . Kurzia Bazzania Gray Mastigobryum Nees Plants small to large, 1–6  mm wide, usually deep green to olive green when fresh, becoming brown or reddish-brown on age, creeping, dichotomously branched, with long ventral flagella. Stems rigid, consisting of thick-walled cells; hyalodermis lacking. Branching lateral-terminal (Frullania-type) and ventral-intercalary (Bazzania-type). Leaves incubous, wide-spreading, straight to falcate, ovate-triangular to ovate-oblong to ovate-rectangular, apex truncate and usually divided into 2–3(−4) teeth, rarely entire, dorsal and ventral leaf margins entire or, occasionally, toothed. Cells with small or large trigones, cuticle smooth or finely papillose, cells in the lower half of the leaf middle often larger, resembling a vitta, sometimes with a true, sharply delimited vitta of 4–6 rows of much larger and longer cells sharply separated from the rest of the leaf cells; oil bodies large, few segmented, sometimes homogeneous. Underleaves well developed, distant or imbricate, apex and lateral margins entire to shallowly lobed to toothed, underleaf bases cuneate to auriculate, free or connate to leaf bases on one or both sides; underleaf cells chlorophyllose or in part without chlorophyll. Dioicous. Gametoecia and sporophytes infrequent. Vegetative reproduction common, by caducous leaves. About 100 species worldwide, 26 in Colombia and Ecuador. Bazzania is readily recognized by the dichotomously branched plants, incubous leaves with a 2–3-toothed apex (rarely entire), well-developed underleaves, and long flagelliform

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branches sprouting from the axils of the underleaves. The neotropical species of Bazzania remain poorly known in spite of the detailed studies by Fulford (1946, 1959, 1963); identifications of herbarium specimens are frequently erroneous. The present treatment follows Gradstein (2017b). Bazzania species are highly variable as pointed out by Spruce (1885), Fulford (1963), Schuster (1966), Jones (1975), and others. Particularly troublesome are the small, “hygromorphic” phenotypes from shaded places with distant leaves and small underleaves (Jones 1975). These modifications are common and can be difficult to identify. Collections of such specimens should be carefully examined and searched for the occurrence of better-developed stems with imbricate leaves that may permit identification of the material. The presence or absence of a vitta in the leaves is an important taxonomic character that should be used with caution. It is present in the common B. gracilis and in the rare B. denticulifera and B. spruceana and consists of a narrow band, ca. 4–6 longitudinal cell rows wide, of very large, quadrate to rectangular cells that are sharply separated from the rest of the leaf cells. The true vitta should not be confused with the “vitta-like” area seen in many species (e.g., B. affinis, B. diversicuspis, B. falcata, B. hookeri, B. jamaicensis, B. longistipula, B. stolonifera) and consisting of larger leaf cells in the mid-lower portion of the leaves. The vitta-like area is usually much broader than the true vitta (sometimes extending almost to the leaf margin) and differs essentially from the true vitta in being not sharply separated from the rest of the leaf cells. The vitta-like area can be quite variable within species being more pronounced in some populations than in others (e.g., in B. hookeri) and is without taxononomic significance. Literature: Gradstein (2017b) 1. Leaf apex entire, rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. placophylla 1. Leaf apex at least in part of the leaves with 2–3 teeth or shallowly lobed . . . 2 2. Leaf apex mostly trifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaf apex bifid or variable (not mostly trifid) . . . . . . . . . . . . . . . . . . . . . . . . 25 3. Dorsal leaf margin toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Dorsal leaf margin entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 4. Leaves with a well delimited vitta made up of 4–6 rows of large, rectangular cells. Plants growing in lowland rainforest of the Chocó . . . . B. denticulifera 4. Leaves without well delimited vitta, sometimes with larger cells suggesting a vitta. Plants growing above 1000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Underleaves sharply toothed all around, green, not made up of hyaline cells (= cells without chlorophyll), border of hyaline cells lacking . . . . . B. denticulata 5. Underleaves not sharply toothed all around, almost fully hyaline or with a border of hyaline cells. Leaves strongly caducous . . . . . . . . . . . . . . . . . B. serrata 6. All or part of the underleaves with a border of hyaline cells (= cells without chlorophyll) or largely made up of hyaline cells. . . . . . . . . . . . . . . . . . . . . . . 7 6. Underleaves without hyaline border and not largely made up of hyaline cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 7. Leaf cells with small trigones, cell lumen ± quadrate or rectangular. Leaves usually strongly caducous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. affinis

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7. Leaf cells with large trigones, cell lumen angular or rounded . . . . . . . . . . . . . 8 8. Hyaline underleaf cells much thinner-walled than the chlorophyllose underleaf cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Hyaline underleaf cells not thinner-walled than the chlorophyllose cells . . . 11 9. Underleaf bases free. Plants very common . . . . . . . . . . . . . . . . . B. stolonifera 9. Underleaf bases connate with leaves on one or both sides, plants very rare in the northern Andes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Underleaf bases slightly connate with leaves on one side. Underleaves wider than long. [Caribbean species]. . . . . . . . . . . . . . . . . . . . . . . . . . . B. eggersiana 10. Underleaf bases connate with leaves on both sides. Underleaves usually longer than wide. [Patagonian species]. . . . . . . . . . . . . . . . . . . . . . . . . . . B. peruviana 11. Underleaves as long as wide, with a narrow border of hyaline cells all around. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. chilensis 11. Underleaves at least in part longer than wide, with a hyaline border or the upper half of underleaves hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pallidevirens 12. Underleaves divided to almost the middle into 4 narrowly lanceolate lobes. Plants from from the tepui region of Venezuela; not known in Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. chimantaensis Fulford 12. Underleaves not divided to almost the middle into 4 narrowly lanceolate lobes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Underleaves wider than long, reniform. Underleaf apex and margins fully entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. jamaicensis 13. Underleaves as wide as long or longer than wide. Underleaf apex shallowly lobed or toothed, not fully entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Underleaf bases auriculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Underleaf bases not auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 15. Lateral margins of underleaves with 1–2 very large, lobe-like teeth (5–10 cells long, 5–8 cells wide). Auricles of underleaves sometimes overlapping. Leaves without broad vitta-like area. Plants rare. . . . . . . . . . . . . . . . . . B. acanthostipa 15. Lateral margins of underleaves entire or with small, 1–3-cell-long teeth. Auricles of underleaves not overlapping. Leaves with or without broad vittalike area. Plants very common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Underleaf auricles toothed and usually with a small foliose appendage. Leaves ± falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. falcata 16. Underleaf auricles entire, without foliose appendage. Leaves falcate or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (B. hookeri) 17 17. Leaf cells mammillose. Underleaves ca. 1.5× stem width, longer than wide. Plants only known from the Cordillera del Condor, Ecuador. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. hookeri var. mamillosa 17. Leaf cells not mammillose. Underleaves usually more than 1.5× stem width, shape variable. Plants very common . . . . . . . . . . . . . . . B. hookeri var. hookeri 18. Underleaves strongly elongate, about 2× longer than wide. Underleaf apex deeply 4-lobed. Plants rare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. elongata 18. Underleaves 1–1.5× longer than wide. Underleaf apex entire or shallowly 2–4lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

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19. Leaves with a well delimited vitta made up of 5–6 rows of large, rectangular cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. spruceana 19. Leaves without well delimited vitta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. All or part of the underleaves longer than wide . . . . . . . . . . . . . . . . . . . . . . 21 20. Underleaves rounded-subquadrate, not longer than wide . . . . . . . . . . . . . . . 22 21. Leaves very long and narrow, ca. 3× longer than wide, ovate-lanceolate. Cells near leaf apex very large, ca. 45 μm long. Plants rare. . . . . . . B. crassidentata 21. Leaves less elongate, 1.5–2× longer than wide. Cells near leaf apex smaller. Plants common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. longistipula 22. Lateral margins of underleaves recurved, underleaf apex truncate-emarginate. Plants from Mexico and Venezuela, not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . B. acuminata (Lindenb. & Gottsche) Trevis. 22. Lateral margins of underleaves plane, underleaf apex irregularly lobed. . . . 23 23. Ventral leaf base strongly dilated, auriculate. . . . . . . . . B. hookeri var. hookeri 23. Ventral leaf base not or little dilated, not auriculate . . . . . . . . . . . . . . . . . . . 24 24. Underleaves contiguous to imbricate, ca. 2× stem width . . . . . . . B. aurescens 24. Underleaves distant, scarcely wider than the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . shade form of B. longistipula 25. Underleaves 1–2  mm wide, (sub)orbicular. Underleaf bases auriculate. Plants robust, 3–5  mm wide. (Leaf apex with 2–3 very shallow lobes or entire). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. canelensis 25. Underleaves smaller. Underleaf bases not auriculate. Plants smaller, 1–3 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 26. Leaf apex entire to very shallowly bifid. Leaves strongly caducous (or fragmenting). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. gracilis 26. Leaf apex mostly bifid or entire to shallowly bifid to trifid. Leaves not or slightly caducous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27. Leaf apex on single stems varying from undivided to bifid to trifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. diversicuspis 27. Leaf apex mostly bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28. Lateral margins of underleaves ± toothed. Leaf margins between the apical teeth crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. phyllobola 28. Lateral margins of underleaves entire. Leaf margins between the apical teeth entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 29. Well-developed leaves (2.5–)3–4× longer than wide . . . . . . . . . . . . . . B. bidens 29. Well-developed leaves 2–2.5× longer than wide. . . . . . . . . . . . . . . . . . . . . . 30 30. Trigones very large, strongly bulging. Apical leaf cells 25–36 μm. Plants growing above 2000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. roraimensis 30. Trigones smaller, not strongly bulging. Apical leaf cells smaller. Plants growing below 2000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. cuneistipula 1. Bazzania acanthostipa Spruce

Fig. 21F

On bark and rock in montane forest, forming large tufts, 1500–1900  m. Colombia: Ant, Cho, Hui, Mag, Val. Ecuador: ZC (Churchill et al. 2009) General distribution: tropical Andes.

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Bazzania acanthostipa is a rare Andean species from above 1500 m. The species is readily recognized by the very large, imbricate underleaves with 1–3 large, lobelike teeth (5–10 cells long, 5–8 cells wide) on the lateral margins. The leaves are falcate, and the ventral leaf bases are strongly expanded-auriculate and undulatetoothed. Bazzania acanthostipa is morphologically close to B. falcata, but in the latter species, the teeth on underleaf margins are smaller (or absent). 2. Bazzania affinis (Lindenb. & Gottsche) Trevis. Bazzania lindigii (Steph.) Spruce Bazzania taleana (Gottsche) Fulford

Fig. 21G, H

On bark in montane forest, 800–2900 m. Colombia: Ama, Cas, Cun, Hui (as B. taleana), Mag, Met, Qui, Ris, San, Tol. Ecuador: Lo, ZC. General distribution: common and widespread in tropical America. Bazzania affinis is widespread in the neotropics and readily distinguished by the delicate plants (ca. 2–3  mm wide) with trifid leaves made up of thin-walled or slightly thick-walled, quadrate to subrectangular cells, without or with small trigones. A further character is the small, distant to subimbricate underleaves made up largely, or in part, of hyaline cells without chlorophyll. Sometimes the hyaline cells are largely restricted to the underleaf margin, forming a border. The underleaves are free or attached to the leaves on one side. The leaves are often strongly caducous and may possess a broad, ill-defined vitta-like region. Bazzania affinis is morphologically close to B. serrata, but the latter species has finely serrate leaf margins. Bazzania taleana is a phenotype of B. affinis with hyaline cells highly variable in the underleaves, forming a border or being distributed across most of the underleaf (Gradstein 2017b). 3. Bazzania aurescens Spruce Bazzania quadricrenata (Steph.) Steph.

Fig. 21I

On rock, logs, and tree bases in lowland and montane forests, 100–2000  m. Colombia: Nar, San, Vau. Ecuador: Bo, Ch, MS, Or, ZC. General distribution: West Indies, northern South America. Bazzania aurescens is an uncommon neotropical species that is recognized by ovate-oblong, ± straight, trifid leaves and subquadrate, contiguous to imbricate underleaves with an irregularly lobed apex and cuneate bases, which are free or shortly connected with leaves on one side. The leaf cells are ca. 20–34 μm long near apex, the trigones are large and often fused, the ventral leaf bases are not or scarcely dilated, and the underleaves lack a hyaline border. Bazzania aurescens somewhat resembles B. hookeri, but in the latter species, the underleaf bases are auriculate and the ventral leaf bases are expanded-auriculate. Bazzania aurescens may also be confused with delicate phenotypes of B. longistipula, but the latter plants can be separated from B. aurescens by the small, distant underleaves (± imbricate in B. aurescens).

210

4. Bazzania bidens (Gottsche & Lindenb.) Trevis. 

 

Keys and Descriptions

Fig. 21J

On rock, logs, and tree bases in lowland and montane forests, 200–3400  m. Colombia: Nar, San, Vau. Ecuador: Bo, Ch, ZC. General distribution: tropical America. Bazzania bidens is a common neotropical species distinguished by strongly elongate, bifid leaves (ca. 3–3.5× longer than wide) and small, distant underleaves (scarcely wider than the stem) with entire margins and an undulate to shallowly lobed apex. A broad, vitta-like area is usually present in the leaves. Bazzania bidens is morphologically close to B. phyllobola, but in the latter species, the leaf margins between the apical teeth are crenulate (entire in B. bidens) and the underleaves are broader, with ± toothed lateral margins. Bazzania bidens also resembles B. cuneistipula, but in the latter species, the leaves are less elongate (2–2.5× longer than wide) and the apical teeth smaller (to 5 cells long). 5. Bazzania canelensis (Steph.) Fulford On bark, soil, and rock in montane forests and páramo, ca. 1000–3750  m. Colombia: Cau, Hui, Ris. Ecuador: Lo, Pa (Canelos), Tu, ZC. General distribution: northern Andes, Honduras (fide Tropicos). Bazzania canelensis is a poorly known Andean species that is distinguished by robust plants with leaf tips varying from entire to shallowly 2–3-lobed and the large, suborbicular underleaves (1–2 mm wide) with subentire margins and small to large auricles. Bazzania canelensis closely resembles B. placophylla and may be a synonym of the latter, but the leaf apex in B. placophylla is fully entire, rounded. The species also approaches B. hookeri, but in the latter species, the leaf apex is distinctly trifid. Most of the reports of B. canelensis are from páramo although the type (Canelos, Spruce s.n.) is from submontane forest. The herbarium material of the species needs to be checked. 6. Bazzania chilensis (Steph.) Fulford Bazzania bogotensis (Steph.) J.B.Jack & Steph. In montane forest and páramo, 1800–3650 m. Colombia: Boy, Cau, Cho, Cun, Hui, Ris. Ecuador: unknown locality (Gradstein 1999). General distribution: Andes. Bazzania chilensis is an uncommon Andean species that is recognized by rounded-subquadrate underleaves with a narrow, 1–2(−4)-cell-wide border of hyaline cells all around the underleaf. The species resembles B. stolonifera, but in the latter species, the hyaline cells are conspicuously thinner-walled than the chlorophyllose cells and the hyaline border is usually wider, to 8 cells wide. Bazzania chilensis is very close to B. pallidevirens, but in the latter species, the underleaves tend to become longer than wide (never longer than wide in B. chilensis) and the hyaline cells may at least in some underleaves not be restricted to the border but may be occupying the entire upper half of the underleaf.

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7. Bazzania crassidentata Fulford In montane forest and páramo, 1500–3650  m. Colombia: Cau, Hui, Val. General distribution: Colombia, Venezuela (Guayana Highland). Bazzania crassidentata is a rare neotropical species characterized by strongly elongate, ovate-lanceolate leaves (ca. 3× longer than wide), large leaf cells (to 45 μm long near apex), and underleaves often longer than wide. 8. Bazzania cuneistipula (Gottsche & Lindenb.) Trevis. Mastigobryum brevifolium Gottsche

Fig. 21K

On trees and logs in lowland and montane rainforest and scrub, 200–2000(−2900?) m. Colombia: Ama, Ant, Boy, Ces, Cun, Qui, Ris, San, Tol, Vau. Ecuador: Lo, Or, Pa, ZC. General distribution: tropical America. Bazzania cuneistipula is a common, delicate neotropical species recognized by plants usually less than 2 mm wide; leaves bifid, 2–2.5× longer than wide and somewhat falcate; and rather small leaf cells (18–24 μm at leaf apex) with trigones that are not strongly bulging. The species is morphologically close to B. roraimensis; for differences see under the latter species. 9. Bazzania denticulata (Lindenb. & Gottsche) Trevis. Bazzania planiuscula (Lindenb. & Gottsche) Trevis. Bazzania rusbyi Spruce On bark, rotten wood, and soil in montane forest and scrub, 1000–3100  m. Colombia: Cas, Cun, Hui, Met, Ris, San. Ecuador: ZC. General distribution: tropical America. Bazzania denticulata is recognized by the strongly toothed underleaves with numerous sharp, narrowly triangular to ciliate teeth all around the margins. Moreover, the dorsal and ventral leaf margins are usually toothed in the upper half, and the ventral leaf bases are usually dilated and toothed (but sometimes entire). The species seems to be widespread in the neotropics. 10. Bazzania denticulifera Mägd. Pendent on bark in wet lowland rainforest along river, 75 m. Colombia: Cho (Mägdefrau 1983). General distribution: only known from the Chocó. Bazzania denticulifera is a rare Colombian endemic that is distinguished by the trifid leaves with toothed margins, an expanded ventral leaf base and a distinct vitta of 4–6 rows of large, rectangular cells. The underleaves are small, distant, subquadrate (about twice as wide as the stem) with a toothed apex, entire lateral margins, and subauriculate bases. By the toothed leaf margins, B. denticulifera resembles B. denticulata, but the latter species lacks a vitta and the underleaves are sharply toothed ± all around. In its distinct vitta, B. denticulifera resembles B. spruceana, but in the latter species, the leaf margins are entire, the ventral leaf bases are not dilated, and the underleaf bases are not auriculate.

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Keys and Descriptions

11. Bazzania diversicuspis Spruce On trees in rainforests, 100–500(−1000?) m. Colombia: Ama, Ant, Caq, Put, Vau (Campos et  al. 2015). Ecuador: ZC (?). General distribution: northern South America. Bazzania diversicuspis is a rare Amazonian taxon that has frequently been confused with other neotropical Bazzania species. All records of B. diversicuspis from the Andean region must be verified and may be misidentifications. The species is characterized by the rather small plants (2–3 mm wide) with highly variable leaf apices, varying from entire to bifid to trifid on the same stem. The underleaves are rather small, 1.5–2× wider than the stem, with an irregularly lobed apex, a shallowly curved insertion line, and cuneate to rounded bases. 12. Bazzania eggersiana (Steph.) Pagán On tree bases in montane forest, 1900–2500 m. Colombia: Mag (van Reenen et  al. 1984). General distribution: Greater Antilles, northern Andes (Colombia, Peru). Bazzania eggersiana is a Caribbean species that has been recorded once from the Sierra Nevada de Santa Marta (van Reenen et al. 1986) and from Peru (SchultzeMotel and Menzel 1987). It is readily recognized by the small plants (2–3 mm wide) with wider than long underleaves that are slightly connate to leaf bases on one side of the stem (free on the other side) and possess a narrow hyaline border made up of thin-walled cells. Bazzania eggersiana approaches B. stolonifera, but the latter species is more robust, the underleaves and the hyaline border in B. stolonifera are broader, and the underleaf bases are free. 13. Bazzania elongata Fulford On tree trunks in montane forest, 2400  m. Colombia: Hui (Bischler 1962d). General distribution: Colombia, Venezuela (Guayana Highland). This rare species stands out by strongly elongate underleaves (ca. 2× longer than wide) with a deeply 4-lobed apex and by rather large leaf cells, more than 35 μm long in the apical part of the leaf. 14. Bazzania falcata (Lindenb.) Trevis. Bazzania ancistrodes Spruce Bazzania boliviana (Steph.) Fulford Bazzania macrostipula Fulford Bazzania robusta Spruce Bazzania schlimiana sensu Fulford (non typus)

Fig. 21L, M

On bark, rotten wood, and rock in montane forest and páramo, forming large tufts, (100–)1000–3600  m (records from below 1000  m may be erroneous and should be verified). Colombia: Ant, Boy, Cau, Cho, Cun, Hui, Mag, Met, NSa, Ris, San, Tol, Val. Ecuador: Lo, Pa, Tu, ZC.  General distribution: common and widespread in tropical America.

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Bazzania falcata is a common, robust neotropical species that is distinguished by large underleaves with ± toothed auricles and the usual presence of a foliose appendage at the base of each auricle. In some populations the appendage may be absent, however. The leaves are ± falcate, the dorsal leaf base is strongly dilated, the ventral leaf base is expanded-auriculate and usually with undulate to toothed margins, and the underleaf margins are entire or finely toothed. Bazzania falcata is morphologically close to B. hookeri, but in the latter species, the underleaf auricles are entire and lack a foliose appendage. The type material of B. schlimiana belongs to B. hookeri, but Fulford’s (1963) description of B. schlimiana refers to B. falcata. Bazzania macrostipula (Fulford 1959) is a phenotype of B. falcata with leaves with small apical teeth. 15. Bazzania gracilis (Hampe & Gottsche) Steph. Fig. 21N Bazzania bidens sensu Spruce (non typus) Mastigobryum heterophyllum Steph. Type: Gualaquiza, “in silva ad rivum Salado ad truncorum arboris,” Mai 1909, Allioni s.n., Bryotheca Levier 6577 (holotype G-00066890!), syn. nov. Not a synonym of B. phyllobola (Fulford 1963). On bark, logs, and rock in lowland and montane forests up to the páramo, 100–3300 m. Colombia: Ama, Ant, Ces, Cho, Hui, Mag, Met, Ris (Gradstein 8360, 8369, 8425, COL, GOET), San, Tol, Val, Vau. Ecuador: Az, ZC. General distribution: widespread in tropical America. Bazzania gracilis is a common neotropical species that is readily distinguished by the small plants (1–2 mm wide) with strongly caducous or fragmenting, entire to shallowly bifid leaves with short teeth (1–4 cells long). Sometimes, the shoots are almost completely devoid of leaves. A vitta-like area is often present in the leaves. Bazzania herminieri (Steph.) Pagán, described from the Lesser Antilles, is probably a phenotype of B. gracilis with falcate leaves. 16a. Bazzania hookeri (Lindenb.) Trevis. var. hookeriFig. 21O–Q Bazzania arcuata (Lindenb. & Gottsche) Trevis. Bazzania breuteliana (Lindenb. & Gottsche) Trevis. Bazzania chimborazensis Spruce Bazzania cuervi (Gottsche) Steph. Bazzania gottscheana Lindenb. Bazzania glaziovii (Gottsche) Fulford Bazzania flavicans Spruce Bazzania heteroclada Spruce Bazzania latidens (Gottsche) Fulford Bazzania longa sensu Fulford (non typus) Bazzania schlimiana (Gottsche) Fulford Bazzania speciosum Steph. Bazzania teretiuscula (Lindenb. & Gottsche) Trevis. Bazzania viridissima Spruce Mastigobryum ecuadorense Steph.

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Keys and Descriptions

On trees, logs, rock, and soil in lowland and montane forests up to páramo, 100–4000 m. Colombia: Ama, Ant, Boy, Caq, Cas, Cau, Cho, Cun, Guajira, Hui, Mag, Met, Nar, NSa, Put, Qui, Ris, San, Tol, Val, Vau. Ecuador: Az, Ch, Ga, Im, MS, Na, Or, Pa, Pi, Tu, ZC.  General distribution: widespread in tropical America. Bazzania hookeri is the most common neotropical species in the genus. It is very variable and has been described under many different names (see Gradstein 2017b). Bazzania hookeri is characterized by the elongate (ca. 1.6× longer than wide), trifid, straight to falcate, spreading or secund leaves with small to rather large apical teeth and a strongly expanded-auriculate base with (sub)entire margin and by the more or less imbricate, subquadrate-rounded to longer than wide underleaves (rarely wider than long) with a shallowly lobed-toothed apex, (sub)entire lateral margins, and auriculate bases with entire margins (occasionally with 1–3 small teeth) and without appendage. The cuticle varies from finely papillose to quite smooth. A broad, vittalike area of larger cells is often present in mid-lower portion of the leaf. Considerable variation is observed in the size of the plants, the size of the underleaves (0.6–1.2 mm wide), and the size of the auricles at underleaf bases. In robust plants the leaves are often secund and falcate, and the underleaves large and imbricate, with large auricles. In smaller plants and shade forms, the leaves are usually more spreading outward, and the underleaves smaller and the auricles small or sometimes even lacking. Plants with reduced auricles may be recognized by the characteristic expanded-auriculate leaf base. Bazzania hookeri approaches B. falcata and some phenotypes may be difficult to identify. However, the entire underleaf auricles of B. hookeri usually permit to separate this species from B. falcata. A phenotype with strongly falcate leaves and small underleaf auricles has erroneously been called “B. longa,” a species described from Mauritius. Bazzania longa does not occur in the neotropics and is a synonym of the widespread paleotropical B. praerupta (Reinw., Blume & Nees) Trevis. (Gradstein 2017b). 16b. Bazzania hookeri var. mamillosa (Gradst. & A.R.Benitez) Gradst. & A.R.Benitez Bazzania arcuata (Lindenb. & Gottsche) Trevis. var. mamillosa Gradst. & A.R.Benitez On moist soil and on bases of dwarf trees in rather low and open, woody vegetation, 2300 m. Ecuador: ZC (Gradstein and Benitez 2014b). General distribution: only known from the type from southern Ecuador. Bazzania hookeri var. mamillosa is a rare taxon that is only known from Cerro Plateado, highest peak in the Cordillera de Condor. It stands out by the strongly falcate, trifid leaves with conspicuously mammillose leaf cells and a smooth cuticle and by the underleaves only ca. 1.5× wider than the stem and slightly longer than wide. Bazzania hookeri var. mamillosa was originally described as a variety of B. arcuata, but as the latter species proved to be a synonym of B. hookeri, the var. mamillosa has been transferred to B. hookeri. It differs from typical B. hookeri by the narrow underleaves and mammillose leaf cells but otherwise fits B. hookeri

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quite well. Further collections of this interesting taxon are necessary to confirm its status. 17. Bazzania jamaicensis (Lehm. & Lindenb.) Trevis. Bazzania fendleri (Steph.) Fulford

Fig. 22A

On bark, dead wood, and soil in montane forest and shrubby páramo, 1500–3500 m. Colombia: Ant (cf.), Boy, Cas, Cau, Cho, Cun, Hui, Mag, Ris, San, Tol. Ecuador: Na, ZC. General distribution: tropical America. Bazzania jamaicensis is a characteristic neotropical species that is readily distinguished by the distant, reniform underleaves (1.2–1.5× wider than long) with fully entire margins, with a rounded, entire apex and with cordate-auriculate underleaf bases, which are free from the leaves or slightly connate on one side. The leaves are trifid and usually somewhat falcate and secund, the ventral leaf bases are often conspicuously expanded (sometimes not or only slightly expanded), and the leaf cells are rather small, ca. 20–25 μm long near the apex, with small trigones. In robust specimens from Podocarpus National Park, southern Ecuador, the underleaves have a brownish border of thick-walled cells in 1–2 layers; the leaves have a broad, illdefined vitta area; and the dilated ventral leaf bases are sometimes folded backward, forming a small sack (Burghardt 6894, 7121, 7143, PC, QCA). These specimen may represent a separate taxon; this needs more study. 18. Bazzania longistipula (Lindenb.) Trevis. Bazzania angustifalcata Herzog Bazzania decidua Spruce Bazzania tricuspidata (Steph.) Fulford Mastigobryum columbicum Steph.

Fig. 22B, C

On bark, rotten wood, rock, or soil in montane forest and páramo, 750–3500 m. Colombia: Ant, Boy, Cas, Cau, Ces, Cho, Cun, Guajira, Hui, Mag, Met, Nar, NSa, Ris, San, Tol, Val. Ecuador: Lo, Na, Pa, Su, Tu, ZC. General distribution: common and widespread in tropical America. Bazzania longistipula is a common and variable neotropical species that is characterized by trifid leaves with sharp apical teeth (to 6 cells long) and with or without expanded ventral leaf base, leaf cells with medium-sized trigones and a more or less smooth cuticle, and usually longer than wide underleaves with a shallowly and irregularly lobed apex, entire margins, cuneate to rounded (not auriculate) bases and a straight insertion line. The leaves are often falcate and caducous and may possess an ill-defined, vitta-like area. In small shade forms (e.g., the type of B. tricuspidata), the leaves may be quite distant, and the underleaves small, distant, and mostly subquadrate. Such phenotypes can be confused with B. aurescens, but the leaves and underleaves in the latter species are normally (sub)imbricate and the underleaves are never longer than wide. By careful examination of the shade forms of B. longistipula, some longer-than-wide underleaves can usually be found. Bazzania angustifalcata (“B. angustifolia”; Uribe and Gradstein 1998) is a phenotype of B. longistipula with expanded-auriculate ventral leaf bases.

Fig. 22 (A) Bazzania jamaicensis, underleaf, 15×. (B and C) Bazzania longistipula. (B) Habit in ventral view, 8×. (C) Underleaf, 15×. (D) Bazzania placophylla, leaves and underleaf, 8×. (E) Bazzania stolonifera, habit in ventral view showing underleaves with broad hyaline margin, 8×. (F and G) Kurzia capillaris subsp. verrucosa. (F) Habit, ventral view. (G) Leaf lobe. (H) Kurzia brasiliensis, leaf, 180×. (I) Lepidozia alstonii, leaf, 75×. (J and K) Lepidozia auriculata. (J). Leaf, 45×. (K) Apex of leaf lobe, 350×. (L). Lepidozia andicola, apex of leaf lobe, 350×. (M and N) Lepidozia inaequalis. (M) Leaves and underleaf, 33×. (N) Apex of leaf lobe, 350×. (O and P) Lepidozia incurvata. (O) Habit, 35×. (P) Leaf, 60×. (Q) Lepidozia macrocolea, leaves and underleaf, 43×. (R) Lepidozia reptans, habit in ventral view, 15×. (A–E, H–R from Fulford 1966; rest from Gradstein et al. 2001)

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19. Bazzania pallidevirens (Steph.) Fulford On bark and moist cliffs in lowland and montane rainforests, from sea level to 2800  m; common in Amazonia. Colombia: Ama, Boy, Caq, Cun, Hui, Ris, Val, Vau. Ecuador: Ch, Lo. General distribution: tropical South America. Bazzania pallidevirens is recognized by the falcate, narrowly elongate leaves made up of cells with conspicuous trigones and the quadrate to slightly longer than wide underleaves with a hyaline border or hyaline in the entire upper half. The species is morphologically close to B. chilensis, but the leaves in the latter species are less elongate and not falcate; the underleaves are quadrate-rounded, not longer than wide; and the hyaline cells are restricted to a very narrow border (1–2 cells wide) along the underleaf margins. 20. Bazzania peruviana (Nees) Trevis. Bazzania creberrima (Steph.) S.W.Arnell Bazzania lechleri (Steph.) Spruce Bazzania skottsbergii (Steph.) Fulford (see Váňa and Engel 2013) On tree trunks, rock, and soil in moist evergreen forests. Ecuador: without locality (Jameson s.n.; Fulford 1963 as “Peru”); as pointed out by León-Yánez et al. (2006), all specimens collected by Jameson are from Ecuador, not from Peru. General distribution: tropical Andes (very rare), Patagonia, Tristan da Cunha. Bazzania peruviana is a common Patagonian species that has been recorded from Ecuador in the nineteenth century, but not recently (Fulford 1963). The species resembles B. stolonifera in the underleaves with a hyaline margin of thin-walled cells but differs from the latter in the underleaf bases being distinctly connate with leaves on both sides (bases free in B. stolonifera) and the margin of the leaf apex being finely denticulate between the three teeth (entire in B. stolonifera). 21. Bazzania phyllobola Spruce Bazzania bidens (Gottsche & Lindenb.) Trevis. var. dissodonta Spruce On bark of trees in lowland and montane forest, from sea level to 3000  m. Colombia: Ama, Ant, Caq, Cau, Ces, Cho, Cun, Hui, Mag, NSa, San, Val Vau. Ecuador: MS (type of M. heterophyllum), ZC. General distribution: tropical South America. Bazzania phyllobola is a common tropical South American species that is distinguished by bifid (occasionaly trifid) leaves with rather large apical teeth (ca. 6–9 cells long) with crenulate margins. The species is morphologically close to B. bidens but differs from the latter in the crenulate apical leaf margins and the larger underleaves, which are distinctly wider than the stem and have ± toothed lateral margins. 22. Bazzania placophylla (Taylor) Grolle

Fig. 22D

On soil in páramo and in Polylepis pauta forest, 3500–4000 m. Colombia: Cau, Cho, Hui, Ris. Ecuador: Az (V. Cadena 108, QCA), Pi. General distribution: northern Andes.

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Keys and Descriptions

Bazzania placophylla is a rare northern Andean páramo species and the only neotropical member of Bazzania with fully entire, rounded leaf apices. The plants are rather robust, ca. 3–4 mm wide, and the underleaves are large, orbicular, fully entire and with cordate-auriculate bases with small auricles. Bazzania canelensis approaches B. placophylla, but the leaf apex in B. canelensis is mostly shallowly 2–3-lobed, occasionally entire. Bazzania placophylla is exclusive to páramo. 23. Bazzania roraimensis (Steph.) Fulford On trees and logs in montane forest and shrubby páramo, (1900–)2500–3650 m. Colombia: Cau, Hui, NSa. General distribution: West Indies, northern South America, southeastern Brazil. Bazzania roraimensis is habitually similar to B. cuneistipula but differs by slightly larger plant size (ca. 2  mm wide), larger leaf cells (cells near leaf apex 25–36 μm), very large, bulging trigones, and occurence at higher elevation. Possibly, B. roraimensis is a robust, high-elevation phenotype of B. cuneistipula. 24. Bazzania serrata Fulford On tree trunks in upper montane cloud forests, 2150–3000 m. Colombia: Boy (Álvaro Alba et al. 2007), Cas (Gradstein 12700, UPTC), NSa, San. General distribution: only known from the Eastern Cordillera of Colombia. Bazzania serrata is a rare Colombian endemic that is recognized by the strongly caducous leaves with serrate (finely toothed) margins. The underleaves have a hyaline border or are almost fully hyaline, and the leaf cells are usually larger in midleaf, suggesting a vitta. Bazzania serrata is morphologically close to B. affinis but differs from the latter by the serrate leaf margins. 25. Bazzania spruceana Steph. On trees in wet lowland and montane rainforests, 2650–2950  m. Colombia: Boy, San, Val (Bischler 350, COL). Ecuador: ZC (Parolly et al. 2004). General distribution: northern Andes (Colombia to Peru), Brazil. Bazzania spruceana is a rare Andean species that is readily distinguished by the delicate (to 2 mm wide) plants with a true, 5–6 cell rows wide vitta of very large cells in the leaves. The leaves have three small teeth at the apex (2–5 cells long), entire margins, and a straight ventral leaf base; the leaf cells are small (16–22 μm near leaf apex), ± evenly thick-walled, without distinct trigones (except in the vitta cells) and strongly papillose; and the underleaves are small, subquadrate, 1.5–2× stem width, slightly lobed at apex, with entire margins and cuneate to rounded bases that are free from the leaves. The underleaves are fully chlorophyllose and lack a hyaline border. 26. Bazzania stolonifera (Sw.) Trevis. Bazzania leptostipa Spruce Bazzania liebmanniana (Lindenb. & Gottsche) Trevis. Bazzania vincentina (Lehm. & Lindenb.) Spruce Mastigobryum azuayense Steph.

Fig. 22E

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On bark, rotten wood, rock, and soil in montane forest, 500–3000 m. Colombia: Ant, Boy, Cho, Cun, Hui, Mag, Met. Ecuador: Ch, MS, Or, Pa, Tu. General distribution: common and widespread in tropical America. Bazzania stolonifera is a common neotropical species that is characterized by underleaves bordered by hyaline cells with rather thin walls and small trigones that are much smaller than those of the chlorophyllose cells. The hyaline border is 1–8 cells wide. The plants are rather robust, 3–5 mm wide; the leaves are trifid, distinctly elongate (ca. 2× longer than wide) and somewhat falcate; and the underleaves are quadrate-rounded, with an entire or shallowly lobed apex, entire lateral margins, a straight or shallowly curved insertion line, and cuneate to slightly rounded bases that are not connate to the leaf base. Bazzania stolonifera is one of the few neotropical species of Bazzania that is often fertile and may freely produce sporophytes. In most or other neotropical Bazzania species, sporophytes are rare or unknown. The type of Mastigobryum azuayense is not from Azuay, as suggested by the epithet, but from Morona Santiago (Gualaquiza, leg. M. Allioni). Indeed, numerous specimens collected by the reverend Michel Allioni in the surroundings of Gualaquiza and Bomboiza that were subsequently included the Bryotheca E. Levier have been mislabelled as being from “Azuay.” Excluded Records Bazzania heterostipa (Steph.) Fulford  – Colombia: San (Campos and Uribe 2006). The material belongs to B. affinis. Bazzania longa (Nees) Trevis.  – Colombia: recorded from numerous departments (Gradstein and Uribe 2016a). Bazzania longa is a misinterpreted species that is excluded from the neotropical flora (Gradstein 2017b). The type specimen is from Africa (Mauritius), not from Martinique (Fulford 1963), and belongs to B. praerupta (Reinw., Blume & Nees) Trevis., a species widespread in the paleotropics. The neotropical specimens identified as B. longa usually belong to B. hookeri; all neotropical records must be revised. Bazzania tayloriana (Mitt.) Kuntze  – Colombia: Cun (Fulford 1946), Val (Bischler 1962d). Species endemic to New Zealand; the specimen from Valle belongs to B. hookeri, that from Cundinamarca (“Andes Bogotenses,” 4000 ft., Weir s.n., NY) was correctly identified but had been mislabeled and must have originated from New Zealand (Gradstein 2017b). Bazzania tricrenata (Wahlenb.) Lindb. – Ecuador: ZC (Parolly and Kürschner 2004a). Holarctic species; the Ecuadorian material belongs to B. aurescens (LeónYánez et al. 2006). Kurzia G.Martens Microlepidozia (Spruce) Jørg. Plants very small, 0.5–2 cm long and 0.3–0.4 mm wide, dark green to brown, creeping, irregularly pinnate. Branches Frullania-type and Microlepidozia-type on

220

 

Keys and Descriptions

alternate sides, often flagelliform toward the tip; ventral-intercalary branches also present, these sometimes stoloniform. Stems without or with a weak hyalodermis. Leaves very small, transverse, rarely succubous, deeply divided into four subulate (rarely narrow-triangular) lobes, the lobes straight or incurved, 2–3(−4) cells wide at the base, margins entire. Cells thick-walled, without trigones, cuticle papillose; oil bodies finely granular. Underleaves about half the size of the leaves, deeply 2–4lobed, the segments often unequal in length. Dioicous. About 40 species worldwide, three in Colombia and Ecuador. Kurzia is closely related to Lepidozia but differs from the latter genus by (1) plants very small, less than 0.5 mm wide, usually brown; (2) leaves transverse, rarely succubous, deeply divided into 4 subulate (or triangular) lobes; and (3) leaf cells thick-walled, cuticle usually papillose (with small, rounded papillae). A further peculiarity of Kurzia is the occurrence of Frullania-type branches on one side of the stem and Microlepidoziatype branches on the other side. Literature: Fulford (1966 as Microlepidozia), Pócs (2012) 1. Leaf lamina at least 3 cells high (measured from leaf base to the deepest sinus between the lobes). Leaf lobes 3–4 cells wide at the base. Leaves succubous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. brasiliensis 1. Leaf lamina only 1–2 cells high. Leaf lobes 2(−3) cells wide at the base. Leaves transverse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Cells of leaf lobes rectangular, 1.5–3× longer than wide . . . . . . . K. flagellifera 2. Cells of leaf lobes quadrate, about as large as wide . . . . . . . . (K. capillaris) 3 3. Uniseriate part of leaf lobes less than 1/2× lobe length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. capillaris subsp. paramicola 3. Uniseriate part of leaf lobes 1/2–2/3× lobe length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. capillaris subsp. capillaris 1. Kurzia brasiliensis (Steph.) Grolle

Fig. 22H

On shaded, humic soil in shrubby subpáramo, 3250  m. Colombia: Boy, San (Pócs 2012). General distribution: Brazil, Colombia. Kurzia brasiliensis is a very rare South American species that is recognized by quadrifid, succubous leaves with a distinct lamina, being divided to the middle into four triangular lobes. In the other two Kurzia species, the leaves are transverse without distinct lamina, being divided to near the base into four subular lobes. 2a. Kurzia capillaris (Sw.) Grolle subsp. capillarisFig. 22F, G On trunk bases, rotten wood, rock, humus, or damp soil, usually in deep shade, in montane forest areas, scrub, and páramo, 1500–3800 m. Colombia: Ant, Boy, Cau, Cho, Cun, Hui, Met, Nar, Ris, San. Ecuador: Lo, ZC (Nöske et  al. 2003). General distribution: tropical America, Africa. Kurzia capillaris subsp. capillaris is a common, montane Afro-American taxon that usually grows in dense, low, dull green to brownish mats, in strongly shaded and humid sites. The species is distinguished by the transverse, (3-)4-lobed leaves

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with subulate lobes, the lobes being 2 cells wide, made up of (sub)quadrate cells with thick walls and with a smooth or papillose cuticle and typically curved toward the stem. The underleaves are much smaller than the leaves, and the lobes are 2 cells wide (= biseriate) in the lower half and 1 cell wide (= uniseriate) in the upper half, the biseriate portion being maximally half the length of the lobe, often less. Based on the cuticle, Pócs (2012) recognized two varieties in tropical America, K. capillaris subsp. capillaris var. capillaris with a  ±  smooth cuticle and var. verrucosa (Steph.) Pócs with a papillose cuticle. The plants from Colombia and Ecuador normally have a papillose cuticle – even though the degree of papillosity may vary – and belong to var. verrucosa. 2b. Kurzia capillaris subsp. paramicola Pócs On peaty soil in páramo, 3500–3800  m. Colombia (Pócs 2012): Boy, Cun. General distribution: only known from the Eastern Cordillera of Colombia. Kurzia capillaris subsp. paramicola is a rare Colombian endemic that has been collected a few times in the páramos of the Cordillera Oriental. The subspecies differs from subsp. capillaris by the ± straight leaf lobes (not clearly curved toward the stem) and, especially, the longer biseriate portion of the underleaf lobes, being longer than the uniseriate portion. 3. Kurzia flagellifera (Steph.) Grolle On shaded rotten wood, leaf litter, soil, and rock in montane forest, 1600–3750 m. Colombia: Cun, Mag, Qui (Lopez s.n., HUQ), Ris. Ecuador: Az, Lo, ZC (SchäferVerwimp et al. 2013a). General distribution: tropical America. Kurzia flagellifera is very similar to K. capillaris but differs by rectangular leaf lobe cells, 1.5–3× longer than wide. Lepidozia (Dumort.) Dumort. Plants small to rather large but always narrow, 1–10 cm long and 0.5–1.5 mm wide, typically pale green, deep green to greenish-brown, rarely dark brown, creeping, sometimes long pendent, 1–3-pinnate. Branches mostly Frullania-type, often ending in slender flagella; ventral-intercalary branches also present, these sometimes stoloniform. Leaves small, incubous, usually concave, divided into 4(−6) equal or unequal lobes, the lobes plane or incurved, margins entire or with a few teeth. Cells with uniformly thickened walls, trigones ± lacking, cuticle usually smooth; oil bodies finely granular. Underleaves about half the size of the leaves, rarely larger, divided into 4 equal lobes. Dioicous, rarely monoicous. A genus of about 75 species worldwide, 13 in Colombia and Ecuador. Lepidozia is readily recognized by the whitish-green to greenish-brown, 1–3-pinnate plants with small, concave leaves divided into 4–6 lobes. The branches become progressively narrower toward the tips and often end in a flagellum.

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Keys and Descriptions

Literature: Fulford (1966), Pócs (in prep.) 1. Margins of leaf lobes pinnately ciliate by long multicellular teeth. Plants resembling Trichocoleaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pinnaticruris 1. Margins of leaf lobes entire or irregularly toothed. Plants not resembling Trichocoleaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves very small, scale-like, less than 0.4 mm long, highly distant . . . . . . . 3 2. Leaves larger, not scale-like, more than 0.4 mm long, distant to imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 3. Leaves appressed to the stem, quadrifid to 1/4–1/3. . . . . . . . . . . . . . . . . . . . . 4 3. Leaves spreading away from the stem, quadrifid to 1/2 . . . . . . . . . . . . . . . . . . 5 4. Leaves concave, leaf tips incurved, almost touching the stem. Plants common, 2–3-pinnate, without dichotomous branching . . . . . . . . . . . . . . . . L. incurvata 4. Leaves ± flat, leaf tips not incurved. Plants rare, 1-pinnate, the primary branches frequently dichotomously branched . . . . . . . . . . . . . . . . . . . . . L. subdichotoma 5. Plants hanging from trees, pale green to light brown, common in the West Indies, rare in the Andes. Stems to 15  cm long, with very long branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. patens 5. Plants creeping, in small pale green mats. Stems 1–6  cm long, with short branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. squarrosa 6. Leaves deeply concave, dorsally strongly expanded, the expanded part almost as wide as the rest of the leaf. Rather robust plants from high elevation . . . . . 7 6. Leaves concave or not, dorsally not strongly expanded . . . . . . . . . . . . . . . . . . 8 7. Leaf lobes ending in a row of narrow-rectangular cells, cells in the leaf tips 3–4× longer than wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. andicola 7. Leaf lobes ending in a row of quadrate to subrectangular cells, cells in the leaf tips 1–2× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. auriculata 8. Leaves shallowly lobed to 1/4(−1/3) of leaf length. Leaf insertion sublongitudinal. Plants monoicous, frequently fertile, widespread in the Holarctic, rare in the Andes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. reptans 8. Leaves more deeply lobed. Leaf insertion subtransverse. Plants dioicous, frequently sterile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Leaf margins undulate. Plants robust, brown, leaves more than 1  mm long, deeply split into 4 equal lobes. Plants rare (high Andes) . . . . . . . . . . L. alstonii 9. Leaf margins not undulate, plants different. . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Dorsal leaf margin toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10. Dorsal leaf margin entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11. Tips of stem leaves strongly incurved, usually touching the stem in at least part of the leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. macrocolea 11. Tips of stem leaves erect or slightly incurved, not touching the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cupressina 12. Leaf lobes ending in a long row of narrow-rectangular cells, cells in the leaf tips 3–4× longer than wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. inaequalis 12. Leaf lobes ending in a row of shorter cells, cells in the leaf tips 1–2× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Tips of stem leaves strongly incurved, usually touching the stem in at least part of the leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. dendritica

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13. Tips of stem leaves erect or slightly incurved, not touching the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cupressina 1. Lepidozia alstonii Fulford

Fig. 22I

On bark and rock in upper montane forest and páramo, 2100–3500 m. Colombia: Boy, Ris, San. Ecuador: Pi (Fulford 1966). General distribution: northern Andes. Lepidozia alstonii is a rare northern Andean páramo species, distinguished by the robust, brownish plants (to 5 cm long) with deeply 4-lobed leaves, undulate lobe margins, and 1–2 teeth near the leaf base. The underleaves are almost as large as the leaves and similar. The species may be confused with Lepicolea due to its rather large size but is readily separated from Lepicolea by the almost entire margins of the leaf lobes (densely ciliate-laciniate in Lepicolea) and the leaf cells with uniformly thickened walls, without trigones (with large trigones in Lepicolea) (Fulford 1966). 2. Lepidozia andicola Beauverd Lepidozia appendiculata Steph.

Fig. 22L

On tree bases, logs, humus, and soil in upper montane forest and páramo, 2650–3600  m. Colombia: Boy, Cau, Cun, Val. Ecuador: Az (Fulford 1966). General distribution: tropical Andes. Lepidozia andicola is very similar to L. auriculata and grows in the same general habitat but differs by more strongly elongate leaf tip cells (3–4× longer than wide). The species seems to be rarer than L. auriculata. 3. Lepidozia auriculata Steph.

Fig. 22J, K

On tree bases, logs, humus, soil, and rock in upper montane forest, Polylepis forest, and páramo, occasionally at lower elevation in montane rainforest, (2100–) 3000–4050 m. Colombia: Ara, Boy, Cau, Cun, Met. Ecuador: Az, Co, Lo, Na, Pi, ZC. General distribution: northern Andes (Colombia to Peru). Lepidozia auriculata is a rather robust, high-Andean species that is recognized by the deeply concave leaves with a strongly expanded dorsal part of the leaf lamina. The expanded part is undivided and almost as wide as the rest of the leaf and sometimes has an auricle (toothed or entire) at the dorsal base. Lepidozia auriculata is close to L. andicola which occurs in the same habitat, but the leaf tip cells in L. andicola are more strongly elongate, 3–4× longer than wide (maximally 2× longer than wide in L. auriculata). The correct author citation for the species should be “Mitt. ex Steph.” or, in shortened version, “Steph.” (not “Mitt.” as in Söderström et al. 2016). 4. Lepidozia cupressina (Sw.) Lindenb. Lepidozia bogotensis Steph. Lepidozia heterophylla Steph. Lepidozia brasiliensis Steph. (fide T. Pócs) Lepidozia caespitosa Spruce (fide T. Pócs) Lepidozia lindigiana Steph. (fide T. Pócs)

Fig. 23A–C

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Fig. 23 (A–C) Lepidozia cupressina. (A) Habit, ventral view. (B) Leaf. (C) Habit with short ventral female branch, ventral view. (D) Lepidozia squarrosa, leaves and underleaf, 60×. (E) Lepidozia subdichotoma, branching pattern. (F and G) Micropterygium pterigophyllum. (F) Habit, ventral view. (G) Stem portion with leaf, ventral view. (H) Micropterygium leiophyllum, habit in ventral view, 75×. (I) Micropterygium carinatum, habit in ventral view, 75×. (J) Micropterygium duidae, habit in ventral view, 75×. (K) Micropterygium trachyphyllum, habit in ventral view, 75×. (D, E, H–K from Fulford 1966; rest from Gradstein et al. 2001)

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Lepidozia muenchiana Steph. (fide T. Pócs) Lepidozia wallisiana Steph. (fide Gradstein and Uribe 2016a) Lepidozia moritziana Steph., syn. nov. Type: Venezuela, Mérida, Moritz 74 (G-00069699!) Lepidozia tenuicuspis Steph., syn. nov. Type: Ecuador, Morona Santiago, Gualaquiza, “in silva ad rivum Sapotilla,” 9 Aug 1909, Allioni s.n., Bryotheca Levier 6561, c.spor. (lectotype, designated here, G-00112407!); ibid., “in silva ad rivum Salado,” May 1909, Allioni s.n., Bryotheca Levier 6559 (syntype G-00043706!) Lepidozia tunguraguae Steph., syn. nov. Type: Colombia, Lindig s.n. (G-00069663!) [not from Tungurahua, Ecuador] On bark, logs, rock, humus, and soil in montane forests and páramo, 1000–4500 m. Colombia: Ant, Ara, Boy, Cau, Ces, Cho, Cun, Hui, Mag, Met, Nar, NSa, Qui, Ris, San, Tol. Ecuador: Az, Lo, MS, Na, Pa, Pi, Tu, ZC. General distribution: widespread in tropical America and Africa, also in western Europe. Lepidozia cupressina is a common and widespread Afro-American species that has been described under many different names. It is distinguished by the spreading, deeply 4-lobed leaves (0.5–0.9  mm long) with acute to finely acuminate tips and ± entire margins (occasionally with a few small tooth on the dorsal margin). The leaves are concave and asymmetrical with the dorsal side somewhat expanded, and the lobe tips are straight or slightly curved, never strongly incurved as in L. dendritica and L. macrocolea. The lobes are equal or unequal in length, and the uniseriate tips are 1–7 cells long and consist of quadrate to subrectangular cells that are 1–2× longer than wide. Prof. T. Pócs has kindly confirmed the synonymy of L. brasiliensis, L. caespitosa, L. lindigiana, and L. muenchiana under L. cupressina. Judging from the description and illustration in Fulford (1966), L. wallisiana is a further synonym. Lepidozia pseudocupressina Schiffn. (= L. allionii Steph.), described from southern Brazil and recorded from Ecuador (Az, MS), resembles robust phenotypes of L. cupressina but differs in the ciliate perianth mouth (cilia 2–5 cells long). In L. cupressina the perianth mouth is usually subentire. Possibly, L. pseudocupressina is a variety of L. cupressina. 5. Lepidozia dendritica Spruce Lepidozia peruviensis Steph., syn. nov. Type: Peru, Sandia, Weberbauer 820 (G-00112583!) Lepidozia quitensis Steph. On tree bases, rock, humus, and soil in montane forests and in páramo, 750–3700 m. Colombia: Cun, Hui, Qui, San (van der Hammen 1678, det. T. Pócs, U), Tol; in addition recorded from Boy, Cau, Mag and Nar as L. peruviensis. Ecuador: Lo (Burghardt 6935, QCA), MS, Pa, Tu, ZC (Burghardt 6988, QCA). General distribution: northern Andes. Lepidozia dendritica resembles L. macrocolea but is more robust, and the dorsal leaf margin is entire.

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Keys and Descriptions

6. Lepidozia inaequalis (Lehm. & Lindenb.) Lehm. & Lindenb. Fig. 22M, N Lepidozia cupressina (Sw.) Lindenb. var. tenuicuspis Spruce (Lepidozia tenuicuspis [Spruce] Spruce) Lepidozia tenuicola Spruce On bark, rock, and rotten logs in montane forest, 750–2000  m. Ecuador: Az (Spruce 1885), MS.  General distribution: southeastern Brazil, Central Andes (southern Ecuador to Bolivia). Lepidozia inaequalis is a mainly Brazilian species recognized by the finely acuminate leaf lobes terminating in a longly uniseriate tip, made up of a row of 4–8 narrowly rectangular cells that are 3–4× longer than wide. The species is near to L. cupressina, but in the latter the cells in the leaf tips are shorter, 1–2× longer than wide. 7. Lepidozia incurvata Lindenb.

Fig. 22O, P

On bark in montane cloud forest and páramo, 1800–3600 m. Colombia: Ara, Boy, Cas, Cau, Cho, Cun, Hui, Met, NSa, Qui, Ris, San. Ecuador: Ca, ZC. General distribution: Central America, tropical Andes, southern Brazil. Lepidozia incurvata is a common Andean species that is readily distinguished by the thread-like, 2–3-pinnately branched plants with minute, distant leaves. The leaves are shallowly 4-lobed (to 1/3) and strongly concave with incurved tips. 8. Lepidozia macrocolea Spruce Fig. 22Q Lepidozia armata Steph. (fide T. Pócs) Lepidozia karstenii Steph. var. standleyi Herzog, syn. nov. (fide T. Pócs) On tree bases, rock, humus, and soil in montane forests and páramo, 1500–4000 m. Colombia: Ara, Boy, Cal, Cas, Cau, Cun, Hui, Mag, Met, Ris, Tol. Ecuador: Ca, Lo, Na, Pi, Tu, ZC. General distribution: tropical America. Lepidozia macrocolea is a common neotropical high-montane species that is distinguished by the deeply concave leaves with strongly incurved tips, which are often touching the stem, and with toothed lateral margins. The species is morphologically close to L. dendritica, but in the latter species, the dorsal leaf margin is entire. The species may also be confused with L. cupressina, but in the latter species, the leaf tips are plane or only slightly incurved. 9. Lepidozia patens Lindenb. Lepidozia commutata Steph. On tree trunks and branches in montane forests, 700–3100 m. Colombia: Boy, Cho, Mag, NSa, Ris. Ecuador: Lo (Arnell 1962). General distribution: widespread and common and widespread in the West Indies, also in Central America; rare in the northern Andes. Lepidozia patens is a characteristic Caribbean species that is recognized by the long pendent, thread-like stems with very small, distant leaves that are spreading away from the stem and are deeply divided into 4 subequal, diverging lobes with entire margins. The leaves of Lepidozia patens are quite similar to those of

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L. squarrosa, and the two species may be confused but can be separated by their different growth habit, L. patens being a pendent plant with long stems and branches, whereas L. squarrosa is a creeping plant with short stems and branches. In its pendent growth and minute leaves, L. patens resembles L. incurvata and L. subdichotoma, but the latter two species have appressed leaves. 10. Lepidozia pinnaticruris Steph. On bark in lowland and montane rainforests, 80–2400 m. Colombia: Cho, Nar, Ris, Val. Ecuador: Ch (Fulford 1966), Na (Cerro Huancamayos, Oellgaard et al. 35893, AAU, PC). General distribution: northern Andes and Chocó biogeographic region. Lepidozia pinnaticruris is a robust Chocó-northern Andean species that is readily distinguished by the peculiar, deeply 4-fid leaves with (2-)pinnately ciliate lobes. Each leaf lobe has long, subopposite, simple or branched ciliate teeth on its margins. The plants are to 10  cm long and 1.5  mm broad and are regularly bipinnately branched. According to Fulford (1966): “this large and beautiful species looks like a large Trichocolea rather than a Lepidozia. However, in Trichocolea the leaves are succubous and the cilia of the leaves and underleaves are made up of very long cells, two or three times as long as the cells of the cilia of L. pinnaticruris.” In addition, the cuticle of leaf cells is striate-papillose in Trichocolea and smooth in L. pinnaticruris. 11. Lepidozia reptans (L.) Dumort.

Fig. 22R

On tree bases, logs, tree stumps, rock, and soil in upper montane forest, 2600–3100 m. Colombia: Ant, Cho, Cun, Ris, Tol. General distribution: common and widespread in the temperate regions of the Northern Hemisphere; scattered in Central America, the West Indies, Venezuela, and Colombia; also in tropical Asia. Lepidozia reptans is a widespread Holarctic species that extends southward to the northern Andes. It is the only monoicous species in the neotropics and frequently fertile. The species is readily recognized by the almost longitudinally inserted leaves, which are slightly longer than wide and are rather shallowly divided (to maximally 1/3 of leaf length) into 3 or 4 lobes. In other neotropical species, the leaves are subtransverse and are usually more deeply divided (except in L. incurvata). The sinus between the underleaf lobes in L. reptans is U-shaped. 12. Lepidozia squarrosa Steph.

Fig. 23D

On bark, logs, and rock in montane forest, 1800–3600 m. Colombia: Ara, Boy, Cas (Gradstein 12703, PC), Cun, Mag, Met, San. Ecuador: Lo (Schäfer-Verwimp et  al. 2013a), ZC (páramo of Saraguro and Yacuambi, Benitez & Gradstein s.n., HUTPL). General distribution: northern Andes, Central America (Costa Rica, Panama). Lepidozia squarrosa is a tiny northern Andean species with distant, squarrose leaves that are little wider than the stem. In its small leaves, the species is similar to L. patens; differences are discussed under the latter species.

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Keys and Descriptions

13. Lepidozia subdichotoma Spruce

Fig. 23E

On bark, humus, and soil in montane forests, 1200–3150 m. Colombia: Cauca (Cleef 4996, det. T. Pócs, U), Hui (Cleef 5059, det. T. Pócs, U). Ecuador: Az, Pa, Lo, Tu, ZC.  General distribution: tropical Andes (Colombia to Bolivia), Costa Rica. Lepidozia subdichotoma is an uncommon Andean species that resembles L. incurvata by the thread-like, filiform stems with very small, distant, appressed leaves. The two species differ in branching pattern: L. incurvata is 2–3-pinnate with rather short branches and no dichotomies, while L. subdichotoma is laxly 1-pinnate with rather long primary branches that are frequently dichotomously branched. Further Record Lepidozia serpens Spruce – Ecuador: Tu (Fulford 1966, p. 213). The identity of this species needs study. Micropterygium Gottsche Plants small- to medium-sized, 0.5–2  mm wide, light yellow-green to brown, ascending from a branched rhizome system, irregularly branched. Branches intercalary (lateral and ventral), often stoloniform. Stems with a thick-walled, brown, 1–2-layered cortex surrounding smaller, thin-walled medulla cells. Leaves transverse or succubous, insertion line extending to the dorsal stem-midline, orbicular to lanceolate, ± concave to complicate-bilobed and keeled with a wing at least in the upper half, the wing sometimes reduced, apex undivided or very short-bifid, margins entire or toothed. Cells usually small, 10–20 μm wide, mostly thick-walled, plane or mammillose, and sometimes with a large papilla on the outer surface, cuticle smooth or finely papillose; oil bodies granular. Underleaves present, smaller to almost as large as the leaves. Dioicous. Gametoecia as in the family. A neotropical genus (ca. 20 spp.), ten species in Colombia and Ecuador. Micropterygium is recognized by the peculiar, folded, transverse to succubous leaves with a more or less winged keel. The rigid stems, which are little branched and stolon-like at the base, and the small, ± thick-walled leaf cells are further characteristics of this genus. The underleaves are quite variable in size, and in some species they gradually decrease in size toward to the stem apex. Literature: Fulford (1966), Schuster (2000a), Uribe and Linares (2015) 1. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf margins toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Underleaves distinctly decreasing in size toward the upper portion of the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. carinatum 2. Underleaves not decreasing in size toward the upper portion of the stem . . . . 3 3. Plants very small, 0.4–0.8 mm wide. [Leaf apex acute. Leaves with a low wing in the upper half. Leaf cells plane. Guyana Highlands, found once in the Cordillera del Condor]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. duidae

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3. Plants larger, more than 1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaf apex rounded. Leaves with a broad wing from base to apex. Leaf cells not mammillose, without large papillose projection on the outer surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lechleri 4. Leaf apex acute. Leaves with a narrow wing in the upper half. Leaf cells mammillose, part of the cells with a large papillose projection on the outer surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. campanense 5. Underleaves distinctly decreasing in size toward the upper portion of the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Underleaves not or hardly decreasing in size toward the upper portion of the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Leaf cells strongly mammillose and often with a large papillose projection on the outer surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. parvistipulum 6. Leaf cells not mammillose, without large papillose projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pterygophyllum 7. Leaf cells narrowly elongate, ca. 4–9× as long as wide . . . . M. longicellulatum 7. Leaf cells (sub)quadrate, 1–1.5× as long as wide . . . . . . . . . . . . . . . . . . . . . . 8 8. Leaf cells strongly mammillose (dorsal surface), often with a large papillose projection on the outer surface . . . . . . . . . . . . . . . . . . . . . . . . M. trachyphyllum 8. Leaf cells not mammillose, without large papillose projection on the outer surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Leaves deeply concave, complicate-bilobed. Cells in the upper part of leaf 10–16 μm in diameter. Plants montane . . . . . . . . . . . . . . . . . M. reimersianum 9. Leaves plane or slightly concave in the upper half, not complicate-bilobed. Cells in the upper part of leaf 18–20 μm. Plants of lowlands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. leiophyllum 1. Micropterygium campanense Reimers On tree bases, logs, rock, and humid soil in montane rainforest, ca. 1000–2200 m. Ecuador: ZC (Nöske et  al. 2003). General distribution: Peru, Ecuador, Venezuela (Guayana Highland), southeastern Brazil (Minas Gerais). Micropterygium campanense is a rare tropical South American species that is characterized by deeply concave leaves with entire margins (occasionally with a small tooth), an acute apex, a narrow wing in the upper half, and more or less mammillose leaf cells. Part of the leaf cells, especially those near to the wing in the uppér half of the leaf, may bear a large papillose projection on the outer surface. The underleaves are large, broader than the stem, and do not decrease in size toward the stem apex. Micropterygium campanense is somewhat similar to M. trachyphyllum, but the latter species differs in having toothed leaf margins. 2. Micropterygium carinatum (Grev.) Reimers Fig. 23I Micropterygium exalatum Steph., syn. nov. Type: Dominica, without locality, Elliott 2233 (G-00112613!) On bark, logs, and rock in submontane and montane rainforest, (350–) 500–2000 m. Colombia: Ant, Cho, Nar, Ris, San. General distribution: West Indies, Costa Rica, Colombia, Venezuela (Guayana Highland).

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Keys and Descriptions

Micropterygium carinatum is a characteristic Caribbean species that is distinguished by deeply concave, complicate-bilobed leaves with entire margins, acute tips, and a short wing in the upper half, leaf cells not mammillose and underleaves usually decreasing in size toward the stem apex. As already suggested by Fulford (1966) and Gradstein (1989a), M. exalatum does not differ in any respect from M. carinatum and is a synonym. 3. Micropterygium duidae Reimers On moist soil in rather low and open montane scrub, ca. 2300 m. Ecuador: ZC (Gradstein and Benitez 2014c). General distribution: Venezuela (Guayana Highland), southern Ecuador (Cordillera del Condor). Micropterygium duidae is a very distinct species that is recognized by the very small, brownish plants (to maximally 0.8  mm wide) with deeply concave leaves with a low wing in the upper half, entire margins, and large underleaves, which are only little smaller than the leaves. The leaf cells are usually finely verruculose and not mammillose. The species is known from a few localities in the Guayana Highland and from Cordillera del Condor (Cerro Plateado). The distribution of M. duidae lends further support to the floristic similarity of the Guayana Highland and the Condor region (see also Neill 2005). 4. Micropterygium lechleri Reimers On bark, logs, and soil in submontane and montane rainforests, to 1700  m. Colombia: Ama, Boy, Caq, Met, San. Ecuador: ZC (Schäfer-Verwimp et  al. 2013a). General distribution: tropical Andes (Colombia to Bolivia), Brazil. Micropterygium lechleri is a rare Andean species that is characterized by concave leaves with entire margins, rounded tips, and a long wing from base to apex, leaf cells not mammillose and underleaves large, not decreasing in size toward the stem apex. 5. Micropterygium leiophyllum Spruce

Fig. 23H

On rotten wood and trees in Amazonian rainforest, 100–250 m. Colombia: Ama, Caq, Gua, Put, Vau; to be expected in Ecuador. General distribution: Amazonia, Costa Rica. Micropterygium leiophyllum is a common Amazonian species that is readily distinguished by plane to slightly concave leaves with toothed margins and a short wing in the upper half, leaf cells not mammillose, ca. 18–20 μm in diameter in the upper part of the leaf, and underleaves large, as long as wide or longer than wide, not significantly decreasing in size toward the stem apex. 6. Micropterygium longicellulatum Uribe & E.L.Linares On trees and logs in montane cloud forests, 1900–2400  m. Colombia: Cho (Uribe and Linares 2015). General distribution: only known from Colombia (Chocó). Micropterygium longicellulatum is only known from Cerro del Torrá (Chocó) and stands out by strongly elongate leaf cells, about 4–8× longer than wide (Uribe

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and Linares 2015). The plants are robust, 3–4  mm wide; the leaves are strongly concave and complicate-bilobed, ovate-lanceolate, with toothed margins, a narrow wing, and an acute apex, and the underleaves are large, ca. 2/3× the length of the leaves and not decreasing in size to the stem apex. 7. Micropterygium parvistipulum Spruce On rotten wood and trees in lowland and submontane rainforest, 100–700  m. Colombia: Ama, Caq, Put, Vau (Campos et  al. 2015). General distribution: Amazonia, Guianas, Panama (Dauphin et al. 2015). Micropterygium parvistipulum is an Amazonian species characterized by complicate-bilobed leaves with toothed margins and a long wing, and underleaves decreasing in size toward the stem apex. The leaf cells are mammillose (dorsal surface) and often bear a large papillose projection on the outer surface. The species seems to be close to M. trachyphyllum (see under the latter). 8. Micropterygium pterygophyllum (Nees) Trevis. Micropterygium pterygophyllum var. gracile Reimers

Fig. 23F, G

On trees, logs, rock, and soil in lowland and submontane rainforests, from sea level to 700 m. Colombia: Ama, Caq, Cho, Put, Vau (Campos et al. 2015); to be expected in Ecuador. General distribution: Amazonia, southeastern Brazil. Micropterygium pterygophyllum is very similar to M. leiophyllum but differs in the underleaves distinctly decreasing in size toward the stem apex. The leaf cells are not mammillose. The species is very common in the lowland rainforests of upper Amazonia, where it occurs on almost every rotten log (W. R. Buck, pers. comm.). 9. Micropterygium reimersianum Herzog On soil in montane rainforest, ca. 1500–2000 m. Colombia: Cho, San. Ecuador: MS (El Quimi reserve, among the type of Riccardia verticillata, Zapata 541a, QCA, PC), ZC. General distribution: Venezuela (Guayana Highland), Colombia, Ecuador, eastern Brazil (Bahia). Micropterygium reimersianum is a rare montane northern South American species characterized by deeply concave, complicate-bilobed leaves with toothed margins, acute apex, a narrow wing in the upper half, leaf cells not mammillose and without papillose projection on the outer surface, and underleaves large, suborbicular, about as long as wide, not decreasing in size toward the stem apex. The species approaches M. leiophyllum but clearly differs from the latter in the deeply concaveconduplicate leaves. Moreover, M. leiophyllum is a lowland species, whereas M. reimersianum grows in montane forest, above 1000 m. 10. Micropterygium trachyphyllum Reimers

Fig. 23K

On tree bases, logs, rock. and humus in lowland and montane rainforests, from sea level to 1900 m. Colombia: Ama, Cun, Hui, San, Vau. Ecuador: ZC (Parolly et al. 2004). General distribution: widespread in tropical America; very common in Amazonia and the Guianas, more scarce in the Andes.

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Keys and Descriptions

Micropterygium trachyphyllum is a widespread neotropical species that is characterized by concave, complicate-bilobed leaves with toothed margins, an acute apex, a long wing, and mammillose leaf cells frequently with a large papillose projection on the outer surface. The underleaves are large, broader than the stem, and are not decreasing in size toward the stem apex. Micropterygium trachyphyllum is quite similar to M. parvistipulum, but the underleaves in M. parvistipulum are strongly decreasing in size toward the stem apex. Micropterygium trachyphyllum also approaches M. leiophyllum, but the leaves in the latter species are more flat, not distinctly complicate-bilobed, the keel and wing are less well developed, and the leaf cells in M. leiophyllum are not mammillose and without papillose projection. Moreover, M. leiophyllum is largely restricted to Amazonian rainforest. Monodactylopsis (R.M.Schust.) R.M.Schust. Regredicaulis Fulford Plants minute, creeping, filamentous, ca. 0.2  mm wide, whitish, scarcely branched. Stems very thin, 2 cells wide dorsally, made up of 4–5 cells in cross section, with a hyalodermis. Branches ventral-intercalary. Leaves rudimentary, consisting of 1–2 cells capped by a slime papilla. Underleaves made up of 2 very small, adjacent cells. Monoicous. Perianth long, 3-keeled, mouth with a few long lacinia. A neotropical genus with a single species, characterized by the minute, filamentous plants with reduced leaves and underleaves. Literature: Fulford (1966 as Regredicaulis), Schuster (2000a). 1. Monodactylopsis monodactyla (Spruce) R.M.Schust. Regredicaulis monodactylus (Spruce) Fulford Monodactylopsis minima (R.M.Schust.) R.M.Schust.

Fig. 24B

On decaying wood, soil, and tree bases in lowland rainforest, from sea level to 300 m. Colombia: Ama, Caq, Cho, Put, Ris, Vau (Campos et al. 2015). General distribution: scattered in northern South America, Costa Rica, and the West Indies. Monodactylopsis monodactyla is a minute neotropical lowland rainforest species characterized by the less than 0.2-mm-wide whitish plants with rudimentary leaves, consisting of a row of 1–2 cells capped by a slime papilla, and with stems only 2 cells wide. The plants grow mostly on rotten wood and are easily overlooked because of their very small size. Because of the extreme reduction of the gametophyte, the taxonomic position of M. monodactyla is enigmatic, and the species has been placed in four different genera. Mytilopsis Spruce Plants small, ca. 1 mm wide, whitish-green to brown, ascending from a branched rhizome system, irregularly branched. Branches ventral-intercalary, often stoloniform. Stems with a very strongly thick-walled, yellow brown, 1-layered cortex surrounding smaller, thin-walled medulla cells. Leaves transverse, with a long insertion

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Fig. 24 (A) Mytilopsis albifrons, habit in ventral view. (B) Monodactylopsis monodactyla, habit in dorsal view. (C) Paracromastigum granatense, leaves and underleaf, 40×. (D and E) Paracromastigum pachyrhizum. (D) Habit, ventral view. (E) Midleaf cells. (F) Pseudocephalozia quadriloba, habit in ventral view. (G–I) Pteropsiella serrulata. (G) Habit with ventral intercalary branch, ventral view. (H) Thallus margin, showing long papilla, 200×. (I) Portion of thallus with gynoecium. (J) Telaranea diacantha, habit with gynoecium in dorsal view. (K and L) Telaranea nematodes. (K) Habit, ventral view. (L) Leaf. (C from Fulford 1966; rest from Gradstein et al. 2001)

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Keys and Descriptions

line extending to the dorsal stem-midline and almost to the ventral midline, widespreading and boat-shaped, sharply keeled, the keel with or without a rudimentary wing, apex bifid to maximally 1/4 of leaf length or less, margins crenulate-denticulate, bases broadly auriculate, clasping the stem. Cells small, 15–20  μm, thickwalled, mammillose, cuticle papillose. Underleaves lacking. Dioicous. Gametoecia as in the family. A neotropical genus with a single species, related to Micropterygium but differing by the symmetrically complicate leaves with 2 equal lobes that are clasping the stem, the shortly bifid leaf apex, the absence of underleaves on mature shoots, and the purely ventral branching. Literature: Fulford (1966), Schuster (2000a) 1. Mytilopsis albifrons Spruce

Fig. 24A

On soil and rock in moist montane rainforest, 1200–2650 m. Colombia: Ant, Ris, San. Ecuador: ZC (Nöske et al. 2003). General distribution: scattered in tropical America. Mytilopsis albifrons is readily recognized by the small, whitish-green to brownish plants without underleaves and with transverse, folded, boat-shaped leaves, which are very shortly bifid and divided into 2 equal lobes. Neolepidozia Fulford & J.Taylor Telaranea sect. Neolepidozia (Fulford & J.Taylor) J.J.Engel & G.L.Merr. Plants delicate, to 1  mm wide, pale green, 1–3-pinnate. Branches mostly Frullania-type or Bazzania-type. Stems with a hyalodermis. Leaves incubous, partially divided into 3–4 almost uniseriate lobes. Cells usually large, longer than wide, walls uniformly thin, cuticle smooth; oil bodies finely granular. Underleaves similar to leaves but smaller. Dioicous or monoicous. Perianths longly cylindrical, mouth short-toothed or with long lacinia. A mostly southern-temperate genus of about 30 species (Söderström et al. 2016), one in Ecuador. Neolepidozia is recognized by the delicate, pinnate plants with leaves split to about half into 3–4 somewhat diverging filaments, large thin-walled leaf cells, underleaves about half the size of the leaves, and stems with a distinct hyalodermis. The genus was included in Telaranea by Schuster (2000a) and Engel and Smith Merrill (2004) but reinstated by Cooper (2013) based on molecular evidence. Neolepidozia is readily separated from Telaranea by the presence of a discrete leaf lamina. Literature: Schuster (2000a as T. subg. Neolepidozia), Engel and Smith Merrill (2004 as T. sect. Neolepidozia), Cooper (2013) 1. Neolepidozia rectangularis (R.M.Schust.) E.D.Cooper Telaranea rectangularis R.M.Schust.

Fig. 25A, B

On humus in upper montane scrub, 2550 m. Ecuador: ZC (Schäfer-Verwimp et al. 2013a); to be expected in Colombia. General distribution: northern Andes (Venezuela, Ecuador).

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Fig. 25 (A and B) Neolepidozia rectangularis. (A) Leaf, 86×. (B) Underleaf, 86×. (C and D) Zoopsidella integrifolia. (C) Habit, ventral view. (D) Leaf apex with two large slime papillae. (E) Zoopsidella serra, habit in dorsal view, 70×. (F–H) Clasmatocolea vermicularis. (F) Habit, lateral view. (G) Habit, ventral view. (H) Underleaf. (I and J) Cryptolophocolea martiana var. martiana. (I) Habit with gynoecium, dorsal view. (J) Habit, ventral view. (K) Cryptolophocolea connata, connate underleaves, 20×. (L) Cryptolophocolea guadeloupensis, leaves in dorsal view, 20×. (M) Heteroscyphus contortuplicatus, habit in dorsal view, 20×. (N and O) Heteroscyphus thraustus. (N) Leaves, dorsal view, 20×. (O) Cells at leaf margin, 275×. (A, B redrawn from Schuster 2000a; E from Fulford 1968; K–O from Fulford 1976; rest from Gradstein et al. 2001)

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Keys and Descriptions

Neolepidozia rectangularis is a rare northern Andean species that is readily distinguished by the delicate, pale green, densely 2-pinnate, feather-like plants with leaves split to about the middle into 3–4 somewhat diverging filaments. The filaments are 4–6 cells long and 1 cell wide except at the base, and the undivided leaf lamina is 4–5 cells high. The plants from Ecuador are female; the perianth is longcylindrical, the mouth is short-toothed, and the teeth consist of 1–2 narrowly rectangular cells. Paracromastigum Fulford & J.Taylor Bonneria Fulford & J.Taylor, Leucosarmentum Fulford Plants very small, 1–1.5  mm wide, pale green to yellow or greenish-brown, creeping or ascending from a stoloniform base, irregularly branched. Branches terminal and ventral, sometimes flagelliform. Stems with a weak hyalodermis. Leaves suberect to wide-spreading, transverse or succubous, plane or concave, ovate to elliptical, narrowed to the base, 2–3-lobed to leaf middle or more, the lobes triangular to lanceolate. Cells quadrate to rectangular, with thin or uniformly thickened walls, cuticle finely papillose; oil bodies lacking or finely granular. Underleaves similar to the leaves, slightly smaller. Gametoecia as in the family. A southern-temperate genus of about 15 species, 2 in Colombia and Ecuador. Paracromastigum is recognized by the small, pale green to greenish-brown plants with deeply 2–3-lobed, succubous to almost transverse leaves and large, bifid underleaves. The plants may superficially resemble Telaranea, but the underleaves are larger and the leaf lobes are at least in part more than 1 cell wide. Like in the family, the rhizoids are developed in tufts near the bases of the underleaves, and the gametoecia are produced on short ventral branches. Literature: Fulford (1966 as Bonneria, Leucosarmentum, Paracromastigum), Schuster (2000a), Gradstein and Costa (2003) 1. Leaf lobes diverging outward, narrowly triangular . . . . . . . . . . . P. granatense 1. Leaf lobes erect or somewhat connivent, not diverging, ovatelanceolate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pachyrhizum 1. Paracromastigum granatense (Gottsche) R.M.Schust. Bonneria granatensis (Gottsche) Fulford & J.Taylor

Fig. 24C

On soil on road cut banks in montane forest environments, 300–3300  m. Colombia: Cun. Ecuador: EO (Reserva Buenaventura, 300–400 m, Benitez et al. 1316, HUTPL), ZC (Schäfer-Verwimp et  al. 2013a). General distribution: northern Andes, Costa Rica. Paracromastigum granatense is a rare and very delicate northern Andean species that is distinguished by the almost transverse leaves divided to about half their length into 2–3 narrowly triangular, diverging lobes. The underleaves are similar to the leaves but about half to 3/4 their size. The plants superficially resemble Telaranea but the leaf lobes are more than 1 cell wide.

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2. Paracromastigum pachyrhizum (Nees) Fulford Fig. 24D–E Paracromastigum bifidum (Steph.) R.M.Schust. (Leucosarmentum bifidum [Steph.] Fulford) Leucosarmentum portoricense Fulford On steep, moist earth and on rock, frequently on road cut banks, 1100–3300 m. Colombia: Cun, Ris. Ecuador: Az, Lo, ZC.  General distribution: tropical America. Paracromastigum pachyrhizum is a characteristic pale green species of shaded, road cut banks in moist montane forest areas. Characteristics are the ovate to elliptical, succubous leaves, which are divided to about half their length into 2–3 erect or somewhat connivent, ovate-lanceolate lobes. The underleaves are similar to the leaves but about half their size, and the leaf cells vary from quadrate to rectangular, becoming longer in more elongate leaves. The variation in leaf shape and cell size can be observed within single populations. Paracromastigum bifidum, described from the West Indies and often accepted as a separate species, is a phenotype of P. pachyrhizum with rather narrow, elongate-elliptical leaves and rectangular cells (Gradstein and Costa 2003). Pseudocephalozia R.M.Schust. Plants small, 0.3–2 mm wide, light green to pale brown, ascending to erect from a stoloniform base, irregularly branched. Branches mostly intercalary (lateral and ventral), rarely terminal (Frullania-type, Microlepidozia-type). Stems with a hyalodermis. Leaves transverse, insertion line extending to the dorsal stem-midline, concave, obtrapezoid, quite variable in size, shallowly 2–4-lobed (to 1/4 of leaf length or less), segments triangular, margins entire, lamina in lower half of leaf 1–5-stratose. Cells ± thin-walled, cuticle smooth to weakly papillose; oil bodies coarsely granular, 2–3 per cell, or lacking. Underleaves similar to the leaves but usually smaller. Dioicous. Gametoecia as in the family. A southern-temperate genus (6 spp.), one species in Colombia and Ecuador. For the main features of the genus, see under the species. Literature: Schuster and Engel (1974), Schuster (2000a) 1. Pseudocephalozia quadriloba (Steph.) R.M.Schust.

Fig. 24F

On moist, peaty soil in páramo, often among Sphagnum, 3200–4100  m. Colombia: Boy, Cau, Cun. Ecuador: Az, Ca, Pi, ZC.  General distribution: Costa Rica to Patagonia and Tristan da Cunha. Pseudocephalozia quadriloba is a characteristic species of marshes and bogs in the high Andes and is readily recognized by the small, pale green to brownish, somewhat swollen plants with shallowly 2–4-lobed, transverse leaves and underleaves, and a stoloniform stem base. The leaves are more than 1 cell thick in the lower half.

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Keys and Descriptions

Pteropsiella Spruce Plants thalloid, small, 1–3  cm long, 3–5  mm wide, pale green, creeping, usually simple (± unbranched), resembling a small Symphyogyna but with short, leafy sexual branches. Branches ventral-intercalary. Stems retained as a narrow midrib, hyalodermis present. Thallus margin entire or toothed, sometimes bordered with thickened cells, with elongate slime papillae. Cells without trigones, cuticle weakly papillose. Underleaves reduced to a few cells, with rhizoids in bundles at the base. Dioicous or monoicous. Perianths long-cylindrical, mouth with long, bristle-like lacinia. Setae very thin, of only 8 outer rows of cells and 4 inner rows. A small neotropical genus of two species, one in Colombia and Ecuador. A very peculiar plant, related to Zoopsidella but thalloid, with leafy sexual branches. The thalloid habit is probably due to fusion and reduction of leaves on either side of the stem. The remnants of the leaves (= leaf tips) are marked by sausage-shaped slime papillae on the thallus margin, like in Zoopsidella. The sporophytes are also similar to those of Zoopsidella. Literature: Fulford (1968), Schuster (2000a) 1. Pteropsiella metzgeriiformis R.M.Schust. Pteropsiella serrulata Steph., nom. inval.

Fig. 24G–I

On shaded, moist sand in savanna and along rivulets in Amazonia, also on decaying wood, 100–450  m. Colombia: Ama (Pinzón et  al. 2003), Vau (Schultes & Cabrera 14187, CINC, PC). General distribution: Amazonia (Brazil, Guyana, Venezuela, Colombia). Pteropsiella metzgeriiformis is a very characteristic Amazonian species recognized by the small, pale green, thalloid plants with a narrow midrib, a border of 1–2 rows of small cells, irregularly crenate-serrate margins, ventral stoloniform branches, and leafy gametoecia produced on short, ventral branches. The perianth is long-cylindrical, crowned by long, bristle-like lacinia and beset with numerous mucilage papillae. Pteropsiella frondiformis Spruce, a rare species from the upper Rio Negro and the Guayana Highland, is close to P. metzgeriiformis but differs in having subentire thallus margins and margin cells scarcely smaller than the rest of the thallus cells (Fulford 1968). Contrary to Fulford (1968) and Gradstein and Costa (2003), the name Pteropsiella serrulata was not validly published by Stephani. Telaranea (Spruce) Schiffn. Arachniopsis Spruce Plants very small, filamentous, less than 1 mm wide, pale green, 1–3-pinnate or irregularly branched. Branches mostly Frullania-type or Bazzania-type. Stems with a distinct hyalodermis. Leaves transverse or oblique, deeply divided into 2–4 almost uniseriate filaments, lamina reduced, rarely consisting of only 1 filament (T. pecten). Cells usually large, longer than wide, walls uniformly thin, cuticle smooth; oil

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bodies finely granular. Underleaves smaller than the leaves, deeply divided into 2–4 filaments, or reduced. Dioicous or monoicous. Perianths longly cylindrical, mouth usually with long, bristle-like lacinia. A mostly southern-temperate genus of about 30 species (Söderström et al. 2016), 4 in Colombia and Ecuador, in two sections, sect. Telaranea and sect. Tenuifoliae (R.M.Schust.) J.J.Engel & G.L.Merr. The genus is heterogeneous and needs monographic revision (Cooper 2013). Telaranea is recognized by the leaves divided to the base into 1–4 hair-like filaments made up of rectangular cells with thin, smooth walls and the stems with a distinct hyalodermis made up of large, thin-walled epidermis cells that are much larger than the inner cells. Underleaves may be well developed and up to 1/2 the leaf length or are reduced. Telaranea may be confused with Blepharostoma (Pseudolepicoleaceae), but the latter genus has larger underleaves, about as large as the leaves, lacks a hyalodermis, and has somewhat thick-walled cells with a finely striate-papillose cuticle (smooth in Telaranea). Due to the thickened cell walls, the leaves in Blepharostoma are usually rather stiff and dull in color. Those of Telaranea, on the other hand, are quite flexible and glistening-hyaline. Literature: Engel and Smith Merrill (2004), Pócs (2006), Cooper (2013) 1. Leaves consisting of only 1 uniseriate lobe (= filament). . . . . . . . . . . T. pecten 1. Leaves 2–4-lobed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves divided to the base into 2–4 filaments, leaf lamina lacking . . . . . . . . . 3 2. Leaves divided to ca. 1/2 into 3–4 filaments, leaf lamina 4–5 cells high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neolepidozia rectangularis 3. Stem leaves 3–4-lobed. Underleaves well developed, 1/4–1/2 the length of the leaves, 2–3-lobed, the filaments made up of a row of 2–3 narrow rectangular cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. nematodes 3. Stem leaves 2-lobed. Underleaves reduced, consisting of 2 short, adjacent cells each capped with slime papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Filaments (6–)8–10 cells long. Plants autoicous . . . . . . . . . . . . . . . T. sejuncta 4. Filaments 4–6 cells long. Plants dioicous . . . . . . . . . . . . . . . . . . . T. diacantha 1. Telaranea diacantha (Mont.) J.J.Engel & G.L.Merr. Arachniopsis diacantha (Mont.) Howe

Fig. 24J

On rotten wood, bark, and humus in lowland and lower montane rainforest, from sea level to 2000 m. Colombia: Ama, Ant, Ces, Cho, Mag, Put, Qui, Ris, San, Vau. Ecuador: Or, ZC. General distribution: tropical America, Africa. Telaranea diacantha (sect. Tenuifoliae) is a common species that is distinguished by the alga-like habit, with hair-like leaves divided to the base into 2 long filaments, each consisting of a row of 4–6 rectangular cells, and reduced underleaves consisting of only 2 adjacent cells, each cell capped by a slime papilla. The species is morphologically close to T. sejuncta, but the latter species is autoicous (T. diacantha is dioicous) and the leaf filaments in T. sejunta are longer (see key). Telaranea diacantha was treated in a broad sense in the catalogue of Colombia (Gradstein and Uribe 2016a) with T. sejuncta as synonym. However, Engel and Smith Merrill

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Keys and Descriptions

(2004) and Pócs (2006) had resurrected T. sejuncta as a good species. The distributions of T. diacantha and T. sejuncta in Colombia are unclear, and all herbarium collections of the two species need revision. 2. Telaranea nematodes (Austin) M.A.Howe Telaranea bicruris (Steph.) M.Howe Telaranea chaetophylla (Spruce) Schiffn. Telaranea sejuncta sensu Fulford (non typus)

Fig. 24K, L

On soil, litter, humus, rotten wood, bark, and rock in montane forests and páramo, (150–)400–4250  m. Colombia: Ama, Ant, Boy, Cas, Ces, Cho, Cun, Hui, Mag, Met, Put, Ris, San, Tol, Val, Vau. Ecuador: Ch, EO, Ga, MS, Pi, Tu, ZC. General distribution: tropical America, tropical Africa, western Europe. Telaranea nematodes (sect. Telaranea) is recognized by the leaves split almost to the base into 3–4 filaments, each filament constisting of a row of 4–7 narrowly rectangular cells (2-cellular at the base) with thin, smooth walls, and by the presence of small underleaves, about 1/4–1/2 the leaf length and split to the base into 2–3 filaments, each filament being 2–4 cells long. The present definition of T. nematodes follows Fulford (1966 as T. sejuncta), with T. chaetophylla and T. bicruris as synonyms. Engel and Smith Merrill (2004) treated the latter two species as separate taxa. 3. Telaranea pecten (Spruce) J.J.Engel & G.L.Merr. Arachniopsis pecten Spruce On bark, rotten wood, and rock in Amazonian rainforest, 135–215 m. Colombia: Caq, Vau (Campos et al. 2014); to be expected in Ecuador. General distribution: northern Amazonia, Guyana, West Indies. Telaranea pecten (sect. Tenuifoliae) is a very distinct neotropical species that is distinguished by the leaves consisting of only one filament, each filament made up of a row of 3–7 narrowly rectangular cells (Fulford 1966; Schuster 2000a). 4. Telaranea sejuncta (Ǻngstr.) S.W.Arnell Telaranea coactilis Spruce) J.J.Engel & G.L.Merr. (Arachniopsis coactilis Spruce) On rotten wood, bark, and humus in lowland and lower montane rainforest, from sea level to 1900 m. Colombia: Ama, Mag, Vau. Ecuador: ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical America, Africa. Characteristic of T. sejuncta are the leaves divided to the base into 2 long filaments, each consisting of a row of (6–)8–10 rectangular cells, and the reduced underleaves consist of only 2 adjacent cells, each cell capped by a slime papilla. The species is morphologically close to T. diacantha, but the latter species is dioicous (T. sejuncta is autoicous) and the leaf filaments in T. diacantha are shorter and consist of 4–6 cells (Engel and Smith Merrill 2004). Fulford (1966) placed T. sejuncta in the synonymy of T. nematodes, but Engel and Smith Merrill (2004) accepted it as a good species, close to T. coactilis. The

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latter species, however, is doubtfully different from T. sejuncta and not recognized here (Gradstein and Uribe (2016a) erroneously placed T. coactilis in the synonymy of T. diacantha). Telaranea sejuncta was not recognized in the catalogue of Colombia; its distribution in the country remains unclear and needs study. Zoopsidella R.M.Schust. Plants small, usually less than 1  mm wide, whitish-green, creeping, irregularly branched. Branches ventral-intercalary, occasionally Frullania-type; ventral branches usually stoloniform. Stems thin, dorsal side 2 cells wide, hyalodermis present. Leaves very flat and wide-spreading, succubous and almost horizontally inserted on the lateral side of the stem (dorsal side of the stem leaf-free), quadrate to elongate, bifid or undivided, upper margins often with (1–)2 large, sausageshaped or globose slime papillae. Cells very large, ca. 60–80 μm long in midleaf, thin-walled, without trigones, cuticle smooth; oil bodies small, finely segmented. Underleaves reduced to a transverse row of 3–4 large cells. Dioicous or monoicous. Perianths long-cylindrical, mouth with bristle-like lacinia or cilia, uniseriate part of cilia and lacinia 2–8 cells long. Setae very thin, of only 8 outer rows of cells and 4 inner rows. A neotropical-Asiatic genus of about five species, three in Colombia and Ecuador. Zoopsidella is readily distinguished by the very flat spreading leaves, attached to the lateral side of the stem, with frequently two large, sausage-shaped slime papillae on the upper margins. The slime papillae mark the tips of the 2 leaf lobes potentially produced in this genus. In Z. serra and Z. grahamii, large sausage-shaped slime papillae are lacking; instead, the leaf tips may terminate in a small, globose hyaline papilla (seen in Z. grahamii). Literature: Fulford (1968), Gradstein and Ilkiu-Borges (2009), Pócs (2019b) 1. Leaves narrowly rectangular, 2× longer than wide. Leaf apex bifid to 1/5. In upper montane elfin forest above 2000 m . . . . . . . . . . . . . . . . . . . . Z. grahamii 1. Leaves ovate-quadrate to ovate-oblong, 1–1.5× longer than wide. Leaf apex entire or bifid. In lowland and lower montane rainforests below 2000 m. . . . 2 2. Leaves entire or shallowly retuse. Leaf apex with 1–2 large sausage-shaped papillae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. integrifolia 2. Leaves regularly bifid to 1/4–1/3. Leaf apex without sausage-shaped papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. serra 1. Zoopsidella grahamii Pócs On shaded, humic soil in very humid upper montane elfin forest, 2500–2650 m. Ecuador: Lo, ZC (Benitez and Gradstein 2011 as Z. caledonica). General distribution: Peru, southern Ecuador. Zoopsidella grahamii was first recorded from southern Ecuador (based on sterile material) as Z. caledonica (Steph.) R.M.Schust., a species known from Australasia (Benitez and Gradstein 2011). Pócs (2019b) gathered fertile material of the species

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Keys and Descriptions

in Peru and found that the Andean plant differed from Z. caledonica in several features and represents a separate species. Both species have horizontally spreading, narrowly rectangular leaves with a short-bifid apex (to 1/5 of leaf length), each tip ending in a globose hyaline papilla. Zoopsidella grahamii differs from Z. caledonica in the more delicate, only 0.7–0.9-mm-wide plants (1.1–1.35  mm wide in Z. caledonica) and the shorter leaves, 250–300 μm long and 2× longer than wide (600–700 μm long and 3× longer than wide in Z. caledonica). In addition, the uniseriate portions of the perianth mouth cilia are longer in Z. grahamii, being 6–7 cells long, as compared to 4 cells long in Z. caledonica (Pócs 2019b). 2. Zoopsidella integrifolia (Spruce) R.M.Schust. Fig. 25C, D Zoopsidella dichotoma R.M.Schust., nom. inval., syn. nov. Material: Colombia, Huila, road to Palermo, Punto Viejo, in Quercus forest, 2100  m, Schuster 99-1322 (F-CO172193!, dupl. [Schuster “1335” pro err.] F-CO172192!) Zoopsidella macella (Spruce) R.M.Schust. On rotten wood and rock in lowland and montane rainforest, 20–1600  m. Colombia: Ama, Cau (Isla Gorgona, Schuster s.n., COL), Ces, Cho, Hui, Ris, San, Vau; to be expected in Ecuador. General distribution: tropical America. Zoopsidella integrifolia is a morphologically highly variable neotropical species; probably most neotropical Zoopsidella species are synonyms of Z. integrifolia (see Gradstein and Ilkiu-Borges 2009). The plants vary from 0.5 to 1.5 mm wide, the stems have 6–12 rows of epidermal cells (and few to numerous rows of medullary cells), Frullania-type branches are present or absent, and stoloniform or flagelliform ventral branches are common. The leaves are quadrate to ovate to oblong, ca. 1–1.5× longer than wide, and the leaf apex varies from rounded to truncate to shortbifid; this variation can be seen in a single plant. The leaf cells are very large (ca. 60–80 μm long in midleaf; as in the genus); the apex usually has 1–2 large, sausagelike papillae; and underleaves are reduced. The plants may be autoicous or dioicous; autoicous plants from Colombia were named Z. dichotoma or Z. macella (the latter species was considered dioicous by Schuster (1993), but Spruce described it as autoicous). The male bracts and bracteoles are divided to about the middle into long-lanceolate lobes with acute to short acuminate tips and entire to crenate margins. 3. Zoopsidella serra (Spruce) R.M.Schust. Regredicaulis serrus (Spruce) Fulford

Fig. 25E

On rotten wood and rock in everwet coastal Pacific rainforest, 20–270  m. Colombia: Cau, Cho. General distribution: northern Amazonia, Chocó. A rare Amazonian-Chocó species characterized by wide-spreading, flat, regularly bifid stem leaves (to 1/3) that are ovate-quadrate in shape, 1–1.5× longer than wide, with very large cells (as in the genus) and without large sausage-shaped papillae at the leaf tips. The underleaves are reduced to a transverse row of 3–4 large adjacent cells, and the branches are frequently flagelliform.

Lophocoleaceae

243

Lophocoleaceae Plants green to brown or reddish-brown, creeping to ascending. Branches variable, Frullania-type and intercalary (lateral and ventral); stolons or flagella usually lacking. Stems without hyalodermis, cortex usually not differentiated. Leaves succubous, usually almost horizontal in position, insertion line usually reaching dorsal stem-midline, bifid or undivided, leaf margins entire or toothed. Cells thin-walled, with or without trigones, cuticle usually smooth; oil bodies granular, rarely homogeneous. Underleaves present, small or large, usually bifid, often toothed and connate with the leaves. Rhizoids usually in bundles from underleaf bases, rarely scattered. Gametoecia on leading shoots or on short branches. Sporophytes surrounded by a terete or 2–3-keeled perianth. Setae of numerous cells (cross section). Capsule walls 3–8-layered. Vegetative reproduction rare, by gemmae or caducous leaves. About 20 genera worldwide. The principal features of Lophocoleaceae are (1) leaves often almost horizontally inserted; (2) underleaves well developed, often united on one or both sides with leaf bases; (3) rhizoids in tufts at underleaf bases; and (4) sporophyte surrounded by a terete or 2–3-keeled perianth. The family was previously included in Geocalycaceae, but molecular analysis has shown that the two families are distinct and only remotely related. Morphologically they differ by the presence of a perianth in Lophocoleaceae and a marsupium in Geocalycaceae. Geocalycaeae are not known from Colombia and Ecuador. The genera of Lophocoleaceae are morphologically not sharply separated, and the current generic classification of the family is largely based on molecular evidence (e.g., Hentschel et al. 2006; Söderström et al. 2013a). 1. Leaf surface covered by numerious small, spinose teeth . . . Lophocolea muricata 1. Leaf surface smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves apex entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaf apex bifid or toothed or variable (entire to toothed on one stem) . . . . . . 6 3. Leaves broadly reniform, 1.5–1.8× wider than long, stiffly appressed to each other. Plants brown, rare, in páramo . . . . . . . . . . . . . . . . . . . . . . . . Platycaulis 3. Leaves ovate-orbicular to oblong, as wide as long or longer than wide, appressed or spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Underleaves variable, entire to bifid on the same stem and usually narrower than the stem, oblong to lanceolate. Leaf cells thin-walled, ± without trigones. Plants pale green, rather small, less than 1.5 mm wide, growing in or near running water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clasmatocolea vermicularis 4. Underleaves not entire and bifid on the same stem. Leaf cells usually with trigones. Plants green to brown, small or large. . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Underleaves free from the leaves or narrowly connate on one side. Leaf apex rounded to retuse. Perianth with 3 sharp keels . . . . . . . . . . . . . . . . Lophocolea 5. Underleaves connate with leaves on 1–2 sides. Leaf apex rounded. Perianth with 0–2 keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptoscyphus

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Keys and Descriptions

6. Leaves ± transverse, strongly caducous. Plants minute, yellowish-green, ca. 0.5 mm wide, on bark in upper montane cloud forest and páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophocolea fragmentissima 6. Leaves distinctly succubous, not caducous. Plants different. . . . . . . . . . . . . . 7 7. Underleaves free from the leaves or narrowly connate on one side . . . . . . . . . 8 7. Underleaves connate with leaves on both sides. . . . . . . . . . . . . . . . . . . . . . . . 9 8. Leaf apex entire or bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophocolea 8. Leaf apex with 3 or more cilia (rarely only 2). . . . . . . . . . . . . . . Leptoscyphus 9. Plants fertile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Plants sterile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 10. Gametangia on very short ventral branches, hidden under the leaves. Perianth without keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroscyphus 10. Gametangia on long branches, not hidden under the leaves. Perianth 2–3-keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Leaves constantly bifid. Trigones absent. . . . . . . . . . . . . . . . Cryptolophocolea 11. Leaves with 3–15 cilia or varying from entire to 2-toothed. Trigones present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptoscyphus 12. Leaf apex varying from entire to retuse to 2-toothed . . . . . . . . . . . . . . . . . . . 13 12. Leaf apex mostly with 2 or more teeth or cilia . . . . . . . . . . . . . . . . . . . . . . . 14 13. Underleaves very small, about as wide as the stem, narrowly attached to leaves on one side. Leaf cells fully thin-walled, without trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroscyphus contortuplicatus 13. Underleaves larger, ca. 2× wider than the stem, attached to leaves on both sides. Leaf cells with small trigones. Plants common, variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptoscyphus amphibolius 14. Leaf apex with 2 teeth. Underleaves with 2–6 teeth or cilia (rarely more) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Leaf apex with 3 or more teeth or cilia. Underleaves with numerous cilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15. Leaves narrowed to the apex. Leaf cells ± without trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptolophocolea 15. Leaves not narrowed to the apex. Leaf cells with distinct trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroscyphus marginatus 16. Underleaves 2–3× wider than the stem, subquadrate, deeply bifid. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptoscyphus trapezoïdes 16. Underleaves larger, 3–5× wider than the stem, reniform, not or shallowly bifid. Plants monoicous or dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroscyphus Clasmatocolea Spruce Plants small, 1–2  mm wide, pale green, creeping, sparingly branched. Branches Frullania-type or intercalary, lateral or ventral. Leaves succubous, ± concave to almost flat, somewhat vertically oriented, ovate-subquadrate, apex rounded or truncate, margins entire. Cells thin-walled, trigones absent or very small in the neotropical species, cuticle smooth; oil bodies very finely granular, 2–3 per cell. Underleaves small and variable, ovate to lanceolate to subulate, undivided or short-bifid, margins

Lophocoleaceae

245

entire. Dioicous. Antheridia on elongate shoots. Perianths rare, strongly inflated, urn-like, with a wide, open mouth. A southern-temperate genus of about 20 species, 1  in the tropical Andes. For main features of Clasmatocolea in Colombia and Ecuador, see under the species. Literature: Engel (1980) 1. Clasmatocolea vermicularis (Lehm.) Grolle  Clasmatocolea heterostipa Spruce Leioscyphus chamissonis (Lehm. & Lindenb.) Spruce Lophocolea elata Gottsche Notoscyphus lindmanii (Steph.) Schiffn.

Fig. 25F–H

On moist soil and rock, in rivulets or in gulleys near running water, sometimes in hot springs, mostly in upper montane environments and páramo, 1800–4500  m. Colombia: Boy, Cal, Cau, Cun, San. Ecuador: Ca, Lo, Na, Pi, Tu, ZC. General distribution: temperate regions of the Southern Hemisphere, in the neotropics in southeastern Brazil and the tropical Andes northward to Costa Rica and the Dominican Republic. Clasmatocolea vermicularis is a widespread semi-aquatic species that is distinguished by (1) plants small, pale green; (2) leaves somewhat distant, flat or concave, ovate-subquadrate, apex rounded to truncate; (3) leaf cells thin-walled, cuticle smooth; and (4) underleaves small but conspicuous, not wider than the stem, longer than wide, lanceolate or oblong, entire or bifid (often entire to bifid on a single stem). The plants grow on soil or rock in or near running water and are usually sterile. Cryptolophocolea L.Söderstr. et al. Lophocolea sect. Connatae Lindenb., Lophocolea martiana complex (Fulford 1976) Plants medium-sized, 2–4  mm wide, (pale) green, becoming brown with age, creeping, irregularly branched. Branches lateral, Frullania-type or intercalary. Leaves succubous, alternate, ovate-rectangular to narrow-rectangular, apex bifid, margins entire or with a few cilia, leaf surface smooth. Cells thin-walled, trigones lacking or small, cuticle smooth; oil bodies usually finely granular. Underleaves attached to leaves on either side, 2–4-lobed. Monoicous. Gametoecia on elongate shoots. Perianths 3-keeled, long-exserted beyond the bracts. A genus of about 23 species, 3 in Colombia and Ecuador. Cryptolophocolea is quite similar to Lophocolea but differs by underleaf bases connate with the leaves on both sides. The genus was traditionally included in Lophocolea, but molecular studies indicated that the group is a separate lineage and merits recognition at genus level (Hentschel et al. 2006; Söderström et al. 2013a). Literature: Fulford (1976 as Lophocolea martiana complex), Söderström et al. (2013a) 1. Teeth at leaf apex large, the tips ending in a row of 5–8 cells; sinus between the 2 apical teeth deeply U-shaped . . . . . . . . . . . . . . . . . . . . . . . C. guadalupensis

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Keys and Descriptions

1. Teeth shorter, tips ending in a row of 1–4 cells; sinus between the 2 teeth shallowly U-shaped or truncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Underleaves bifid, with an additional tooth (2–5 cells long) on the outer margins much shorter than the 2 lobes . . . . . . . . . . . . . . . . . . . . . . . . . . C. connata 2. Underleaves bifid to quadrifid, with 1(−2) additional, long ciliate teeth on the outer margins about as long as the 2 lobes . . . . . . . . . . . . . . . (C. martiana) 3 3. Ventral leaf margin entire . . . . . . . . . . . . . . . . . . . . . C. martiana var. martiana 3. Ventral leaf margin with 1–2 teeth or cilia . . . . . C. martiana var. perissodonta 1. Cryptolophocolea connata (Sw.) L.Söderstr. & VáňaFig. 24K Lophocolea connata (Sw.) Nees. Type: Jamaica, Swartz s.n. (lectotype S; isolectotypes PC-0102442!, PC-Mont.-0723569!) Lophocolea canelensis Spruce Lophocolea granatensis Gottsche, syn. nov. Type: Colombia, Cundinamarca, Bogotá, “in silva alta prope Aserradero, ubi ad arborum trunco, mense julio 1860,” Lindig 1737 (lectotype, designated here, PC-0102439!; isolectotype G-279161/1) On bark, dead wood, or soil in montane forests and Polylepis forests up to the páramo, 800–4000  m. Colombia: Ant, Ces, Cun, Hui, Mag, Met, Qui, Ris, Tol. Ecuador: Na (Polylepis pauta forest, Benitez & León-Yánez 1301, HUTPL), Pa, Tu, ZC. General distribution: tropical America. Cryptolophocolea connata is habitually similar to C. martiana, but the underleaves in C. connata are deeply bifid with an additional, shorter tooth (2–5 cells long) on the outer margins. In C. martiana the outer teeth are often almost as long as the 2 central teeth-like lobes, many underleaves thus appearing quadrifid. Moreover, C. connata occurs at higher elevations. Lophocolea granatensis is a phenotype of C. connata with relatively short, ovate-rectangular leaves. Gottsche in his original description of L. granatensis already noted the close affinity to C. connata. The collection Lindig 1737 is designated as lectotype of L. granatensis because it has androecia and gynoecia and was cited as “type” by Fulford (1976). 2. Cryptolophocolea guadalupensis (Steph.) L.Söderstr. & VáňaFig. 24L Lophocolea guadalupensis Steph. On trees, 600–3300 m. Colombia: Boy, Cun, Hui. Ecuador: Pa, ZC. General distribution: tropical America. Cryptolophocolea guadelupensis is recognized by the ± opposite, long-rectangular leaves which are bifid to 1/5–1/4 with a deeply U-shaped sinus and 2 longly acuminate lobes, each with 5–8-cell-long uniseriate tip. The underleaves are small, 2–4-fid, and broadly connate to the leaves on either size. Cryptolophocolea guadelupensis may be confused with C. martiana, but the apical teeth in C. martiana are smaller and the sinus separating the 2 teeth is truncate to shallowly U-shaped (see key). Cryptolophocolea guadelupensis may also be confused with Lophocolea bidentata, but the latter species differs in having alternate, ovate-rectangular leaves (not opposite long-rectangular), a narrower sinus between the 2 apical teeth, and, especially, free underleaf bases which are not attached to the leaves.

Lophocoleaceae

247

3a. Cryptolophocolea martiana (Sw.) L.Söderstr., Crand-Stotl. & Stotler var. martiana  Fig. 24I, J Lophocolea martiana Nees Lophocolea pertusa Taylor On rotten logs, rock, and soil in lowland and montane rainforests, from sea level to 2000(−3000) m. Colombia: Ant, Boy, Caq, Cas, Ces, Cho, Cun, Hui, Mag, Met, Qui, Ris, San, Vau. Ecuador: ZC (Parolly et al. 2004). General distribution: tropical America, Africa. Cryptolophocolea martiana is a very common and widespread Afro-American species that is readily distinguished by the subrectangular leaves with a ± broadly truncate apex with 2 rather small teeth and the underleaves with 4 long, ciliate teeth. In Ecuador, C. martiana has only been recorded from Zamora Chinchipe but should be more widespread in the country. 3b. Cryptolophocolea martiana var. perissodonta (Spruce) Gradst. Lophocolea martiana var. perissodonta Spruce (Lophocolea perissodonta [Spruce] Steph.) Lophocolea proteus Herzog. Type: Colombia, without locality, Killip 14330 (JE!) On rotten logs in lowland and montane rainforests, from sea level to 1700 m. Colombia: Ama, Caq, Cun, Met, San, Vau. General distribution: tropical America. Differs from var. martiana in the frequent presence of 1–2 short teeth or cilia on the ventral leaf margin. In addition, the underleaves in var. perissodonta have sometimes more than 4 cilia. Lophocolea proteus, characterized by ventral leaf margins with an occasional tooth (see Herzog 1955, Fig. 4A), is a synonym (Gradstein and Uribe 2016a). Although no locality is given for L. proteus in the protologue, the type is probably from Mesa de los Santos, Santander, based on Killip’s collecting numbers. Heteroscyphus Schiffn. Plants medium-sized to large, 2–5 mm wide, green to brown, creeping, irregularly branched. Branches usually ventral-intercalary. Leaves succubous, ± flat, alternate to opposite, ovate-subrectangular, apex entire, short-bifid or with 3 or more teeth, dorsal margin usually entire, ventral margin entire or toothed or ciliate, plane or undulate. Cells thin-walled or with distinct trigones, cuticle smooth or weakly papillose; oil bodies unknown. Underleaves bifid, margins usually toothed or ciliate, bases connected to the leaves on one or two sides. Monoicous, rarely dioicous. Gametoecia on very short ventral or lateral branches hidden behind the stem leaves. Male branches tiny, catkin-like. Perianths inflated, smooth, without keels, the mouth 3-lobed-ciliate. A large genus of about 65 species, mostly in Southeast Asia, 5 species in Colombia and Ecuador. Heteroscyphus is recognized by the underleaves connate with the leaves on both sides, usually ventral branching (rarely lateral), gametoecia

248

 

Keys and Descriptions

on very short sexual branches and usually hidden behind the leaves, and perianths without keels. Sterile material may be difficult to distinguish from Cryptolophocolea, but in the latter genus, branching is usually lateral only. Literature: Fulford (1976) 1. Leaf apex and ventral leaf margin entire or with 1–2 small teeth. Underleaves small, 1–2× wider than the stem (3–4× stem width in H. thraustus, couplet 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf apex and ventral leaf margin with several long teeth. Underleaves larger, 3–6× stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Plants delicate, less than 2 mm wide, growing along rivers. Underleaves narrowly connate to leaves on one side only. Leaf cells fully thin-walled, without trigones, cuticle finely papillose . . . . . . . . . . . . . . . . . . . . H. contortuplicatus 2. Plants larger, more than 2  mm wide. Underleaves connate to leaves on both sides. Leaf cells with or without trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaf cells without trigones. Leaves narrowed to the apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptolophocolea 3. Leaf cells with trigones. Leaves narrowed or not . . . . . . . . . . . . . . . . . . . . . . 4 4. Trigones large, swollen. Leaves narrowed to the apex . . . . . . . . . H. thraustus 4. Trigones small, not swollen. Leaves not narrowed to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. marginatus 5. Underleaves strongly crispate-ciliate, margins irregularly recurved and cilia pointing in different directions. Trigones, large, swollen . . . . . H. polychaetus 5. Underleaves ciliate but not crispate, ± flat. Trigones small or large . . . . . . . . 6 6. Trigones large, swollen. Leaves with 3–5 teeth. . . . . . . . . . . . . . . H. thraustus 6. Trigones absent or very small. Leaves with 3–10 teeth. . . . . . H. polyblepharis 1. Heteroscyphus contortuplicatus (Nees & Mont.) Grolle Lophocolea contortuplicata (Nees & Mont.) Fulford Lophocolea brachydonta Spruce (fide Fulford 1976) Lophocolea irrigata Spruce (fide Fulford 1976)

Fig. 25M

On tree bases, decaying wood, rock, and soil, frequently on river banks, in montane environments, 900–2700 m. Colombia: Boy, Cun, Mag, Ris. Ecuador: Pa, Tu, ZC.  General distribution: northern Andes, Central America, Dominican Republic, southeastern Brazil. Heteroscyphus contortuplicatus is a little-known, semi-aquatic neotropical species that is distinguished by the rather small plants (1–1.5 mm wide) with lax and rather flat, dull green leaves with an entire (rounded to truncate), shallowly 2-lobed or 2-toothed leaf apex, fully thin-walled leaf cells, and a weakly papillose cuticle. The underleaves are very small, hardly wider than the stem, deeply bilobed, with 1–3 small teeth on outer margins and the underleaf bases narrowly connected to the leaves on one side. The plants are dioicous, and gynoecia and androecia are produced on very short branches, as in Heteroscyphus (Grolle 1984a). Heteroscyphus contortuplicatus may be confused with Lophocolea aberrans; for differences see under the latter species. According to Grolle (1984a), the synonymy of L. brachydonta and L. irrigata needs verification.

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2. Heteroscyphus marginatus (Steph.) Fulford Chiloscyphus bomboizanus Steph. On bark in montane forest, 1000–3300 m. Colombia: Ant, Boy, Cau, Cun, Hui, Met, Tol. Ecuador: MS (type), Pa, Tu, ZC.  General distribution: tropical Andes, Central America (Panama, Costa Rica). Heteroscyphus marginatus is a rather common Andean species that is characterized by (1) plants medium-sized to large plants (2–5  mm wide), with rather flat, symmetrical, subrectangular leaves with a broad apex, leaf apex not narrower than the leaf base; (2) leaf apex with 1–2 small teeth (rarely trifid), margins entire with 1–2 small teeth; (3) leaf cells with small trigones; and (4) underleaves small (ca. 2× stem width), deeply bifid, connate to the leaves on both sides and with 1(−3) teeth on the outer margins. Heteroscyphus marginatus resembles H. thraustus, but the leaves in the latter species are narrowed to the apex (leaf base distinctly wider than leaf apex) and the trigones are large, swollen. Sterile, bifid plants with entire leaf margins may be confused with Cryptolophocolea, but in the latter genus, the leaf cells are fully thinwalled, without trigones, and the leaves are somewhat narrowed to the apex. 3. Heteroscyphus polyblepharis (Spruce) Schiffn.

Fig. 26A, B

On bark, rock, and soil in upper montane forest and páramo, 1500–3700  m. Colombia: Boy, Cau, Cho, Hui, Ris, San (Gil 2926, UPTC), Tol. Ecuador: Lo, Pa, Pi, Tu. General distribution: northern Andes, Guayana Highland (Venezuela), Guatemala, Brazil (? see Gradstein and Costa 2003). Heteroscyphus polyblepharis is a common northern Andean species that is recognized by the flaccid, pale green plants (2–5 mm wide) with broadly ovate-triangular to subrectangular leaves with 3–10 ciliate teeth on the apex and ventral margin, thin-walled cells without or with very small trigones, and large, reniform, flat underleaves with numerous long ciliate teeth on the margins (usually more than 10, sometimes fewer) and with connate bases. 4. Heteroscyphus polychaetus (Spruce) Hentschel & Heinrichs Lophocolea polychaeta Spruce On shaded rock and humus in montane forest, 1000–2500 m. Ecuador: Pa, Tu, ZC. General distribution: northern Andes (Ecuador, northern Peru). Heteroscyphus polychaetus is very distinct northern Andean species that is distinguished by the robust, greenish to yellowish-brown plants with convex, asymmetrically triangular leaves tapering to a very narrow apex. The dorsal leaf margin is straight and fully entire, whereas the ventral margin and apex are curved and strongly ciliate-laciniate (apex with 2–3 lacinia). The leaf cells have large, swollen trigones, and the underleaves are large, imbricate, much wider than the stem, connate with the leaves and with strongly ciliate-crispate margins, which are irregularly and narrowly recurved with the cilia pointing in different directions. Heteroscyphus polychaetus may be confused with H. polyblepharis and Leptoscyphus trapezoïdes, which also have ciliate leaves and underleaves. However,

Fig. 26 (A and B) Heteroscyphus polyblepharis. (A) Habit, ventral view. (B) Portion of shoot with small male branch and female branch, ventral view. (C) Leptoscyphus amphibolius, cells near leaf margin, 275×. (D) Leptoscyphus autoicus, habit with 2 perianths, lateral view, 20×. (E and F) Leptoscyphus cleefii. (E) Portion of stem with leaf, ventral view, 25×. (F) Underleaf, 45×. (G–I) Leptoscyphus cuneifolius subsp. fragilis. (G and H) Habit, ventral view, 18×. (I) Midleaf cells, 350×. (J and K) Leptoscyphus gibbosus. (J) Habit, ventral view. (K) Habit with perianth, lateral view. (L) Leptoscyphus jackii, leaf, 35×. (N) Leptoscyphus porphyrius, midleaf cells, 275×. (O) Leptoscyphus spectabilis, habit in ventral view, 20×. (C, E–I from Fulford 1976; D from Engel and Gradstein 2003; L–O from Fulford 1976; rest from Gradstein et al. 2001)

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the underleaves in the latter two species are not crispate and the trigones are smaller. Moreover, L. trapezoïdes has a much broader leaf apex and much smaller and less ciliate underleaves. 5. Heteroscyphus thraustus (Spruce) Fulford

Fig. 25N, O

On moist rock, 1800–2100 m. Colombia: Boy, Hui, Ris. Ecuador: Tu (Fulford 1976). General distribution: northern Andes (Colombia, Ecuador). Heteroscyphus thraustus is a little-known northern Andean species that is characterized by the medium-sized plants (2–3 mm wide) with opposite, subrectangular leaves with shallowly bifid apices and 1–3 additional long teeth on the ventral leaf margin and leaf cells with large, swollen trigones. The underleaves are ca. 3–4× stem width and rather deeply bifid, connate to leaves on both sides and with 2–5 teeth on each lateral margin. Leptoscyphus Mitt. Leioscyphus Mitt., Physotheca J.J.Engel & Gradst. Plants small to large, 1–5  mm wide, greenish-brown to dark brown, black, or reddish-brown, creeping, irregularly branched. Branches intercalary (ventral or lateral), rarely Frullania-type. Leaves succubous, strongly concave or almost flat, alternate to ± opposite, ovate-orbicular to oblong, as long as wide or longer than wide, undivided in neotropical species, apex rounded or truncate, rarely retuse, entire, margins usually entire, occasionally with teeth or cilia. Cells usually with large trigones, occasionally with small trigones (e.g., L. amphibolius); cuticle smooth or papillose; oil bodies finely granular. Underleaves usually connected to the leaves, rarely free, bifid (rarely undivided), often toothed or ciliate on the outher margins. Rhizoids in bundles or scattered. Dioicous, very rarely autoicous (L. autoicus). Androecia on elongate shoots or short ventral branches. Perianths on short or long shoots, usually laterally flattened and 2-keeled, occasionally terete or 3-keeled with a low third, ventral keel. Vegetative reproduction by caducous leaves. A genus of about 30 species in the New World, tropical Africa, Australasia, and western Europe, 16 species in Colombia and Ecuador, in several subgenera, including subg. Leptoscyphus, subg. Physoscyphus Grolle, and subg. Spinoscyphus Vanderp., Schäf-Verw. & D.G.Long (Grolle 1962; Vanderpoorten et al. 2010). The genus is readily recognized by the brownish plant color (but often green in L. amphibolius and L. leoniae), the usually undivided leaves with ± entire or ciliate margins, the small to very large trigones, the conspicuous underleaves which are usually connected to the leaves on both sides (rarely free), and the inflated perianth. Literature: Grolle (1962), Fulford (1976), Vanderpoorten et al. (2010) 1. Plants autoicous and always highly fertile, on twigs and branches of shrubs in Ecuadorian páramo, rare. Perianth terete, apex with a small, pore-like mouth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. autoicus 1. Plants dioicous and frequently sterile, common. Perianth laterally compressed or terete, apex with a wide mouth, almost as wide as the perianth . . . . . . . . . 2

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Keys and Descriptions

2. Leaf apex with 1 or more ciliate teeth (occasionally a few leaves entire) . . . . 3 2. Leaf apex mostly entire [when in doubt try both leads]. . . . . . . . . . . . . . . . . . 4 3. Leaf margin bordered by thick-walled cells. Leaves with 1–3(−4) short ciliate teeth at the apex, occasionally some leaves entire. Leaf cells with small trigones. Underleaves free from the leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . L. leoniae 3. Leaf margin not bordered by thick-walled cells. Leaves with (2–)4–15 long ciliate teeth. Leaf cells with small to large trigones. Underleaves usually connected to leaves at the base, rarely free. . . . . . . . . . . . . . . . . . . . . . . . . . L. trapezoïdes 4. Ventral leaf margin with 1 or more teeth, at least in some of the leaves (sometimes only with a short tooth at the base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Ventral leaf margin fully entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5. Trigones of leaf cells large, bulging, cell-lumina stellate. Teeth on leaves, when present, restricted to the ventral leaf base. Ventral leaf base auriculate . . . . . . 6 5. Trigones small, not bulging, cell lumina rounded. Teeth restricted or not restricted to the ventral leaf base, upper ventral leaf margin sometimes with 1 or more teeth. Ventral leaf with or without auricle . . . . . . . . . . . . . . . . . . . . . 7 6. Plants robust, (3–)4–5 mm wide. Ventral leaf base entire or with 1 small tooth. Leaf cells 30–40 μm in diameter in the upper half of the leaf. . . L. hexagonus 6. Plants smaller, 2–3 mm wide. Ventral leaf base with 1–3 large teeth. Leaf cells 20–30 μm in the upper half of the leaf . . . . . . . . . . . . . . . . . . . . . . . . . L. jackii 7. Leaves asymmetrical, ventral margin expanded (more strongly arched than the dorsal margin) and often recurved in the lower half. Plants of páramo bogs (Colombia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cleefii 7. Leaves subsymmetrical, ventral margin not expanded, plane. Plants of upper montane forest (southern Ecuador) . . . . . . . . . . . . . . . . . . . . . . . L. spectabilis 8. Plants tiny, less than 2 mm wide. Underleaves undivided or bifid. Cells in the upper part of leaf 15–24 μm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Plants larger. Underleaves bifid. Cells in the upper part of leaf larger, (20–) 25–35 μm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. All underleaves bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. intermedius 9. Underleaves mostly undivided (some may be bifid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cuneifolius subsp fragilis 10. Underleaves small, 1–2× stem width, bifid to more than half of underleaf length. Trigones small or large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10. Underleaves larger, 2–5× stem width, less deeply bifid, to maximally half of underleaf width. Trigones large. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 11. Trigones small, cell-lumina rounded to slightly angular. . . . . . . . . . . . . . . . . 12 11. Trigones large, bulging, cell-lumina sinuate to stellate. . . . . . . . . . . . . . . . . 13 12. Ventral leaf margin revolute. Underleaf lobes longly ciliate, terminating in a uniseriate row of more than 10 cells. Underleaf bases free. Plants rare, endemic to Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. revolutus 12. Ventral leaf margin plane. Underleaf lobes shortly ciliate, terminating in a row of up to 5 cells. Underleaf bases usually connected with leaf bases. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. amphibolius 13. Leaf apex undivided, rounded. Perianth mouth ± entire . . . . . . . L. porphyrius

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13. Leaf apex emarginate to short-bifid with rounded lobes. Perianth mouth coarsely toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. obcordatus 14. Outer margins of underleaves ± entire. Underleaves emarginate to short-bifid (to 1/4 of underleaf length). Cuticle smooth. . . . . . . . . . . . . . . . . L. physocalyx 14. Outer margins of underleaves with 1 or more longly ciliate teeth. Underleaves more deeply bifid (to 1/3). Cuticle striate-papillose or smooth. . . . . . . . . . . 15 15. Ventral leaf bases with a small auricle. Plants robust, 4–5 mm wide. Plants from the high Andes, above 2500 m. . . . . . . . . . . . . . . . . . . . . . . . . . . L. hexagonus 15. Ventral leaf bases without auricle. Plants smaller, less than 4 mm wide. . . . 16 16. Underleaf bases shortly connate with leaves on both sides. Plants widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. gibbosus 16. Underleaf bases free. Plants very rare, on rock in páramo of Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sotiauxii 1. Leptoscyphus amphibolius (Nees) Grolle Leioscyphus galipanus Gottsche Lophocolea pycnorrhiza Spruce

Fig. 26C

On soil and trunk bases in montane forest up to the páramo, ca. 800–4000 m. Colombia: Boy, Ces, Cun, Hui, Mag, Met, Ris, San. Ecuador: Im, Lo, MS, Tu, ZC. General distribution: tropical America. Leptoscyphus amphibolius (subg. Physoscyphus) is a common and variable neotropical species that is distinguished by the rather large, glossy green to brownish plants (to 4 mm wide) with ovate-orbicular to elongate, entire-margined leaves with the ventral margin usually more strongly arched than the dorsal margin, with a rounded or truncate leaf apex which is usually entire, exceptionally with 1–2 teeth (to short-bifid), and thin-walled cells with very small trigones and a smooth cuticle. The gynoecia in L. amphibolius are on long or short branches, and the perianth is usually 3-keeled near the apex, as in Lophocolea (Grolle 1962). The underleaves of L. amphibolius are similar to those of L. porphyrius (small, 1–2× wider than the stem, deeply bifid to over 1/2, with a few teeth on the outer margins, usually connate with leaf bases), and these two species may be confused. They are separated, however, by the smaller plant size, the much larger trigones, and the more symmetrical leaves of L. porphyrius, with the ventral margin not more strongly arched than the dorsal margin. Moreover, the leaves in L. porphyrius are frequently caducous while being never caducous in L. amphibolius. Phenotypes of L. amphibolius with entire to 2-toothed leaf apices (e.g., Meta, Schultes 11711, CINC, PC) resemble Cryptolophocolea connata, but the latter species is more pale-colored (not brownish), the leaf apex is constantly short-bifid (varying from entire to 2-toothed in L. amphibolius), and the leaf cells in C. connata are fully thin-walled (with small trigones in L. amphibolius). 2. Leptoscyphus autoicus (J.J.Engel & Gradst.) Vanderp. & Gradst. Physotheca autoica J.J.Engel & Gradst.

Fig. 26D

On trunks and twigs in upper montane forest and shrubby páramo, 2300–4000 m. Ecuador: Az (páramo de Patococha), Lo, Na, ZC. General distribution: only known from Ecuador.

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Keys and Descriptions

Leptoscyphus autoicus (subg. ?) is the only monoicous species in the genus and always freely fertile. It is readily recognized by the numerous terete, bladder-like perianths with a very small opening (mouth) at the tip and long male branches. The sterile leafy shoots are rather inconspicuous, to 1.3 mm wide, with rounded, entire leaves, cells with large trigones and very small, bifid underleaves. Because of its unusual perianth, autoicy, and various unusual sporophytic features (Engel and Gradstein 2003), L. autoicus was originally placed in a separate genus, Physotheca, but molecular analysis showed that it belongs in Leptoscyphus (Vanderpoorten et al. 2012). 3. Leptoscyphus cleefii Fulford

Fig. 26E, F

In páramo bogs, 3200–4100 m. Colombia: Ara, Boy, Cau, Cho, Cun, Met, Ris. Ecuador: Pi (páramo de Papallacta; Gradstein and Benitez 2017). General distribution: northern Andes (Colombia, Ecuador). Leptoscyphus cleefii (subg. Spinoscyphus) is a characteristic of páramo bogs and distinguished by leaves with 1–4 long cilia on the lower half of the ventral leaf margin; a few scattered cilia may be present higher up along the ventral margin, up to the apex. The plants are flaccid, brown to blackish in color and rather small (ca. 1.5–2.5 mm wide), and the leaves are asymmetrically ovate-orbicular with a rounded apex and an expanded ventral margin that is often recurved near the base. The leaf cells are thin-walled with small trigones and a smooth to finely papillose cuticle, and the underleaves are small (to 2× stem width), deeply bifid and with a long tooth on the outer margins. Leptoscyphus cleefii approaches L. spectabilis, but the latter species clearly differs by the ± symmetrical leaves with plane ventral margin. Moreover, L. spectabilis does not grow in páramos bogs. 4. Leptoscyphus cuneifolius (Hook.) Mitt. subsp. fragilis (J.B.Jack & Steph.) GrolleFig. 26G–I Mylia fragilis (J.B.Jack & Steph.) S.W.Arnell On twigs and branches of shrubs in montane forest and páramo, 2500–4400 m. Colombia: Ant, Boy/Cas (Gradstein 12705 p.p. among Lejeunea acanthogona, PC), Cal, Cau, Cho (van Reenen 5047, U), Cun, Tol. Ecuador: Az, Pi, ZC. General distribution: tropical America, Patagonia. Leptoscyphus cuneifolius (subg. Anomylia) is the smallest species in the genus. The plants are thread-like, 2–8  mm long and usually less than 1  mm wide, and reddish-brown to dark brown in color. The leaves are distant, subquadrate to somewhat obcuneate and frequently caducous; the leaf apex is truncate to short-bifid; the leaf cells are rather small, 16–20(−25) μm wide in the upper part of the leaves; and the underleaves are mostly undivided (a few underleaves may be bifid), lanceolate, and attached to leaf bases on one side or free. Leptoscyphus cuneifolius is subdivided into two subspecies, subsp. cuneifolius (North America, Europe, SW Asia) and subsp. fragilis (tropical America, Patagonia), based on differences in plant color, leaf size, and presence or absence of gametoecia (Grolle 1962).

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Leptoscyphus cuneifolius may be confused with juvenile plants of L. porphyrius, which are strikingly similar to L. cuneifolius. The report of L. cuneifolius from the Galápagos Islands (Arnell 1961) is L. porphyrius (Grolle 1962, p. 31). 5. Leptoscyphus gibbosus (Taylor) Mitt.

Fig. 26J, K

On bark in montane forest, scrub, and páramo, 700–3900 m. Colombia: Ant, Cho, Hui, Ris, San (Gil 2081, UPTC), Tol. Ecuador: Im (Mojanda), ZC (?). General distribution: West Indies, southern Central America, northern Andes. Leptoscyphus gibbosus (subg. Leptoscyphus) is a Caribbean species morphologically very close to L. hexagonus but differing from the latter in smaller plant size (less than 4 mm wide) and ventral leaf bases without auricle. Morphologically, the two species deserve to be combined (perhaps at subspecific level), but in a molecular-phylogenetic analysis, they were quite distant (Vanderpoorten et al. 2010). The occurrence of L. gibbosus in Ecuador needs confirmation; most Ecuadorian records have proved to be L. hexagonus (Schäfer-Verwimp et al. 2013a). 6. Leptoscyphus hexagonus (Nees) Grolle Chiloscyphus purpureus Steph. On bark in upper montane forest, subalpine elfin forests, and shrubby páramo, 2500–4000 m; the species is very common in Polylepis pauta forests near Papallacta pass, east of Quito (Gradstein and León-Yánez 2018). Colombia: Cho (Gradstein et  al. 2018c). Ecuador: Az, Ca, MS, Pi, ZC.  General distribution: northern Andes (Colombia to Peru). Leptoscyphus hexagonus (subg. Leptoscyphus) is a robust northern Andean species that is recognized by (1) plants robust, 4–5 mm wide, glossy brown; (2) leaves strongly convex, erect and appressed or spreading, asymmetrical and somewhat expanded ventrally, apex rounded, ventral leaf base auriculate and with a tooth, occasionally entire; (3) leaf cells 30–40 μm in diameter in the upper part of the leaf, trigones large, bulging, cuticle fully smooth, oil bodies grayish, rounded to oval, 4–10 per cell, finely granular, outer wall smooth; and (4) underleaves large, bifid to 1/4–1/3, lobes narrowly acuminate lobes, outer margins with 1 or several long teeth, bases narrowly connate with leaves on both sides. Leptoscyphus hexagonus is closely related to L. jackii, L. gibbosus, L. physocalyx, and L. sotiauxii (Vanderpoorten et  al. 2010). Leptoscyphus jackii differs in smaller plant size (2–3 mm wide), smaller leaf cells (20–30 μm), and the presence of 1–3 long, laciniate teeth at the ventral leaf base, whereas L. gibbosus, L. physocalyx, and L. sotiauxii differ in smaller plant size and fully entire ventral leaf bases, without auricle and without tooth. Leptoscyphus physocalyx moreover has more shallowly bifid underleaves with subentire margins. The record of L. hexagonus from Galápagos Islands (Arnell 1961) belongs to L. porphyrius (Grolle 1962, p. 48). 7. Leptoscyphus intermedius Grolle On bark in montane forest and páramo, 1800–4200  m. Colombia: Cun, Qui (páramo El Campanario, Gradstein et  al. s.n., HUQ). Ecuador: Na (páramo de Oyacachi, on Polylepis pauta, Benitez & León-Yánez 1303, HUTPL), Pi, ZC. General distribution: northern Andes, Guayana Highland, Honduras.

256

 

Keys and Descriptions

Leptoscyphus intermedius (subg. Leptoscyphus) is very similar to L. cuneifolius, but the underleaves in L. intermedius are bifid, not undivided, and the plants are slightly larger, more than 1 mm wide. The perianth is fully inflated, terete. 8. Leptoscyphus jackii (Steph.) Grolle

Fig. 26L

On bark and wet rock in upper montane forest, Polylepis forests, and páramo, 2750–4400 m. Colombia: Ant (type), Cho, Cun, Ris. Ecuador: Ca, Na (páramo de Oyacachi, on Polylepis pauta, Benitez & León-Yánez 1304, HUTPL), Pi (SchäferVerwimp et al. 2013a). General distribution: northern Andes. Leptoscyphus jackii (subg. Leptoscyphus) is a northern Andean species close to L. hexagonus but differing in smaller plant size (2–3 mm wide), smaller leaf cells (20–30 μm), and, especially, the presence of 1–3 long teeth at the ventral leaf base (Grolle 1962). 9. Leptoscyphus leoniae Gradst. On shaded, rotten bark in upper montane forest and Polylepis forest, 2880–4000 m. Ecuador: Lo (Schäfer-Verwimp & Nebel 31759, QCA), Na (páramo de Oyacachi, on Polylepis pauta, Benitez & León-Yánez 1305, HUTPL), Pi (type). General distribution: only known from Ecuador. Leptoscyphus leoniae (subg.?) is a rare Ecuadorian endemic that grows on the shaded underside of old trunks of Polylepis pauta at the forest line. Characteristics are the prostrate, glossy green to greenish-brown plants with suborbicular, widely spreading leaves (sometimes standing up) with a 1–2 cells wide, colorless or brownish border of thick-walled cells and a broadly rounded apex with 1–3(−4) short, widely spaced cilia. Occasionally part of the leaves are entire, without any cilia (seen in Benitez & León-Yánez 1305). The leaf cells are thin-walled, with very small trigones and a smooth cuticle, and the underleaves are scarcely wider than the stem and free from the leaves. Leptoscyphus leoniae is morphologically close to L. trapezoïdes, but the leaves of the latter species lack a leaf border and have more numerous and longer cilia, the leaf cells in L. trapezoïdes usually have conspicuous trigones and a striate-papillose cuticle (sometimes smooth), and the underleaves are usually wider than the stem and connate, rarely free (Gradstein and León-Yánez 2018). 10. Leptoscyphus obcordatus (Spruce) Grolle Leioscyphus obcordatus Spruce On soil over rock in upper montane scrub and páramo, 3300–3500 m. Colombia: Cau, Cho. Ecuador: Ca, Pi. General distribution: tropical Andes. Leptoscyphus obcordatus (subg. Leptoscyphus) is readily distinguished by the emarginate to short-bifid leaf apex with broad, rounded lobes. In all other neotropical Leptoscyphus species, the leaf apex is rounded and entire or with a few cilia. Leptoscyphus obcordatus is close to L. porphyrius but is readily separated from the latter by the characters given in the key.

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11. Leptoscyphus physocalyx (Hampe & Gottsche) Gottsche Leioscyphus campanulata Steph. On bark and soil over rock in upper montane forest and páramo, 3000–4000 m. Colombia: Ant, Boy, Cho, Cun, Qui (páramo El Campanario, Gradstein et al. s.n., HUQ), Ris. Ecuador: Pi (Taylor 1846), ZC (páramo of Saraguro and Yacuambi, Benitez et al. 1310, HUTPL). General distribution: tropical Andes. Leptoscyphus physocalyx (subg. Leptoscyphus) is a characteristic Andean species that is distinguished by the 2–4-mm-wide, brownish plants with large, shallowly emarginate to short-bifid underleaves (bifid to maximally ¼) and  ±  entire outer margins. The ventral leaf margin is fully entire and the leaf cells have large trigones. Leptoscyphus physocalyx may be confused with L. hexagonus, but the ventral leaf base in L. hexagonus is usually auriculate and with a tooth (sometimes entire) and the underleaves have several long-ciliate teeth on the outer margins. 12. Leptoscyphus porphyrius (Nees) Grolle Clasmatocolea fragillima Spruce Leioscyphus fragilifolius (Taylor) Spruce Leioscyphus schizostomus Spruce Leptoscyphus liebmannianus (Lindenb. & Gottsche) Mitt. Mylia minuta Herzog Mylia quitoensis (Mont.) S.W.Arnell

Fig. 26N

On soil, bark, rock, and rotten wood in the understory and the canopy of lowland and montane rainforests and in páramo, from sea level to 4300 m. Colombia: Ant, Ara, Boy, Cal, Cau, Ces, Cun, Hui, Mag, Met, Nar, Ris, San, Tol. Ecuador: Az, Co, Ga, Im, Lo, MS, Na, Or, Pi, Su, Tu, ZC. General distribution: widespread in tropical America; also on the Azores (L. porphyrius subsp. azoricus [H.Buch & Perss.] Vanderp. & Heinrichs). Leptoscyphus porphyrius (subg. Leptoscyphus) is one of the most common neotropical species of Leptoscyphus, with a remarkably wide elevational range. It is recognized by the rather small, glossy green to brownish plants (1.5–3 mm wide) with suberect to wide-spreading, ± symmetrically ovate-orbicular leaves with a rounded to emarginate apex and leaf cells with large trigones. The underleaves are small (1.5–2× stem width), deeply bifid to over 1/2, often with a few teeth on the outer margins, and connected at the base to the lateral leaves. The leaves in L. porphyrius are frequently caducous. 13. Leptoscyphus revolutus Burghardt On bark in upper montane forest and scrub, 3600–3800 m. Ecuador: Pi (volcán Fuya Fuya; Burghardt et  al. 2019). General distribution: only known from Ecuador. Leptoscyphus revolutus (subg. Leptoscyphus) is a rare Ecuadorian endemic species that is recognized by the 2.5–3-mm-wide plants with concave leaves, entire leaf margins, strongly recurved ventral margin, small trigones, and tiny, deeply bifid

258

 

Keys and Descriptions

underleaves that are about as wide as the stem, not connected to leaf bases, with longly ciliate lobes terminating in a uniseriate row of up to 12 cells and with a few ciliate teeth at the base (Burghardt et al. 2019). Leptoscyphus revolutus resembles L. amphibolius and L. porphyrius in the shape of the underleaves, but in the latter two species, the ventral leaf margin is ± flat and the underleaf bases are usually connected with the lateral leaves. Leptoscyphus porphyrius, moreover, has large, bulging trigones. 14. Leptoscyphus sotiauxii Vanderp., Schäf.-Verw. & D.G.Long On soil on steep rocky slope in páramo, 4030 m. Ecuador: Na (Vanderpoorten et al. 2010). General distribution: only known from the type locality in Ecuador. Leptoscyphus sotiauxii (subg. Leptoscyphus) is a little-known páramo endemic that resembles L. gibbosus in the asymmetrical leaves with entire ventral base, large trigones, and large underleaves with longly acuminate-ciliate lobes and 1 or more long linear teeth on the outer margins. Leptoscyphus sotiauxii differs from L. gibbosus by the underleaf bases free from the leaves (shortly connate in L. gibbosus) and the fully smooth cuticle (striate-papillose toward the base in L. gibbosus). The species is also very close to L. hexagonus, but in the latter species, the ventral leaf base has a small auricle and frequently a tooth, and the leaf cells are wider. 15. Leptoscyphus spectabilis (Steph.) Grolle Heteroscyphus arnellii Fulford

Fig. 26O

On humic soil in humid upper montane forest, 2850–2900  m. Ecuador: Lo (Podocarpus National Park; Gradstein and Benitez 2017). General distribution: southeastern Brazil, southern Ecuador. Leptoscyphus spectabilis (subg. Spinoscyphus) is a rare Brazilian species that has been found in southern Ecuador. It closely resembles L. amphibolius but differs by the presence of a few linear teeth (1–3) on the ventral leaf margin. 16. Leptoscyphus trapezoïdes (Mont.) L.Söderstr. Fig. 27A, B Lophocolea trapezoïdes Mont. Lophocolea breutelii Gottsche Lophocolea columbica Gottsche (incl. vars.) Lophocolea effusidens Steph. Lophocolea orbigniana Nees & Mont., syn. nov. Type: Bolivia (“Peruvia”), Yungas, Chupé, d’Orbigny 183 (lectotype, designated here, PC-Mont.-0705343!; isolectotype PC-Mont.-0705342!) Lophocolea quadridentata Spruce (Chiloscyphus quadridentatus [Spruce] J.J.Engel & R.M.Schust.), syn. nov. Type: Bolivia, “Near Yungas, 4000  ft., 1885,” Rusby s.n., ex herb. Pearson (syntype, MANCH!, c. gyn. juv.) On bark, rock, leaf litter, humus, and soil in montane rainforests and páramo, 1000–4000 m. Colombia: Ant, Boy, Cal, Cas, Cau, Ces, Cho, Cun, Hui, Mag, Nar, Qui, Ris, San. Ecuador: Az, Cañ, Ch, Co, Ga, Pi, Tu, ZC. General distribution: tropical America; also in the East African high mountains when Lophocolea

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259

Fig. 27 (A and B) Leptoscyphus trapezoïdes. (A) Habit, ventral view. (B) Midleaf cells. (C) Lophocolea bidentata, habit in ventral view, 20×. (D) Lophocolea liebmanniana, habit in ventral view. (E) Lophocolea muricata, portion of stem with leaf in dorsal view. (F) Platycaulis renifolia, portion of stem with leaf in lateral view, showing underleaf with rhizoid bundle, 19×. (G and H) Plagiochila adianthoides. (G) Habit, dorsal view, 18×. (H) Leaf margin with tooth and border of elongate cells. (I and J) Plagiochila aerea. (I) Habit, dorsal view, 30×. (J) Midleaf cells. (K) Plagiochila amicta, habit in dorsal view, 6×. (L and M) Plagiochila alternans. (L) Leaf. (M) Midleaf cells. (C redrawn from Fulford 1976; F redrawn from Schuster 1995b; K redrawn from Heinrichs 2002a, b; L, M from Müller et al. 1999; rest from Gradstein et al. 2001; Gradstein and Ilkiu-Borges 2009)

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Keys and Descriptions

muhavurensis (S.W.Arnell) Pócs proves to be synonymous with Leptoscyphus trapezoïdes (T. Pócs, pers. comm.). Leptoscyphus trapezoïdes (subg.?) is a common and highly variable, neotropical-montane species that is distinguished by the green to yellowish-brown to brown plants with ovate-orbicular to trapezoidal leaves with (2–)4–15 cilia on the margins, especially at the apex and in the upper half but sometimes all around the leaves. The leaf cells have rather small to large trigones and a smooth to finely striate-papillose cuticle, and the cells at the leaf margin are of the same size as the inner cells or somewhat larger, but they are not thick-walled than the inner leaf cells. The underleaves are 1–2× stem width, subquadrate, deeply bifid and with few to many cilia on the outer margins. The underleaf bases are usually connected to the leaves on one or both sides (occasionally free). The perianths in L. trapezoïdes are distinctly trigonous, with two 2-winged, laciniate lateral keels and a low, rounded ventral keel. Because of its trigonous perianth, L. porphyrius was previously placed in Lophocolea, in spite of its yellowish-brown, typical Leptoscyphus-like color. Molecular analysis has confirmed that L. porphyrius belongs in Leptoscyphus. The species is defined here in a broad sense, including forms with few cilia and with many cilia, all around the leaf. Phenotypes with few cilia were described as Lophocolea quadridentata, while those with cilia all around the leaf were named Lophocolea orbigniana. Leptoscyphus trapezoïdes is close to L. leoniae, an Ecuadorian endemic from Polylepis forest, but the latter species clearly differs in its brownish leaf border of thick-walled cells and fewer and shorter cilia. Leptoscyphus trapezoïdes may be confused with Heteroscyphus polychaetus, which resembles L. trapezoïdes in the long-ciliate leaf margins. However, H. polychaetus clearly differs from L. trapezoïdes in the triangular shape of its leaves, tapering to a narrow apex, and the strongly crispate-ciliate underleaves. Lophocolea (Dumort.) Dumort. Campanocolea R.M.Schust. Plants small to large, (0.6–)1–5  mm wide, (pale) green, creeping, irregularly branched. Branches Frullania-type or intercalary, mostly lateral, rarely ventral. Leaves succubous, alternate, ovate-orbicular to rectangular, apex bifid or with a few teeth or cilia, margins entire or toothed to ciliate, leaf surface smooth or roughened by tooth-like papillae in L. muricata. Cells usually thin-walled, hexagonal-subisodiametrical, trigones lacking or small, cuticle smooth or papillose; oil bodies usually finely granular. Underleaves free from the leaves, bifid, often toothed. Dioicous, monoicous, or heteroicous. Gametoecia on elongate shoots. Perianths 3-keeled, longly exserted beyond the bracts. A genus of about 90 species worldwide, 6 in Colombia and Ecuador. The main features of Lophocolea are (1) plants pale green to green, not brown; (2) leaves bifid, margins entire or with small teeth; (3) leaf cells with small trigones; (4) underleaves free from the leaves; and (5) gynoecia on long stems or branches, perianth trigonous. The genus has sometimes been united with Chiloscyphus Corda, but

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molecular study shows that the two groups should be kept apart (Söderström et al. 2013a). Chiloscyphus does not occur in the neotropics. Lophocolea resembles Cryptolophocolea and Heteroscyphus but differs in having free or very narrowly connate underleaves (broadly connate in Cryptolophocolea and Heteroscyphus) and from Heteroscyphus furthermore by gametoecia on elongate branches and perianths deeply trigonous. Lophocolea may be confused with Neesioscyphus (Balantiopsidaceae), but the latter genus is readily separated from Lophocolea by purely ventral branching (branching mostly lateral in Lophocolea) and the conspicuously elongate, rectangular leaf cells with a densely striate-papillose cuticle. Moreover, rhizoids and parts of the whole plant are sometimes reddish in color in Neesioscyphus (never reddish in Lophocolea). Literature: Fulford (1976), Söderström et al. (2013a), Dauphin et al. (2018) 1. Leaf surface covered by numerous teeth-like projecting cells. . . . . L. muricata 1. Leaf surface smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves transverse, deeply bifid to more than 1/2 of leaf length, usually caducous. Plants minute, ca. 0.5 mm wide, growing on bark in upper montane cloud forest and páramo, above 3000 m . . . . . . . . . . . . . . . . . . . . . L. fragmentissima 2. Leaves almost longitudinally inserted, succubous, bifid to 1/4–1/3, not caducous but sometimes with fragile tips. Plants usually more than 0.5 mm wide (sometimes very small, flagelliform in L. tenerrima), growing on soil, rock, and rotten wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaf tips fragile, usually broken and disfigured. Cuticle on pale, flagelliform branches finely punctate-papillose, on well-developed plants smooth. Plants growing on soil under rock in deep shade or in rock fissures, above 2000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. tenerrima 3. Leaf tips not fragile. Cuticle smooth or papillose . . . . . . . . . . . . . . . . . . . . . . 4 4. Plants very small, 0.7–1.2  mm wide. Leaf cells small, 13–23  μm wide in midleaf. Leaf margins with 1 or more teeth (sect. Microlophocolea). . . . . . . 5 4. Plants and leaf cells larger. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . 6 5. Leaf margin with 1(−2) teeth near apex, otherwise entire. Caducous leaves present. Plants from Costa Rica (Cocos Island); not known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophocolea fragrans (Moris. & De Not.) Gottsche, Lindenb. & Nees subsp. cocosana G.Dauphin, Gradst. & M.I.Morales 5. Leaf margin with 5–20 teeth. Caducous leaves absent. Plants from lowland and lower montane rainforests, below 2000 m. . . . . . . . . . . . . . . . L. liebmanniana 6. Leaf apex constantly bifid, lobes acuminate. Underleaves mostly free from the leaves. Cuticle smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bidentata 6. Leaf apex varying from entire to short-bifid, lobes rounded or acute. Underleaves mostly narrowly attached to leaves on one side. Cuticle smooth or finely papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Cuticle finely papillose. Leaf cells without trigones. Underleaves 1–1.5× stem width. Gynoecia on a very short branch. Perianth without keels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroscyphus contortuplicatus

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Keys and Descriptions

7. Cuticle smooth. Leaf cells with small trigones. Underleaves 1.5–3× stem width. Gynoecia on a long shoot. Perianth with 3 sharp keels. . . . . . . . . . L. aberrans 1. Lophocolea aberrans Lindenb. & Gottsche Type: Mexico, Veracruz, “apud Mirador in terra,” Mar 1842, Liebmann 74, c. gyn. (lectotype, designated here, C!) On soil and rotten bark in montane rainforest and páramo, 500–4000  m. Colombia: Boy (4000 m, Bischler 2998, PC), Ces, Cun, Mag. Ecuador: Pa (Spruce 1885). General distribution: Mexico, Honduras, northern Andes (Colombia, Ecuador), Chile (Easter Island). Lophocolea aberrans is a little-known neotropical species known from only few collections. The main characters are (1) plants small, to 1.5 mm wide, dioicous; (2) stem leaves subquadrate-suborbicular, about as long as wide, apex varying from broadly rounded to shallowly retuse to short-bifid, lobes broadly rounded to obtuse to acute; (3) leaf cells with small trigones and a smooth cuticle; and (4) underleaves deeply bifid, to 2/3–3/4 of underleaf length, 1.5–3× wider than the stem, outer margins with 1 short tooth, underleaf bases usually narrowly connate with leaf bases on one side, occasionally free. Contrary to the description in Fulford (1976), the mature leaves of L. aberrans are not longer than wide, the cuticle is not verruculose, the underleaves are wider than the stem and deeply bifid (to 2/3–3/4), and the inner female bracteole is more shallowly bifid than the underleaves. Sterile L. aberrans may be confused with Heteroscyphus contortuplicatus; the latter species differs in fully thin-walled leaf cells (without trigones), finely papillose cuticle, and smaller underleaves, maximally 1.5× width and with 1–3 teeth on outer margins. Moroever, H. contortuplicatus is a rheophyte. Fertile H. contortuplicatus is readily separated from L. aberrans by gametoecia on a very short branches and perianths without keels. Lophocolea aberrans approaches the widespread southern-temperate L. semiteres (Lehm.) Mitt., but the latter species is more robust (1.5–3 mm wide); the leaves in L. semiteres are more narrowed to the apex, ± trapezioid; and the underleaves frequently possess several teeth on the outer margins (Grolle 1985; Paton 1999). 2. Lophocolea bidentata (L.) Dumort. Lophocolea blepharoptera Spruce Lophocolea coadunata (Sw.) Nees Lophocolea cuspidata (Nees) Limpr. Lophocolea diaphana Spruce Lophocolea latifolia Nees Lophocolea longiflora Hampe Lophocolea mandonii Steph. Lophocolea mascula Gottsche Lophocolea subtrilobata Herzog

Fig. 27C

On soil, rotten wood, and rock in rather open locations in montane rainforest areas and páramo, occasionally on bark, sometimes growing over rock in rivers and periodically submerged, 100–4450 m. Colombia: Boy, Cau, Ces, Cun, Hui, Mag,

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Met, Put, Qui, Ris, San, Tol. Ecuador: Az, Cañar, Ch, Co, Lo, MS, Na, Or, Pi, Tu, ZC. General distribution: subcosmopolitan. Lophocolea bidentata is a very common and variable, monoicous or dioicous (heteroicous) species that is readily recognized by flaccid, light green plants growing in loose mats among vegetation, with regularly bifid leaves (bifid to 1/4) with acuminate lobes with 3–6-cell-long uniseriate tips and a deeply U-shaped or V-shaped sinus between the lobes. The underleaves are small, bisbifid (ca. 0.7–1.5× stem width), free from the leaves, and the leaf cells are thin-walled, ± isodiametrical, ca. 25–30 μm in midleaf, with minute trigones and a smooth cuticle. Some authors have used the name L. coadunata, a species described from Jamaica by Swartz (type S-B25183!), for the neotropical plants (e.g., Fulford 1976), but the late J.  Váňa and I have been unable to find any morphological character separating L. bidentata from L. coadunata (Gradstein and Váňa 1987). Stotler and Crandall-Stotler (2017) suggested that L. coadunata is monoicous but the type of L. coadunata is dioicous and most of the neotropical populations that I have examined were dioicous, even though Fulford (1976) described the species as monoicous or dioicous. Culture studies have shown that asexual, unisexual, and bisexual plants (described as Lophocolea cuspidata) can develop in culture from a single clone of L. bidentata (Vogelpoel 1982). An integrative morphological-molecular study into the relationships between the monoicous and dioicous populations of L. bidentata, based on a broad, worldwide sampling and including specimens from all over the neotropics, has not been done and would make a worthwile subject for a thesis. 3. Lophocolea fragmentissima R.M.Schust. Campanocolea fragmentissima (R.M.Schust.) R.M.Schust. On mossy tree trunks and on twigs and branches of treelets and shrubs in upper montane cloud forest, in Polylepis forests, and páramo, 3000–4100 m. Colombia: Boy, Cas, Cun, Ris, Tol. Ecuador: Az, Ca, Lo, Na, Pi, ZC (páramo of Saraguro and Yacuambi, Benitez & Gradstein s.n., HUTPL). General distribution: northern Andes, Costa Rica, Dominican Republic. Lophocolea fragmentissima is a common, tiny epiphyte in scrubby vegetation near the forest line in the northern Andes. The species is readily distinguished by the small, less than 1-mm-wide yellowish-green plants with ± distant, deeply bifid, transversally inserted leaves that are usually freely caducous (rarely not caducous; seen in material from the páramo of Saraguro and Yacuambi) and with large, inflated perianths. The cells are very thin-walled, with small trigones, and the underleaves are bifid and about half the size of the leaves. 4. Lophocolea liebmanniana Gottsche

Fig. 27D

On bark and living leaves, mostly on tree bases, in the shaded understory of lowland and lower montane rainforests, 200–2000 m. Colombia: Cho, Cun, Mag, Put, Qui, Ris. Ecuador: Or, ZC. General distribution: tropical America. Lophocolea liebmanniana (sect. Microlophocolea) is a characteristic shade epiphyte of lowland and lower montane rainforests that is recognized by the small, dull green, heteroicous plants (less than 1 mm wide) with a bifid apex, irregularly toothed

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Keys and Descriptions

margins with 1–3-cell-long teeth and small, subquadrate leaf cells, ca. 13–18 μm in diameter. The species is related to L. muricata, but in the latter species, the entire leaf surface is densely toothed-spinose; in L. liebmanniana the leaf surface is smooth. 5. Lophocolea muricata (Lehm.) Nees Lophocolea hirta Steph.

Fig. 27E

On bark, decaying wood, and living leaves in montane forests up to the forest line, 1000–3800 m. Colombia: Ant, Boy, Cau, Ces, Cho, Cun, Hui, Mag, Met, Qui, Ris, Tol. Ecuador: Az, Lo, Pi (Mojanda, on Polylepis, 3800 m, León-Yánez s.n., QCA), ZC. General distribution: temperate regions of the Southern Hemisphere and tropical mountains. Lophocolea muricata (sect. Microlophocolea) is a small shade epiphyte of montane forests that is readily recognized by the small, glossy green to dark green, creeping plants (0.5–1.5 mm wide) with densely spinose leaf surfaces. The leaf cells are small, like in L. liebmanniana. The plants are monoicous. 6. Lophocolea tenerrima Spruce Lophocolea erosa Gradst., nom. inval. On moist soil in deep shade, usually on steep road banks under rock or in rock fissures in upper montane environments, 2400–3700 m. Ecuador: Ca, Co, Pi (type); to be expected in Colombia. General distribution: Ecuador, Costa Rica. Lophocolea tenerrima is a small, little-known, high-montane species that was first collected by Richard Spruce on Mt. Pichincha and was more recently detected in several localities in Ecuador as well as in Costa Rica by the author and provisionally named L. erosa (in León-Yánez et  al. 2006). Spruce (1885), in his original description of L. tenerrima, was somewhat unsure whether the species belonged to Lophocolea: “This little plant is so disfigured by the apices of the leaves being dissolved into propagula as to be difficult to describe. Its characters are somewhat anomalous. The toothed leaves seem to place it near Lophocolea liebmanniana; but neither that nor any other Lophocolea has assurgent leaves” (Spruce 1885, p. 439). Recently, gynoecial plants (but without mature perianths) were collected in Costa Rica (Dauphin et al. 2018); the occurrence of gynoecia on long shoots supported the placement of the species in Lophocolea. A further gynoecial population, again without mature perianths, was found in Ecuador among the unidentified liverwort collections kept in herbarium QCA (Pichincha, road Aloag to St. Domingo km 8, 3000 m, 7 Apr 1982, Meenks 256, with Riccardia wallisii). By the highly irregular leaf apices, L. tenerrima cannot be confused with any other species of the genus. The plants are dioicous, delicate (1–2 mm wide), green to pale brown, the leaves are distant and subrectangular, the underleaves are about as wide as the stem and free from the leaves, the cell walls are ± thin-walled, and the leaf and underleaf margins are usually furnished with several irregular and usually broken, unicellular to multicellular teeth (occasionally entire). Plants growing in deep shade are often flagelliform; the cuticle of stems and leaves of these flagelliform plants is typically densely punctate-papillose. Well-developed plants, however, usually have a smooth cuticle.

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Further Record Lophocolea foliicola Spruce – Ecuador: Tu (Spruce 1885). The material could not be found in MANCH (L. Loughtman, pers. comm.). Excluded Records Lophocolea leptantha (Hook.f. & Taylor) Gottsche, Lindenb. & Nees (Lophocolea brasiliensis Schiffn.)  – Colombia: Mag (Winkler 1976), Qui (Orrego and Uribe 2004) and various other departments (herb. COL). The Colombian material was misidentified and belongs to L. bidentata (J. Uribe, pers. comm.). Lophocolea lindmanii Steph. – Colombia: Mag (Winkler 1976). A southern subtropical species (southern Brazil, Bolivia); the Colombian material is probably misidentified. Lophocolea widgrenii Steph.  – Colombia: Mag (Winkler 1976). A dubious Brazilian species, known only from the type from Caldas; the record has been excluded from Colombia by Uribe and Gradstein (1998). Platycaulis R.M.Schust. Plants medium-sized, green to dark brown, simple or sparingly branched. Stems rigid, with a thick-walled, brown, 1-layered cortex, all cells narrowly elongate. Branches Frullania-type and lateral-intercalary. Leaves alternate, transverse and laterally appressed, flat, broadly reniform, ca. 1.5–1.8× wider than long, ventral base widely expanded across the stem, the expanded ventral portion of the leaf about as the broad as the dorsal lamina; leaf undivided, margins entire, dorsal base longly and broadly decurrent. Cells with large trigones and papillose cuticle; oil bodies unknown. Underleaves small, hidden between the leaves, ca. 1 mm long and wide, wider than the stem, deeply bifid, each outer margin with 2–3 cilia, rhizoids (when present) in bundles from underleaf bases. Perianths unknown. A neotropical genus with a single species, close to Leptoscyphus but differing by broadly reniform leaves, ca. 1.5–1.8× wider than long. Possibly, Platycaulis is a synonym of Leptoscyphus. Literature: Schuster (1995b) 1. Platycaulis renifolia R.M.Schust.

Fig. 27F

In wet páramo, under shrubs, and in bogs among cushions of Oreobolus obtusangulus, 3200–3600  m. Colombia: Boy (Gil-Novoa et  al. 2015). Ecuador: Ca (Benitez et  al. 2012). General distribution: northern Andes (Venezuela to Ecuador). Platycaulis renifolia is a rare northern Andean páramo species that is distinguished by the brownish plants with transverse, broadly reniform, flat leaves (more than 1.5× wider than long) that are ventrally widely expanded across the stem, the expanded ventral part being about as large as the dorsal lamina. As a result, the stem

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is positioned behind the mid-portion of the leaf. The leaf apex is rounded, the margins are entire and flat, and the dorsal base is long and broadly decurrent. The plants resemble Plagiochila dependula but differ by the presence of small underleaves and the rhizoids originating in bundles from underleaf bases (underleaves absent and rhizoids scattered or absent in P. dependula).

Plagiochilaceae A family of about five genera, represented in Colombia and Ecuador by one genus, Plagiochila. Gradstein et al. (2001) listed three further genera of Plagiochilaceae from Ecuador and Colombia, Plagiochilion S.Hatt., Steereochila Inoue, and Szweykowskia Gradst. & M.E.Reiner. Steereochila and Szweykowskia are now considered synonyms of Plagiochila, while Plagiochilion is a synonym of Chiastocaulon Carl, characterized by ventral branching and restricted to Asia (Heinrichs 2002a). The neotropical species of the former genus Plagiochilion, P. bryhnii and P. heteromallum, lack ventral branching and are members of Plagiochila (Heinrichs 2002a). Plagiochila (Dumort.) Dumort. Steereochila Inoue, Szweykowskia Gradst. & M.E.Reiner Plants small to very robust, (0.5–)1–10 mm wide, green to brown, creeping to ascending or pendent, often with a rhizome-like creeping base. Stems rigid, usually with a brown cortex of thick-walled cells in several layers. Branches Frullania-type or lateral-intercalary. Leaves succubous, alternate, rarely opposite, orbicular to ovate to oblong to almost linear, usually asymmetrical, with reflexed dorsal margin and decurrent bases, apex undivided or 2–3-lobed, margins usually toothed to ciliate, especially along the apical and ventral margins, teeth at the apex often larger, margins rarely entire. Cells variable; oil bodies usually granular. Underleaves absent or very small. Rhizoids few, scattered. Dioicous. Gametoecia on elongate shoots. Androecia in long, terminal spikes. Sporophytes surrounded by a perianth, perianths laterally compressed, with a wide, truncate mouth, unfertilized perianths often with innovations. Setae of numerous cells (cross section). Capsule walls thick, 4–9-layered. Spores unicellular or multicellular due to endosporous germination. Elaters with 1–2(−3) spirals. Vegetative reproduction by cladia, caducous leaves or leaf fragmentation. Plagiochila with about 400–450 species worldwide and ca. 120 in the neotropics (Heinrichs 2002a) is probably the largest genus of the liverworts and exhibits an extraordinary morphological variation in spite of its rather simple basic structure. More than 200 species have been recorded from Colombia and Ecuador, but the majority of them are now synonyms (e.g., Gradstein 2015a, 2016a; Grolle and Heinrichs 1999; Heinrichs 2002a, b; Heinrichs et al. 1998, 1999, 2000, 2001, 2002a; Müller et  al. 1999). In this treatment 65 species are accepted for Colombia and Ecuador, in nine sections, sect. Adianthoideae Lindenb., sect. Arrectae Carl, sect.

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Cucullatae Schiffn., sect. Denticulatae Schiffn., sect. Fuscoluteae Carl, sect. Glaucescentes Carl, sect. Heteromallae Lindenb., sect. Hylacoetes Carl, and sect. Vagae Lindenb. (Heinrichs 2002a; Söderström et  al. 2015b; Gradstein 2016a). A few insufficiently known taxa are listed in “Further records”; most of these will probably turn out to be synonyms. The principal features of the genus Plagiochila are (1) leaves succubous and with a decurrent dorsal base and a recurved dorsal margin, leaf margin frequently toothed; (2) underleaves absent or very small; (3) stems rigid, brown, with a thick-walled cortex; (4) male spikes very long; and (5) perianths flattened, with a wide, truncate, and often ciliate mouth. The leaf surface may be horizontally spreading or ventrad. When ventrad, the leaf surface is somewhat tilted downward toward the ventral side of the plant. The ventral leaf bases in Plagiochila are sometimes shouldered or ampliate. When shouldered, the leaf margin at the ventral base is a little bit extended parallel to the stem before curving outward away from the stem. When ampliate, the ventral leaf bases are expanded across and beyond the stem; when strongly expanded, the ventral leaf bases are concealing the stem and are frequently curved downward, forming a crest. Plagiochila is always dioicous. Vegetative reproduction is common, by small plantlets (cladia) originating from the ventral surface of leaves, by caducous leaves, or by leaf fragmentation. In many species the leaf cells become somewhat larger and more elongate toward the leaf base. In members of the sect. Arrectae, the basal area of larger, elongate cells is very conspicuous and may extend up to the leaf middle. The area of larger, elongate leaf cells in Plagiochila is usually called “vitta” (e.g., Heinrichs 2002a; Gradstein 2016a), but here the term “vitta-like area” is used (see Bazzania) because the vitta in Plagiochila is not sharply separate from normal leaf cells and merges gradually into the rest of the leaf cells. In contrast, a true vitta consists of a line or narrow band of large cells sharply separated from the rest of the leaf cells. Notes of caution: 1. Leaf characters are examined on mature, vegetative stem leaves, not on branch leaves or leaves near the gynoecium. Branch leaves may not allow to distinguish between species, e.g., between P. gymnocalycina and P. simplex. Juvenile plants without mature stem leaves cannot be identified with certainty. 2. Branching type should be determined based on vegetative branches arising from parts of the stem away from the gynoecium and not based on branches originating below the gynoecium (= innovations). Selected literature: Heinrichs et  al. (2002a), Heinrichs et  al. (1998, etc.), Gradstein (2015a, 2016a) 1. Leaves sac-like, strongly convex, dorsal and ventral margins strongly curved downward. Plants robust, 6–10 mm wide, yellowish-brown . . . . . P. cucullifolia 1. Leaves not sac-like. Plants small or large . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Stem apex with conspicuous, yellowish-green clusters of flagelliform branchlets (resembling miniature cauliflower), producing numerous minute lanceolate caducous leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ecuadorica

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Keys and Descriptions

2. Stem apex without yellowish-green clusters of flagelliform branchlets . . . . . 3 3. Ventral leaf margin strongly undulate from the base to the middle of the leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fastigiata 3. Ventral leaf margin not undulate or slightly undulate at the base only . . . . . . 4 4. Upper half of leaves frequently broken due to leaf fragmentation . . . . . . . . . 5 4. Upper half of leaves not broken. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 5. Branches predominantly terminal. (Ventral leaf bases distinctly ampliate. Plants robust, 5–7 mm wide) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. buchtiniana 5. Branches intercalary or lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Leaves 2–4-lobed, margins entire or with a few small teeth. Plants small, 1–2.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Leaves not lobed, margins with 2–50 teeth. Plants small or large. . . . . . . . . . 8 7. Leaves distant, 2–3-lobed to 1/4–1/3 of leaf length. Midleaf cells 20–32  μm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cuneata 7. Leaves subimbricate, deeply 3–4-lobed to 1/3–1/2 of leaf length. Midleaf cells larger, 25–43 μm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. loriloba 8. Unfragmented leaves more than 2× longer than wide. Perianth base “naked,” inner bracts inserted at some distance from the perianth and not enveloping the lower half of the perianth. (Leaves ventrad, leaf base not ampliate) . . . . . . . P. gymnocalycina 8. Unfragmented leaves less than 2× longer than wide. Perianth base not “naked,” inner bracts enveloping the lower half of the perianth . . . . . . . . . . . . . . . . . . . 9 9. Leaves transverse to obliquely spreading. Leaf base with a conspicuous vittalike area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bifaria 9. Leaves obliquely to widely spreading. Leaf base without vitta-like area . . . 9a 9a. Ventral leaf base distinctly ampliate. Leaf margin slightly bordered by thickwalled cells. Plants montane . . . . . . . . . . . . . . . . . . . . . . . . . . . P. adianthoides 9a. Ventral leaf base not or scarcely ampliate. Leaf margin not bordered by thickwalled cells. Plants mostly in lowlands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9b 9b. Midleaf cells 30–45 μm wide, thin-walled, with very small trigones. Leaf apex with long, spinose teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subplana 9b. Midleaf cells smaller, ca. 20–30 μm wide, with distint trigones. Leaf apex short triangular teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. disticha 10. Leaves with a brownish border of thicker-walled cells and with numerous long, linear, often brown-colored teeth. (Leaves ovate-oblong, strongly ventrad, with a conspicuous vitta-like area, leaf cells small, 15–25 μm) . . . . . P. pachyloma 10. Leaves without brownish border of thicker-walled cells, plants different. . . 11 11. Stems bluish near stem apex. Leaves with long-linear teeth all around the margins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. longispina 11. Stems not bluish near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Stem leaves entire or with only 1–3(−4) teeth or lobes . . . . . . . . . . . . . . . . . 13 Note: juvenile plants of several species may have very small, bifid or trifid leaves; such juvenile plants cannot be identified with this key. 12. Stem leaves mostly with more than 3 teeth or lobes . . . . . . . . . . . . . . . . . . . 39 13. Stem leaves linear-lanceolate, (5–)6–10× longer than wide, entire or with 1(−2) small teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. rudischusteri

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1 3. Stem leaves not linear-lanceolate, less than 6× longer than wide . . . . . . . . . 14 14. Cuticle papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Cuticle smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 15. Plants robust, stems 4–25  cm long. Leaves entire, transverse, wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. dependula 15. Plants small, stems 1–2 cm long. Leaves bifid, obliquely to widely spreading, longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Leaves narrowly oblong, ca. 2× longer than wide, bifid . . . . . . . . . . . P. tenuis 16. Leaves ovate to ovate-oblong, 1–1.5(−1.7)× longer than wide, bifid and usually with a few additional small teeth. . . . . . . . . . . . . . . . . . . . . . . . . P. papillifolia 17. Leaves of main stems transverse to obliquely spreading. Plants of páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Leaves of main stems obliquely to widely spreading [when in doubt try both leads]. Plants of forests and páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 18. Margin of leaf apex revolute. Leaves wider than long . . . . . . . . . P. revolvens 18. Margin of leaf apex ± plane. Leaves as wide as long or longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19. Plants very small, dark brown, leaves less than 1 mm long, without vitta-like area. Dorsal leaf base very shortly decurrent . . . . . . . . . . . . . . . . P. cuatrecasii 19. Plants larger, pale yellowish brown, leaves more than 1 mm long, base with a short vitta-like area. Dorsal leaf base shortly or longly decurrent . . . . . . P. cleefii 20. Ventral leaf bases ampliate. Leaves ovate to ovate-triangular . . . . . . . . . . . . . . 21 20. Ventral leaf bases not ampliate. Leaves ovate to oblong to ligulate . . . . . . . 28 21. Leaf cells mostly 15–35 μm wide in midleaf . . . . . . . . . . . . . . . . . . . . . . . . . 22 21. Leaf cells mostly more than 35 μm wide in midleaf . . . . . . . . . . . . . . . . . . . 27 22. Dorsal leaf bases shortly decurrent. Leaves ventrad . . . . . . . . . . . . . . . . . . . 23 22. Dorsal leaf bases longly decurrent. Leaves not ventrad [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. Leaves transverse to obliquely spreading, ovate-orbicular to ovate-oblong. Leaf apex toothed. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bifaria 23. Leaves widely spreading, ovate-triangular. Leaf apex entire or bifid. Plants of the West Indies, not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bicornis Hampe & Gottsche (Fig. 28B) 24. Dorsal surface of stem with small, toothed paraphyllia near the dorsal leaf base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. paraphyllina 24. Dorsal surface of stem without paraphyllia. . . . . . . . . . . . . . . . . . . . . . . . . . 25 25. Ventral leaf bases with 1-cell-wide cilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . robust montane form of P. disticha 25. Ventral leaf bases without cilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 26. Dorsal leaf bases very longly decurrent, fully concealing the stem. Leaves ovate-triangular. Plants very robust, 5–10 mm wide, growing above 2500 m, mostly in cloud forests and páramo. . . . . . . . . . . . . . . . . . . . . . . . P. fuscolutea 26. Dorsal leaf bases less longly decurrent, not fully concealing the stem. Leaves ovate-oblong. Plants less robust, in lowland and montane rainforests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. raddiana 27. Ventral leaf base shortly decurrent, the decurrent part 1 cell wide. Leaves ovateorbicular, with a broad rounded apex . . . . . . . . . . . . . . . . . . . . . . . P. ensiformis

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27. Ventral leaf base longly and broadly decurrent, the decurrent part several cells wide. Leaves ovatre-triangular, with a rather narrow apex . . . . . . . P. guevarai 28. Plants 3–6 mm wide, leaves oblong to ligulate . . . . . . . . . . . . . . . . . . . . . . . 29 28. Plants smaller, maximally 2.5 mm wide, leaves ovate to oblong . . . . . . . . . 33 29. Midleaf cells conspicuously elongate, many cells at least 2× as long as wide. Leaf apex with 2–3 large teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 29. Midleaf cells shorter, 1–1.5(−1.7)× as long as wide. Leaf apex entire or with a few small teeth, occasionally bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 30. Branching intercalary. Leaf apex bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 30. Branching mostly terminal. Leaf apex entire or with a few small teeth, not bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 31. Leaves 1.3–2× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. macra 31. Leaves more than 2× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . poor phenotype of P. gymnocalycina 32. Leaves ovate-lingulate, horizontally spreading. Leaf bases longly decurrent. Ventral leaf base with a shoulder. . . . . . . . . . . . . . . . . . . . . . . . . . P. patula s.l. 32. Leaves narrowly oblong, ventrad. Leaf bases shortly decurrent. Ventral leaf base without shoulder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. macrifolia 33. Leaf apex entire. [Plants very small, 0–1(−1.5) mm wide. Leaves ovate-orbicular or subquadrate, apex ± truncate. Leaf bases hardly decurrent. In páramo, above 3000 m] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cuatrecasii 33. Leaf apex bifid or trifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 34. Cuticle finely papillose. Leaves ca. 2× longer than wide . . . . . . . . . . . P. tenuis 34. Cuticle smooth. Leaves 1–2× longer than wide . . . . . . . . . . . . . . . . . . . . . . . 35 35. Leaves fragmenting, part of the leaf lamina and/or teeth broken off . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cuneata 35. Leaves caducous (dropping off entirely) or vegetative reproduction absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 36. Leaf cells ca. 2× longer than wide. . . . . . . . . . . . . small phenotype of P. aerea 36. Leaf cells 1–1.5× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 37. Mature stem leaves oblong, 1.7–2.5× longer than wide . . . . . . . P. bicuspidata 37. Mature stem leaves ovate to ovate-oblong, 1–1.5× longer than wide . . . . . . 38 38. Leaf base with a vitta-like area. Ventral leaf margin often with more than 1 tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. punctata 38. Leaf base without vitta-like area. Ventral leaf margin with 0–1 tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. exigua 39. Leaf margin toothed near the dorsal base (to be observed on vegetative leaves, not on leaves near the gynoecia). Plants robust. . . . . . . . . . . . . . . . . . . . . . . 40 39. Leaf margin not toothed near the dorsal base. Plants small or robust. . . . . . 51 40. Leaves ovate-orbicular, with 100–250 small teeth all around the leaf . . . . P. ovata 40. Leaves with less than 100 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 41. Branching mostly terminal. Teeth very long, to 15 cells long. Underleaves present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. longispina 41. Branching intercalary. Teeth short or long. Underleaves ± lacking. . . . . . . . 42 42. Dorsal surface of stem with small, toothed paraphyllia (sometimes only few) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. paraphyllina

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4 2. Dorsal surface of stem completely smooth, without paraphyllia . . . . . . . . . . 43 43. Leaves 1–1.4× longer than wide. Plants montane . . . . . . . . . . . . . . . . . . . . . 44 43. Leaves 1.3–2× longer than wide. Plants of lowlands and montane . . . . . . . . 45 44. Leaves subopposite. Dorsal and ventral leaf bases shortly decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. turgida 44. Leaves alternate. Dorsal and ventral leaf bases longly decurrent ��������� P. patriciae 45. Ventral leaf base toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 45. Ventral leaf base entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 46. Ventral leaf base longly decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 46. Ventral leaf base shortly decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . P. husnotii 47. Leaf base with vitta-like area. . . . . . . . . . . . . . . . . . . . . . . . . . . . P. breuteliana 47. Leaf base without vitta-like area . . . . . . . . . . . . . . . . . . . . . . . . . . P. vincentina 48. Ventral leaf base ampliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 48. Ventral leaf base not ampliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 49. Dorsal leaf margin toothed only near the base, otherwise entire. Apical and ventral leaf margins with 0–6(−10) teeth. Plants of humid Polylepis pauta forest near the forest line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pautaphila 49. Dorsal leaf margin toothed throughout. Apical and ventral leaf margin with more than 10 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. dominicensis 50. Leaves ovate-oblong to ovate-triangular, apex rounded. Male branches in clusters of 2–6. Plants of lowland and montane rainforests. . . . . . . P. dominicensis 50. Leaves ± rectangular, apex truncate. Male branches single. Plants very common in lowland rainforest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subplana 51. Terminal branches present, but sometimes only few and seen only in the upper parts of the plant (in addition to intercalary branches). Asexual reproduction by small small plantlets (propagules) on ventral leaf surfaces or absent (rarely by caducous leaves, couplet 66) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 51. Terminal branches absent. Asexual reproduction by caducous leaves, by leaf fragmention, or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 When branches and asexual reproduction are absent, try both leads. Note that identification of unbranched, sterile plants may not always be possible with this key. 52. Terminal branches forming an irregularly pinnate pattern . . . . . . . . . . . . . . . 53 52. Terminal branches dichotomous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 53. Leaves subimbricate to imbricate, 1.1–1.6× longer than wide, widest near the base (below 1/3 of leaf length). Ventral leaf base weakly to strongly ampliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. deflexirama 53. Leaves distant, 1.8–2.5× longer than wide, widest at 1/3–1/2 of leaf length. Ventral leaf base not ampliate . . . . . . . . . . . . . . . . . . . . . . . . . . P. tamariscina 54. Ventral leaf bases clearly ampliate, partly or completely concealing the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 54. Ventral leaf bases not or scarcely ampliate, not concealing the stem . . . . . . 60 55. Ventral leaf base entire, longly decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 55. Ventral leaf base toothed, shortly or longly decurrent . . . . . . . . . . . . . . . . . . 57 56. Leaves 1–1.4× longer than wide, often fragmented. Underleaves present. Plants rare, in high-Andean forest and páramo (above 3000 m) . . . . . . P. buchtiniana

272

 

Keys and Descriptions

56. Leaves 1.5–2.5× longer than wide, not fragmented. Underleaves absent or very small. Plants common in lowland and montane rainforest areas (up to 3300 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. raddiana 57. Midleaf cells 20–35(−40) μm wide. Androecia not in a cluster . . . . . . . . . . 58 57. Midleaf cells large, 35–55 μm wide. Androecia usually in a cluster of 2–8 male branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 58. Leaf apex bifid by 2 large teeth. Leaves elongate-triangular, with 15–60 teeth . . . . . . . . . . . . . . . . . . . . P. cristata (phenotype with terminal branching) 58. Leaf apex not bifid. Leaves ovate-lingulate, with 3–30 teeth . . . . . . . . . . . . 59 59. Teeth on ventral leaf base linear (1 cell wide, except at the base). Underleaves present, 0.5–1.5 mm long, ciliate. Ventral leaf base shortly decurrent. Plants of lowland and lower montane rainforest . . . . . . . . . . . . . . . . . . . . . . . . P. disticha 59. Teeth on ventral leaf base at least in part triangular (more than 1 cell wide for part of their length). Underleaves absent or vestigial, to 0.6 mm long, without or with few cilia. Ventral leaf base shortly or longly decurrent. Plants of lowland rainforest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. montagnei 60. Ventral base of mature leaves with linear teeth . . . . . . . . . . . . . . . . P. disticha 60. Ventral base of mature leaves without teeth . . . . . . . . . . . . . . . . . . . . . . . . . . 61 61. Leaves 1–1.5× longer than wide, horizontally spreading . . . . . . . . . . . . . . . 62 61. Leaves 1.5–2.4× longer than wide, horizontally spreading or ventrad [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 62. Ventral leaf base with a shoulder (basal ventral margin somewhat extended parallel to the stem before curving outwards). Underleaves absent. Stems not turning blackish-brown. Leaves ovate. Leaf cells with distinct trigones. Oil bodies colorless. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. laetevirens 62. Ventral leaf base without shoulder. Underleaves usually present (small, to 1 mm long). Stems turning blackish-brown. Leaves ovate-oblong. Leaf cells without or with very small trigones. Oil bodies brown . . . . . . . . . . . . . . . P. diversifolia 63. Leaf bases ± longly decurrent. Leaves with a shoulder at the ventral base (basal ventral margin somewhat extended parallel to the stem before curving outward). Plants rather robust, 4–6 mm wide . . . . . . . . . . . . . . . . . . . . . . P. patula 63. Leaf bases shortly decurrent. Leaves without shoulder. Plants more delicate, 3–4(−5) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 64. Leaves horizontally spreading, rectangular, margins conspicuously parallel (dorsal and ventral margins almost straight). Plants common in Brazil, not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . P. crispabilis Lindenb. 64. Leaves ± ventrad, narrowly oblong, margins not conspicuously parallel (dorsal margin ± straight, ventral margin curved). . . . . . . . . . . . . . . . . . . . . . . . . . . 65 65. Leaf margin bordered by thick-walled cells. Asexual reproduction by small plantlets (propagules) developed on ventral leaf surfaces. . . . . P. distinctifolia 65. Leaf margin not bordered by thick-walled cells. Asexual reproduction by caducous leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 66. Leaves with 2 large teeth at apex and smaller teeth on the ventral margin. Midleaf cells about 2× as long as wide. Plants very common and variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aerea

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273

66. Leaves with a few small teeth near apex, ventral margin entire. Midleaf cells shorter, 1–1.7× as long as wide. Plants very rare, only known from Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. macrifolia 67. Leaves (sub)opposite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 67. Leaves alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 68. Plants delicate, 2–3 mm wide. Ventral leaf base not ampliate. Leaf bases distinctly connate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bryhnii 68. Plants more robust, 4–10  mm wide. Ventral leaf base ampliate. Leaf bases approximate but not distinctly connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 69. Leaves longer than wide. Leaf margin not bordered by thick-walled cells. Midleaf cells (25–)30–50  μm wide. Androecia in a cluster of 2–8 male branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. macrostachya 69. Leaves about as long as wide. Leaf margin frequently bordered by thick-walled cells. Midleaf cells 25–35 μm wide. Androecia not in a cluster . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. heteromalla 70. Ventral leaf base ± ampliate or with a high shoulder, at least in older, mature leaves. Plants rather robust, (3–)5–10 mm wide . . . . . . . . . . . . . . . . . . . . . . 71 70. Ventral leaf base not ampliate and usually without high shoulder [when in doubt try both leads]. Plants small or large . . . . . . . . . . . . . . . . . . . . . . . . . . 90 71. Mature stem leaves mostly with less than 7 teeth . . . . . . . . . . . . . . . . . . . . . 20 71. Mature stem leaves mostly with more than 7 teeth [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 72. Midleaf cells 35–55  μm wide. Androecia usually in a cluster of 2–8 male branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 72. Midleaf cells smaller, 15–35(−40) μm wide. Androecia not in a cluster . . . 78 73. Cuticle of leaves papillose. Plants of Bolivia, SE Brazil, and E Africa; not yet known from Ecuador and Colombia . . . . . . . . . . . . . . . . . . . P. boryana Steph. 73. Cuticle of leaves smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 74. Leaves 0.9–1.3× longer than wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 74. Leaves more elongate, 1.3–1.8× longer than wide . . . . . . . . . . . . . . . . . . . . 77 75. Leaves laterally appressed, with 3–25 long-linear teeth (teeth more than 6 cells long). Plants very rare, growing on branches of shrubs in subpáramo of southern Ecuador . . . . . . . . . . . . . . . . . . . . . . Plagiochila cucullifolia var. anomala 75. Leaves wide spreading, with few or many teeth . . . . . . . . . . . . . . . . . . . . . . 76 76. Teeth small, maximally 4 cells long, not becoming distinctly longer at the leaf apex, maximally 30(−40) per leaf, absent in the lower half of the dorsal leaf margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ensiformis 76. Teeth larger, 2–8(−11) cells long, becoming longer at the leaf apex, up to 60 per leaf, frequently present in the lower half of the dorsal leaf margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. canelensis 77. Teeth on leaf margin narrowly triangular, mostly more than 1 cell wide. Leaves ovate-oblong, with a broad apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. amicta 77. Teeth on leaf margin linear, mostly 1 cell wide (except at the base). Leaves usually ovate-triangular with a narrow apex, rarely ovate-oblong with a broad apex (in P. superba var. macrotricha) . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. superba 78. Leaves 1–1.3(−1.4)× longer than wide, ventrad . . . . . . . . . . . . . . . . . . . . . . 79

274

 

Keys and Descriptions

78. Leaves 1.3–2.5× longer than wide, horizontally spreading or ventrad [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 79. Leaves with (85–)100–250 short, linear teeth and a 2–3-cell-wide border of thick-walled cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ovata 79. Leaves with fewer teeth, border absent or 1 cell wide . . . . . . . . . . . . . . . . . . 80 80. Dorsal leaf base longly and broadly decurrent. Leaves with 35–65 short-linear teeth. Border absent. Robust plants (ca. 5 mm wide) of Polylepis forests above 3500 m in southern Ecuador (Cajas) . . . . . . . . . . . . . . . . . . . . . . . . P. priceana 80. Dorsal leaf base shortly decurrent. Leaves with 10–35(−40) linear to triangular teeth. Border present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 81. Leaf base without vitta-like area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. macra 81. Leaf base with a short vitta-like area. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 82. Midleaf cells 30–40 μm wide. Leaves rather widely spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. trichostoma 82. Midleaf cells smaller, 15–30 μm wide. Leaves transverse to obliquely spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bifaria 83. Leaves ovate-oblong, with a broad apex and frequently with a border of thickwalled cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 83. Leaves mostly elongate ovate-triangular, with a rather narrow apex and without border of thick-walled cells. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 84. Leaf border yellowish. Leaf base without vitta-like area. Teeth linear to lineartriangular. Cuticle fully smooth. . . . . . . . . . . . . . . . . . . . . . . . . P. adianthoides 84. Leaf border colorless or brown. Leaf base with a vitta-like area. All teeth linear. Cuticle finely striate-papillose (especially on the teeth) . . . . . . . P. pachyloma 85. Mature stem leaves 1.8–2.5× longer than wide, ventral leaf base toothed or entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 85. Mature stem leaves less than 1.8× longer than wide, ventral leaf base entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 86. Ventral leaf base entire, longly decurrent. Leaf apex not with 1–2 conspicuously larger teeth . . . . . . . P. raddiana (phenotype with intercalary branching) 86. Ventral leaf base toothed, decurrent or not. Leaf apex usually with 1–2 conspicuously larger teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 87. Plants robust, 4–8 mm wide. Ventral leaf base strongly ampliate, fully covering the stem and forming a high crest. Ventral leaf base not or scarcely decurrent. Plants common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cristata 87. Plants delicate, 2–3 mm wide. Ventral leaf base scarcely ampliate, not covering the stem, crest absent. Ventral leaf base conspicuously decurrent. Plants of West Indies and Western Cordillera of Colombia, rare . . . . . . . . . . . . . . . . P. arcuata 88. Leaf base with a large vitta-like area. Leaves contiguous or subimbricate, ventrad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. deflexa 88. Leaf base without or with a weak vitta-like area. Leaves imbricate, horizontally spreading or slightly ventrad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 89. Dorsal leaf bases very longly decurrent, completely concealing the stem. Leaf apex bifid. Plants very robust, 5–11 mm wide. Plants common in upper montane cloud forests and páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fuscolutea

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275

  89. Dorsal leaf bases not very longly decurrent, not concealing the stem. Leaf apex not bifid. Plants less robust, 2.5–5 mm wide. Plants of montane forests up to 2500 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grandicrista   90. Mature stem leaves conspicuously elongate, mostly more than 2× as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91   90. Mature stem leaves less elongate, mostly less than 2× as long as wide [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98  91. Midleaf cells conspicuously elongate, many cells at least 2× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92   91. Midleaf cells shorter, 1–1.5(−1.7)× as long as wide [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93   92. Leaves widest at the base. Leaf apex deeply split into 2–4 long, ± linear teeth with a broad base, the teeth fragile, 1/3–2/3 of leaf length. . . . . P. tabinensis  92. Leaves widest in the middle. Leaf apex with shorter teeth, the teeth not fragile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aerea   93. Leaf cells finely papillose. Plants less than 2 mm wide . . . . . . . . . . P. tenuis   93. Leaf cells smooth. Mature plants usually more than 2 mm wide . . . . . . . . 94   94. Leaves horizontally spreading. Perianth not “naked,” inner bracts inserted at perianth base and covering the lower half of the perianth. . . . . . . . . . . . . . 95   94. Leaves ventrad. Perianth “naked” (inner bracts inserted at some distance from the perianth base and not covering the lower half of the perianth) or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97   95. Midleaf cells large, 30–45  μm wide, very thin-walled, with small trigones. Leaves ± rectangular, apex with long spinose teeth . . . . . . . . . . . P. subplana   95. Midleaf cells smaller, ca. 20–30 μm wide, with small to large trigones. Leaves narrowly oblong or rectangular, apex with short teeth . . . . . . . . . . . . . . . . 96   96. Leaves narrowly oblong, teeth restricted to the upper part of the leaf (absent in the lower part). Plants when fresh with peppermint smell . . . . . P. rutilans   96. Leaves ± rectangular, teeth not restricted to the upper part of the leaf (except in young plants). Plants when fresh without peppermint smell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. disticha (phenotype with intercalary branching)   97. Leaves widest at the base, elongate triangular. Ventral leaf base with a high shoulder and conspicuously and narrowly decurrent. Perianth not “naked.” Plants of the West Indies and the Western Cordillera of Colombia, rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. arcuata  97. Leaves widest in the middle, narrowly oblong. Ventral leaf base without high shoulder, not or scarcely decurrent. Perianth “naked.” Plants common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. gymnocalycina   98. Leaf cells finely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99   98. Leaf cells smooth (occasional slightly papillose in the lower half of the leaf: P. bifaria, couplet 103) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101   99. Plants very small, 0.5–1.5 mm wide. Leaves less than 1 mm long, bifid and with 1–5 teeth on the ventral margin. Leaf base without vitta. In upper montane cloud forests and páramo. . . . . . . . . . . . . . . . . . . . . . . . . . P. papillifolia

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Keys and Descriptions

  99. Plants larger, 2–4 mm wide. Leaves larger, not or slightly bifid, ventral margin with more than 5 teeth. Leaf base with a vitta. From sea level to the páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 100. Leaves with a border of thicker-walled cells. Teeth linear . . . . P. pachyloma 100. Leaves without border of thicker-walled cells. Teeth narrowly triangular (rarely linear) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. stricta 101. Leaves transverse to obliquely spreading, orbicular to ovate-oblong. Leaf base with a distinct vitta-like area. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 101. Leaves obliquely to widely spreading. Leaf base with or without vitta-like area [when in doubt try both leads]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 102. Leaves strongly caducous, branches frequently denuded . . . . . . . P. punctata 102. Leaves not caducous, branches not denuded . . . . . . . . . . . . . . . . . . . . . . 103 103. Leaf cells thin-walled, with very small trigones. Leaves with 2–5(−7) small teeth, the teeth only 1–2(−3) cells long, apical teeth not larger. Plants very rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subundulata 103. Leaf cells with conspicuous trigones. Leaves with 5–20 teeth, the teeth 2–10(−15) cells long, apical teeth usually larger. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bifaria 104. All teeth of mature leaves broken, leaves looking battered. Juvenile leaves with very long, simple to pinnately branched teeth. Plants in lowland rainforests of the Chocó. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. eggersii 104. Teeth of mature leaves not broken or only some teeth broken. Juvenile leaves without very long pinnately branched teeth . . . . . . . . . . . . . . . . . . . . . . . 105 105. Leaves horizontally spreading, not caducous. Trigones absent or small. Plants in lowland and montane forests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 105. Leaves ± ventrad, caducous or not. Trigones well developed, small or large. Plants mostly in montane forests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 106. Ventral leaf base longly decurrent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 106. Ventral leaf base not or very shortly decurrent . . . . . . . . . . . . . . . . . . . . . 108 107. Leaves with 0–15 teeth. Leaf cells small, 20–25 μm wide in midleaf. Plants of Amazonian lowland forest . . P. patula (phenotype with intercalary branching) 107. Leaves with 20–100 small teeth. Leaf cells larger, 25–40 μm wide in midleaf. Plants of montane forests above 1800 m . . . . . . . . . . . . . . . . . . . P. alternans 108. Leaf cells 30–40 μm wide in midleaf, trigones absent or very small. Leaves with (5–)15–40-cell-long teeth. Plants of lowland and submontane forests, below 1800 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. subplana 108. Leaf cells smaller, less than 30 μm wide in midleaf, trigones well developed. Leaves with less than 15 short teeth. Plants of lowland forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. disticha (phenotype with intercalary branching) 109. Leaves not caducous. Perianth base covered by bracts. . . . . . . . . . . P. macra 109. Leaves frequently caducous. Perianth base covered by bracts or “naked” (bracts at some distance from the perianth) . . . . . . . . . . . . . . . . . . . . . . . . 110 110. Leaf base with a vitta-like area. Leaves 0.9–1.2(−1.5)× longer than wide. Perianth not “naked,” bracts inserted at the perianth base and covering the lower half of the perianth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. punctata

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110. Leaf base without vitta-like area. Leaves more than 1.2× longer than wide. Perianth ± “naked,” inner bracts inserted at some distance from the perianth base and not covering the lower half of the perianth. . . . . . . . . . . . P. simplex 1. Plagiochila adianthoides (Sw.) Lindenb. Plagiochila densispina Steph. Plagiochila esmeraldana Steph. Plagiochila eximia Spruce Plagiochila glomerulifera Herzog Plagiochila keckiana Steph. Plagiochila notidophila Spruce Plagiochila procera Lindenb. Plagiochila saladona Steph.

Fig. 27G, H

On bark, twigs, and humic soil in montane forests, (300–)1500–3400  m. Colombia: Ant, Boy, Cau, Cho, Cun, Hui, Mag, NSa, Ris, San, Tol. Ecuador: Az, Ch, LR, MS, Na, Pa, Pi, Su, Tu, ZC. General distribution: widespread in tropical America. Plagiochila adianthoides (sect. Adianthoideae) is a common neotropical species that is recognized by (1) plants robust, 5–10 mm wide, branching intercalary; (2) leaves ovate-oblong with a broad apex, 1.3–1.8× longer than wide, ampliate or with a high shoulder, margins usually with a yellowish border of elongate thick-walled cells (but border sometimes only weakly developed), with ca. 15–50 linear to lineartriangular teeth, the teeth not larger at apex, lower half of dorsal margin without teeth, leaf bases rather shortly decurrent; (3) leaf cells rather small, 20–35 μm wide in midleaf, with small to large, bluntly radiate trigones, becoming thicker-walled and with larger trigones along the leaf margin, vitta-like area lacking, cuticle smooth; (4) androecia intercalary or terminal, occasionally two male spikes in a terminal cluster; and (5) vegetative reproduction by leaf fragmentation (seen in Chocó, Gradstein 8932, COL). Plagiochila adianthoides resembles P. superba and P. amicta but differs from the latter two in having smaller leaf cells, male branches intercalary or terminal and usually not in a cluster, and the presence of yellowish leaf border of thick-walled cells. Sometimes, however, the leaf border is only weakly developed or almost lacking. Plagiochila adianthoides may also be confused with P. trichostoma; for differences see under the latter species. 2. Plagiochila aerea Taylor Plagiochila alpina Gottsche Plagiochila bursata (Desv.) Lindenb., nom. illeg. Plagiochila bursata var. andina Spruce Plagiochila cobana Steph. fo. linearis Herzog Plagiochila macvicarii Steph. Plagiochila quitensis Steph. Plagiochila pungens Steph. Plagiochila rutilans Lindenb. var. aequatorialis Spruce

Fig. 27I, J

On bark of trees, rock, and soil in submontane and montane rainforests and cloud forests, at forest margins, and in open, scrubby vegetation, occasionally in the

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Keys and Descriptions

lowlands, (100–)500–3500  m. Colombia: Ama, Ant, Ara, Boy, Cau, Cho, Hui, Mag, Nar, Qui, Ris, Val, Vau. Ecuador: Ch, EO, Es, Ga, Lo, LR, MS, Na, Pa, Pi, Su, Tu, ZC. General distribution: common and widespread in tropical America but rare in Brazil. Plagiochila aerea (sect. Fuscoluteae) is a common neotropical species that is distinguished by (1) plants small to large, (1–)3–6(−8) mm wide, branching intercalary, sometimes a few terminal branches present as well; (2) leaves distant to imbricate, ventrad to almost horizontally spreading, narrowly oblong to ligulate, ca. 2.5–6× longer than wide, not ampliate, widest in the middle, unbordered, apex with 1–2 long, triangular to narrowly lanceolate teeth, ± bifid, ventral margin with 0–5 smaller teeth mostly in the upper half, leaf bases shortly decurrent, entire; (3) leaf cells rectangular, (1.5–)2–2.5× longer than wide in midleaf, 30–60  ×  15–30  μm, with radiate and often confluent trigones on longer walls, cuticle smooth to slightly rough; and (4) vegetative reproduction by caducous leaves. Plagiochila aerea is a highly variable species (Grolle and Heinrichs 1999). Plants with ligulate, bifid leaves may resemble P. bicuspidata and P. cuneata but the latter species are smaller and have short leaf cells, and the leaves of P. cuneata are fragmenting, not caducous. Large plants with several teeth resemble P. distinctifolia, P. macrifolia, and P. rutilans, but the latter species have short leaf cells and shorter teeth. Moreover, the leaves in P. distinctifolia are usually bordered. Plagiochila aerea is related to P. tabinensis, but the latter species is a more robust, brownish plant with leaves widest at the base and the apex deeply split into longlinear, fragile, lobe-like teeth. 3. Plagiochila alternans Lindenb. & Gottsche Plagiochila calomelanos Spruce Plagiochila columbica Gottsche Plagiochila grandifolia Steph. Plagiochila oblita Steph. Plagiochila rosariensis Steph. Plagiochila saettonii Steph. Plagiochila semidentata Steph.

Fig. 27L, M

On humic soil and moist rock in humid montane forests, 1500–3300  m. Colombia: Boy, Cal, Cau, Cun, Mag, Nar, Ris, San, Tol. Ecuador: Ca, Lo, MS, Na, Pi, Tu, ZC. General distribution: tropical America. Plagiochila alternans (sect. Denticulatae) is a rather common neotropical species that is recognized by (1) plants 4–10  mm wide, branching intercalary; (2) leaves horizontally spreading, symmetrically ovate-oblong, 1.2–2× longer than wide, not ampliate, unbordered, with 20–70(−100) teeth in the upper half and sometimes down to the ventral base, lower half of dorsal margin entire, teeth linear, 1–4 cells long; leaf bases longly and narrowly decurrent; and (3) cells ca. 25–40 μm wide in midleaf, isodiametric to elongate, without or with very small trigones, cuticle smooth. Plagiochila alternans is somewhat similar to P. subplana, but the latter species has more rectangular leaves with a truncate apex, a very shortly decurrent ventral leaf base, and less densely toothed leaf margins with much longer teeth. Moreover, P. subplana occurs at much lower elevation, usually below 1000  m. Plagiochila

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alternans is related to P. ovata, but leaves in P. ovata are orbicular (not ovateoblong), distinctly ventrad and toothed all around by numerous small, linear teeth (more than 100), and the leaf cells have large trigones. Moreover, P. ovata occurs at higher elevation, above 3000 m (Müller et al. 1999). 4. Plagiochila amicta Steph. Plagiochila douinii Steph., Plagiochila subaequalis Steph.

Fig. 27K

On bark, rock, or soil in montane forest and scrubby subpáramo, 1400–3100 m. Colombia: Ant, Cun, Mag. Ecuador: Tu, ZC.  General distribution: tropical America. Plagiochila amicta (sect. Hylacoetes) is an uncommon neotropical species that is distinguished by (1) plants large, 7–10 mm wide, branching intercalary; (2) leaves ovate-oblong with a broad apex, 1.2–1.8× longer than wide, shouldered or ampliate, unbordered, with ca. 6–40 triangular teeth, teeth to 6 cells long, bases rather shortly decurrent; (3) cells large, 35–55 μm wide in midleaf, slightly elongate, with small, radiate trigones, cuticle smooth; and (4) androecia in clusters of male branches at the plant apex. Plagiochila amicta is similar to P. superba var. macrotricha in leaf shape but differs in the shape of the marginal teeth, which are mostly linear (not triangular) in P. superba (Heinrichs 2002a). Plagiochila boryana, an Afro-American species recorded from Bolivia and SE Brazil, closely resembles P. amicta but differs by having a papillose cuticle. 5. Plagiochila arcuata Lindenb. Type: St. Vincent, unknown collector [Guilding? ], ex herb. Hooker (isotype PC-Mont.!, c. gyn.) On pole in wet, liverwort-rich submontane rainforest of the Cordillera Occidental, 1200 m. Colombia: Ris (Mistrato, trail Jeguadas to Puerto de Oro, 27 Jul 1992, Gradstein 8574, GOET, PC). General distribution: West Indies, western Colombia. Gradstein (2015a) considered the Caribbean P. arcuata (sect. Fuscoluteae) a slender phenotype of P. cristata, but comparison of fresh material from Colombia of P. arcuata with the type shows that P. arcuata deserves recognition. Characteristic features of the species are (1) plants delicate, 2–3 mm wide, branching intercalary; (2) stem leaves narrowly elongate-triangular, conspicuously narrowed to a truncate and usually bifid apex, 2–2.7× longer than wide (branch leaves smaller and shorter, ovate-oblong, 1.5–2× longer than wide), unbordered, with ca. 20–30 linear to narrowly triangular teeth, the teeth 3–6 cells long, the 2 upper tooth cells 5–7× longer than wide, leaf apex with 1–2 longer teeth (to 8 cells long), dorsal base entire and rather longly decurrent, ventral base with a high shoulder (scarcely ampliate) and narrowly decurrent, the decurrent part 0.1–0.15 mm long; (3) cells ca. 20–30 μm wide in midleaf, somewhat elongate, with rather large, radiate trigones, cuticle smooth; and (4) androecia intercalary. Plagiochila arcuata differs from P. cristata in smaller plant size (2–3 mm wide) and shouldered ventral leaf bases, which are conspicuously but narrowly decurrent

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Keys and Descriptions

and not or slightly ampliate, reaching only to the middle of the ventral stem base and leaving the ventral stem surface well visible. Plagiochila cristata is more robust (4–8 mm wide) and has scarcely decurrent and strongly ampliate ventral leaf bases, reaching across and beyond the ventral stem surface, covering the stem surface entirely, and forming a high crest. 6. Plagiochila bicuspidata Gottsche Plagiochila inouei Grolle Plagiochila nudiuscula Inoue, nom. illeg.

Fig. 28A

On bark and rock in humid upper montane forests, Polylepis forest, and páramo, 3000–4560  m on the mainland; at lower elevation on the Galápagos Islands, 750–800 m. Ecuador: Az (Gradstein and Benitez 2017), Ca, Ga (Inoue 1977 as P. nudiuscula), Na (on Polylepis pauta, Gradstein 12803, QCA), Pi (Mojanda lake, on Polylepis pauta, León-Yánez s.n., QCA); to be expected in Colombia. General distribution: scattered in the mountains of Central America (Mexico, Costa Rica) and the tropical Andes (Venezuela, Ecuador; new to Peru: La Convención, S. Teresa, lower Phaccaq valley, 4560  m, Sylvester s.n., PC); at lower elevation on the Galápagos Islands and in Chile. Plagiochila bicuspidata (sect. Heteromallae) is a small high-montane species recognized by (1) plants small, 1.5–2.5 mm wide, pale green to yellowish-green to brown, branching intercalary; (2) leaves distant, obliquely spreading, ventrad, narrowly oblong, 1.7–2.5× longer than wide, sometimes caducous, emarginate or short-bifid to 1/5(−1/3), lobe apices rounded to subacute, margins entire or dorsal leaf margin with a short tooth, leaf bases not or little decurrent; (3) cells 25–33 μm wide in midleaf, 1–1.5× longer than wide, trigones small to conspicuous, vitta-like area lacking, cuticle smooth; and (4) vegetative reproduction by caducous leaves. The species is remarkably common in humid Polylepis forests of Ecuador. By the small plants with oblong, bifid leaves, P. bicuspidata resembles P. cuneata, P. tenuis, and bifid phenotypes of P. aerea. Plagiochila cuneata differs from P. bicuspidata in the fragmenting leaf lobes, P. tenuis in the papillose cuticle, and P. aerea in the more elongate leaf cells and the sharply acute-acuminate leaf lobes (= teeth). Plagiochila bicuspidata may also be confused with P. exigua, but the leaves of P. exigua are less elongate, 1–1.5× longer than wide, and the leaf cells are smaller, less than 25 μm in midleaf. According to Groth et al. (2004), fresh plants of P. bicuspidata may emit a peppermint smell. 7. Plagiochila bifaria (Sw.) Lindenb.

Fig. 28C–E

Plagiochila arnelliana Steph., syn. nov. Type: Ecuador, Chimborazo, Spruce s.n. (G-00064088!) Plagiochila arrecta Gottsche Plagiochila atrovirens Taylor Plagiochila biceps Steph. Plagiochila centrifuga Spruce Plagiochila compressula (Nees) Lindenb. Plagiochila echinella Gottsche (?) (type not found) Plagiochila exesa Lindenb. & Gottsche

Fig. 28 (A) Plagiochila bicuspidata, habit in dorsal view, 33×. (B) Plagiochila bicornis, habit in ventral view. (C–E) Plagiochila bifaria. (C and D) Leaves, showing vitta-like area at the base (D: 18×). (E) Midleaf cells. (F and G) Plagiochila bryhnii. (F) Habit in ventral view, showing deflexed leaves (type of P. reclinata Herzog), 20×. (G) Leaf, 20×. (H and I) Plagiochila canelensis. (H) Habit, dorsal view. (I) Leaf. (J) Plagiochila cleefii, habit in lateral view, 17×. (K and L) Plagiochila cuatrecasii. (K) Leaves, 10×. (L) Habit with perianth, lateral view, 10×. (M and N) Plagiochila cucullifolia var. cucullifolia. (M) Habit with perianth, dorsal view. (N) Habit, ventral view. (O) Plagiochila cucullifolia var. anomala, 10×. (P) Plagiochila deflexa, 7×. (Q and R) Plagiochila deflexirama. (Q) Habit, dorsal view. (R) Branching pattern (pinnate). (A redrawn from Inoue 1977; B, H, I redrawn from Heinrichs 2002a, b; C, E redrawn from Heinrichs et al. 1998; F, G redrawn from Herzog 1932; K, L redrawn from Robinson 1967; L, M redrawn from Inoue 1987; M, N from Gradstein et al. 2001; O redrawn from Heinrichs et al. 2003; P, Q redrawn from Heinrichs et al. 2002b)

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Keys and Descriptions

Plagiochila fragilis Taylor Plagiochila geniculata Lindenb. Plagiochila gradsteinii Inoue Plagiochila horrida Gottsche Plagiochila hystrix Steph. Plagiochila implexa Lindenb. & Gottsche Plagiochila increscentifolia Spruce Plagiochila jaramilloi H. Rob. Plagiochila neorichardsii Inoue Plagiochila panamensis Inoue, syn. nov. Type: Panama, Chiriquí, Fortuna, 1150 m, “on living tree in forest,” Spörle & Aranda 1.33 (TNS n.v.) Plagiochila pinnata Spruce Plagiochila pinnatidens Steph. Plagiochila spinifera Ångstr. Plagiochila subrotundifolia Steph., syn. nov. Type: Ecuador, Mt. Tungurahua, Spruce s.n. (G-00061601!) Plagiochila trifida Steph. Common on bark, twigs, decaying wood, humic soil, and rock in montane forests, scrub, and páramo, 500–4200 m. Colombia: Ama, Boy, Ces, Cun, Hui, Mag, Nar, Qui, Vau. Ecuador: Az, Ca, Ch, Co, Ga, Lo, Na, Pa, Pi, Tu, ZC. General distribution: tropical America and coastal areas of western Europe. Plagiochila bifaria (sect. Arrectae) is a very common and widespread species that is distinguished by (1) plants 1–3 mm wide, green to brownish, arising from a rhizomatous base, stem apex conspicuously curved downward, branching sparse to regularly pinnate, purely intercalary; (2) leaves transverse and laterally appressed or obliquely spreading, asymmetrically ovate-orbicular to ovate-oblong, 0.8–1.5× longer than wide, often fragmenting with upper parts of leaf or teeth broken, not or slightly ampliate, unbordered, dorsal margin ± entire and straight, strongly recurved, ventral margin strongly arched and plane, with 3–40 short or long, narrowly triangular to linear to lobe-like teeth (2–11 cells long), the teeth at the leaf apex usually larger; leaf bases shortly to longly decurrent; (3) cells 15–30 μm wide in midleaf, subisodiametric (except at base), trigones conspicuous, vitta-like area present, cuticle smooth or faintly papillose; (4) rhizoids sometimes present in small tufts on the ventral stem surface at the ventral leaf base; and (5) vegetative reproduction by leaf fragmentation and fragmenting teeth. Plagiochila bifaria is highly variable and has been described under many different names (Heinrichs et  al. 2004a). Plants with broken teeth have been called P. exesa, those with strongly fragmented leaves P. gradsteinii and P. panamensis (the latter species is synonymized here based on the detailed original description and illustration; Inoue 1989b). Plagiochila bifaria is closely similar to P. pachyloma, and these two species have sometimes been confused, but P. pachyloma has a leaf border of thicker-walled cells, which is often brownish in color, and purely linear teeth, while the teeth in P. bifaria are more variable in shape, triangular to linear to lobe-like. Plagiochila bifaria also resembles robust phenotypes of P.

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punctata with suberect leaves, but the latter species clearly differs by copious production of caducous leaves. In P. bifaria, leaves are not caducous. Moreover, the dorsal leaf base in P. punctata is scarcely decurrent. Phenotypes of P. bifaria with laterally appressed leaves may be confused with Adelanthus lindenbergianus, but in the latter species, the dorsal leaf margin is curved inward, not outward, and the teeth are much smaller; both species have a short vitta-like area. Plagiochila bifaria also resembles Plagiochila retrorsa Gottsche from Mexico and Central America, but the leaves in P. retrorsa are more ovate-triangular and toothed all around, also on the dorsal margin (Rycroft et al. 2001). 8. Plagiochila breuteliana Lindenb. Plagiochila breuteliana var. acutifolia Herzog Plagiochila breuteliana var. novo-granatensis Gottsche Plagiochila paucispinula Herzog On bark and soil in lowland and montane rainforest, 200–2700 m. Colombia: Ant, Boy, Caq, Cho, Cun, Guj, Mag, Ris, Val. Ecuador: ZC (Heinrichs 2002a). General distribution: tropical America. Plagiochila breuteliana (sect. Hylacoetes) is a widespread neotropical species that is recognized by (1) plants large, 5–8  mm wide, branching intercalary; (2) leaves triangular to oblong with a narrow or broad apex, 1.4–2× longer than wide, shouldered to ampliate, unbordered, with 20–80 triangular to linear teeth, the teeth to 10 cells long, dorsal and ventral leaf bases toothed and longly decurrent; (3) cells large, 35–55  μm wide in midleaf, slightly elongate, with small, radiate trigones, vitta-like area present, cuticle smooth; and (4) androecia usually intercalary (not in clusters). Plagiochila breuteliana is very similar to P. vincentina; both species have ampliate leaves with toothed and longly decurrent bases (both dorsal and ventral). The two species differ mainly in the presence of a short vitta-like area in the leaves of P. breuteliana (Heinrichs 2002a). 9. Plagiochila bryhnii Steph.  Fig. 28F, G Plagiochilion bryhnii (Steph.) Inoue Plagiochila reclinata Herzog (Plagiochilion reclinata [Herzog] R.M.Schust.) On moist earth banks in montane forest, 2000–2600  m. Colombia: San, Val. General distribution: tropical Andes. Plagiochila bryhnii (sect. Heteromallae) is a rare Andean species that is distinguished by (1) plants delicate, 2–3 mm wide, branches lateral-intercalary; (2) leaves opposite with connate dorsal and ventral bases, distant, secund, asymmetrically ovate, 1.1–1.3× longer than wide, not ampliate, unbordered, with 7–14 teeth along the ventral and apical margins, teeth at apex often larger, dorsal base longly decurrent, ventral base shortly decurrent; (3) cells 15–30 μm wide in midleaf, walls flexuose and with distinct trigones, vitta-like area absent, cuticle smooth; (4) perianth base stalked and “naked,” not covered by bracts; and (5) vegetative reproduction by caducous leaves.

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Keys and Descriptions

Plagiochila bryhnii is the only neotropical Plagiochila species with connate dorsal and ventral leaf bases. Opposite leaves are shared with P. macrostachya and P. heteromalla, but in the latter two species, the leaf bases are not connate, the plants are more robust (more than 4 mm wide), and the leaves are usually ampliate and with more than 15 teeth. In addition, P. heteromalla differs in having bordered leaves and P. macrostachya by larger leaf cells. 10. Plagiochila buchtiniana Steph. Plagiochila friabilis Inoue On bark or soil in Polylepis forest, scrub, and páramo, 3100–4000 m. Ecuador: Az, Co (Llanganates, Burghardt 6764, QCA), Im (Schäfer-Verwimp et al. 2006), Pi (León-Yánez s.n., QCA), ZC (Reserva Biologica San Francisco, Burghardt 7141, QCA); to be expected in Colombia. General distribution: Central Andes (Ecuador to Bolivia). Plagiochila buchtiniana (sect. Glaucescentes) is a rare central Andean species from high elevation, ranging northward to Ecuador. The species is recognized by (1) plants large, 4–7 mm wide, pale yellowish-green to brownish, branching predominantly terminal; (2) leaves horizontally spreading, usually strongly fragmenting, broadly ovate-triangular, 1.1–1.4× longer than wide, ampliate and recurved, apical and ventral margins toothed by about 5–18 teeth, bases entire, longly and broadly decurrent; (3) cells 22–45 μm wide in midleaf, subisodiametrical, without or with small trigones, vitta-like area absent, cuticle smooth, oil bodies brown; (4) underleaves conspicuous, to 1.2 mm long, triangular, ciliate; and (5) vegetative reproduction by leaf fragmentation. Plagiochila buchtiniana is closely related to P. diversifolia but is readily separated from the latter in the pale yellowish-brown color, the fragmenting leaves, and the strongly ampliate and longly and broadly decurrent ventral leaf bases (Heinrichs et al. 2000). The species may be confused with P. raddiana but differs from the latter in shorter and broader, fragmenting leaves (less than 1.5× longer than wide), conspicuous underleaves, and brown oil bodies (colorless in P. raddiana). 11. Plagiochila canelensis Steph.

Fig. 28H, I

On bark and soil in upper montane forest and páramo, 2500–4000 m. Colombia: Cau, Cho, Cun. Ecuador: Lo, Na, Pa, Pi (páramo de Papallacta, Burghardt 7443, 7446, QCA), Tu. General distribution: tropical Andes; distribution incompletely known. Plagiochila canelensis (sect. Hylacoetes) is a poorly known high-Andean species that is very close to P. superba but differs in the shorter and broader leaves, about as long as wide (1–1.2× longer than wide). The species can be confused with toothed phenotypes of P. ensiformis, but the latter species differs in having smaller teeth, 1–2(−4) cells long, and not becoming longer toward the leaf apex. In P. canelensis the teeth are much longer, longly linear like in P. superba, 2–8(−11) cells long, being longest at the leaf apex. Moreover, in P. canelensis teeth may be present

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in the lower half of the dorsal leaf margin; in P. ensiformis the lower half of the dorsal leaf margin is entire (Heinrichs 2002a). 12. Plagiochila cleefii Inoue

Fig. 28I

On soil and moist rock in humid páramo and Polylepis forests, occasionally growing submerged, 3150–4300  m. Colombia: Boy (Bischler 2860, 2870, PC), Cun (Aguirre 1860, COL), Met (type). Ecuador: Az, Co (páramo de Llanganates, Burghardt numerous collections, PC, QCA), Lo, Na, Pi. General distribution: northern Andes (Venezuela [new, Pócs 5034/X, PC], Colombia, Ecuador). Plagiochila cleefii (sect. Arrectae) is a characteristic northern Andean páramo species that is distinguished by (1) plants 1–2.5 mm wide, pale yellowish-brown, little branched, apex curved, branching intercalary, rhizomatous base lacking; (2) leaves transverse to obliquely spreading, obovate, (1.0–)1.1–1.3× longer than wide, apex rounded to emarginate to short-bifid, ± plane, apical margins entire or with 1–2(−3) small teeth, dorsal and ventral margins entire, dorsal margin straight, ventral margin curved and plane, dorsal and ventral base shortly or longly decurrent; (3) cells 20–30 μm wide in midleaf, subisodiametric, trigones small to large, not elongate, short vitta-like area present, cuticle smooth; and (4) vegetative reproduction by caducous leaves. Plagiochila cleefii may be confused with P. revolvens and P. punctata; P. revolvens differs in the wider than long leaves with more strongly recurved margins and P. punctata in the toothed ventral leaf margin, scarcely decurrent leaf bases, and more strongly vegetative reproduction by caducous leaves. Plagiochila vittiana Inoue described from Mexico is very close to P. cleefii but differs in having orbicular leaves with a very shortly decurrent dorsal base and entire to finely toothed margins. 13. Plagiochila cristata (Sw.) Lindenb. On bark and rock in humid montane forests, 1500–2300  m. Colombia: Ant, Cho, Cun, Mag, Qui, Ris, San, Tol, Val. Ecuador: Pa, Tu, ZC. General distribution: widespread in tropical America. Plagiochila cristata (sect. Adianthoideae) is a common neotropical species that is recognized by (1) plants robust, 4–8 mm wide, branching intercalary, occasionally terminal; (2) leaves narrowly elongate-triangular, conspicuously narrowed to a truncate and usually bifid apex, 1.8–2.5× longer than wide, unbordered, with ca. 15–60 triangular to linear teeth, apex with 1–2 larger teeth, dorsal base entire, decurrent, ventral base strongly ampliate, forming a high crest, not or very shortly decurrent; (3) cells ca. 20–30 μm wide in midleaf, somewhat elongate, with rather large, radiate trigones, cuticle smooth; and (4) androecia intercalary. In its narrowly elongate-triangular leaves with 1–2 large, apical teeth and a strongly ampliate ventral base forming a high crest, P. cristata can hardly be confused with any other species except possibly P. arcuata. The latter species, however,

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Keys and Descriptions

is a smaller plant with scarcely ampliate and conspicuously decurrent ventral leaf bases (see also under P. arcuata). 14. Plagiochila cuatrecasii H.Rob. 

Fig. 28K, L

On soil in humid páramo, 3600–4200 m. Colombia: Boy, Met, Ris. Ecuador: Az, Pi (Gradstein and Benitez 2017). General distribution: northern Andes (Colombia, Ecuador). Plagiochila cuatrecasii (sect. Arrectae) is one of the smallest species in the genus and a rare páramo element. Characteristics are (1) plants less than 1  mm wide, branching intercalary, rhizomatous base lacking; (2) leaves distant, ovateorbicular to subquadrate, apex rounded to retuse, margins entire, plane or dorsal margin slightly recurved, bases hardly decurrent; (3) cells ca. 20–25 μm wide in midleaf, trigones large, cuticle smooth. In its entire leaves, P. cuatrecasii approaches P. cleefii and P. revolvens, which are also characteristic of páramo. The latter two species, however, are larger (plants more than 1 mm wide), the dorsal leaf bases are longly decurrent, and a short vitta-like area is present in the leaves. Plagiochila cleefii, moreover, is pale yellowish-green to yellowish-brown in color, not dark brown, and often bifid and has caducous leaves. Plagiochila cuatrecasii is confusingly similar to Leptoscyphus cuneifolius, which may occur in the same habitat. However, L. cuneifolius has underleaves. 15a. Plagiochila cucullifolia J.B.Jack & Steph. var. cucullifoliaFig. 28M, N Szweykowskia cucullifolia (J.B.Jack & Steph.) Gradst. & M.E.Reiner On bark of trees and shrubs in humid cloud forest areas and bogs, 1300–3000 m. The species usually grows in rather open habitats, e.g., at forest margins and on road sides in somewhat disturbed environments. Colombia: Ant, Cho, Hui, Nar. Ecuador: Na, ZC. General distribution: northern Andes, Costa Rica. Plagiochila cucullifolia var. cucullifolia (sect. Hylacoetes) is a striking northern Andean-Costa Rican species that is immediately distinguished by the swollen yellowish-brown plants with sac-like leaves. Further characters include (1) plants robust, 6–10 mm wide, ± unbranched; (2) leaves saccate-suborbicular, strongly convex, dorsal and ventral margins curved downward and with numerous (to 45) longlinear teeth, the teeth 5–20 cells long and mostly 1 cell wide; (3) leaf cells very large, ca. 50–80 μm wide in midleaf, cuticle smooth; and (4) androecia terminal to intercalary, not in clusters. 15b. Plagiochila cucullifolia var. anomala Heinrichs & Gradst.

Fig. 28O

On twigs and branches of low shrubs in open, wet subpáramo vegetation, 2900–3000 m. Ecuador: ZC (Heinrichs et al. 2003). General distribution: only known from the type locality in southern Ecuador. Plagiochila cucullifolia var. anomala is only known from subpáramo in the San Francisco Biological Reserve and resembles var. cucullifolia in the characteristic yellowish-brown color but lacks saccate leaves. Instead, the leaves are rather flat, laterally appressed, with plane ventral margin and recurved dorsal margin. Moreover, the plants are smaller (4–6 mm wide) and the leaf margins have fewer teeth (3–25).

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In spite of its very different morphology, var. anomala is genetically very close to var. cucullifolia (Heinrichs et al. 2003) and may therefore be considered an abnormal phenotype of the latter. Its unusual morphology may be largely due to environmental modification in the harsh páramo environment. 16. Plagiochila cuneata Lindenb. & Gottsche Plagiochila rara Gottsche On bark and rock in humid montane forest, on the mainland (Colombia) found at 2600 m, on the Galápagos Islands at 500–800 m. Colombia: Cun (type of P. rara). Ecuador: Ga (Isabela, Pinta, San Cristóbal, Santa Cruz). General distribution: Mexico to Bolivia. Plagiochila cuneata (sect. Heteromallae) is an uncommon neotropical-montane species that is distinguished by (1) plants 1–1.5 mm wide, with a peppermint smell when fresh, branching intercalary; (2) leaves rather distant, elongate-obcuneate to narrowly oblong, deeply bifid to trifid to 1/4–1/3 of leaf length, sometimes with an additional tooth near the apex, usually strongly fragmenting, upper part of leaves often broken, margins entire, bases shortly decurrent; (3) cells 20–32 μm wide in midleaf, 1–1.5× longer than wide, with distinct trigones, cuticle smooth; (4) perianth base free, not covered by bracts; and (5) vegetative reproduction by leaf fragmentation. Plagiochila cuneata may be confused with P. aerea, but the leaves in the latter species are caducous, not fragmenting, the cells are more elongate in midleaf (1.5–2.5× longer than wide), and the plants lack a peppermint smell. Both species are common on the Galápagos Islands and have been confused there (Grolle and Heinrichs 1999). Plagiochila cuneata is closely related to P. loriloba; both species have a peppermint smell and strongly fragmenting leaves. However, P. loriloba is a more robust and genetically distinct plant, to 2.5 mm wide, with imbricate, deeply 3–4-lobed leaves and larger cells, 25–43 μm wide in midleaf (Groth et al. 2002). Small forms of P. cuneata resemble P. bicuspidata, but the leaves in latter species are bifid (never trifid) and not fragmenting. 17. Plagiochila deflexa Mont. & Gottsche Fig. 28P Plagiochila chimborazensis Spruce, syn. nov. Type: Ecuador, Mt. Chimborazo, Spruce s.n. (lectotype, designated by Heinrichs et al. (2002a), MANCH-cc7553) Plagiochila ecuadorensis Steph., syn. nov. Type: Ecuador, Morona Santiago, above El Rosário, 1700 m, Allioni s.n., Bryotheca Levier 6363 (G-00061382!, c. gyn.) Plagiochila mutila Gottsche (?) Plagiochila subtrinitensis Herzog Plagiochila tovarina Gottsche var. trianae Gottsche, syn. nov. Type: Colombia, without loc., Triana s.n. (isotype G-00282660!, ster.) On bark and rock in montane rainforest, growing procumbent or pendent, ca. 1500–2000 m. Colombia: without loc. (type of P. tovarina var. trianae). Ecuador: Ca (Heinrichs et  al. 4425, GOET), Ch, MS, Pi (Gradstein 12631, QCA), ZC

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Keys and Descriptions

(Holm-Nielsen et al. 4071, PC). General distribution: Central America, Greater Antilles, northern Andes (Venezuela to Ecuador), Hawaii. Plagiochila deflexa (sect. Heteromallae) is a neotropical-Hawaiian disjunct that is recognized by the medium-sized plants (ca. 4–5  mm wide) with intercalary branching; rather distant and ventrad, ovate-oblong to elongate-triangular leaves with a narrowly truncate apex and ampliate and longly decurrent ventral bases; leaf margins with rather few teeth (frequently with 2 larger teeth at apex) and without border; midleaf cells subrectangular (ca. 1.5–2× longer than wide), ca. 25–35 μm wide and with conspicuous, radiating trigones; and the presence of a large vitta-like area. The perianth base is usually covered by bracts; Heinrichs et  al. (2002a) described the perianth of P. chimborazensis as “naked,” but his illustration (Plate 46B) shows a perianth covered by bracts in the lower half. Plagiochila deflexa was described from Colombia as P. tovarina var. trianae and from Ecuador as P. ecuadorensis and P. chimborazensis. Plagiochila mutila described from Venezuela may be a further synonym of P. deflexa; I have not seen the type, but a specimen from Mérida collected by Goebel was identified by Stephani as P. mutila, with the annotation “stimmt gut mit Gottsche’s Pflanze von Moritz gesammelt” [fits the type of G. mutila well]. Plagiochila deflexa is related to P. trichostoma and P. macra, but P. trichostoma differs in less elongate and more densely imbricate leaves and absence of 2 larger teeth at leaf apex, while P. macra differs in the lack of a vitta-like area. Stem leaves of P. deflexa are weakly ampliate, but branch leaves may not be ampliate and may resemble leaves of P. simplex or P. gymnocalycina. The latter two species, however, have naked perianths and leaves without vitta-like area. 18. Plagiochila deflexirama Taylor Fig. 28Q, R Plagiochila andicola Mont. & Gottsche, syn. nov. Type: Ecuador, near Quito, Jameson s.n. (isotype G-00282870!, ster.) Plagiochila bicaudata Steph., syn. nov. Type: Ecuador, Morona Santiago, Bomboiza, 30 Jan 1910, Allioni 215 (lectotype, designated here, G-00128747!) Plagiochila bryopterioides Spruce Plagiochila dilatata Steph. Plagiochila filicina Herzog Plagiochila relicta Steph., syn. nov. Type: Colombia, Cundinamarca, Pacho, Lindig s.n. (G-00064906!) Plagiochila subdeflexiramea Herzog Plagiochila trianae Gottsche, syn. nov. Type: Colombia, without loc., Triana s.n. (lectotype, designated here G-00282659!, c. andr. & gyn.) Plagiochila trollii Herzog On bark and rotten wood in montane rainforest and cloud forest, (300–) 900–2800 m. Colombia: Ant, Boy, Cas, Cun, Hui, Qui, Put, Tol. Ecuador: Az, EO (300 m), Lo, MS, Na, Pi, ZC. General distribution: Mexico to Bolivia. Plagiochila deflexirama (sect. Vagae) is a common Andean species that is distinguished by (1) plants medium-sized to large, 3–5 mm wide and 15 cm long, upper part of shoot (bi)pinnate by terminal branches, lower part often more sparsely

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branched and sometimes with a few intercalary branches; (2) leaves (sub)imbricate, ± ventrad, ovate to oblong, 1.1–1.8× as long as wide, widest near the base, conspicuously narrowed to apex, weakly to strongly ampliate, ventral bases concealing the stem or not, apical and ventral margin sharply toothed, with (2–)10–25 teeth, apex sometimes bifid, dorsal leaf base rather longly decurrent, swollen, ventral leaf base shortly to rather longly decurrent; (3) cells 15–28 μm wide in midleaf, isodiametric to elongate, trigones conspicuous, elongate, on longer walls often subconfluent, vitta-like area absent, cuticle smooth; and (4) vegetative reproduction by small plantlets (propagula) on ventral leaf surfaces. Among the neotropical Plagiochila species with predominantly terminal branching, a (bi)pinnate branching pattern is only seen in P. deflexirama and P. tamariscina; all other species are dichotomously branched. The pinnate branching pattern is particularly evident in mid-portions of the plant; in the lower parts, branches may be scarce, and in the upper part, the branching can sometimes be dichotomous. Plagiochila tamariscina is close to P. deflexirama but differs in being more delicate and by leaves distant, more narrowly elongate and not ampliate (but sometimes shouldered), widest in the middle or the lower third. Plagiochila andicola, P. bryopterioides, and P. longitexta are phenotypes of P. deflexirama with weakly ampliate leaves. Plagiochila trianae Gottsche is a further synonym; this species was based on sterile specimen collected by Lindig and a fertile one collected by Triana. The original syntypes are destroyed, but an isosyntype of the Triana collection (with gynoecia and androecia) is in G and is designated here as lectotype. The male plant is very slender and unusually little branched, but its leaves are essentially similar to those of the female plant. 19. Plagiochila dependula Taylor Jamesoniella dependula (Taylor) Steph. Plagiochila pendula Steph.

Fig. 29A, B

On humic soil, rock, and leaning trunks in wet, subalpine forests near the forest line and in wet páramo, 3150–4300 m. Colombia: Cau, Cho, Hui, Met, Nar, Ris. Ecuador: Az, Ca, Co, Lo, Na, Pi, ZC.  General distribution: tropical Andes (Colombia to Bolivia). Plagiochila dependula (sect. Fuscoluteae) is a characteristic Andean páramo species that is recognized by (1) plants medium-sized to robust, stems very long (to 25 cm long), almost unbranched; (2) leaves transverse and laterally appressed, reniform with the leaf base widely expanded across and beyond the stem, 1.5–2.5× longer wide, apex rounded, margins entire, flat; (3) cells 30–40 μm wide in midleaf, isodiametric to elongate, trigones large and swollen, leaf base with a broad, illdefined vitta-like area, cuticle papillose; and (4) perianth mouth entire. Because of its entire leaves, P. dependula was sometimes placed in Jamesoniella (= Syzygiella), but the species clearly differs from the latter genus in the flattened perianth. 20. Plagiochila disticha (Lehm. & Lindenb.) Lehm. & Lindenb. Plagiochila blepharobasis Herzog

Fig. 29C–E

Fig. 29 (A and B) Plagiochila dependula. (A) Habit, dorsal view, 11×. (B) Midleaf cells. (C–E) Plagiochila disticha. (C) Habit in ventral view, showing ciliate leaf base. (D) Branching pattern (dichotomous). (E) Leaf base with two cilia. (F–H) Plagiochila distinctifolia. (F) Habit, dorsal view, 11×. (G) Leaf, 13×. (H) Leaf margin, showing border of thick-walled cells, 200×. (I–K) Plagiochila diversifolia. (I) Habit, dorsal view, 10×. (J) Leaf, 10×. (K) Midleaf cells, 300×. (L and M) Plagiochila dominicensis. (L) Leaf. (M) Midleaf cell. (N) Plagiochila exigua, habit in dorsal view, 60×. (O and P) Plagiochila ensiformis. (O) Leaves toothed. (P) Leaves entire. (Q and R) Plagiochila eggersii. (Q) Mature leaf with broken teeth, 25×. (R) Upper portion of young leaf with teeth, 70×. (A, B redrawn from Grolle 1972; C from Gradstein and Ilkiu-Borges 2009; E from Heinrichs and Gradstein 2000; F–H redrawn from Schuster 1980a; I–K redrawn from Inoue 1977; L, M, O, P redrawn from Heinrichs 2002a; N redrawn from Inoue 1980; Q, R redrawn from Inoue 1988)

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Plagiochila falcatoserrata Carl Plagiochila fimbristipula Spruce Plagiochila hortorum Spruce Plagiochila juruensis Steph. Plagiochila orbigniana Nees & Mont. Plagiochila pensilis Spruce Plagiochila serrata (Roth) Lindenb. Plagiochila thysanotis Spruce (incl. vars.) On bark and rock in lowland and lower montane rainforests, 100–1800  m. Colombia: Ama, Caq, Met, Nar, Put, Ris, Val, Vau. Ecuador: Es, MS, Na, Or, Pa, Pi, ZC. General distribution: tropical America. Plagiochila disticha (sect. Vagae) is a widespread and variable neotropical species that is distinguished by (1) plants 3–5  mm wide, mostly dichotomous with terminal branching, but sometimes unbranched or irregularly pinnate with intercalary branches only; (2) leaves horizontally spreading, ovate-lingulate or narrowly rectangular, 1.5–2.5× longer than wide, ventral base ampliate or not ampliate, when ampliate often forming a ventral crest, leaf apex subtruncate, with 1–5 rather small teeth, dorsal leaf margin entire, ventral leaf margin entire or with a few teeth especially at the base, teeth on ventral base of mature leaves long-linear, of 1 long row of cells, only the base of the tooth more than 1 cell wide; dorsal base in mature leaves shortly to longly decurrent, the decurrent part strongly swollen, ventral base not or shortly decurrent; (3) cells 20–30 μm wide in midleaf, isodiametric to elongate, trigones conspicuous, cells sometimes thicker-walled toward the apical margins (P. blepharobasis), cuticle smooth; (4) underleaves often conspicuous (but sometimes lacking), 0.5–1.5 mm long, margins strongly ciliate; and (5) vegetative reproduction by small plantlets (propagula) on ventral leaf surfaces. Plagiochila disticha is very similar to P. montagnei; both species have terminal branching, ovate-rectangular leaves with a long-decurrent dorsal base, and toothed ventral leaf bases. However, the teeth on ventral leaf bases in P. montagnei are usually triangular and more than 1 cell wide for most of their length, while being usually linear, of 1 row of cells for most of their length, in P. disticha. Moreover, the underleaves in P. disticha are more conspicuous, and the ventral underleaf bases always shortly decurrent. In P. montagnei the underleaves are rudimentary, and the ventral underleaf bases shortly to longly decurrent (Heinrichs and Gradstein 2000). Lowland forms of P. disticha are often not or scarcely ampliate, but robust montane forms may have strongly ampliate leaf bases, forming a high crest. The robust montane forms of P. disticha have been described as P. pensilis Spruce and P. falcatoserrata Carl (ventral leaf margin toothed) or as P. blepharobasis Herzog (ventral leaf margin entire except at the base). Plagiochila disticha is abundant in Amazonian rainforest of Ecuador (Yasuni), and juvenile or rhizomatic plants of the species are commonly seen on tree trunks. The juvenile plants are very tiny and have small, bifid leaves, resembling Lophocolea, while rhizomatic plants stand out by ovate or shortly rectangular leaves (as found on creeping rhizomes of mature plants) with almost entire ventral leaf bases, shortly to longly decurrent dorsal bases, and rounded leaf apices without or with a few

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Keys and Descriptions

irregular small teeth. Branching in these rhizomatic plants is purely intercalary. These phenotypes of P. disticha may be difficult to identify in the absence of mature plants. 21. Plagiochila distinctifolia Lindenb. Plagiochila diffusa Steph. Plagiochila hypantra Spruce Plagiochila lindigiana Gottsche Plagiochila miqueliana Lehm. & Lindenb.

Fig. 29F–H

On bark and rotten logs in lowland and montane rainforests, from sea level to 2700 m. Colombia: Cun, Mag, Put (Fernandez 423, COL). Ecuador: EO (Reserva Buenaventura, Benitez et  al. 1317, HUTPL), MS, Pi (Buck 10344, QCA), ZC. General distribution: tropical and subtropical America. Plagiochila distinctifolia (sect. Vagae) is a little-known neotropical species that is recognized by (1) plants delicate, 2–4  mm wide, dichotomous to subpinnate, branches predominantly terminal, intercalary branches few on older or broken stem portions; (2) leaves mostly distant (some leaves may be slightly imbricate), ± ventrad, ovate-oblong to lingulate, ca. 1.5–2.5× longer than wide, not ampliate, subsymmetrical, dorsal margin straight, ventral margin slightly arched, margins bordered by elongate, somewhat thick-walled cells, toothed in the upper half with 4–15 triangular teeth, often 2–3 larger teeth present at apex (especially in young leaves), dorsal base rather longly decurrent, ventral leaf base shortly decurrent; (3) cells 15–30 μm wide in midleaf, subisodiametrical, trigones small, cuticle smooth; and (4) vegetative reproduction by small plantlets (propagula) on ventral leaf surfaces. Plagiochila distinctifolia closely resembles the rare P. macrifolia from Ecuador, but the leaves in the latter species are not bordered and much less toothed. Plagiochila distinctifolia may also be confused with P. rutilans, but the latter species has intercalary branching, horizontally spreading leaves (not ventrad) without border, and a strong peppermint smell (fresh plants). 22. Plagiochila diversifolia Lindenb. & Gottsche Plagiochila dotensis Herzog Plagiochila galapagona Inoue Plagiochila glaucescens Steph. Plagiochila praetermissa Gottsche Plagiochila subedentata Steph.

Fig. 29I–K

On bark of trees and shrubs, occasionally on rock or moist soil, in montane forests, 500–3000 m. Colombia: Ant, Boy, Cun, Mag, Ris. Ecuador: Ch, Es, Ga, Lo, LR, Na, Or, Pa, Su, ZC. General distribution: Mexico, Central America, tropical South America. Plagiochila diversifolia (sect. Glaucescentes) is a common neotropical species that is distinguished by (1) plants rather robust, 5–9 mm wide, green to brownishgreen with dark brown stems especially in older parts, dichotomous in the upper half by terminal branches, older parts with a few intercalary branches; (2) leaves

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horizontally spreading, ovate-oblong, 1.1–1.8× longer than wide, not ampliate and without high shoulder, apex broadly rounded, margins toothed in the upper half by about 8–30 small to large teeth, teeth sometimes extending to the ventral base, dorsal margin entire except in the upper half, dorsal base rather longly and narrowly decurrent, ventral leaf base shortly decurrent; (3) cells 20–45 μm wide in midleaf, subisodiametrical, without or with small trigones, cuticle smooth, oil bodies brown; and (4) underleaves usually present, to 1 mm long, deeply bifid. Plagiochila diversifolia may be confused with P. laetevirens, but the latter species is less robust, with green to pale brown stems, shorter, ovate-subrectangular leaves with a truncate apex, a high shoulder and shortly decurrent bases, leaf cells with conspicuous trigones and colorless oil bodies, no underleaves, and vegetative reproduction by small plantlets (propagula) on ventral leaf surfaces. Plagiochila diversifolia may also be confused with forms of P. deflexirama with scarcely ampliate ventral leaf bases, but branching in P. diversifolia is dichotomous while being mostly pinnate in P. deflexirama. Moreover, P. diversifolia usually has small underleaves (lacking in P. deflexirama). A phenotype of P. diversifolia from southern Ecuador with minutely toothed leaves (teeth of only 1–2 cells) was erroneously identified as P. martiana (Nees) Lindenb. (Benitez and Gradstein 2011). 23. Plagiochila dominicensis Taylor Plagiochila pichinchensis Taylor var. basidentata Herzog Plagiochila virens Spruce

Fig. 29L, M

On bark and soil in lowland and montane rainforests, 100–2500 m. Colombia: Cho, Mag, Ris, Vau (Schultes & Cabrera 13860, CINC, PC). Ecuador: LR, Pa, Pi. General distribution: tropical America. Plagiochila dominicensis (sect. Hylacoetes) is a rather common neotropical species that is recognized by (1) plants large, 5–10 mm wide, branching intercalary; (2) leaves asymmetrically elongate-triangular to oblong, 1.3–2× longer than wide, shouldered or somewhat ampliate, unbordered, with 25–65 triangular to linear teeth, bases shortly decurrent, dorsal base usually densely toothed, ventral base entire; (3) cells 30–55 μm wide in midleaf, elongate, with conspicuous, radiate trigones, vittalike area absent, cuticle smooth; and (4) androecia in clusters of 2–6 male branches at stem apex. Plagiochila dominicensis resembles P. breuteliana, P. vincentina, and P. turgida by the toothed dorsal leaf base, but the latter three species are separated from P. dominicensis by thre more strongly ampliate ventral leaf bases. Moreover, in P. breuteliana and P. vincentina, the ventral leaf base is toothed and longly decurrent (entire and shortly decurrent in P. dominicensis), and the male branches are not arranged in clusters at the stem apex. Plagiochila dominicensis may also be confused with P. subplana, but in the latter species, the leaves are rather symmetrically rectangular to oblong, with a ± truncate apex, and the leaf cells are (sub)isodiametrical, without or with very small trigones (Heinrichs 2002a). 24. Plagiochila ecuadorica (Inoue) L.Söderstr. Steereochila ecuadorica Inoue (Plagiochila dimorpha Lindenb. var. ecuadorica [Inoue] Heinrichs)

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Keys and Descriptions

On bark of low trees and shrubs in upper montane cloud forests and bogs, usually in rather open places, 1500–2900  m. Ecuador: Ca, Na, Pi; to be expected in Colombia. General distribution: tropical Andes (Bolivia, Ecuador), Costa Rica. Plagiochila ecuadorica (sect. Hylacoetes) is a characteristic Andean-Costa Rican species that is immediately distinguished by the yellowish-green clusters of flagelliform branchlets at shoot apices (producing caducous leaves), resembling miniature heads of cauliflower. The yellowish-green color of these clusters contrast sharply with the dull olive green leafy shoots. Because of its peculiar apical clusters, this species was considered a separate genus, Steereochila (Inoue 1987). Further characters of the species are (1) plants 4–6  mm wide, branching intercalary; (2) leaves subopposite, somewhat ventrad, ovate-triangular to ovate-oblong, 1.1–1.5× longer than wide, not ampliate, margins toothed in the upper half of the leaf, bases longly decurrent dorsally and shortly decurrent ventrally; and (3) cells 30–50 μm wide in midleaf, cuticle smooth. In its large-celled, subopposite leaves, P. ecuadorica resembles P. macrostachya, but the latter species lacks clusters of caducous-leaved branchlets. 25. Plagiochila eggersii Inoue Plagiochila pinnatispina Gradst. (in sched.)

Fig. 29Q, R

On bark of living and dead trees in undisturbed and disturbed lowland rainforests, 30–400  m. Colombia: Cho (Gradstein 2016a). General distribution: Dominica (type), Chocó biogeographic province (Colombia, Panama), eastern Brazil (Bahia). Plagiochila eggersii (sect. Arrectae) is a rare and delicate neotropical species that is recognized by the small plants, 1–2 cm long and 1–2 mm wide, with ventrad and somewhat secund, ovate-obcuneate leaves with highly fragile teeth (10–40 per leaf). The teeth are simple or pinnately branched and 15 cells long on young leaves at the stem apex and generally broken and only 1–3 cells long on older leaves further down the stem. The leaf cells are slightly elongate, 35–55  μm long and 25–33  μm wide in midleaf, with small, radiate trigones, an ill-defined vitta-like area, and a smooth cuticle. The peculiar, pinnately branched teeth of young leaves set this species well apart from other Plagiochilas, which generally have simple teeth. In its ventrad, ovateobcuneate leaves, small plant size, and fragile teeth, P. eggersii approaches P. cuneata but the latter species has trifid leaves with or without additional teeth and unbranched teeth. Fragile teeth occur also in various other Plagiochila species, such as P. bifaria, P. ensiformis, and P. gymnocalycina (var. surinamensis), but the teeth in these species are simple, not branched. 26. Plagiochila ensiformis Taylor Plagiochila azuayensis Steph. Plagiochila jamesonii Taylor

Fig. 29O, P

On moist soil, rock, and tree bases in humid montane forest environments, in Polylepis forests, and in páramo, (1250–)2000–4100 m. Colombia: Boy, Cau, Cun, Hui, Put, Qui, Ris, San, Tol, Val. Ecuador: Az, Ca, Im, Lo, Na, Pi, Tu, ZC. General distribution: northern Andes.

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Plagiochila ensiformis (sect. Hylacoetes) is a characteristic robust northern Andean species that is distinguished by (1) plants robust, 5–9 mm wide, pale yellowish-green to yellowish-brown to dark brown, branches intercalary; (2) leaves ovate-orbicular, 0.8–1.2(−1.3)× longer than wide, ampliate, apex rounded, margins entire or with a few small teeth (1–15) at apex, occasionally teeth more numerous (to 40 per leaf) and extending along the ventral margin and the upper half of the dorsal margin (but absent on the lower half of the dorsal margin), teeth short, maximally 4 cells long, sometimes broken; dorsal leaf base shortly or longly decurrent, ventral leaf base shortly decurrent; (3) leaf cells large, 35–55 μm wide in midleaf, somewhat elongate, with rather large trigones, cuticle smooth; and (4) androecia intercalary or in clusters of male branches at the plant apex. In its ampliate leaves with relatively few, short teeth, P. ensiformis resembles P. guevarai and P. fuscolutea, but P. guevarai differs in the longly and broadly decurrent ventral leaf bases, while P. fuscolutea has smaller leaf cells and always very longly decurrent dorsal leaf bases. In P. ensiformis the dorsal leaf base varies from shortly to longly decurrent. Plagiochila ensiformis may resemble P. canelensis in leaf shape, and these two species may be confused (Heinrichs 2002a). In P. canelensis, however, the teeth on leaf margins are longer, to 8(−11) cells long, and more numerous, to ca. 60 per leaf and frequently occurring on the lower half of the dorsal margin. In P. ensiformis the lower half of the dorsal leaf margin is always entire. 27. Plagiochila exigua (Taylor) Taylor Fig. 29N Plagiochila corniculata auct. (non typus) Plagiochila tridenticulata auct. (non typus) Tylimanthus bispinosus J.B.Jack & Steph. (Acrobolbus bispinosus [J.B.Jack & Steph.] Steph.), syn. nov. Type: Colombia, Santander, Ocaña, 1874, Wallis s.n. (lectotype, designated here, G-00128780!; isolectotype G-00120032!, in mixture) On bark and soil in montane forests and páramo, 2000–4000 m. Colombia: Boy (Gradstein 12713, UPTC), Cun, Ris, Tol. Ecuador: Az (páramo de Cajas, Burghardt 7453, 7472, QCA), Im, Pi (Burghardt 6652, QCA), ZC. General distribution: oceanic-montane regions of tropical America and Africa, extending into warm-temperate oceanic regions of the Northern Hemisphere. Plagiochila exigua (sect. Heteromallae) is a delicate, widespread but uncommon species that is readily distinguished by the rather small, 1–2-mm-wide, green to brown plants with leaves ventrad, short-ovate to ovate-oblong (1–1.5× longer than wide), bifid (or trifid) to 1/5–1/3, apex acute to acuminate, margins entire or with a tooth on the upper ventral margin, bases not or shortly decurrent. The leaf cells are isodiametric-quadrate to slightly elongate, 17–25 μm wide in midleaf, 1–1.5× longer than wide, with small to large trigones and a smooth cuticle. A vitta-like area is lacking. Asexual reproduction by caducous leaves is present or abent. In a collection from Ecuador (laguna de Mojanda, Gradstein et al., QCA), female and male plants were growing mixed and were conspicuously different in size and color, with female plants being rather robust, 2 mm wide and green, while male plants were much more slender, 1–1.2 mm wide and brown.

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Keys and Descriptions

Plagiochila exigua resembles few-toothed phenotypes of P. punctata; however the latter species has a vitta-like area and usually several teeth on the ventral margin. Plagiochila exigua may also be confused with P. bicuspidata; for differences see under P. bicuspidata. Schuster (2001a) provided a detailed description of P. exigua under the name Acrobolbus bispinosus. 28. Plagiochila fastigiata Lindenb. & Gottsche On tree trunk, 1680 m. Colombia: Cau (Gradstein and Feuillet 2016). General distribution: Mexico, Costa Rica (Holz et al. 2001), Colombia. Plagiochila fastigiata (sect. Vagae) is a little-known Central American species that has been recorded once from Cauca. The species is quite similar to P. raddiana but differs in the shorter and broader leaves, 1–1.3× longer than wide (1.5–2.5× longer than wide in P. raddiana), and the ventral leaf margin being strongly undulate from the base to at least the middle of the leaf. In P. raddiana the ventral leaf margin is not undulate or slightly undulate only at the base. In both species branching is terminal, and the ventral leaf bases are strongly ampliate, entire, and longly decurrent. Plagiochila fastigiata is closely related to Plagiochila corrugata (Nees) Mont. & Nees from eastern Brazil, Bolivia, and East Africa, but in the latter species, the ventral leaf base is toothed (Heinrichs et al. 2004b). 29. Plagiochila fuscolutea Taylor Plagiochila gymnostoma J.B.Jack & Steph. Plagiochila jelskii Loitl. Plagiochila potrerillana Steph. Plagiochila scopulosa Steph. Plagiochila triangulifolia Steph. Plagiochila wolframii Inoue

Fig. 30A

On humic soil and bark in upper montane cloud forests, elfin forests, and páramo, often growing in large cushions, 2500–4100  m. Colombia: Ant, Boy, Cal, Cau, Cho, Cun, Mag, Met, Nar, Ris, Tol, Val. Ecuador: Az, Ca, Co, Lo, MS, Na, Pi, Tu, ZC. General distribution: tropical Andes. Plagiochila fuscolutea (sect. Fuscoluteae) is a characteristic and common species of upper montane cloud forests and páramo that is recognized by (1) plants robust, 5–11 mm wide when flattened, branches intercalary; (2) leaves spreading or transverse and laterally appressed, ovate-triangular with a very broad base and narrow apex, 0.7–1.5× longer than wide, shouldered or ampliate, apex rounded to acute, entire or 2–3-toothed, ventral and dorsal margin entire, ventral margin entire or with 1–12 teeth, dorsal base very longly decurrent, concealing them stem surface, ventral leaf base shortly to longly decurrent; (3) cells 20–35 μm wide in midleaf, somewhat elongate, with large, swollen trigones, vitta-like area absent, cuticle smooth or weakly rough; (4) underleaves and paraphyllia absent; and (5) perianth mouth entire or with a few teeth. Plagiochila fuscolutea is close to P. paraphyllina and P. longiramea Steph. from Bolivia, but the latter two species have stem paraphyllia (Heinrichs 2002a). In its ovate-triangular, subentire leaves, P. fuscolutea also resembles P. ensiformis and

Fig. 30 (A) Plagiochila fuscolutea, habit in dorsal view. (B and C) Plagiochila grandicrista. (B) Portion of stem with leaf, dorsal view. (C) Midleaf cell. (D) Plagiochila guevarai, leaf. (E and F) Plagiochila gymnocalycina. (E) Habit, dorsal view. (F) Habit, ventral view. (G–I) Plagiochila laetevirens. (G) Branching pattern. (H) Leaf, 8×. (I) Leaf with propagules, 8×. (J) Plagiochila longispina, leaf. (K and L) Plagiochila loriloba. (K) Habit in dorsal view. (L) Midleaf cell. (M) Plagiochila macrostachya, habit in dorsal view. (N) Plagiochila macra, habit in dorsal view. (O) Plagiochila montagnei, habit in ventral view. (P, Q) Plagiochila ovata. (P) Leaf, 12×. (Q) Midleaf cell. (R and S) Plagiochila papillifolia. (R) Leaf cell with finely papillose surface. (S) Habit, dorsal view. (A–D, M, N redrawn from Heinrichs 2002a; G–I redrawn from Heinrichs et al. 2002b; J, P, Q redrawn from Robinson 1967; K, L redrawn from Groth et al. 2002; R, S redrawn from Heinrichs et al. 2002c; rest from Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009)

298

 

Keys and Descriptions

P. guevarai, but the latter two species have larger leaf cells (35–60  μm wide in midleaf) and dorsal leaf bases less longly decurrent. 30. Plagiochila grandicrista Steph.

Fig. 30B, C

On tree trunks and logs in humid montane forests, 1800–2400  m. Colombia: Cun, Ris (Gradstein 8705, COL), Val. General distribution: northern Andes, Central America. Plagiochila grandicrista (sect. Adianthoideae) is a rare northern Andean species that is distinguished by (1) plants 2.5–5 mm wide, branches intercalary; (2) leaves ovate-triangular, 1.3–1.8× longer than wide, unbordered, strongly ampliate, ventral side of stem invisible, covered by a high crest formed by the expanded ventral leaf bases, apex truncate, margins toothed in the upper half with ca. 5–20 narrowly triangular teeth, bases entire, dorsal leaf base longly decurrent, ventral base shortly decurrent; (3) cells ca. 25–35 μm wide in midleaf, somewhat elongate, with rather large, swollen trigones, vitta-like area lacking, cuticle smooth; and (4) androecia intercalary. Plagiochila grandicrista is close to P. adianthoides but differs from the latter in the unbordered, ovate-triangular leaves with fewer teeth (only in the upper half). Plagiochila grandicrista also approaches P. cristata and P. deflexa, but P. cristata has much more elongate leaves (more than 1.8× longer than wide) with 2 large teeth at apex and a toothed ventral base, while P. deflexa differs in leaves with a distinct vitta-like area, a usually bifid apex, and leaf bases only weakly ampliate (stem without a high ventral crest formed by the leaf bases) (Heinrichs et al. 2002a). Plagiochila grandicrista is also close to P. olivacea Steph. from southeastern Brazil, but the ventral leaf bases in P. olivacea are only weakly ampliate and a ventral crest is lacking (Gradstein 2015c). 31. Plagiochila guevarai H.Rob. 

Fig. 30D

On bark and humic soil in páramo, 3300–3800 m. Colombia: Cun, Met, Ris, Tol. Ecuador: Ca, Co, Na. General distribution: northern Andes (Colombia, Ecuador). Plagiochila guevarai (sect. Hylacoetes) is a characteristic northern Andean páramo species that is recognized by (1) plants robust, 5–9  mm wide, branches intercalary; (2) leaves ovate-triangular, narrowed to the apex from a very broad base, 0.8–1.2× longer than wide, strongly ampliate, with a high ventral crest formed by the expanded ventral leaf bases, margins entire or with a few teeth (1–6) at apex, dorsal and ventral leaf bases entire, longly and broadly decurrent; (3) leaf cells 35–60 μm wide in midleaf, somewhat elongate, with rather large, swollen trigones, vitta-like area absent, cuticle smooth; and (4) androecia intercalary. In its subentire leaves, P. guevarai resembles P. ensiformis and P. fuscolutea; for differences see under P. ensiformis. 32. Plagiochila gymnocalycina (Lehm. & Lindenb.) Mont. & Nees Plagiochila demissa Gottsche (?) (type not found)

Fig. 30E, F

Plagiochilaceae

299

Plagiochila gymnocalycina var. surinamensis (Molk.) Heinrichs & D.S.Rycroft, syn. nov. Type: Suriname, without collector, ex herb. Sande Lacoste (L!) Plagiochila grateloupii Mont. Plagiochila polopolensis Herzog Plagiochila rutilans Lindenb. var. angustifolia Herzog Plagiochila wolfii Inoue, syn. nov. Type: Colombia, Amazonas, Río Caqueta, Isla Mocorroy, Wolf & Hofstede 1482 (holotype COL; isotype U!) On bark in lowland, submontane, and lower montane rainforest, 200–1500(−2000) m. Colombia: Ama, Caq, Cho, Nar, San, Vau (Schultes & Cabrera 11933, CINC, PC). Ecuador: Az, EO (Reserva Buenaventura, Benitez et al. 1318, HUTPL), MS, Or (Estación Tiputini, Oct 2018, Gradstein, Deleg & Bader s.n., HUTPL), Pa. General distribution: widespread in tropical America. Plagiochila gymnocalycina (sect. Heteromallae) is a common neotropical species of rather low elevations (mostly below 1500  m) that is distinguished by (1) plants 4–5 mm wide, branching intercalary; (2) leaves distant to subimbricate, ventrad, asymmetrically and rather narrowly oblong-rectangular, 2.1–3.5× longer than wide, widest in the middle, not ampliate, dorsal margin ± straight, ventral margin curved, with (2–)6–18 teeth in the upper half, apex occasionally with 2 larger teeth, bases short-decurrent, ventral leaf base slightly recurved; (3) cells 18–30 μm wide in midleaf, isodiametrical to somewhat elongate, with conspicuous radiate/trabeculate trigones, vitta-like area lacking, cuticle smooth; (4) perianth base “naked,” inner bracts inserted at some distance from the perianth and not enveloping the lower half of the perianth; and (5) vegetative reproduction by caducous leaves, caducous teeth, and fragmentation of the leaf lamina. Plagiochila gymnocalycina is close to P. simplex; for differences see under P. simplex. Plagiochila gymnocalycina has sometimes been considered a synonym of P. rutilans, but the latter species differs in the horizontally spreading leaves (not ventrad), perianths enveloped by bracts, and plants with a characteristic peppermint smell when fresh. Plagiochila gymnocalycina var. surinamensis (= P. wolfii) is characterized by fragile teeth; in other respects the plant is identical to P. gymnocalycina, also genetically and chemically (Heinrichs et al. 2006). Caducous teeth occur in several Plagiochila species (e.g., P. bifaria, P. ensiformis) and are without taxonomic relevance (except in P. eggersii, which has specialized, pinnate caducous teeth). 33. Plagiochila heteromalla (Lehm. & Lindenb.) Lindenb. Plagiochilion heteromallum (Lehm. & Lindenb.) Hässel Plagiochila oresitropha Spruce Plagiochila ovifolia Steph. Plagiochila paludosa Steph., syn. nov. Type: Ecuador, Morona Santiago, Río Salado, Allioni s.n., Bryotheca Levier 6358 (lectotype, designated by J.  Heinrichs in sched., G-00121456!; four additional paralectotypes present in G) Plagiochila sylvicultrix Spruce

300

 

Keys and Descriptions

On bark in humid montane forests and páramo, 1200–3900 m. Colombia: Cal, Mag. Ecuador: Lo (Parque Nac. Podocarpus, Burghardt 7158, PC, QCA), MS, Tu, ZC. General distribution: tropical Andes, Costa Rica. Plagiochila heteromalla (sect. Heteromallae) is a little-known Andean species that is readily recognized by the (sub)opposite and usually strongly secund leaves (opposite leaf bases ± touching each other, especially the ventral leaf bases, but not connate) with thick-walled cells along the leaf margin which may form a distinct border and leaf bases with an ill-defined vitta-like area. The plants are mediumsized to large, 4–7  mm wide, with intercalary and sometimes a few terminal branches, and the leaves are short-ovate, 1–1.3× longer than wide, shouldered to ampliate, toothed all around except on the lower half of the dorsal margin, with 16–44 short, narrowly triangular to linear teeth and with the bases rather shortly and narrowly decurrent. The cells are 25–35 μm wide in midleaf, slightly elongate, with conspicuous, simple to radiate trigones, a smooth cuticle, and homogeneous oil bodies. The perianth is elongate-cylindrical with a narrow stalk at the base, and the perianth base is covered by bracts or not. Vegetative reproduction has not been observed. Plagiochila heteromalla may be confused with P. macrostachya, which also has opposite leaves, but the latter species has more elongate leaves (1.2–1.6× longer than wide) with larger cells and without vitta-like area and leaf cells not thickwalled along the margin. Plagiochila heteromalla also approaches P. trichostoma, but the leaves in the latter species are not (or almost) opposite and less strongly secund, and a leaf border is usually absent. Spruce (1885) described two varieties in P. heteromalla from Ecuador, P. heteromalla var. latifolia Spruce and P. heteromalla var. densifolia Spruce, both from Mt. Tungurahua and occurring together with the typical variety. The status of these varieties needs study. 34. Plagiochila husnotii Steph. On bark in everwet lowland rainforest along the Pacific coast, 30 m. Colombia: Cho (Heinrichs 2002a). General distribution: West Indies, coastal areas of northern South America. Plagiochila husnotii (sect. Hylacoetes) is a Caribbean species that is distinguished by (1) plants large, 5–9 mm wide, branching intercalary; (2) leaves elongate-triangular to narrowly ovate-oblong, ca. 1.8–2.1× longer than wide, weakly ampliate, margins with 20–60 cells long, triangular to linear teeth, dorsal and ventral bases strongly toothed and shortly decurrent; (3) cells 30–50  μm wide in midleaf, slightly elongate, with small radiate trigones, no vitta-like area, and a smooth cuticle; and (4) androecia intercalary or in clusters of male branches at the stem apex. Plagiochila husnotii is very similar to P. dominicensis, but the ventral leaf bases in the latter species are entire (strongly toothed in P. husnotii). 35. Plagiochila laetevirens Lindenb. Plagiochila bancroftii Steph. Plagiochila binominis Gottsche Plagiochila boissieri Steph.

Fig. 30G–I

Plagiochilaceae

301

Plagiochila cipaconensis Steph. Plagiochila contigua Gottsche Plagiochila flabellifrons Spruce Plagiochila impluviata Spruce Plagiochila micropteryx Gottsche Plagiochila pachoensis Steph. Plagiochila pastasensis Steph., syn. nov. Type: Ecuador (“Brazil”), Río Pastaza, Spruce s.n. (G-00067767!, c. gyn.) Plagiochila pseudopatula Herzog Plagiochila tocarema Gottsche On bark and rock in montane rainforests, usually in rather open places, also on isolated trees in meadows, (300–)600–3000  m. Colombia: Boy, Cun, Ris, Tol. Ecuador: EO (300 m), MS, Pa (type), Pi, Tu, ZC. General distribution: widespread in tropical America. Plagiochila laetevirens (sect. Vagae) is a common and variable neotropical species that is recognized by (1) plants soft-textured, medium-sized, 3–4  mm wide, dichotomous to irregularly pinnate, branching predominantly terminal, toward the shoot base sometimes with a few intercalary branches; (2) leaves horizontally spreading to slightly ventrad, short-ovate to subrectangular to ovate-triangular, 1–1.5× longer than wide, shouldered to weakly ampliate, unbordered, apex truncate, apical and ventral leaf margin with few to many, 5–25 coarse, triangular teeth, dorsal base shortly to longly decurrent, ventral base recurved, shortly decurrent; (3) cells ca. 20–30 μm wide in midleaf, subisodiametrical, with conspicuous, triangular trigones, cuticle smooth; (4) underleaves completely lacking; and (5) vegetative reproduction by numerous small plantlets (propagula) on ventral leaf surfaces. Plagiochila laetevirens is related to P. distinctifolia and P. patula, but the latter two species have more elongate leaves (more than 1.5× longer than wide). Moreover, P. distinctifolia is more delicate and has bordered leaves without a high shoulder. Plagiochila laetevirens may also be confused with P. diversifolia, but the latter species is more robust and has darker brown stems, leaves without high shoulder, larger leaf cells, brown oil bodies, and usually small underleaves. 36. Plagiochila longispina Lindenb. & Gottsche Fig. 30J Plagiochila matanga Steph., syn. nov. Type: Ecuador, Azuay, páramo de Matanga, Allioni 679 (holotype G-00064204!) On humid rock, soil, and trunk bases in very humid upper montane forest and páramo, 2700–4050 m. Colombia: Ara, Boy, Cal, Cun, Met, Ris, Tol, Val. Ecuador: Az/MS, Ca, Ch, Co, Na, Pi, Tu. General distribution: tropical Andes, Costa Rica, Azores. Plagiochila longispina (sect. Glaucescentes) is a very distinct high-montane species that is readily distinguished by the bluish stems (in upper part of shoot) and the leaves with numerous long-linear teeth all around the margins. The plants are rather robust, 5–7 mm wide, flaccid, stems in the upper part often bluish, irregularly pinnate to ± dichotomous, with terminal and intercalary branching. The leaves are horizontally spreading, ovate to ovate-oblong, not ampliate, with usually rather longly

302

 

Keys and Descriptions

decurrent bases and the leaf cells subisodiametrical, 25–40  μm wide in midleaf, with rather small, triangular trigones, a smooth cuticle, and brown oil bodies. The underleaves are conspicuous, to 1 mm long, ciliate. Vegetative reproduction has not been observed. Plagiochila longispina is closely related to P. diversifolia, but the stems in the latter species are never bluish and the teeth are shorter and mainly restricted to the ventral and apical leaf margins (Heinrichs et al. 2000). 37. Plagiochila loriloba Carl Fig. 30K, L Plagiochila cuneata Lindenb. & Gottsche var. loriloba (Carl) Herzog Habitat in Colombia unknown; in Costa Rica found on branches of shrubs in the understory of upper montane forest at 2900–3300 m (Holz et al. 2001). Colombia: Cau (Puracé; type). General distribution: Colombia, Costa Rica. Plagiochla loriloba (sect. Heteromallae) is a little-known neotropical species that has been recorded once from Colombia (type). The species is characterized by (1) plants 1.5–2.5 mm wide, with a peppermint smell when fresh, branching intercalary; (2) leaves subimbricate, obliquely to widely spreading, elongate-obcuneate, deeply 3–4-lobed with diverging lobes, usually strongly fragmenting, upper part of leaves often broken, margins entire, bases shortly decurrent; (3) cells 25–43  μm wide in midleaf, 1–1.5× longer than wide, with distinct trigones, cuticle smooth; (4) perianth base free, not covered by bracts; and (5) vegetative reproduction by leaf fragmentation. The main characters of Plagiochila loriloba are the deeply 3–4-lobed, strongly fragmenting leaves, and the peppermint smell of the fresh plants. The species is closely related to P. cuneata (Groth et al. 2002); for differences with P. cuneata, see under the latter species. 38. Plagiochila macra Taylor  Fig. 30N Type: Ecuador, Pichinchina “on mosses,” 22 Dec 1846, Jameson s.n. (lectotype, designated here, FH-Taylor!, c. spor.; isolectotype G-00280494) Plagiochila abscedens Gottsche Plagiochila heterophylla Lehm., syn. nov. Type: Costa Rica, Ǿrsted s.n. (lectotype, designated by Heinrichs et al. (2002a), S-B58739!) Plagiochila tunguraguensis Spruce On bark, rock, and soil in humid montane forest areas and scrub, (400–) 1150–3400  m. Colombia: Boy (Gradstein 10002, COL), Mag, Vau (Schultes & Cabrera 17495, cf.). Ecuador: Az, Ca, Lo, Na, Pa, Pi, Su, Tu, ZC. General distribution: Mexico to Ecuador, Dominican Republic, western Europe. Plagiochila macra (sect. Fuscoluteae) is a medium-sized neotropical-western European species (2–6 mm wide) that is recognized by (1) branching predominantly intercalary (rarely terminal); (2) leaves wide-spreading and somewhat ventrad, sometimes secund, ovate-oblong to ovate-triangular (1.3–2× longer than wide), unbordered, not or slightly ampliate, shortly to moderately decurrent, with 2–15(−20) rather short teeth along the ventral margin (mostly in the upper half) and at the apex, often with 2 larger teeth at the apex; (3) leaf cells 20–35 μm wide in

Plagiochilaceae

303

midleaf, somewhat elongate, with small to rather large, elongate trigones, without or with a very short vitta-like area and a ± smooth cuticle; (4) female bracts covering the base of the perianth; and (5) vegetative reproduction not observed (Heinrichs 2002a as P. heterophylla). Plagiochila macra somewhat resembles P. simplex, but the latter species has caducous leaves and “naked” perianths, with the bracts positioned at some distance below the perianth and covering the perianth base. The type of P. macra, which consists of a single fertile plant, represents a phenotype with secund leaves and rather small trigones. 39. Plagiochila macrifolia Taylor Plagiochila macrifolia var. taylorii Spruce On bark in montane forest. Ecuador: Az, Ch (type), Pi. General distribution: only known from Ecuador. Plagiochila macrifolia (sect. Vagae) is a poorly known species that is only known with certainty from Ecuador (Spruce 1885). A record from Colombia (Wolf 1993) belongs to P. alternans (Gradstein 2016a). Plagiochila macrifolia is distinguished by the rather small plants (3–4  mm wide) with terminal-dichotomous branching, leaves narrowly oblong (2–2.5× longer than wide), distant, ventrad, not ampliate, unbordered, with 2–4 small teeth at the apex, margins entire and bases shortly decurrent. The leaf cells are isodiametric to elongate, 30–50 × 25–30 μm in midleaf, 1–1.7× longer than wide, with elongate, subconfluent trigones, leaf margin cells more narrowly elongate but not more strongly thickened than inner cells, and cuticle smooth. Underleaves are fully lacking. According to Spruce (1885), the perianth is “naked” and not covered by bracts. Plagiochila macrifolia closely resembles P. distinctifolia, but the latter species usually has a leaf border of thick-walled cells and a toothed ventral leaf margin. Plagiochila macrifolia may also be confused with P. aerea, but the apical teeth in P. aerea are much larger, the ventral leaf margin is ± toothed, the leaf cells in P. aerea are much more elongate, and branching is mostly intercalary (occasionally terminal). The status of Plagiochila macrifolia var. angustifolia Spruce (1885) needs study. 40. Plagiochila macrostachya Lindenb. Fig. 30M Plagiochila contingens Gottsche Plagiochila flaccida Lindenb. Plagiochila jamesonii Taylor var. secundifolia Spruce Plagiochila leptophylla Spruce Plagiochila meridana Gottsche, syn. nov. Type: Venezuela, Mérida, Moritz s.n., ex herb. Musei Berol. (lectotype, designated here, G!; isolectotypes (3) G!) Plagiochila meridana var. funkii Gottsche, syn. nov. Lectotype (designated here): Venezuela, Caracas, Funk & Schlim 340, ex herb. Jack (G s.n.!). Syntype: ibid., Funk & Schlim 343 (G-025273!) Plagiochila sacramenti Steph. Plagiochila zygophylla Spruce

304

 

Keys and Descriptions

On bark and rock in humid lowland and montane rainforests and in the páramo, from sea level to 4000 m. Colombia: Ant, Boy, Cas, Cho, Cun, Qui, Ris. Ecuador: Ca, Ch, MS, Na, Pa, Tu, ZC.  General distribution: widespread in tropical America. Plagiochila macrostachya (sect. Hyalocoetes) is a common neotropical species that is distinguished by (1) plants large, 5–10 mm wide, branching intercalary; (2) leaves horizontally spreading or secund, (sub)opposite, ovate-oblong or ovate-triangular, with a broad apex, 1.2–1.6× longer than wide, shouldered or ampliate, unbordered, with ca. 10–40 triangular to linear teeth along the ventral, apical, and upper dorsal margin, bases ± shortly decurrent; (3) cells 25–50  μm wide in midleaf, slightly elongate, with small to large trigones, without or with a weakly developed vitta-like area, cuticle smooth; (4) androecia intercalary or in clusters of male branches at the plant apex, male bracts dorsally not overlapping; and (5) vegetative reproduction sometimes by caducous leaves. In its subopposite leaves, P. macrostachya resembles P. heteromalla, but the leaves in the latter species are about as long as wide and never caducous, the leaf margin is usually bordered by thick-walled cells, and the leaf cells are smaller. 41. Plagiochila montagnei Nees Fig. 30O Plagiochila bogotensis Gottsche Plagiochila florida Spruce Plagiochila hypnoides Lindenb. Plagiochila nova Steph., syn. nov. Type: Ecuador, Morona Santiago, Gualaquiza, Bomboiza, “frequens ad terram et ad truncos putrescentes in Granadillas,” 2500 m, 15 Nov 1910, Allioni 709 (lectotype, designated here, G-000061534! Icon. Ined. 11214). A syntype (Allioni 691, G-00113038!) belongs to P. macrostachya. On bark and rock in lowland rainforests, plantations, and isolated trees, from sea level to 500 m, very common in Amazonia. Colombia: Ama, Ant, Caq, Cun, Mag, Met, Put, Vau. Ecuador: Gu, EO, Es, LR, Na, Or, Pa, Su, ZC (?). General distribution: tropical and subtropical America. Plagiochila montagnei (sect. Vagae) is a common neotropical lowland rainforest species that is recognized by (1) plants 3–5  mm wide, dichotomous by terminal branching; (2) leaves widely and horizontally spreading, ovate-lingulate, 1.5–2.5× longer than wide, strongly ampliate (dilated ventral leaf bases forming a crest), ventral leaf margin plane or undulate, apex and basal part of ventral leaf margin with 10–30 teeth, ventral base shortly or longly decurrent, with linear to narrowly triangular teeth (in part more than 1 cell wide), dorsal base longly and broadly decurrent; (3) cells 20–30 μm wide in midleaf, isodiametric to elongate, trigones conspicuous but not swollen, cuticle smooth; (4) underleaves rudimentary, to maximally 0.5 mm long; and (5) vegetative reproduction by small plantlets (cladia) on ventral leaf surfaces. Plagiochila montagnei is close to P. disticha and P. raddiana; for differences see under the latter species.

Plagiochilaceae

42. Plagiochila ovata Lindenb. & Gottsche Plagiochila cordilliera Steph. Plagiochila minutidens Steph. Plagiochila multispina Steph. Plagiochila soratensis Steph.

305

Fig. 30P, Q

On humic soil, rock, or bark in upper montane and subalpine forest and in scrubby páramo, ca. 3000–4200 m. Colombia: Boy, Cas, Cun, Mag, Met. Ecuador: Az, Co, Im, MS, Na, Pi, ZC. General distribution: tropical Andes, Costa Rica. Plagiochila ovata (sect. Denticulatae) is a characteristic high-Andean species that is readily distinguished by the ovate-orbicular leaves with numerous (ca. 100–250) short, linear teeth, 1–3(−4) cells long, all around the margin, and a leaf border of thick-walled cells. None of the other neotropical species of Plagiochia has so many teeth on leaf margins. The plants are robust, 4–8 mm wide; branching is intercalary; the leaves are obliquely spreading and ventrad, with a longly decurrent dorsal base and a shortly decurrent ventral base; and the leaf cells are isodiametric to elongate, 25–40 μm wide, with small to large trigones and a smooth cuticle. Plagiochila ovata is related to P. alternans, but the latter species occurs at lower elevation; the leaves in P. alternans are horizontally spreading (not ventrad) and distinctly longer than wide with a broadly rounded apex, toothed only in the upper half (less than 100 teeth); and the leaf cells are thin-walled with very small trigones (Müller et al. 1999). Plagiochila ovata is also close to P. priceana, and the two species grow side by side in the type locality of P. priceana. The latter species is readily distinguished from P. ovata in the different leaf shape (ovate-triangular, with a narrower apex and broadly decurrent dorsal base) and leaves with much fewer teeth, 35–65 per leaf. Moreover, P. priceana lacks a leaf border of thick-walled cells. 43. Plagiochila pachyloma Taylor Plagiochila cuervina Gottsche Plagiochila homochroma Spruce Plagiochila wallisiana Steph. On bark in montane forest and páramo, (1000–)2000–4100 m. Colombia: Boy, Mag, Nar, Qui, Ris, San, Tol. Ecuador: Az, Ca, Co, Im, Lo, Na, Pi, Su, Tu, ZC. General distribution: northern Andes, Costa Rica. Plagiochila pachyloma (sect. Arrectae) is a characteristic, medium-sized highAndean species that is readily recognized by the rather dark green plants with ventrad, transverse to rather widely spreading, ovate-triangular to ovate-oblong leaves with an ampliate base, a 1–2-cell-wide border of thicker-walled cells, and numerous long, linear teeth (up to 50) on the ventral leaf margin and apex and sometimes also on the dorsal margin. The teeth consist of a 1–6-cell-long row of narrowly elongate cells with a 1–2-cell-wide base; the border and teeth are usually yellowish to deep brown in color, occasionally colorless; and the border cells are frequently somewhat larger than the inner leaf cells. In plants with a brownish border, the teeth tend to be pigmented as well. The leaf cells are subisodiametrical and small, 15–25  μm in diameter in midleaf, with variable, small to large trigones and a more or less

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Keys and Descriptions

striate-papillose cuticle. A vitta-like area of larger and thinner-walled cells is present at the leaf base. Branching is intercalary, and vegetative reproduction is by leaf fragmentation and caducous teeth. Plagiochila pachyloma approaches P. bifaria, but the leaves in the latter species lack a conspicuous border of thicker-walled cells and the teeth are much more variable in shape, not strictly linear, and with a wider, 1–4(−6)-cell-wide base. A phenotype of P. pachyloma with an uncolored leaf border and a toothed dorsal leaf margin has been described as P. homochroma Spruce. 44. Plagiochila papillifolia Steph. Plagiochila caespiticia Herzog, nom. inval. Plagiochila deciduifolia Steph. Plagiochila verruculosa R.M.Schust.

Fig. 30R, S

On bark and rock in upper montane cloud forest up to the páramo, 2150–3600 m. Colombia: Ant, Boy/Cas (Gradstein 12702, PC), Cau, Cun, Qui (páramo El Campanario), Ris, San. Ecuador: Az, Na, Tu, ZC. General distribution: tropical America, Azores. Plagiochila papillifolia (sect. Arrectae) is a widespread species that has probably been overlooked due to its small size. Characteristics are (1) plants 0.5–1.5  mm wide and 1–2.5 cm long, branching intercalary; (2) leaves obliquely spreading, ventrad, ovate to ovate-oblong, usually less than 1 mm long, 1.0–1.7× longer than wide, apex truncate, with 2 long, spaced teeth, ventral margin with 1–10 shorter teeth, the teeth usually triangular, sometimes linear (see below), cells of the uniseriate part of the teeth elongate, leaf bases slightly decurrent; (3) leaf cells slightly longer than wide, 18–25 μm wide in midleaf, trigones well developed, triangular to nodulose, cuticle densely papillose, vitta-like area absent; and (4) vegetative reproduction by caducous leaves (Heinrichs et al. 2002b). Plagiochila papillifolia varies considerably in leaf size and dentition (Heinrichs et al. 2002b). According to the latter authors, the marginal teeth in this species are usually triangular in shape, but plants from mossy elfin forest in the páramo of Yacuambi, southern Ecuador (Benitez & Gradstein s.n., HUTPL), had leaves with 4(−5) linear teeth made up of a row of 2–3 narrowly elongate cells. By its linear teeth, these plants looked like a dwarf form of P. pachyloma. Possibly, these plants represent an undescribed species; this needs further study. Plagiochila papillifolia may be confused with P. tenuis and P. stricta, which also have a papillose cuticle. However, P. tenuis has more elongate leaves (at least 2× longer than wide), whereas P. stricta is a much more robust plant (>2 mm wide) with leaf bases decurrent and with a short, vitta-like area. 45. Plagiochila paraphyllina Herzog On bark and soil in upper montane forest and páramo, 2300–3700 m. Colombia: Cun, Mag. Ecuador: Ca (Heinrichs 2002a). General distribution: northern Andes (Colombia, Ecuador). Plagiochila paraphyllina (sect. Hylacoetes) is a rare northern Andean species that resembles P. fuscolutea but differs from the latter in toothed dorsal leaf bases

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and the presence of small, toothed paraphyllia on the dorsal surface of the stem, near the leaf bases or scattered over the stem surface. Plagiochila paraphyllina is very similar to P. longiramea Steph. from Bolivia, but in the latter species, the dorsal leaf bases and the paraphyllia are entire. 46. Plagiochila patriciae Heinrichs & H.Anton On bark in upper montane forest near the forest line, 3500 m. Colombia: Ant (Heinrichs 2002a). General distribution: Colombia, Costa Rica. Plagiochila patriciae (sect. Hylacoetes) is a rare northern Andean-Costa Rican species that is distinguished by (1) plants large, 5–7 mm wide, branches intercalary; (2) leaves ovate-triangular, 1–1.4× longer than wide, strongly ampliate-cristate, with ca. 10–30 teeth on the ventral and apical margins and sometimes at the dorsal base, bases longly decurrent, the decurrent part of the ventral base conspicuously toothed; (3) leaf cells large, ca. 30–60 μm wide in midleaf, somewhat elongate, with rather large, swollen trigones, cuticle smooth; and (4) androecia in clusters of male branches at the stem apex. In its rather short and broad leaves with longly decurrent bases, P. patriciae resembles P. guevarai which occurs in the same habitat. However, the leaves in P. guevarai have much fewer teeth (1–6), and the decurrent part of the ventral leaf base is entire. 47. Plagiochila patula (Sw.) Lindenb. Fig. 31A, B Plagiochila barutana Gottsche Plagiochila beskeana Steph. Plagiochila dichotoma sensu Spruce (non typus) Plagiochila discreta Gottsche, syn. nov. Type: Colombia, Cundinamarca, Bogotá, Aserradero, Lindig 1729c (lectotype, designated here, C-00282666!) Plagiochila dubia Lindenb. & Gottsche Plagiochila fallax Lindenb. & Hampe Plagiochila interjecta Gottsche, syn. nov. Type: Colombia, Cundinamarca, Bogotá, near Tocarema, Lindig 1705g (lectotype, designated here, G-0781918!, c. gyn.) Plagiochila intricata Steph. Plagiochila martiana (Nees) Lindenb. Plagiochila montana Spruce Plagiochila rigidula Hampe, syn. nov. Type: Venezuela, Tovar, Moritz 92 (91?) (isotypes G-00280620!, G-00280619!, G-00282601!) Plagiochila truncatella Gottsche Plagiochila xanthochroma Spruce On bark and soil in lowland and montane rainforest areas, 200–2800  m. Colombia: Cun,? Put (Robinson 1967 as P. aspleniformis), Ris, Val, Vau (Schultes & Cabrera 15276, CINC, PC). Ecuador: Lo, MS, Or, ZC.  General distribution: widespread in tropical and subtropical America. Plagiochila patula (sect. Vagae) is a common and variable neotropical species that is recognized by (1) plants rather robust, 4–6  mm wide, branching usually

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Fig. 31 (A and B) Plagiochila patula. (A) Habit, dorsal view. (B) Branching pattern. (C) Plagiochila raddiana, leaves. (D) Plagiochila punctata, habit in dorsal view, showing denuded branches. (E) Plagiochila rutilans, habit in dorsal view. (F and G) Plagiochila simplex. (F) Habit, ventral view. (G) Habit, dorsal view. (H) Plagiochila subplana, habit in ventral view. (I) Plagiochila superba, leaf. (J and K) Plagiochila tabinensis. (J) Leaf. (K) Cells in lower half of leaf. (L and M) Plagiochila tenuis. (L) Habit in dorsal view. (M) Midleaf cells, showing finely papillose surface. (N) Plagiochila trichostoma, leaf. (O) Plagiochila vincentina, leaf. (A–C redrawn from Heinrichs and Renker 2001; D redrawn from Heinrichs et al. 2005; E–H from Gradstein and Ilkiu-Borges 2009; I, O redrawn from Heinrichs 2002a; J, K redrawn from Schuster 2001a; L, M redrawn from Inoue 1977; N redrawn from Heinrichs et al. 2002a)

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terminal and dichotomous, sometimes purely intercalary (e.g., Orellana, Whitefoord et al. 64, QCA); (2) leaves widely and horizontally spreading, ovate-oblong to lingulate, 1.5–2.5× longer than wide, with 0–15 teeth (mostly near apex), the teeth sometimes very small, apex broadly rounded to truncate, ventral base entire, shouldered, not or only slightly ampliate, usually longly decurrent (occasionally shortly decurrent); (3) cells rather small, 20–25(−30) μm wide in midleaf, isodiametric to elongate, trigones absent or conspicuous but never swollen, cuticle smooth; (4) underleaves absent or rudimentary; and (5) vegetative reproduction by small plantlets (propagula) produced in great abundance on ventral leaf surfaces. The ovate-oblong leaves with a broadly rounded to truncate apex, few teeth, longly decurrent ventral base, and a high shoulder are the main characters of P. patula. The species may be confused with P. crispabilis, a Brazilian species not yet known from Colombia and Ecuador, but in the latter species, the ventral leaf base is very shortly decurrent and a ventral shoulder is lacking. Phenotypes of P. patula with a slightly ampliate ventral base approach P. raddiana, but in the latter species, the ventral leaf bases are always conspicuously ampliate and are usually concealing the ventral surface of the stem. A common phenotype from inundated Amazonian rainforest with a broadly rounded leaf apex and a few very small teeth was identified by Spruce as P. dichotoma (not typus) and was probably recorded from Colombia as P. aspleniformis R.M.Schust. (material not seen) (Robinson 1967). These Amazonian populations are also very similar to P. lingua Steph., a species described from southeastern Brazil. The relationship between P. patula and P. lingua needs further study. The record of P. lingua from Colombia (Gradstein et al. 2016) was erroneous; the plants belong to P. raddiana. Plagiochila discreta, P. martiana, and P. rigidula are phenotypes of P. patula with rather shortly decurrent ventral leaf bases. Plagiochila interjecta was considered a synonym of P. raddiana by Inoue (1989a), but the weakly ampliate leaf bases place P. interjecta in P. patula. 48. Plagiochila pautaphila Gradst. & León-Yánez On trunk bases and large branches of Polylepis pauta in remnant P. pauta forest, 3600–4000 m. Ecuador: Na (páramo de Oyacachi, remnant P. pauta forest, 18 Sept 2019, leg. Gradstein, QCA), Pi (Gradstein and León-Yánez 2018). General distribution: only known from Polylepis pauta forests of Central Ecuador. Plagiochila pautaphila (sect. Hylacoetes) is a robust Ecuadorian endemic that occurs in Polylepis pauta woodlands in the páramos of Papallacta and Oyacachi east of Quito. The species is distinguished by (1) plants robust, 5–15 cm long, 6–12 mm wide, bright yellowish-green, very sparsely branched, branching lateral-intercalary; (2) leaves horizontally spreading to slightly ventrad, ovate-triangular, apex rounded, ventral base strongly ampliate forming a high crest, shortly decurrent, dorsal base very longly and rather broadly decurrent; (3) dorsal base mammillose and finely toothed with 5–15 short-linear teeth, rest of dorsal margin entire or with scattered teeth, apical and ventral margins entire or with a few (1–6(−10)) short-linear teeth, the teeth 1–2(−4) cells long; and (4) leaf cells very large, 60–100 × 50–75 μm in

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Keys and Descriptions

midleaf, rather thin-walled, trigones small, intermediate thickenings absent, vittalike area lacking, cuticle smooth. Plagiochila pautaphila may occur together with P. ensiformis and resembles the latter in size, color, and ± lack of teeth on ventral leaf margins but differs in the ovate-triangular leaves and the toothed dorsal leaf base. The leaves of P. ensiformis are broader, ovate-orbicular, and the dorsal leaf bases are entire. 49. Plagiochila priceana Gradst. & A.R.Benitez On the floor of Polylepis reticulata forests and on shaded soil below shrubs in páramo, 3700–4200  m. Ecuador: Az (Gradstein and Benitez 2017), Co, Na, Pi (Burghardt 2019). General distribution: only known from Ecuador. Plagiochila priceana (sect. Denticulatae) is a distinct Ecuadorian endemic that is recognized by (1) plants robust, 10–13 cm long, ca. 5 mm wide, bright yellowishgreen, very sparsely branched, branching lateral-intercalary; (2) leaves ventrad, broadly ovate-triangular, unbordered, apex rounded, ventral base ampliate but not forming a crest, shortly decurrent, dorsal base very longly and rather broadly decurrent; (3) leaves finely toothed along the ventral margin, the apex and the upper part of the dorsal margin, with 30–65 linear teeth, the teeth 1–4(−5) cells long, consisting of strongly elongate cells, lower half of dorsal margin and ventral base entire; and (4) leaf cells ca. 30–40 μm in midleaf, with small to large, simple to radiate trigones, intermediate thickenings ± absent, border lacking, base with a broad, illdefined vitta-like area, cuticle smooth. Plagiochila priceana approaches P. ovata but differs in having ovate-triangular leaves with much fewer teeth (35–65) and without border of thick-walled cells. 50. Plagiochila punctata (Taylor) Taylor Plagiochila chinantlana Gottsche Plagiochila choachina Gottsche Plagiochila filicaulis Spruce Plagiochila paramicola Herzog Plagiochila patzschkei Steph. Plagiochila stolonifera Lindenb. & Gottsche Plagiochila subrara Herzog

Fig. 31D

On soil, bark, and rock in exposed montane environments and páramo, 1600–4300 m. Colombia: Boy, Cun, Mag, Met, Qui, Ris, Tol. Ecuador: Az, Ch, Co, Im, Na, Tu, ZC. General distribution: tropical America, East Africa, western Europe. Plagiochila punctata (sect. Arrectae) is a very common Afro-AmericanEuropean species that is readily distinguished by the rather small, green to brownish-green plants (1–3 mm wide) with caducous leaves produced mostly on branches, which are often completely denuded except at the tip. Sometimes the whole plant consists of denuded, leafless shoots. The leaves are shortly (ob)ovate to oblong, obliquely to widely spreading or occasionally suberect, distinctly ventrad, not ampliate, little-decurrent, with a toothed apex and ventral margin (with 1–14 small teeth, apical teeth often larger). The leaf cells are isodiametrical to somewhat

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elongate, ca. 20–30  μm wide in midleaf, usually with large, bulging trigones, a short, ill-defined vitta-like area, and a ± smooth cuticle. Plagiochila punctata is related to P. stricta and P. bifaria, but the leaves in the latter two species are not caducous and more distinctly decurrent, especially dorsally. Moreover, the cuticle in P. stricta is distincty papillose (Heinrichs et al. 2005). Small phenotypes of P. punctata from páramo, with subentire ventral leaf margins, were described as P. paramicola and P. subrara. These plants resemble P. cleefii, but in the latter species, the dorsal leaf bases are longly decurrent. 51. Plagiochila raddiana Lindenb. Plagiochila aliena Gottsche Plagiochila anderssonii Ångstr. Plagiochila connivens Gottsche Plagiochila crispatodecurrens Herzog Plagiochila funckiana Steph. Plagiochila gualaquizana Steph. Plagiochila guilleminiana Mont. Plagiochila ludoviciana Sull. Plagiochila perbella Herzog Plagiochila plicata Lindenb. & Gottsche Plagiochila rhizophila Spruce Plagiochila schlimiana Steph. Plagiochila silvatica Gottsche Plagiochila subcristata Gottsche Plagiochila subsimplex Steph. Plagiochila undulifolia Herzog

Fig. 31C

Common on bark and shaded rock (occasionally on soil and leaves) in lowland and montane rainforests, from sea level to 3800 m. Colombia: Ant, Boy, Cas, Caq, Ces, Cho, Cun, Mag, Nar, Put, Qui, Ris, Tol. Ecuador: Az, EO, Ga, Im, MS, Pa, Su, Tu, ZC. General distribution: widespread in tropical and subtropical America. Plagiochila raddiana (sect. Vagae) is a very common and variable neotropical species that has been described under many different names. The species is distinguished by (1) plants 4–6 mm wide, ± dichotomous by terminal branching, a few intercalary branches sometimes present in the lower parts; (2) leaves widely and horizontally spreading, ovate-oblong to ovate-lingulate with a broad and longly decurrent dorsal base, 1.5–2.5× longer than wide, ampliate, apex broadly subtruncate, apical and ventral margin with ca. (0–)3–25 teeth, ventral base entire or, occasionally, with 1–2 small teeth, longly decurrent; (3) cells 20–30 μm wide in midleaf, isodiametric to elongate, trigones conspicuous but not swollen, cuticle smooth; (4) underleaves often conspicuous, 0.5–1.5 mm long, margins strongly ciliate; and (5) vegetative reproduction by small plantlets (propagula) on ventral leaf surfaces. Characteristics of P. raddiana are the entire and longly decurrent ventral leaf bases which are usually strongly expanded (ampliate), concealing the stem. The species is very close to P. patula, which also has a longly decurrent ventral leaf base,

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Keys and Descriptions

but in P. patula the ventral leaf bases are not or only slightly ampliate and are not concealing the stem (Heinrichs and Gradstein 2000; Gradstein 2015c). Plagiochila raddiana is also close to P. disticha and P. montagnei, but in the latter two species, the ventral leaf bases are toothed and shortly decurrent (sometimes in longly decurrent in P. montagnei). Branching in P. raddiana is normally terminal, but some populations may have predominantly intercalary branches. The dentation of the leaves varies greatly and occasionally the leaves may be subentire. Plants from Colombia with subentire leaves were erroneously identified as P. lingua Steph. (Gradstein et al. 2016). 52. Plagiochila revolvens Mitt. On moist rocky soil in superpáramo, 4000–4300 m. Colombia: Met (Nevado de Sumapaz, Cleef 8082, COL). Ecuador: Na, Pi (type). General distribution: northern Andes (Colombia, Ecuador). Plagiochila revolvens (sect. Arrectae) is only known from superpáramo in the northern Andes. The species is readily recognized by the dark brown to blackish plants with long, unbranched stems with a curved apex and imbricate, transverse leaves with strongly revolute margins without any teeth. The leaves are suborbicular, slightly wider than long (ca. 1.2× wider than long); the leaf bases are long decurrent, especially dorsally; a vitta-like area is present at the leaf bases, and the cuticle is smooth. In its transverse leaves with entire margins and a short vitta-like area, P. revolvens is similar to P. cleefii and P. dependula. Plagiochila dependula differs from P. revolvens in the more robust habit, broadly reniform leaves with plane margins, and the papillose cuticle, while P. cleefii differs in its pale color and slightly longer than wide leaves with plane apical margins. Plagiochila revolvens may be confused with Syzygiella sonderi, which is also blackish in color and occurs in superpáramo, but the latter species has plane to incurved dorsal leaf margins (recurved in P. revolvens). 53. Plagiochila rudischusteri H.Rob. On bark of trees in coastal, lower montane rainforest, 1500–1600 m. Colombia: Ris (Uribe and Gradstein 1998). General distribution: Costa Rica, Panama, Caribbean coast of Venezuela (Paria Peninsula), Pacific side of the Western Cordillera of Colombia. Plagiochila rudischusteri (sect. Fuscoluteae) is a very distinct species that is recognized by (1) plants with linear leaves, 3–5 mm wide, pale yellowish-green to brownish, branching sparse, intercalary; (2) leaves distant, transverse, (5–)6–10× longer than wide, entire or with 1–2 small teeth near apex, bases not or hardly decurrent; (3) leaf cells narrowly elongate, 2–5× longer than wide in midleaf, 15–25  μm wide, with large, swollen trigones, cuticle smooth; and (4) vegetative reproduction by caducous leaves (Heinrichs 2002a). Plagiochila rudischusteri is related to P. aerea and P. tabinensis but is readily separated from the latter two species in the linear, transverse leaves without teeth or with 1–2 very small teeth near apex and leaf bases not or hardly decurrent.

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54. Plagiochila rutilans Lindenb. Fig. 31E Plagiochila cobana Steph., syn. nov. Type: Ecuador, Morona Santiago, Rosario, Allioni s.n., Bryotheca Levier 6490 (holotype G-00064144!) Plagiochila divaricata Lindenb. Plagiochila linearis Carl Plagiochila perrottetiana Mont. & Gottsche On bark, rock, rotten wood, and soil in lowland and montane forests and scrub, 200–3200  m. Colombia: Ant, Cau, Ces, Cun, Put, Ris. Ecuador: EO (Reserva Buenaventura, Benitez et al. 1319, HUTPL), Es, MS, Na, Pa, Pi, Su, Tu, ZC. General distribution: widespread in tropical America. Plagiochila rutilans (sect. Heteromallae) is a common neotropical species that is readily recognized by (1) plants 2–6 mm wide, when fresh with a strong peppermint smell, branching intercalary; (2) leaves distant to slightly imbricate, horizontally spreading, narrowly oblong-rectangular with a rounded apex, (1.5–)2–4× longer than wide, not ampliate but often shouldered, with 5–20 short teeth in the upper half, apical teeth not or only slightly larger, bases shortly decurrent and often recurved; (3) cells 1–2× longer than wide in midleaf, 20–35(–40) μm wide in midleaf, isodiametrical to somewhat elongate (1–2× longer than wide), with small to large trigones, vitta-like area lacking, cuticle smooth; (4) perianth base covered by the bracts; and (5) vegetative reproduction by caducous leaves (Heinrichs et al. 2001). Plagiochila rutilans resembles P. aerea, P. distinctifolia, P. macrifolia, and P. gymnocalycina in leaf shape, but the leaves in the latter four species are ventrad and lack a peppermint smell. Moreover, P. aerea has more elongate leaf cells, and P. distinctifolia and P. macrifolia have predominantly terminal branching. Plagiochila cobana reported from Cundinamarca and Valle (Gradstein and Uribe 2016a) is identical to P. rutilans and a synonym. 55. Plagiochila simplex (Sw.) Lindenb. Plagiochila confundens Lindenb. & Gottsche Plagiochila erronea Steph. Plagiochila fendleri Mont. Plagiochila regeliana Steph. Plagiochila umbrosa Steph., nom. illeg.

Fig. 31F, G

On bark and shaded moist rock in humid lowland and montane forest environments, from sea level to the páramo (3700 m). Colombia (new): Cho (Nuqui, “El Amargal,” coastal rainforest, Gradstein 8835, COL, PC). Ecuador: Az (Parque Nacional Cajas, Gradstein 12664, QCA), Na (páramo de Oyacachi, Gradstein 12804, QCA), MS, Or (Mota de Oliveira and ter Steege 2013), Pa, Pi (Río Guajalito, Gradstein 12633, QCA), ZC (Río Nangaritza, Gradstein 12116, PC). General distribution: tropical America. Plagiochila simplex (sect. Heteromallae) is a widespread neotropical species that has been little recorded in the Andes. There are a few recent records from Ecuador (most of them new), and the species is reported here for the first time from Colombia. A previous record from Colombia (Magdalena) belongs to P. tamariscina (Gradstein

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Keys and Descriptions

2016a). Plagiochila simplex is distinguished by (1) plants 1–3(–4) mm wide, branching intercalary; (2) leaves ventrad, somewhat caducous, ovate to oblong to subrectangular, ca. 1.2–2× longer than wide on main stems (sometimes more elongate on branches), with 3–10 teeth in the upper half, apex occasionally with 2 somewhat larger teeth, ventral leaf base not ampliate, bases rather shortly decurrent; (3) cells small, 15–30(−35) μm wide in midleaf, isodiametrical-subquadrate to rectangular, with conspicuous trigones, the trigones sometimes confluent and walls uniformly thickened, vitta-like area absent, cuticle smooth; oil bodies rather small, filling part of the cell lumen, oblong, 3–8 per cell, faintly segmented-papillose (Gradstein 12633); (4) perianth base “naked,” inner bracts inserted at some distance from the perianth and not enveloping the lower half of the perianth; and (5) vegetative reproduction by caducous leaves. Plagiochila simplex is close to P. gymnocalycina and differs from the latter only in the less elongate stem leaves, which are less than 2× longer than wide (Heinrichs et al. 1998). Branch leaves, however, are usually more narrowly elongate and frequently more than 2× longer than wide. For identification, full-grown stem leaves should therefore be examined, not branch leaves. Plagiochila simplex may be confused with P. subplana, but the leaves in P. subplana are horizontally spreading, not ventrad, and not caducous. Moreover, the leaf cells in P. subplana are larger, 30–45 μm wide in midleaf, and more thin-walled, with very small trigones; and the base of the perianth is covered by the bracts. Phenotypes of P. simplex with strongly caducous leaves may be confused with P. punctata, but the latter species has shorter, ovate leaves, 1.9–1.2× longer than wide, with a vitta-like area (vitta-like area lacking in P. simplex). 56. Plagiochila stricta Lindenb. Plagiochila acanthoda Lindenb. & Gottsche Plagiochila asperifolia Steph. var. columbica Herzog Plagiochila elegantula Herzog Plagiochila oxyphylla Spruce On bark, from sea level to 4000 m. Colombia: Cho, Cun, Qui (Gradstein 12632, HUQ). Ecuador: Az, Ca, Co, Ch, Im, Pi, ZC.  General distribution: tropical America, tropical Africa (Madagascar), Macaronesia. Plagiochila stricta (sect. Arrectae) is a widespread peri-African species that is recognized by (1) plants slender and little branched, 2–4 mm wide, branching intercalary; (2) leaves obliquely to widely spreading, distant to subimbricate, asymmetrically ovate-triangular to ovate-oblong, 1.1–1.7× longer than wide, without border of thicker-walled cells, dorsal margin ± entire, ventral and apical margins with 5–20(−50) narrowly triangular to linear teeth, the teeth ca. 2–6 cells long, often 2 larger teeth present at apex (to 11 cells long), dorsal and ventral bases shortly decurrent, ventral base not or slightly ampliate; and (3) leaf cells small, 15–30(−35) μm wide in midleaf, isodiametrical to somewhat elongate, with small to conspicuous trigones, leaf base with a short vitta-like area, cuticle finely striate-papillose; oil bodies large, almost filling up the cell, 4–8 per cell in midleaf, oblong, faintly and coarsely segmented (Gradstein 12632).

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Plagiochila stricta is closely related to P. punctata, but the latter species has caducous leaves produced on flagelliform branches and a smooth cuticle. Plagiochila stricta shares a finely papillose cuticle with P. papillifolia, but the latter species is a very much smaller plant (0.5–1[−1.5] mm wide) with leaf bases scarcely decurrent and without vitta-like area (Heinrichs et  al. 2002b). Plagiochila stricta also approaches P. pachyloma, but the latter species has a distinct border of thickerwalled cells. 57. Plagiochila subplana Lindenb. Plagiochila subplana var. scoposa Herzog Plagiochila amazonica Spruce Plagiochila harlingii S.W.Arnell Plagiochila kegeliana Steph. Plagiochila leptodictyon Herzog

Fig. 31H

Common on bark in shaded places in lowland and submontane rainforests and scrub, 100–1200(−1900) m. Colombia: Ama, Boy, Cho, Gui, Mag, Ris, Val, Vau. Ecuador: Es, Ga, LR, MS, Na, Or, Pa, Pi, Su. General distribution: tropical America. Plagiochila subplana (sect. Cucullatae) is a common neotropical species of low elevations (mostly below 1200 m) that is distinguished by (1) plants 2–7 mm wide, branches intercalary; (2) leaves horizontally spreading, ± rectangular with a broad, truncate apex, occasionally ovate-oblong, 1.3–2.5(−3)× longer than wide, not ampliate, with 5–40 rather longly linear teeth in the upper half or along the entire leaf margin, ventral base slightly recurved, not or shortly decurrent, dorsal base shortly decurrent; and (3) cells 30–45 μm wide in midleaf, subisodiametrical, thinwalled with small, simple-triangular trigones, vitta-like area lacking, cuticle smooth (Heinrichs et al. 1999). The horizontally spreading, ± rectangular leaves with a broad, truncate apex, long spinose teeth, and rather large, translucent cells are typical of P. subplana. Plagiochila subplana may be confused with P. rutilans and P. alternans, but P. rutilans has more narrowly oblong leaves with smaller teeth and smaller cells (20–35 μm wide) and P. alternans has ovate-oblong leaves with a broadly rounded apex, conspicuously decurrent ventral leaf bases and more numerous and much smaller teeth on the leaf margins. Moreover, P. alternans grows at much higher elevation. 58. Plagiochila subundulata Lindenb. On wet rock at the base of small waterfall, 2040 m. Colombia: Mag (Gradstein et al. 2016). General distribution: southeastern Brazil, Colombia. Plagiochila subundulata (sect. Arrectae) is a rare, rheophilous Brazilian species that has been found once in the Sierra Nevada de Santa Marta. Characteristics are (1) plants rather small, ca. 2  mm wide, bright to dark green, leafy stems rigid, upright, arising from a creeping stolon, sparsely branched in the lower half by intercalary branching; (2) leaves obliquely spreading, strongly ventrad-secund, distant to contiguous, convex with recurved dorsal and ventral margins and  ±  plane apex, ovate, 1.2–1.3× longer than wide, not ampliate, apex rounded, apical and upper

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Keys and Descriptions

ventral margin with 2–7 very small, triangular teeth, the teeth 1–2(−3) cells long and sometimes rudimentary or broken, dorsal base shortly decurrent, ventral base longly and very narrowly decurrent; and (3) cells ca. 25–40 μm wide in midleaf, isodiametric to somewhat elongate, toward the margin smaller, quadrate to elongate, toward the base elongate, forming an obscure vitta-like area, walls very thin, trigones absent or minute, cuticle smooth. Plagiochila subundulata is morphologically close to P. bifaria but differs from the latter in having subentire leaves with 2–7 very small teeth and very thin-walled leaf cells without or with minute trigones. 59. Plagiochila superba (Spreng.) Mont. & Nees Plagiochila conspicua Taylor Plagiochila cultrifolia Spruce Plagiochila frontinensis Steph. Plagiochila pichinchensis Taylor

Fig. 31I

On trunk bases, rotten logs, rock, and humic soil in humid montane forests and scrubby páramo, ca. 1000–3200 m. Colombia: Ant, Cau, Ces, Cho, Mag, Nar, Put, Qui, Ris, Val. Ecuador: Ch, Co, EO, Lo, LR, MS, Na, Pi, Tu, ZC. General distribution: throughout tropical America. Plagiochila superba (sect. Hylacoetes) is a common and variable neotropical species that is readily recognized by (1) plants large, 5–11  mm wide, branching intercalary, rarely terminal; (2) leaves wide-spreading, imbricate, ovate-triangular with a narrow, truncate apex (ovate-oblong with a broad, rounded apex in var. macrotricha), ca. 1.2–1.6× longer than wide, ± ampliate, unbordered, with 10–40 mostly linear teeth, the teeth to 11 cells long, ventral and dorsal bases without teeth, dorsal base shortly to rather longly decurrent, ventral base shortly decurrent; (3) leaf cells large, 35–55  μm wide in midleaf, slightly elongate, with conspicuous trigones, vitta-like area lacking, cuticle smooth; and (4) androecia in clusters of male branches at the plant apex. The robust plants with ampliate, ovate-triangular leaves with a narrow apex and numerous linear teeth (to 11 cells long, mostly 1 cell wide), large leaf cells, and entire dorsal leaf bases are unmistakable. Heinrichs et al. (2002a) recognized two varieties, P. superba var. superba with ovate-triangular leaves and a narrow, truncate leaf apex and P. superba var. macrotricha (Spruce) Heinrichs with ovate-oblong leaves and a broad, rounded leaf apex. Both varieties occur in Colombia and Ecuador. Plagiochila superba var. superba may be confused with P. canelensis, but the latter species has shorter and broader leaves that are hardly longer than wide and frequently toothed at the dorsal base. Plagiochila superba var. macrotricha, on the other hand, is very similar to P. amicta in leaf shape but differs from the latter species in the shape of the marginal teeth, which are triangular in P. amicta and linear in P. superba var. macrotricha. 60. Plagiochila tabinensis Steph. Acrobolbus laceratus R.M.Schust.

Fig. 31J, K

On trunks and canopy branches in humid montane forest and in scrub, 1500–3100 m. Colombia: Boy, Cau, Mag, Ris. Ecuador: Lo, Su, ZC. General distribution: tropical Andes.

Plagiochilaceae

317

Plagiochila tabinensis (sect. Fuscoluteae) is characteristic Andean species that is readily distinguished by (1) plants large, 4–10 mm wide, whitish-green to brown, branching intercalary, occasionally terminal; (2) leaves horizontally spreading to weakly ventrad, 2–4× longer than wide, widest at the base, not ampliate, apex deeply split (to 1/3–2/3 of leaf length) into 2–4 fragile, narrowly lanceolate to linear, lobe-like teeth, ventral margin entire or with 1–3 teeth, dorsal margin ± entire, dorsal base shortly decurrent, ventral base shortly (small plants) or longly decurrent (robust plants); and (3) leaf cells distinctly elongate, (1.5–)2–4× longer than wide, ca. 20–30 μm wide in midleaf, with large, radiate to confluent trigones, no vitta-like area, and a slightly rough cuticle. The leaf apex splitting deeply into several long and fragile, linear lobes is the principal character of this species. Plagiochila tabinensis is close to P. aerea, but the latter species is a more delicate plant with leaves widest in the middle and the leaf apex splitting into 2–3 shorter, sharp, lanceolate teeth (to maximally 1/3 of leaf length), not into long-linear lobes. Moreover, the apical teeth of P. aerea are not fragile (Heinrichs 2002a). 61. Plagiochila tamariscina Steph. On tree trunks and rotten wood in lowland and lower montane rainforests, 180–2200  m. Colombia: Boy, Cun, Mag, Val (Herzog 1955 as P. crispabilis). Ecuador: Lo, MS (near Misión Bomboiza, Holm-Nielsen et al. 4404, AAU, PC), Na (slopes of Guagra Urcu, Holm-Nielsen et al. 26836, AAU, PC), ZC. General distribution: scattered in tropical America. Plagiochila tamariscina (sect. Vagae) is a predominantly Caribbean species that is recognized by (1) plants rather delicate, 2.5–5 mm wide, pale yellowish-green, feather-like, (bi)pinnately branched by terminal branching; (2) leaves distant, obliquely spreading, oblong, 1.8–2.5× longer than wide, not ampliate, sometimes with a shoulder at the ventral base, widest in the lower third or middle, ventral and apical margins sharply toothed, bases shortly decurrent; (3) cells 15–25 μm wide in midleaf, elongate, walls with small trigones or ± evenly thickened, cuticle smooth; and (4) vegetative reproduction by small plantlets (propagula) from ventral leaf surfaces. In its pinnate-terminal branching, P. tamariscina is similar to P. deflexirama, but the latter species has broader, subimbricate to imbricate leaves that are widest near the base and weakly to strongly ampliate. 62. Plagiochila tenuis Lindenb. Plagiochila bidens Gottsche Plagiochila scabrifolia Inoue

Fig. 31L, M

On bark, rock, and soil in montane forests and páramo, 1600–4000 m; at lower elevation on Galápagos (750 m). Colombia: Mag, Ris. Ecuador: Ga, Pi (Burghardt 6604, QCA), ZC (Burghardt 6989, QCA). General distribution: widespread in tropical America. Plagiochila tenuis (sect. Arrectae) is a delicate neotropical-montane species distinguished by (1) plants 1–2 mm wide, branching intercalary; (2) leaves narrowly oblong, 2–2.2× longer than wide, sometimes caducous, apex bifid to 1/5, ventral

318

 

Keys and Descriptions

margin entire or with 1–3 small teeth; (3) cells ca. 18–28 μm wide in midleaf, isodiametric to elongate, 1–2× longer than wide, with small trigones, vitta-like area absent, cuticle finely striate-papillose; and (4) vegetative reproduction by caducous leaves. In its bifid leaves with a papillose cuticle, P. tenuis resembles P. papillifolia. However, the leaves in P. papillifolia are shorter, less than 1.8× longer than wide, and more strongly toothed (to 12 teeth). Small bifid leaves also occur in P. aerea, P. bicuspidata, P. cuneata, and P. exigua, but in the latter four species, the cuticle is smooth. 63. Plagiochila trichostoma Gottsche Plagiochila densa Herzog (?) Plagiochila granatensis Gottsche Plagiochila hans-meyeri Steph. Plagiochila ovato-obconica Steph. Plagiochila permista Spruce Plagiochila saltuensis Steph.

Fig. 31N

On bark, soil, rock, and rotten logs in humid montane forests and scrubby páramo, 1500–4000  m. Colombia: Ant, Boy (Bischler 2959, PC), Cau, Cho (Gradstein 8946, COL), Cun. Ecuador: Co (páramo de Llanganates, 4000  m, Burghardt 6756, QCA), Im, Na, Lo, Tu, ZC.  General distribution: tropical Andes, Costa Rica. Plagiochila trichostoma (sect. Heteromallae) is an uncommon Andean species that is recognized by (1) plants rather robust, (3–)4–7 mm wide, branching intercalary, occasionally flagelliform (Bischler 2959); (2) leaves alternate to subopposite, imbricate, ventrad-secund, shortly ovate to ovate-triangular, 0.9–1.3× longer than wide, shouldered or ampliate, not or slightly bordered, toothed all around except on the lower half of the dorsal margin, with (3–)15–40 triangular to linear teeth, the teeth 2–6 cells long, leaf bases shortly decurrent; (3) cells 30–40 μm wide in midleaf, subisodiametrical, with large swollen, trabeculate trigones becoming confluent toward leaf margin and sometimes forming a weak border, leaf base usually with a vitta-like area, cuticle smooth, oil bodies almost homogeneous; (4) perianth base covered by bracts or free; and (5) vegetative reproduction occasionally present, by caducous leaves (Heinrichs et al. 2002a). Plagiochila trichostoma resembles P. heteromalla, but the leaves in the latter species are usually opposite, especially ventrally, and more conspicuously bordered. Plagiochila trichostoma approaches P. adianthoides, but the leaves in P. adianthoides are more elongate (1.3–1.8× longer than wide), not ventrad or secund, and without vitta-like area. Moreover, the teeth on leaf margins in P. adianthoides are mostly linear. Plagiochila trichostoma may also be confused with P. simplex, but the latter species has more elongate leaves, 1.2–2(−2.5)× longer than wide, without ampliate base and lacking a vitta-like region. A phenotype from high elevation in páramo of Ecuador (Napo, 3650 m, Gradstein 12799, QCA) stands out by very large trabeculate trigones and stem leaves with only 3–12 teeth.

Plagiochilaceae

319

64. Plagiochila turgida Herzog Plagiochila longaeva Herzog On bark in humid montane forests up to the páramo, 1800–3500 m. Colombia: Cau, Hui, Ris. Ecuador: Pi, ZC. General distribution: northern Andes, Central America. Plagiochila turgida (sect. Hylacoetes) is an uncommon neotropical species that is similar to P. macrostachya in the subopposite leaves with rather large cells but differs in having less elongate leaves, 1–1.4× longer than wide (1.2–1.6× longer than wide in P. macrostachya) and toothed dorsal leaf bases (entire in P. macrostachya) (Heinrichs 2002a). 65. Plagiochila vincentina Lindenb. Plagiochila depressa Spruce Plagiochila diversispina Steph. Plagiochila hylacoetis Spruce

Fig. 31O

On bark, from sea level to 2200 m. Colombia: Ant, Boy, Caq, Cho, Cun, Mag, Ris, Val, Vau. Ecuador: LR, Ma, Pi. General distribution: widespread in tropical America. Plagiochila vincentina (sect. Hylacoetes) is a common neotropical species that is distinguished by (1) plants rather large, 5–8 mm wide, branching intercalary; (2) leaves elongate-triangular to oblong with a narrow or broad apex, ca. 1.4–2× longer than wide, shouldered to ampliate, unbordered, with 20–80 triangular to linear teeth, the teeth to 10 cells long, dorsal and ventral bases longly decurrent and strongly toothed; (3) leaf cells large, ca. 35–55 μm wide in midleaf, slightly elongate, with small radiate trigones, vitta-like area absent, cuticle smooth; and (4) androecia intercalary. Plagiochila vincentina is quite similar to P. breuteliana; both species have ampliate leaves with toothed and longly decurrent bases (Heinrichs 2002a). However, the leaves in P. breuteliana have a short vitta-like area (vitta-like area lacking in P. vincentina). Possibly, these two species are conspecific. Further Records Plagiochila arcta S.Winkl.  – Colombia: Mag (Winkler 1976). Probably not a good species; the type (ULM?) has not been studied. Plagiochila flavescens (Gottsche, Lindenb. & Nees) Gottsche fo. ampliata Herzog – Ecuador: MS (Harling 2267a). Dubious name; the type has not been seen. Plagiochila humboldtiana Gottsche – “Nova Granada” (Humboldt & Bonpland s.n.). The type has not been located; according to Stephani (1898–1924), it is lost. Plagiochila insularis Steph., nom. inval.  – Ecuador: Ga (Santa Cruz; Arnell 1961). The Galápagos material has not been located, and the identity of this undescribed taxon is considered unclear (Inoue and Gradstein 1980; Gradstein 2009). León-Yánez et al. (2006) listed the species as synonym of P. raddiana with reference to Inoue and Gradstein (1980), but the latter authors did not propose the synonymy. However, since P. insularis was collected from living leaves, the plants

320

 

Keys and Descriptions

might indeed have belonged to P. raddiana, which is the only Plagiochila species found epiphyllous on Galápagos (Gradstein 2009). Moreover, a specimen of P. insularis from Guatemala in NY (H. von Tuerkheim, det. F. Stephani) resembles P. raddiana (fide photograph sent by A. Hagborg, Dec 2017). Plagiochila rufoviridis Spruce – Colombia (Dugas 1928). The type, from Bolivia, could not be found in MANCH (L. Loughtman, pers. comm.). Plagiochila rutilans Lindenb. fo. foliicola Herzog  – Colombia: Cau (Herzog 1955). The type is lost (Heinrichs et al. 2001). Plagiochila sprucei Steph. – Ecuador: Ch (Spruce s.n., type). The type has not been located. Plagiochila thyoides Spruce  – Ecuador: Tu (confluens Río Pastaza and Río Verde, Spruce s.n., type). The type could not be found in MANCH (L. Loughtman, pers. comm.). Excluded Records Plagiochila aspleniformis R.M.Schust. – Colombia: Put (Robinson 1967). The Colombian material of this North American species presumably belongs to P. patula. Plagiochila crispabilis Lindenb. (= Plagiochila jovoensis Steph. = Plagiochila patentissima Lindenb.) – Colombia: Val (Herzog 1955), Mag (Winkler 1976). The material from Valle (Killip 11652, JE) belongs to P. tamariscina, that from Magdalena (Winkler C178, COL) to P. punctata (see Gradstein 2016a). Plagiochila dichotoma (P.Beauv.) Nees & Mont.  – Ecuador (Spruce 1885; Herzog 1952). This African species has frequently been recorded from tropical America, but the neotropical records are considered erroneous (Jones 1962). The records from Ecuador belong to P. patula. Plagiochila irmscheri Steph. – Colombia (Stephani 1898–1924). The type and the original description and illustration are a mixture of P. laetevirens and P. fuscolutea (Heinrichs 2002a). Plagiochila longiramea Steph. (= Plagiochila apicidens Steph.)  – Colombia (Gradstein and Hekking 1979 as P. apicidens), Ecuador (Parolly et al. 2004). The Ecuadorian specimen belongs to P. fuscolutea (León-Yánez et  al. 2006); the Colombian collection has not been checked but may be misidentified. According to Heinrichs et al. (2002a), P. longiramea is endemic to Bolivia.

Pseudolepicoleaceae (Blepharostomataceae) Plants pale green to brown, creeping to ascending, irregularly branched, stolons lacking. Stems with a weakly differentiated cortex of thick-walled cells. Branches Frullania-type, Microlepidozia-type and intercalary (ventral and lateral intercalary). Leaves transverse or succubous, deeply divided into 3–4(−5) uniseriate to

Pseudolepicoleaceae

321

lanceolate segments, margins entire or toothed. Cells normally with uniformly thickened walls and a striate-papillose cuticle; oil bodies granular. Underleaves similar to the leaves. Rhizoids in tufts from underleaf bases. Gametoecia on long shoots. Sporophytes surrounded by a plicate perianth, mouth of the perianth longciliate. Setae rather thin, with 8–20 outer rows of cells. Capsules ellipsoidal, wall 2–5-layered. Vegetative reproduction rare. A small family with eight or nine genera worldwide, three genera at high elevations in tropical America. Pseudolepicoleaceae are mainly distinguished by (1) transverse leaves, which are deeply divided into 3–4 linear, hair-like lobes; (2) underleaves large, similar to leaves; and (3) uniformly thick-walled cells with a striate-papillose cuticle. The family shares many characters with Lepidoziaceae, but the gametoecia in Pseudolepicoleaceae are on leading shoots, not on short ventral branches, and the cuticle is striate-papillose (smooth or rough by small, rounded papillae in Lepidoziaceae). 1. Leaves succubous, divided to ca. 1/2 into narrow lanceolate segments. Segments 2–6 cells wide in the lower half, uniseriate above. Leaf cells with trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetocolea 1. Leaves transverse, divided to more than 1/2 into subulate segments. Segments uniseriate throughout or the base of the segments 2–3 cells wide. Leaf cells with uniformly thickened walls, trigones lacking . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves divided to the base. Plants green . . . . . . . . . . . . . . . . . . Blepharostoma 2. Leaves not divided to the base, with a short lamina. Plants brown, very rare, growing on rotten wood and peaty soil in upper montane cloud forest and subpáramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temnoma Blepharostoma Dumort. Plants very small, to 1 cm long and less than 1 mm wide, pale green, little branched. Branches mostly Frullania-type. Stems without hyalodermis. Leaves transverse, divided to the base into 3–4(−5) stiff, uniseriate segments. Cells longer than wide, the walls uniformly thickened, cuticle finely striate-papillose; oil bodies granular, 3–7 per cell. Underleaves similar to the leaves but with 1 filament less, (2–)3-lobed. Monoicous or dioicous. Perianth surface smooth. Vegetative reproduction rare, by gemmae from upper leaf segments. Widespread in the Holarctic (3 spp.), extending southward into the mountains of Central America and the Andes; one species in Colombia and Ecuador. For main features of the genus, see under the species. Blepharostoma is sometimes placed in a family of its own, Blepharostomataceae (Söderström et  al. 2016), but here the family is retained in Pseudolepicoleaceae following Crandall-Stotler et al. (2009). Literature: Schuster (1966, 1986), Gradstein et al. (2001) 1. Blepharostoma trichophyllum (L.) Dumort.

Fig. 32D–G

On shaded soil in páramo, often under rock, and on rotten wood in Polylepis woodland, 3000–4300 m. Colombia: Boy, Cun, Met, Nar, Ris, Tol. Ecuador: Co (Burghardt 6781, PC, QCA), Lo, Na. General distribution: Holarctic region, Central America, tropical Andes, East Africa.

322

 

Keys and Descriptions

Blepharostoma trichophyllum is a widespread Holarctic species that extends southward to the Andes and the high mountains of East Africa. It is distinguished by its alga-like habit, with leaves and underleaves all of equal length and divided to the base into 3–4 hair-like, 1-cell-wide lobes. The species may be confused with Telaranea, but in the latter genus, the underleaves are much shorter than the leaves or absent, the stem of Telaranea has a distinct hyalodermis (lacking in Blepharostoma), and the leaf cells are thin-walled and smooth (thick-walled and ± striate-papillose in Blepharostoma). The striate papillose cuticle of B. trichophyllum is rather inconspicuous in the Ecuadorian material and is best observed on the leaf base and the stem. Due to the thickened cell walls, the leaves in Blepharostoma are usually rather stiff and dull in color, while those of Telaranea are quite flexible and glistening. A further difference is seen in the position of the gametoecia, which are situated on a very short lateral branch in Telaranea while at the apex of a long shoot in Blepharostoma. Chaetocolea Spruce Plants small, to 5 cm long and 1 mm wide, greenish-brown to dark brown, irregularly branched. Branches terminal (Frullania-type, Microlepidozia-type) and ventral-intercalary. Stems brown, without hyalodermis. Leaves succubous, cuneate to palmate with a narrow base, divided to ca. 1/2 into 3–4 narrow segments, the segments straight or conspicuously diverging, uniseriate above over a length of 4–6 cells, the lower part 2–6 cells wide. Cells with trigones, cuticle densely striatepapillose; oil bodies finely granular, 2–10 per cell. Underleaves large but shorter than leaves, deeply bifid with 2 lateral teeth (to 4-fid). Monoicous. Perianths inflated, urn-shaped, surface with scattered cilia. A genus with a single species, endemic to the northern Andes. For main features of the genus, see under the species. Literature: Schuster (1986, 2002), Gradstein et al. (2001) 1. Chaetocolea palmata Spruce 

Fig. 32H–K

On soil, rotten wood, and over leaf litter in upper montane cloud forest areas and elfin forest, 2400–3300 m. Colombia: Tol (Gradstein et al. 1977). Ecuador: Na (Guagra Urcu, elfin forest near the summit, 3150  m, Holm-Nielsen et  al. 27615, optime, AAU, PC), Tu (type, Spruce 1885). General distribution: northern Andes. Chaetocolea palmata is a rare northern Andean endemic that is known from one collection from Colombia, two from Ecuador, and one from northern Peru (Frey 1987). The species is characterized by (1) leaves palmate, divided to the middle into 3–4 diverging, triangular lobes; (2) leaf cells with conspicuous trigones, cuticle densely covered by striate papillae; and (3) underleaves large, 2(−4)-fid; (4) perianth urn-shaped, surface with scattered cilia.

Pseudolepicoleaceae

323

Fig. 32 (A–C) Pleurozia paradoxa. (A) Habit with sterile perianths, dorsal view. (B) Leaf margin. (C) Leaf. (D–G) Blepharostoma trichophyllum. (D) Habit with perianth, dorsal view. (E) Leaf. (F) Underleaf. (G) Leaf cells, showing papillose cuticle. (H–K) Chaetocolea palmata. (H) Habit with gynoecium, lateral view. (I) Leaves. (J) Underleaf. (K) Leaf lobe. (L–N) Temnoma chaetophyllum. (L) Habit with gynoecium, ventral view. (M) Habit, ventral view. (N) Leaf lobe. (All from Gradstein et al. 2001)

324

 

Keys and Descriptions

Temnoma Mitt. Plants small, to 1 cm long, less than 1 mm wide, greenish-brown to golden-brown, little branched. Branches mostly terminal, Frullania-type and Microlepidozia-type. Stems brown, without hyalodermis. Leaves transverse, divided to 2/3–3/4 into 3–4 stiff, linear segments, the segments uniseriate except at the base where 2–3 cells wide, the lamina 4–5 cells high. Cells with uniformly thickened walls, cuticle of the lamina cells striate-papillose, of the lobe cells ± smooth; oil bodies very small, granular, 5–10 per cell. Underleaves similar to the leaves but somewhat smaller and with a lower lamina (2–3 cells high). Dioicous. Perianth surface smooth. A Southern Hemispheric genus with about ten species, one in the neotropics. For diagnostic features of the genus, see under the species. Literature: Schuster (1986, 2000a, b), Gradstein et al. (2001) 1. Temnoma chaetophyllum R.M.Schust.

Fig. 32L–N

On rotten log in upper montane cloud forest, ca. 3000 m. Colombia: Cas (Uribe and Gradstein 1998). General distribution: northern Andes (Colombia, Venezuela). Temnoma chaetophyllum is a very rare northern Andean species that is known from eastern Venezuela (Schuster 1986) and the department of Casanare (just east of the páramo de Pisba). The species is distinguished by the small, glistening, greenish-brown plants with transverse leaves and underleaves with a short lamina (3–5 cells high) and 3–4 hair-like, ciliate lobes, the cilia ca. 5 cells long. The lamina cells have uniformly thickened walls, without trigones, and are striate-papillose; the cells of the cilia, however, are ± smooth. The underleaves are similar to the leaves but somewhat smaller.

Scapaniaceae (Anastrophyllaceae, Jungermanniaceae p.p., Lophoziaceae) Plants green to brown to red or purple, creeping or erect, simple or irregularly branched. Stems without hyalodermis, cortex weakly differentiated. Branches usually lateral-intercalary, sometimes Frullania-type; stolons lacking. Leaves succubous, alternate with insertion line usually reaching dorsal stem-midline, 2–4-lobed to complicate-bilobed, the lobes equal or unequal, leaf margins entire or toothed. Cells usually with trigones; oil bodies finely granular. Underleaves lacking or very small. Rhizoids scattered. Gametoecia on long shoots. Sporophytes surrounded by a perianth, the perianth inflated with a narrow mouth or dorsiventrally flattened and with a wide mouth, surface plicate or smooth. Setae of numerous cells (cross section). Capsule walls 2–5-layered. Vegetative reproduction common by 1–2-celled gemmae from leaf margins, the gemmae usually rounded or angular to stellate, pale green to red or brown.

Scapaniaceae

325

A family of about 10–20 genera worldwide, about 10 genera occurrring at high elevations in the tropical Andes. The main features of Scapaniaceae include (1) leaves succubous, 2(−4)-lobed, sometimes differentiated into a small dorsal lobe and a large ventral lobe (Scapania, Diplophyllum); (2) underleaves usually absent; (3) rhizoids scattered; (4) gametoecia on long shoots; (5) perianth inflated or flattened; and (6) frequent production of small, 1–2-celled gemmae, which are often angular-stellate in outline. The circumscription of the family is controversial and varies among authors; here it is defined in a broad sense and includes the genera of the former family Lophoziaceae. Three of them, Anastrophyllum, Sphenolobus, and Isopaches, are sometimes placed in a separate family, Anastrophyllaceae. The latter family lacks distinctive morphological features and is characterized by molecular characters only (Söderström et al. 2010) and is therefore not accepted here. 1. Leaves divided into a small dorsal lobe and a large ventral lobe . . . . . . . . . . . 2 1. Leaves not divided into a small dorsal and a large ventral lobe . . . . . . . . . . . 3 2. Ventral leaf-lobe 2× longer than wide. Plants 1–1.5 mm wide, creeping on soil. Perianth inflated, plicate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplophyllum 2. Ventral leaf-lobe 1–1.5× longer than wide. Plants (1–)1.5–5 mm wide. Perianth flat, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scapania 3. Leaf margins on sterile stems usually with a few teeth. Leaf cells large, 30–60 μm in midleaf, thin-walled, with very small trigones. . . . . . . . . . . . . . 4 3. Leaf margins on sterile stems fully entire. Leaf cells smaller. . . . . . . . . . . . . . 6 4. Plants with ventral stolons or with a stoloniform stem base. Oil bodies granular, less than 10 per cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophonardia 4. Stolons absent. Oil bodies homogeneous, 20–50 per cell . . . . . . . . . . . . . . . . 5 5. Leaf cells very large, 40–65 μm in midleaf. Gemmae smooth, 1-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tritomaria capitata 5. Leaf cells smaller, 30–40  μm in midleaf. Gemmae angular-stellate, 1–2celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schistochilopsis incisa 6. Trigones large. Leaf apex acute to acuminate. Plants reddish-brown or purple, rarely green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anastrophyllum 6. Trigones lacking or small. Leaf apex obtuse to subacute. Plants green or brownish, rarely purplish or reddish-brown (S. austroamericanus). . . . . . . . . . . . . . 7 7. Leaf cells with small trigones. Stolons present . . . . . . . . . . . . . . Lophonardia 7. Leaf cells with ± uniformly thickened walls, trigones lacking. Stolons absent. Plants rare, in (super)páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Leaves rather densily imbricate, apices acute. Leaf cells 20–30 μm in midleaf. Gemmae reddish. Plants paroicous, often fertile, with resin smell, above 4000 m (very rare) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isopaches 8. Leaves distant or laxly imbricate, leaf apices acute or obtuse. Leaf cells smaller, 12–20 μm in midleaf. Gemmae absent or reddish-brown. Plants dioicous, frequently sterile, without resin smell. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Leaves succubous. Plants greenish-brown, ca. 1 mm wide. Perianth terete . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocoleopsis (see family Cephaloziellaceae) 9. Leaves transverse. Plants reddish-brown or greenish-brown, 0.3–1 mm wide. Perianth plicate, sometimes with a white mouth . . . . . . . . . . . . . . Sphenolobus

326

 

Keys and Descriptions

Anastrophyllum (Spruce) Steph. Plants small- to medium-sized, 0.5–3  mm wide, dark reddish-brown or purple, sometimes green or brownish-green, creeping or ascending, little branched, shoot apex ascending. Branches variable, terminal and ventro-lateral intercalary. Stems fragile to rigid. Leaves succubous to almost transverse, asymmetrically to subsymmetrically bifid, the dorsal segment transversely inserted and the ventral segment oblique, strongly concave to canaliculate with the tips often somewhat incurved, apices acute to acuminate, rarely obtuse, margins entire. Cells usually with large trigones, cuticle smooth or papillose; oil bodies finely granular. Underleaves lacking. Rhizoids scarce. Dioicous (usually). Perianths 3–6-plicate, usually with a white mouth. Capsule walls (2–)3–5-layered. Gemmae present or absent, when present yellow-brown to wine-red to purplish, angular, produced on apices of upper leaves. About 15 species worldwide, 5  in Colombia and Ecuador. The genus Anastrophyllum is readily recognized by (1) plants reddish-brown to purplish (occasionally green), stem little branched; (2) leaves bifid, concave, almost transverse, leaf tips acute to acuminate; (3) leaf cells usually with large trigones; (4) underleaves absent; and (5) perianth terminal on the stem, longly cylindrical, plicate, frequently with a white-colored mouth. When the leaves are canaliculate (e.g., A. stellatum), the ascending leaf tips may help determine which side of the stem is the dorsal one. Literature: Schuster (2002), Váňa et al. (2013c), Álvaro and Uribe (2017) 1. Leaves shallowly bifid, to maximally 1/5(−1/4) of leaf length. . . . . . . . . . . . 2 1. Leaves more deeply bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Gemmae usually present, forming whitish clusters at the stem apex. Leaves canaliculate, asymmetrically bifid, the dorsal lobe shorter than the ventral lobe, lobes acute. Plants yellow-brown to purple. . . . . . . . . . . . . . . . . . A. stellatum 2. Gemmae absent. Leaves very deeply concave but not canaliculate, symmetrically bifid (very shallowly), the dorsal lobe as long as the ventral leaf lobe, lobes obtuse. Plants dark brown. . . . . . . . . . . . . . . . . . . . . . . . . . A. nigrescens 3. Leaves bifid to half or more of leaf length, margins of leaf lobes overlapping in the sinus. Midleaf cells ± longer than wide. Plants rather robust, 1.5–2.5 mm wide, green to reddish-brown. Epiphyte in montane forest up to the páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. piligerum 3. Leaves bifid to less than half of leaf length, margins of leaf lobes not overlapping in the sinus. Midleaf cells ± isodiametrical. Plants smaller, to 1.5  mm wide, usually purplish or brown (but green when growing in shade). On soil or rock, very common in páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Cells at the base of the leaf lobes 15–20 μm wide. Dorsal leaf base decurrent (but sometimes only very slightly). . . . . . . . . . . . . . . . . . . . . . . . . . A. auritum 4. Cells at the base of the leaf lobes larger, 20–25 μm wide. Dorsal leaf base not decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tubulosum

Scapaniaceae

1. Anastrophyllum auritum (Lehm.) Steph. Anastrophyllum calocystum (Spruce) Steph. Anastrophyllum crebrifolium (Hook.f. & Taylor) Steph. Anastrophyllum leucocephalum (Taylor) Spruce Anastrophyllum leucostomum (Taylor) Steph. Anastrophyllum minutirete Herzog Marsupella pusilla Steph.

327

Fig. 33A–D

On moist soil and rock in páramo and at lower elevation on road banks, occasionally epiphytic, 1800–4500 m. Colombia: Ara, Boy, Cal, Cas, Cau, Cun, Mag, Met, Put, Qui, Ris, San. Ecuador: Az, Ca, Co, Lo, Na, Pi, ZC. General distribution: widespread in tropical high mountains and in southern-temperate regions. Anastrophyllum auritum is one of the most common liverworts of páramo and highly variable; the species has been described under many different names (Váňa et  al. 2013c). The plants are deep purplish in color when growing exposed but brownish-green in the shade, creeping to ascending and quite variable in size, typically measuring 1.5 mm wide but sometimes very small and flagelliform, being less than 0.5  mm wide and with distant leaves. Such very small plants occur mainly above 4000 m. The leaves are deeply concave, bifid to (1/4–)1/3–2/5, with highly variable tips, varying from to acute to finely acuminate and hair-like; the dorsal leaf base is more or less decurrent, and the leaf cells have small to large trigones and a smooth to striate-papillose cuticle. Plants with finely acuminate leaf tips were previously called A. crebrifolium or A. leucocephalum. The very small phenotype of A. auritum may be confused with A. tubulosum and Sphenolobus austroamericanus. In the latter two species, however, the dorsal leaf base is not decurrent. Moreover, the leaf cells in A. tubulosum are larger (see key). 2. Anastrophyllum nigrescens (Mitt.) Steph. Anastrophyllum paramicola R.M.Schust.

Fig. 33E

On soil and rock in páramo, 3000–4300 m. Colombia: Ant, Boy, Cal, Cun, Hui, Mag, Met, Qui, Ris, Tol. Ecuador: Az, Ca, Co, Lo, Na, Pi, ZC. General distribution: tropical Andes (Colombia to Bolivia), Costa Rica. Anastrophyllum nigrescens is a distinct Andean-Costa Rican páramo species that is readily distinguished by deeply concave leaves with a short-bifid apex (bifid to 1/6–1/5 of leaf length) and leaf cells with strongly sinuose walls. The plants are usually brown to dark reddish-brown in color. Plants from Azuay, páramo de Patococha (Gradstein 12677, QCA), stand out by very long stems, to 10 cm long. 3. Anastrophyllum piligerum (Nees) Steph. Anastrophyllum hians Steph.

Fig. 33F

On bark of trees in montane forest up to the páramo, rarely on humic soil or rock, 1500–4000 m. Colombia: Boy (Sierra Nevada del Cocuy, 4000 m, Bischler 2923, PC), Hui, Put, Ris, San. Ecuador: Az, Co, Lo (Burghardt 7118, PC, QCA), Na, ZC. General distribution: pantropical.

328

 

Keys and Descriptions

Fig. 33 (A–D) Anastrophyllum auritum. (A) Habit with gynoecium, lateral view. (B) Portion of stem with leaf, dorsal view. (C) Leaf. (D) Midleaf cells. (E) Anastrophyllum nigrescens, leaf. (F) Anastrophyllum piligerum, leaf. (G) An astrophyllum tubulosum, leaf. (H) Anastrophyllum stellatum, habit in dorsal view. (I and J) Diplophyllum obtusatum. (I) Habit, dorsal view. (J) Gynoecium, ventral view. (K and L) Tritomaria capitata. (K) Leaf (flattened), 18×. (L) Midleaf cells, 200×. (M and N) Isopaches bicrenatus. (M) Habit with gynoecia. (N) Midleaf cells. (O–Q) Scapania portoricensis. (O) Habit, dorsal view. (P) Leaf. (Q) Gemmae. (E, G redrawn from Váňa 1984b; H redrawn from Schuster 1997; K, L redrawn from Grolle 1959; rest from Gradstein et al. 2001; Gradstein and Costa 2003)

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Anastrophyllum piligerum is widespread tropical-montane epiphyte (occasionally on humic soil or rock) that is recognized by the relatively large plants (to 2.5 mm wide) with deeply bifid leaves, to more than half of leaf length, with finely acuminate tips and lobe margins ± overlapping in the sinus between the 2 lobes. The midleaf cells are slightly longer than wide and the perianths are deeply 3–6-plicate. Moreover, the plants are usually green or reddish-brownish in color (rarely purplish) and grow mostly epiphytic; other neotropical Anastrophyllum species are mainly terrestrial. 4. Anastrophyllum stellatum R.M.Schust. Anastrophyllum gradsteinii Váňa, nom. inval. On shaded, moist rock and soil in páramo and in Polylepis forest, 3100–4200 m. Colombia: Boy, Cun, Met, Ris, Tol. Ecuador: Ca, Lo, Pi (Cadena 186, QCA), ZC (páramo of Saraguro and Yacuambi, Benitez et al. 1307, HUTPL). General distribution: tropical Andes (Venezuela to Bolivia), Costa Rica. Anastrophyllum stellatum is a characteristic Andean-Costa Rican páramo species that is distinguished by the presence of whitish clusters of gemmae on the leaves of ascending stem tips. The leaves are wide-spreading, canaliculate with a broadly rounded keel, and shallowly and asymmetrically bifid (to maximally 1/4 of leaf length) with the dorsal lobe smaller than the ventral lobe. The leaf cells have large, bulging trigones. 5. Anastrophyllum tubulosum (Nees) Grolle Anastrophyllum adulterinum (Gottsche) Steph. Anastrophyllum conforme (Lindenb. & Gottsche) Steph. Anastrophyllum leptodictyon Herzog Anastrophyllum pearcei (Steph.) R.M.Schust. Anastrophyllum pusillum (Gottsche) Steph.

Fig. 33G

On soil and rock in páramo and on roadsides, 2000–4500 m. Colombia: Boy, Cal, Cun, Mag, Met, Ris. Ecuador: Az, Ca, Na, Pi, ZC. General distribution: probably throughout tropical America. Anastrophyllum tubulosum is a delicate neotropical species with leafy stems not wider than 1 mm. It has often been confused with small forms of A. auritum but differs from the latter in leaves being not decurrent at the base (more or less decurrent in A. auritum) and leaf cells larger, 20–25 μm wide at leaf lobe bases (less than 20 μm wide in A. auritum). The records of A. tubulosum from the northern Andes need verification. Diplophyllum (Dumort.) Dumort. Plants small, 0.5–1.5 cm long and ca. 1–2 mm wide, pale green to reddish-brown, creeping. Leaves complicate-bilobed with rather long lobes. Ventral lobe widespreading, lingulate, 2–4× as long as wide, apex rounded to apiculate. Dorsal lobe subparallel to the stem, about 1/2 the length of the ventral segment, apex rounded to long-acuminate. Cells small, uniformly thick-walled or with small trigones, a vitta

330

 

Keys and Descriptions

sometimes present, cuticle papillose; oil bodies finely granular. Underleaves absent. Perianths inflated, plicate. Gemmae papillose-stellate. About 16 species worldwide, 1 in Colombia and Ecuador. For main features of the genus, see under D. obtusatum. Literature: Schuster (1974, 2002) 1. Diplophyllum obtusatum (R.M.Schust.) R.M.Schust.

Fig. 33I, J

On moist, loamy soil of steep, cut road banks and along trails, in deep shade, 2650–3400  m. Colombia: Cas, Cun. Ecuador: Ca (Gradstein and Váňa 1987). General distribution: North America, Jamaica, Costa Rica, northern Andes. Diplophyllum obtusatum is a North American species that has been collected a few times in the neotropics, including the northern Andes. As the species occurs in the neotropics exclusively in man-made habitats  – as opposed to a more natural habitat (rock outcrops) in North America – the species may have been introduced in tropical America (Gradstein and Váňa 1987). Diplophyllum obtusatum is recognized by the small, pale green, creeping plants (1–1.5  mm wide) with lingulate leaves (ca. 2–4× as long as wide) divided into a large ventral lobe and a smaller dorsal lobe. The leaf margins are finely denticulate, the leaf apices are rounded, the cuticle is strongly papillose, and a vitta is absent. The plants are monoicous (autoicous or paroicous) and always fertile; the perianth is inflated-(ob)ovoid and plicate. Vegetative reproduction by small, 1-celled, stellate-papillose gemmae is common. Isopaches H.Buch Lophozia (Dumort.) Dumort. subg. Isopaches (H.Buch) R.M.Schust. Plants very small, 0.5–1  mm wide, with a strong resin smell, pale yellowishbrown to reddish, sometimes whitish, creeping, stolons lacking. Stems fragile. Leaves densily imbricate, subtransverse, bifid to 1/3 of leaf length, apices obtuse to subacute, margins entire. Cells small, 20–30 μm in midleaf, walls uniformly thickened, slightly thicker-walled at the angles, trigones not sharply delimited, cuticle smooth; oil bodies finely granular. Underleaves lacking. Rhizoids usually numerous. Paroicous, usually fertile. Perianths plicate. Gemmae usually present at leaf tips, reddish-brown, angular-stellate. A small Holarctic genus of four species, one in Ecuador. For characters of the genus, see under the species. Literature: Schuster (1969, 2002 under Lophozia subg. Isopaches) 1. Isopaches bicrenatus (Hoffm.) H.Buch Lophozia bicrenata (Hoffm.) Dumort.

Fig. 33M, N

On rock and soil in superpáramo, 4300–4500 m. Colombia: Cal (Gradstein et al. 2018c). Ecuador: Na (Benitez et  al. 2012). General distribution: temperate

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331

regions of the Northern Hemisphere, scattered elsewhere: New Zealand, northern Andes, and southeastern Brazil (Serra do Itatiaia). Isopaches bicrenatus is a widespread Holarctic species that has been found twice in superpáramo in the northern Andes. The plants are small, ca. 1 mm wide, creeping and yellowish-brownish to whitish in color, paroicous and often fertile, and emit a characteristic resin smell. The leaves are subtransverse and densily imbricate, bifid to 1/3 with obtuse apices, and masses of reddish-brown gemmae are produced at the stem tips. The leaf cells are ca. 20–30 μm wide in midleaf, and the cell walls are more less uniformly thickened. Isopaches bicrenatus may be confused with Sphenolobus minutus, which may grow in the same habitat and also has reddish-brown gemmae. However, the leaves of S. minutus are more laxly imbricate, complicate-bilobed, and transverse, and the plants lack the resin smell characteristic of I. bicrenatus. Moreover, S. minutus is dioicous and usually sterile. Sphenolobus minutus has been found in superpáramo of the Sierra Nevada de Santa Marta. Lophonardia R.M.Schust. Lophozia subg. Hypolophozia R.M.Schust. Plants glossy green to dark green to brown, creeping, with ventral stolons or a stoloniform stem base. Branching Frullania-type and ventral-intercalary (stoloniform). Leaves subimbricate to distant, bifid to 1/3–1/2, leaf tips obtuse to sharply acuminate, margins entire or toothed. Leaf cells 25–40 μm in midleaf, thin-walled or almost uniformly thick-walled, trigones small, cuticle smooth; oil bodies finely granular. Underleaves lacking. Rhizoids usually numerous. Usually dioicous. Perianths plicate. Capsule walls 2(−3)-layered. Gemmae absent or present, pale green or reddish-brown, 1–2-celled, angular, produced on the apices of upper leaves. A small Afro-American genus of three species, two in Colombia and Ecuador. Lophonardia is exclusively found at high elevation, above 2500 m, and is mainly recognized by the creeping, glossy green to dark green to brown plants with bifid leaves, obtuse to acuminate leaf tips, small trigones, a smooth cuticle, and, especially, the stoloniform stem base. Literature: Schuster (2002 as Lophozia subg. Hypolophozia), Söderström et al. (2013b) 1. Leaf apices acuminate. Leaf margins ± toothed. Gemmae usually present, pale green. Plants light green to pale brown, upper leaves sometimes densely crowded and crisped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. jamesonii 1. Leaf apices obtuse. Leaf margins entire. Gemmae lacking. Plants brownishgreen, upper leaves never densely crowded and crisped . . . . . . . . . L. laxifolia 1. Lophonardia jamesonii (Mont.) L.Söderstr. & VáňaFig. 34D–F Andrewsianthus jamesonii (Mont.) Váňa Andrewsianthus achrous (Spruce) R.M.Schust. Lophozia stolonifera R.M.Schust. (fide Söderström et al. 2013b)

332

 

Keys and Descriptions

On soil and rock in shaded habitats in moist páramo, 3000–4550 m. Colombia: Boy, Cal, Cun, Mag, Met, Qui, Ris, San, Tol. Ecuador: Pi, Tu. General distribution: Mexico, Costa Rica, tropical Andes, East Africa. Lophonardia jamesonii is a characteristic Afro-American páramo species that is recognized by (1) plants light green to pale brown, 1–1.5 mm wide, with a stoloniform stem base; (2) leaves bifid, apex acuminate, margins ± toothed by whitish teeth, especially on upper leaves, the upper leaves sometimes densely crowded and crisped, Fossombronia-like; (3) leaf cells rather large, 30–45 μm in midleaf, thinwalled with small trigones, oil bodies granular, less than 10 per cell; (4) gemmae pale green, 1-celled, in clusters at stem tips (Váňa 1980). The plants are dioicous but are frequently fertile and with perianths. Lophonardia jamesonii was long considered a member of Andrewsianthus but was recently transferred to Lophonardia because of the absence of dorsal flagelliform branches (characteristic of Andrewsianthus) and the presence of a stoloniform stem base, which is typical of Lophonardia (Schuster 2002). The species somewhat resembles Schistochilopsis incisa, which may occur in the same habitat, but the latter species is usually sterile, lacks a stoloniform stem base, is often bluish-green in color, and has very small homogeneous oil bodies, ca. 20–50 per leaf cell. In L. jamesonii the oil bodies are finely granular and fewer per cell, less than 10. 2. Lophonardia laxifolia (Mont.) L.Söderstr. & VáňaFig. 34A–C Lophozia laxifolia Mont. Lophonardia caespitosa R.M.Schust. Lophozia opacula (Spruce) Steph. Lophozia subinflata (Spruce) R.M.Schust. On moist soil (trails, earth banks, lake shores) in rather open, high-montane environments, 2500–4500 m. Colombia: Boy, Cal, Cas, Cau, Cun, Mag, Met, Ris, Tol. Ecuador: Az (Burghardt 7452a, PC, QCA), Pi, Tu, ZC. General distribution: Andes (Venezuela to Tierra del Fuego), South Africa. Lophonardia laxifolia is a widespread high-Andean species characterized by (1) plants small, greenish-brown, creeping, with a stoloniform stem base (sometimes large part of the stem leafless, stoloniform); (2) leaves rather distant, shallowly bifid to 1/4–1/3, subtransverse, as long as wide or somewhat longer than wide, apex obtuse, margins entire, with a weak border of smaller, quadrate cells; (3) leaf cells with small trigones or fully thin-walled, cuticle smooth, oil bodies finely granular (1–7 per cell); and (4) gemmae absent. The species may be confused with Gymnocoleopsis cylindriformis, but the latter species has uniformly thickened cell walls and lacks a stoloniform stem base. Scapania (Dumort.) Dumort. Plants small to very large, 0.5–20 cm long, 1–10 mm wide, pale green to reddish or brown, usually creeping, sometimes pendent, irregularly branched. Branches usually lateral-intercalary, sometimes Frullania-type; stolons lacking. Stems without hyalodermis, cortex weakly differentiated. Leaves complicate-bilobed with a small dorsal lobe and a large ventral lobe (sometimes the 2 lobes subequal), the lobes

Scapaniaceae

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Fig. 34 (A–C) Lophonardia laxifolia. (A) Habit with gynoecium, dorsal view. (B) Leaf. (C) Midleaf cells. (D–F) Lophonardia jamesonii. (D) Habit, dorsal view. (E) Habit with gynoecium, dorsal view. (F) Gemmae. (G and H) Schistochilopsis incisa. (G) Habit, dorsal view. (H) Shoot apex with gemmae. (I and J) Solenostoma amplexifolia. (I) Habit with gynoecium, dorsal view, 12×. (I) Leaves, 12×. (K–M) Solenostoma callithrix. (K) Habit, dorsal view. (L) Leaf. (M) Shoot apex with gynoecium. (N and O) Solenostoma hyalinum. (N) Habit, dorsal view. (O) Portion of stem with leaf, lateral view. (P and Q) Nardia succulenta. (P) Habit, lateral view. (Q) Midleaf cells. (I, J redrawn from Váňa 1974; N, O redrawn from Váňa 1975; rest from Gradstein et al. 2001)

334

 

Keys and Descriptions

connected by a short or long keel, margins entire or toothed. Cells usually with trigones, cuticle smooth or papillose; oil bodies finely granular. Underleaves lacking. Rhizoids scattered. Gametoecia on leading shoots. Perianths dorsiventrally flattened, not plicate. Setae of numerous cells (cross section). Capsule walls 3–7-layered. Vegetative reproduction by small, green to brown or reddish gemmae (1–2-celled) from leaf margins; gemmae smooth or papillose. A large, mainly Holarctic genus (ca. 90 spp.), 2 species in Colombia and Ecuadorm, in 2 subgenera, subg. Macroscapania R.M.Schust. and subg. Scapania (Söderström et al. 2016). Characteristics of Scapania are the conduplicate leaves with a large ventral lobe and a smaller dorsal lobe (but sometimes the 2 lobes almost equal in size) and the flattened, smooth perianth. Literature: Schuster (1974, 2002) 1. Small, brownish-green plant on limestone rock in superpáramo (above 4000 m), 1–2 mm wide. Leaf margins entire. Keel long, ca. 1/2× lobe length. Gemmae ellipsoid, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cuspiduligera 1. Robust, reddish or purplish-green epiphyte in montane cloud forest, more than 2 mm wide. Leaf margins toothed. Keel very short, the 2 leaf lobes almost free. Gemmae stellate, papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . S. portoricensis 1. Scapania cuspiduligera (Nees) Müll.Frib. On limestone rock in superpáramo, 4300  m. Colombia: Met (Gradstein and Váňa 1987). General distribution: widespread in the temperate regions of the Northern Hemisphere; at high elevations in Colombia and East Africa. Scapania cuspiduligera (subg. Scapania) is a Holarctic species from limestone rock that has been found once in tropical America, on the summit of the Nevado de Sumapaz in the Colombian Eastern Cordillera (A. M. Cleef, COL, U). The plants are small, to 1–1.5  mm wide, prostrate; the 2 leaf lobes are subequal and somewhat longer than wide; the keel is long, about 1/2× leaf length; the leaf apices are rounded to apiculate; the leaf margins are ± entire; and the gemmae are ellipsoid, smooth. 2. Scapania portoricensis Hampe & Gottsche Fig. 33O–Q Scapania portoricensis var. roraimensis Herzog Scapania geppii Steph. Scapania minutidens Steph., syn. nov. Type: Ecuador, Morona Santiago, Bomboiza, “in superioribus montis ‘Aguacate,’ ad truncos arborum in silvis m. 2400–2600,” Apr 1910, M. Allioni 291 (holotype G-124058!) Scapania splendida Spruce On bark, rock, and moist humus in montane cloud forests and páramo, 800–4100 m. Colombia: Ant, Ara, Boy, Cas, Cho, Cun, Hui, Mag, Met, Qui, Ris, San, Tol, Val. Ecuador: Az, Ca, Lo, MS, Na, Tu, ZC.  General distribution: widespread in tropical America. Scapania portoricensis (subg. Macroscapania) is a common, robust reddishgreen to reddish-brown epiphyte in neotropical upper montane cloud forest. The stems are up to 20 cm long and to 1 cm wide, and usually unbranched, the dorsal

Scapaniaceae

335

leaf-lobe is convex and much smaller than the ventral lobe, the keel connecting the 2 lobes is very short (dorsal and ventral lobes almost free from each other), the leaf apices are rounded to obtuse, the leaf margins are toothed, and the gemmae are stellate-papillose. The latter character is unique to S. portoricensis; all other species in the genus Scapania have smooth gemmae. Schistochilopsis (N.Kitag.) Konstant. Lophozia subg. Schistochilopsis N.Kitag. Plants medium-sized, ca. 1–2 mm wide, pale green to bluish-green, often somewhat fleshy and growing in rosettes, creeping, basal stolons lacking. Stems fleshy, flattened, branching intercalary, flagelliform branches lacking. Stems fragile. Leaves succubous, asymmetrically 2–4-lobed to 2/5 of leaf length, the dorsal lobe smaller than the ventral lobe, apices acute, margins ± toothed and often somewhat undulate-crisped. Cells 30–40 μm in midleaf, thin-walled with small trigones, cuticle smooth; oil bodies homogeneous, very small, 20–50 per cell. Underleaves lacking. Rhizoids numerous or scarce. Dioicous. Perianths large, plicate. Capsule walls 3–5-layered. Gemmae often present, pale green, 1–2-celled, angular-stellate, produced on the apices of upper leaves. A mainly Holarctic genus of about ten species, one in Colombia and Ecuador. The main features of Schistochilopsis are (1) plants usually pale green to bluishgreen, somewhat fleshy; (2) leaves 2–3-lobed, leaf margins frequently toothed; (3) leaf cells 30–40 μm in midleaf, thin-walled with small trigones, cuticle smooth; (4) oil bodies numerous, 20–50 per cell, homogeneous (character shared with Tritomaria capitata); and (5) gemmae angular-stellate. Literature: Schuster (1969, 2002 as Lophozia subg. Schistochilopsis) 1. Schistochilopsis incisa (Schrad.) Konstant. Lophozia incisa (Schrad.) Dumort.

Fig. 34G, H

On soil over rock on steep, cut road banks and along trails, 2500–4500  m. Colombia: Ara, Boy, Cal, Cun, Met, Ris, Tol. Ecuador: Az (Burghardt 7452, PC, QCA), Ca, Lo/ZC (páramo of Saraguro and Yacuambi, Benitez & Gradstein s.n., HUTPL), Pi. General distribution: widespread in the temperate regions of the Northern Hemisphere; scattered at high elevation in Central America and the tropical Andes, probably introduced (Gradstein and Váňa 1987). Schistochilopsis incisa is a widespread Holarctic species that occurs above 2500  m in the Andes. The species is characterized by the green to bluish-green plants with 2–4-lobed leaves with toothed and somewhat undulate margins and acute apices; the thin-walled leaf cells with numerous small, homogeneous oil bodies (20–50 per cell); and the frequent presence of clusters of pale green gemmae at the leaf tips. The leaves may be densely crowded and crisped in mature plants; such plants may resemble Fossombronia. However, Fossombronia is readily separated from S. incisa by its violet rhizoids and absence of gemmae.

336

 

Keys and Descriptions

Schistochilopsis incisa resembles Lophonardia jamesonii but differs in plant color, in the lack of basal stolons, and the presence of 20–50 homogeneous oil bodies per leaf cell (granular and less than 10 per cell in L. jamesonii). Moreover, L. jamesonii is usually copiously fertile, whereas S. incisa is mostly sterile in the Andes. Sphenolobus (Lindb.) Berggr. Anastrophyllum subg. Crossocalyx (Meylan) R.M.Schust. Plants of páramo, very small, 0.3–1 mm wide, greenish-brown to reddish-brown, creeping, little branched. Branches variable, terminal and intercalary. Stems fragile. Leaves (sub)transverse, subsymmetrically bifid to 1/4–1/2, strongly concave, apices acute, margins entire. Cells very small, 12–20  μm in midleaf, with ± uniformly thickened walls, trigones inconspicuous, cuticle smooth or faintly papillose; oil bodies finely granular. Underleaves lacking. Rhizoids scarce. Dioicous. Perianths plicate, usually with a white mouth. Capsule walls (2–)3–5-layered. Gemmae present or absent, when present yellow-brown to wine-red to purplish, angular, produced on apices of upper leaves. A small Holarctic-neotropical genus of three species, two in Colombia. Sphenolobus is very similar to Anastrophyllum and was sometimes included in the latter genus but differs in leaf cells with ± uniformly thickened walls, without or with very small trigones. Literature: Schuster (1969, 2002 as Anastrophyllum subg. Crossocalyx), Váňa (1980 as Anastrophyllum) 1. Gemmae present, reddish-brown, produced at the tip of erect, small-leaved shoots. Plants greenish-brown or brown . . . . . . . . . . . . . . . . . . . . . . S. minutus 1. Gemmae absent. Plants reddish-brown . . . . . . . . . . . . . . . S. austroamericanus 1. Sphenolobus austroamericanus (Váňa) Váňa Anastrophyllum austroamericanum Váňa On soil in humid páramo, 3750–4400  m. Colombia: Boy (Váňa 1980), Cas (Cleef et  al. 11177, PC). General distribution: tropical Andes (Venezuela, Colombia, Bolivia). Sphenolobus austroamericanus is a rare páramo species that has been found in the Sierra Nevada del Cocuy. The species is distinguished by the tiny, creeping, reddish-brown to blackish-brown plants with leafy shoots about 0.4–0.8 mm wide and with suberect to obliquely spreading, transverse leaves that are deeply concave, bifid to ca. ¼–1/3, with obtuse to subacute lobes, not decurrent bases and small, ± evenly thick-walled cells, ca. 12–20  μm in diameter in the leaf lobes. Like in Anastrophyllum, the apical portion of the perianth is whitish in color (Váňa 1980). Sphenolobus austroamericanus may be confused with small forms of Anastrophyllum auritum, but the latter species has distinct trigones and somewhat decurrent leaf bases. Sphenolobus austroamericanus also resembles S. minutus but differs from the latter in the absence of gemmae, the reddish-brown color, and the presence of stoloniform ventral branches (branching lateral in S. minutus and not stoloniform).

Scapaniaceae

337

2. Sphenolobus minutus (Schreb.) Berggr. Anastrophyllum minutum (Schreb.) R.M.Schust. Anastrophyllum gemmiferum S.Winkl. On soil in superpáramo, 4500  m. Colombia: Mag (Winkler 1976). General distribution: widespread in temperate regions of the Northern Hemisphere; scattered in tropical high mountains. Sphenolobus minutus is a widespread Holarctic species that has been collected once in superpáramo of the Sierra Nevada de Santa Marta. The plants are small, ca. 1 mm wide or less, prostrate, brownish, and the leaves are transverse, deeply hollow and regularly complicate-bilobed, with 2 equal lobes. Clumps of small, reddishbrown gemmae with angular to stellate outline are produced at the stem tips. In its complicate-bilobed, transverse leaves, S. minutus resembles Scapania, but in the latter genus, the 2 lobes are unequal in size and the gemmae are smooth, ellipsoid (except in S. portoricensis; see above). Tritomaria Loeske Heterogemma (Jörg.) Konstant. & Vilnet Plants medium-sized, ca. 1–2 mm wide, fleshy, pale green to purplish or reddish, creeping, ventral stolons lacking. Branching intercalary, flagelliform branches lacking. Stems fragile, homogeneous in cross section, epidermal cells often strongly elongate. Leaves succubous to subtransverse, laxly imbricate, asymmetrically 2–3lobed to 2/5, apices obtuse or acute, margins ± toothed and often undulate-crisped. Cells in T. capitata very large, 40–65  μm in midleaf, thin-walled, trigones very small, cuticle smooth in T. capitata; oil bodies (in T. capitata) very small and numerous, 20–50 per cell, ± homogeneous. Underleaves lacking. Rhizoids numerous or scarce. Dioicous, often sterile. Perianths large, plicate. Capsule walls 3–5-layered. Gemmae pale green to purplish, in L. capitata 1-celled and smooth, produced on apices of upper leaves. Eight species worldwide, one in Colombia. The Colombian species was previously placed in the genus Heterogemma, but recent molecular evidence revealed that Heterogemma is nested in Tritomaria (see Stotler and Crandall-Stotler 2017). For the main characteristics of the genus, see under the species. Literature: Paton (1999 as Lophozia capitata), Schuster (2002 as L. patagonica) 1. Tritomaria capitata (Hook.) Stotler & Crand.-Stotl. Fig. 33K, L Heterogemma capitata (Hook.) Konstant. & Vilnet Lophozia capitata (Hook.) Macoun (Heterogemma capitata [Hook.] Konstant. & Vilnet) Lophozia patagonica Herzog & Grolle (Heterogemma patagonica [Herzog & Grolle] L.Söderstr. & Váňa), syn. nov. In bog in páramo, 3550  m. Colombia: Cun (Gradstein and Hekking 1979 as Lophozia patagonica). General distribution: common in western Europe and eastern North America, rare in southern Patagonia and Tierra de Fuego; one record from southern Brazil (Santa Catarina) and one record from the high Andes of Colombia.

338

 

Keys and Descriptions

Tritomaria capitata is a bipolar species that has been recorded once from the tropics (Colombia, A. M. Cleef, COL, U). The Colombian specimen was originally named Lophozia patagonica, but, according to Schuster (2002), L. patagonica is morphologically identical to L. capitata (= T. capitata). The species is characterized by the laxly imbricate and somewhat crisped, bifid to trifid leaves with very large cells (40–65 μm long in midleaf), with obtuse to acute apices and entire to slightly toothed margins. The stems are usually purplish in color (at least ventrally), and the oil bodies are homogeneous and numerous per cell. Gemmae have not been observed in the Colombian specimen; when present they are 1-celled, smooth, and pale green to violet in color (Paton 1999).

Solenostomataceae Plants small- to medium-sized, green to brown to red or purple. Stems without hyalodermis, cortex weakly differentiated. Leaves succubous, undivided or shallowly bifid, alternate, insertion line reaching the dorsal stem-midline, leaf margins entire. Oil bodies finely granular or smooth. Underleaves absent or present, undivided. Rhizoids scattered, colorless or reddish. Gametoecia on long shoots. Sporophytes surrounded by a short perigynium or marsupium (with adnate bracts) in the lower half and by an inflated perianth in the upper half, perianth deeply plicate, mouth contracted and sometimes beaked. Setae thick, of numerous rows of cells. Capsule walls 2(−4)-layered. Gemmae rare. A family of three to four genera. Solenostomataceae are morphologically very similar to Jungermanniaceae, but molecular evidence has shown that the two groups are unrelated (see Crandall-Stotler et  al. 2009). Solenostomataceae differ from Jungermanniaceae in the deeply plicate perianth (usually smooth in Jungermanniaceae) and the presence of a fleshy perigynium surrounding the lower portion of the sporophyte (perigynium lacking in Jungermanniaceae). The family is represented in Colombia and Ecuador by two genera, Solenostoma Mitt. (12 spp.) and Nardia Gray (1 sp.). Nardia was recently transferred to Gymnomitriaeae on molecular grounds (Váňa et  al. 2015), but here the genus is maintained in Solenostomataceae (following Crandall-Stotler et al. 2009) because of the greater morphological similarity of Nardia to Solenostoma than to Gymnomitriaceae. 1. Underleaves present, large, about half the length of the leaf or more, oblong to lanceolate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nardia 1. Underleaves lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenostoma Nardia Gray Apotomanthus Schiffn. Plants small- to medium-sized, 1–2.5  mm wide, green, gray-green, reddishbrown or purple, ascending to erect, little branched, stolons lacking. Branches terminal, Frullania-type, and ventral-intercalary. Stems fragile. Leaves succubous,

Solenostomataceae

339

usually somewhat spreading, concave and often clasping the stem in the lower half, undivided, ovate-orbicular, leaf apex broadly rounded to retuse, margins entire, flat or incurved, bases short-decurrent. Cells with small- or medium-sized trigones, cuticle smooth; oil bodies finely granular in neotropical species. Underleaves present, small or large, undivided, sometimes almost as long as the leaves but narrow, lanceolate to oblong. Rhizoids numerous, long, colorless. Dioicous but often fertile. Perianths plicate, usually hidden between bracts, the lower half united with the bracts and fleshy, forming a short marsupium. Capsule walls 2-layered. Gemmae lacking. About 15 species wordwide, 1 in Colombia and Ecuador. For main features of the genus, see under the species. Nardia has long been placed in the same family as Solenostoma, but molecular evidence suggests that the genus is closer to Gymnomitriaceae (Váňa et al. 2015). Literature: Engel (1988) 1. Nardia succulenta (Lehm.) Spruce Apotomanthus succulentus (Lehm.) Schiffn. Leioscyphus ecuadorensis Steph. Nardia tortistipula Spruce

Fig. 34P, Q

On moist soil and volcanic rock in shaded or exposed habitats, often near hot springs and fumaroles, or on dripping roadside rock where water is rich in minerals, 1500–3500 m. Colombia: Boy, Cal, Cas, Cau, Cun, Nar, San, Tol. Ecuador: Az, Ca, MS, Na, Pi, Tu. General distribution: Central America, West Indies, northern Andes. Nardia succulenta is a rather widespread neotropical species that forms extensive green to red to blackish carpets on volcanic rock or soil and may grow as a pioneer on fresh lava, where it enhances the formation of soil (Jongmans et  al. 2001). The species is readily recognized by the undivided, ovate-orbicular, succubous leaves, which are concave and somewhat clasping the stem, and the presence of conspicuous, lanceolate to oblong, undivided underleaves (to ca. 1/2 of leaf length). Numerous long and  ±  colorless rhizoids are usually sprouting from the ventral side of the stem. The perianth is reduced, and the lower half is fleshy and united with the bracts, forming a short marsupium. Solenostoma Mitt. Jungermannia p.p. (non typus) Plants small- to medium-sized, 0.6–4 mm wide, pale green to reddish or brown, creeping or ascending, little branched, stolons lacking. Branches usually lateralintercalary. Stems fragile. Leaves succubous, usually somewhat spreading, concave and clasping the stem in the lower half, undivided, ovate-orbicular, rarely lingulate, leaf apex broadly rounded, margins entire, not incurved, bases not or short-decurrent. Cells with usually rather small trigones, cuticle smooth or papillose; oil bodies finely granular. Underleaves lacking. Rhizoids numerous, often reddish or brown, sometimes in fascicles. Dioicous or paroicous. Perianths plicate, long-exserted or

340

 

Keys and Descriptions

short and hidden between the bracts, perianth base united with the bracts and forming a short perigynium. Perianth mouth narrowed into a short beak. Capsule walls 2-layered. Gemmae lacking. A large genus of about 120 species wordwide, most of them in Asia, 6  in Colombia and Ecuador, in two subgenera, subg. Plectocolea Mitt. and subg. Solenostoma (Söderström et al. 2016). Solenostoma has often been treated as a subgenus of Jungermannia but was reinstated based on the molecular evidence (see Váňa and Long 2009). The two groups differ in the perianth, which is plicate and with a short beak in Solenostoma (smooth and without beak in Jungermannia), and the presence or absence of a short, fleshy perigynium at the base of the perianth (present in Solenostoma, absent in Jungermannia). The presence of a perigynium in Solenostoma can be ascertained by the bracts being attached to the base of the perianth (not attached in Jungermannia). Literature: Váňa (1974 as Jungermannia) 1. Leaves wider than long. Plants dioicous, usually sterile . . . . . . . . . S. hyalinum 1. Leaves orbicular or longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants with leafless stolons at the base. (Plants very small, to 1 mm wide, growing upright on rock along rivers below 2000 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kymatocalyx dominicensis (Cephaloziellaceae) 2. Stolons lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Rhizoids colorless, in long bands appressed the ventral stem surface. Perianth cylindrical, hardly narrowed to the mouth. . . . . . . . . . . . . . . . . . . . . S. decolor 3. Rhizoids present or absent, when present colorless or reddish, not in long bands appressed to the ventral stem surface. Perianth ovoid or pyriform, distinctly narrowed to the mouth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaf cells 20–30 μm long in midleaf. Leaves orbicular. Plants paroicous, green to brown. Rhizoids colorless or pale brown. . . . . . . . . . . . . . S. sphaerocarpum 4. Leaf cells larger, (20–)30–80 μm long in midleaf. Leaves orbicular to longer than wide. Plants dioicous (or sterile), green to reddish to purple. Rhizoids often reddish in color . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaves orbicular. Plants very small, to 1  mm wide. Trigones small or lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amoenum 5. Leaves ± longer than wide. Plants larger, more than 1 mm wide. Trigones small or large. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Plants growing submerged in rivulets (above 2500  m). Leaf cells 20–40  μm long in midleaf, trigones lacking or very small. Dorsal leaf base not or scarcely decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jungermannia ovatotrigona 6. Plants not growing submerged. Leaf cells larger, 30–80  μm long in midleaf, trigones present, small or large. Dorsal leaf base distinctly decurrent . . . . . . 7 7. Leaf cells 50–80  μm long in midleaf. Trigones small, not bulging. Leaves 1.0–1.5× longer than wide .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. callithrix 7. Leaf cells 30–50 μm long in midleaf. Trigones large, bulging. Leaves ca. 1.5× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amplexifolium

Solenostomataceae

341

1. Solenostoma amoenum (Lindenb. & Gottsche) Váňa, Hentschel & Heinrichs Jungermannia amoena Lindenb. & Gottsche Plectocolea subamoena S.Winkl. (fide Gradstein et al. 2016) Common on shaded rock and earth banks along roads and rivers, 1400–2500 m. Colombia: Mag, Qui, Ris, Val. Ecuador: Ch, MS ZC (Reserva Biologica San Francisco, Holz 491 p.p., QCA). General distribution: tropical America. Solenostoma amoenum (subg. Solenostoma) is a rather widespread neotropicalmontane species that is recognized by the small, dioicous plants (to 1 mm wide) with rounded leaves and leaf cells 25–40 μm long in midleaf; the cuticle is finely papillose or smooth. 2. Solenostoma amplexifolium (Lehm.) Váňa & Schäf.-Verw. Jungermannia laetevirens Spruce Jungermannia linguifolia Gottsche Jungermannia picta Spruce

Fig. 34I, J

On soil on road banks, 1900–3700 m. Colombia: Cal, Cau, Ces, Cun, Mag, Nar, Ris. Ecuador: Ca, Pi, Tu. General distribution: scattered in tropical America. Solenostoma amplexifolium (subg. Solenostoma) is a rather widespread neotropical-montane species that is distinguished by lingulate, ca. 1.5× longer than wide leaves with a longly decurrent dorsal base, leaf cells 30–50 μm long in midleaf with large, bulging trigones, and a smooth to slightly striate-papillose cuticle. The species closely resembles S. callithrix but differs in smaller leaf cells and larger trigones. 3. Solenostoma callithrix (Lindenb. & Gottsche) Steph. Jungermannia callithrix Lindenb. & Gottsche

Fig. 34K–M

On thin soil over rock, on cut road banks and along rivers, often in large mats, 1200–3300 m. Colombia: Boy, Cal, Cun, Mag, Qui, Ris, Tol, Val. Ecuador: Ca, Lo, Pi, Tu, ZC. General distribution: tropical America. Solenostoma callithrix (subg. Plectocolea) is a common neotropical-montane species that is recognized by the ovate to ovate-oblong leaves (1–1.5× longer than wide) with a decurrent dorsal base, very long leaf cells (50–80 μm long in midleaf) with small trigones and a finely striate-papillose cuticle and by the usually reddish pigmented rhizoids. The oil bodies are 2–6 per cell in midleaf, oblong or rounded, grayish, finely granular-papillose. 4. Solenostoma decolor (Schiffn.) Váňa, Hentschel & Heinrichs Jungermannia decolor Schiffn. Nardia hyalina sensu Spruce (non typus) On wet rock in rivers, 475–3000 m. Colombia: Cau, Cun, Hui, Put, San, Tol. Ecuador: Pi. General distribution: tropical South America. Solenostoma decolor (subg. Plectocolea) is a rare neotropical species that is distinguished by the ovate leaves with somewhat undulate margins, longly decurrent and appressed bands of colorless rhizoids along the ventral stem surface, the leaf cells with conspicuous trigones and a finely papillose cuticle, and the cylindrical perianth with a wide mouth. The records from Colombia should be verified.

342

5. Solenostoma hyalinum (Lyell) Mitt. Jungermannia hyalina Lyell

 

Keys and Descriptions

Fig. 34K–M

On soil-covered shaded rock in rivers and along trails in montane environments, 1000–4000 m. Colombia: Boy, Cun, Mag. Ecuador: Az (León-Yánez s.n., QCA), Co (páramo de Llanganates, 4000  m, Burghardt 6737, PC, QCA), Lo, MS, Pi, ZC. General distribution: widespread in temperate of the Northern Hemisphere; scattered in the tropics. Solenostoma hyalinum (subg. Plectocolea) is a widespread Holarctic species that extends southward to the high Andes. The species differs from other neotropical Solenostoma species in having wider than long leaves. The plants are dioicous and usually sterile. 6. Solenostoma sphaerocarpum (Hook.) Steph. Jungermannia sphaerocarpa Hook. On moist soil and rock along roads, on river banks, and in páramo, 1800–4100 m. Colombia: Ant, Boy, Cal, Cau, Cho, Cun, Mag, Qui, Ris, Tol. Ecuador: Az (S. León-Yánez, QCA), Ca, Na, ZC. General distribution: widespread in temperate regions of the Northern Hemisphere and in tropical mountains; possibly introduced in tropical America (Gradstein and Váňa 1987). Solenostoma sphaerocarpum (subg. Solenostoma) is a widespread Holarctic species that is common along roads in the high Andes (above 2000 m) and in páramo. It is the only paroicous species in the genus Solenostoma in tropical America and is usually freely fertile. The antheridia are found just below the perianth, in the axils of bracts with a strongly swollen base. Further characters of the species include orbicular leaves, rather small leaf cells (25–35  μm in midleaf) with very small trigones, a smooth cuticle, and colorless to pale brownish rhizoids.

Stephaniellaceae Plants worm-like, grayish to pale brown, prostrate, frequently with long ventral stolons, usually unbranched, branching intercalary, rarely terminal. Stems with a thick-walled, brownish cortex of 2–5 cell layers; dorsal side of the stem with green paraphyllia. Leaf insertion dorsally interlocking, extending beyond the dorsal midline of stem. Leaves succubous, alternate, undivided, densely imbricate and usually somewhat falcate, wider than long, apex obtuse to acute to apiculate to acuminate, margins entire or toothed near the apex. Cells without chlorophyll and lacking oil bodies except in a few green cells near the base, uniformly thick-walled, elongate along the ventral margin forming a border, cuticle smooth; oil bodies (very scarce) finely granular, 3–8 per cell. Underleaves lacking. Rhizoids scattered. Gametoecia on leading shoots, gynoecia with folded bracts. Sporophytes surrounded by a rudimentary perianth or in a marsupium. Setae of numerous cells (cross section). Capsules cylindrical, wall 2-layered. Vegetative reproduction not observed.

Stephaniellaceae

343

A small family of two genera, Stephaniella and Stephaniellidium, in the high mountains of tropical America; Stephaniella also in South Africa. The genera were traditionally attributed to Gymnomitriaceae or Arnelliaceae, but these placements are now rejected and the two genera are classified in a separate family (Schuster 2002; Juárez-Martínez et al. 2016). The main features of Stephaniellaceae are (1) worm-like terrestrial plants with densely imbricate leaves; (2) stems usually with long ventral stolons and with linear to lanceolate, chlorophyllose paraphyllia on the dorsal side; (3) leaves succubous, undivided, alternate, dorsal insertion interlocking, leaf cells mostly without chlorophyll, cell walls uniformly thickened; (4) underleaves absent; (5) rhizoids scattered; and (6) capsule cylindrical. 1. Stolons present. Leaves strongly falcate, not plicate. Leaf cells mostly hyaline. Paraphyllia linear to lanceolate, numerous . . . . . . . . . . . . . . . . . . Stephaniella 1. Stolons absent. Leaves not falcate, plicate. Leaf cells ± green. Paraphyllia lanceolate, scarce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stephaniellidium Stephaniella J.B.Jack Plants small, worm-like, grayish to whitish to pale brown to reddish-brown, prostrate, forming low mats, with long ventral stolons. Stems on the dorsal side with numerous green paraphyllia covered by the leaves. Leaves asymmetrically falcate, wider than long to somewhat longer than wide, apex obtuse to acute, margins entire or finely toothed near apex. Cells without chlorophyll except near leaf base, elongate along the ventral margin forming a border. Gynoecia with folded bracts and a rudimentary perianth; marsupium lacking. An Afro-American genus of four species, occuring at elevations above 3000 m, three species in Colombia and Ecuador. Stephaniella is one of the most specialized leafy hepatics, adapted to life in extreme alpine tundra conditions. The leaves are very densely imbricate and almost without chlorophyll and are often falcate. They function as a protective shield for the green paraphyllia on the dorsal side of the stem, which have taken over the assimilatory function of the leaves. The long rootlike stolons serve to attach the plants firmly to the soil and prevent them from being blown away by the wind. The plants grow in dense, low mats on soil in open, rather dry páramo or puna habitats and can easily be overlooked because the color of the plants may be little different from that of the soil. Literature: Juárez-Martínez and Delgadillo 2017 1. Paraphyllia 1 cell wide. Leaf apex obtuse to acute . . . . . . . . . . . . . . . . . . . . . 2 1. Paraphyllia at least in part more than 1 cell wide. Leaf apex acute . . . S. uncifolia 2. Leaf apex rounded or obtuse, apical margin entire. . . . . . . . . S. paraphyllina 2. Leaf apex acute, apical margin toothed. . . . . . . . . . . . . . . . . . . . . . . S. rostrata 1. Stephaniella paraphyllina J.B.Jack

Fig. 35A–E

On soil in open or shaded high-montane environments, mostly in páramo, 2700–4500  m. Colombia: Ara, Boy, Cun, Mag. Ecuador: CA, Co, Lo, Na

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Keys and Descriptions

Fig. 35 (A–E) Stephaniella paraphyllina. (A) Habit showing stolons (s), lateral view. (B) Habit, dorsal view. (C) Leaf (p = paraphyllia, r = rhizoids). (D) Paraphyllium. (E) Midleaf cells. (F–I) Stephaniellidium sleumeri. (F) Habit with marsupium, lateral view. (G) Leaf with folds. (H) Paraphyllia. (I) Midleaf cells. (J–L) Leiomitra paraphyllina. (J) Habit. (K) Portion of stem with leaf (p  =  paraphyllium). (L) Leaf cells. (M) Leiomitra flaccida, stem leaf, 60×. (N and O) Trichocolea filicaulis, stem leaves, 60×. (P) Leiomitra sprucei, stem leaf, 60×. (M–P redrawn from Váňa 1975; rest from Gradstein et al. 2001)

Stephaniellaceae

345

(Gradstein 12795, QCA), Pi. General distribution: Mexico to northern Argentina, southeastern Brazil; also in South Africa (Drakensberg). Stephaniella paraphyllina is the most common and widespread species in the genus and is readily distinguished by the rounded or obtuse leaf apex and linear paraphyllia (only 1 cell wide). 2. Stephaniella rostrata U.Schmitt On bare soil in páramo, 3100–4300 m. Colombia: Boy, Cun, Mag. Ecuador (new): Lo (páramo de Saraguro, 3100–3200 m, Nov 2018, Benitez & Gradstein s.n., HUTPL). General distribution: Mexico and northern Andes (Venezuela to southern Ecuador) (Juárez-Martínez and Delgadillo 2017). Stephaniella rostrata is an uncommon páramo species that resembles S. uncifolia in the falcate leaves with an acute, ± toothed apex but differs in having linear paraphyllia (only 1 cell wide). 3. Stephaniella uncifolia S.Winkl. On soil in páramo, 3250  m. Colombia: Boy (Schmitt and Winkler 1968). General distribution: Venezuela to northern Argentina, southeastern Brazil (Gradstein and Costa 2003). Stephaniella uncifolia is a very rare páramo species that is close to S. rostrata but differs in having decurrent leaf bases and mostly foliose paraphyllia (more than 1 cell wide). According to Schuster (2002), the leaves in S. uncifolia may sometimes be slightly plicate. Stephaniellidium Grolle Plants small, worm-like, pale green to reddish-brown, prostrate, forming low mats, without ventral stolons. Stems on the dorsal side with a few foliose paraphyllia covered by the leaves. Leaves symmetrical to asymmetrical, not falcate, longitudinally plicate, wider than long to somewhat longer than wide, apex obtuse to apiculate, margins entire or finely toothed near apex. Cells with chlorophyll except at the apex, elongate along the ventral margin forming a border. Gynoecia with folded bracts and a short marsupium; perianth lacking. A neotropical genus with a single species, endemic to the tropical Andes. The lack of stolons, the chlorophyllose plicate leaves, and the presence of a marsupium instead of a perianth are the principal characters separating this genus from Stephaniella, which is otherwise very similar to Stephaniellidium and grows in the same general habitat. The marsupium of Stephaniellidium was first described by Winkler (1969) based on a specimen from the Sierra Nevada of Santa Marta (Winkler C315a, ULM). I have been able to examine the specimen (packet and slide) thanks to the courtesy of the curator of the herbarium ULM but found that it consisted of a mix of S. paraphyllina and a few degenerated stems of Gongylanthus cf. liebmannianus; no Stephaniellidium was detected in the material. Winkler (1969) described the marsupium of Stephaniellidium as 6–7 mm long, but according to Dr. C. JuárezMartínez (pers. comm.), it is short, ca. 1 mm long.

346

 

Keys and Descriptions

Literature: Juárez-Martínez and Delgadillo (2017) 1. Stephaniellidium sleumeri (Müll.Frib.) Grolle

Fig. 35F–I

On soil in páramo and puna, 3000–4000  m. Colombia: Ces (Winkler 1976). General distribution: Mexico, scattered in the tropical Andes (Venezuela to northern Argentina). Stephaniella sleumeri is known from many collections from Mexico but has been little collected in the Andes; it is known in the northern Andes only from the Sierra Nevada de Santa Marta and the Andes of Mérida (Juárez-Martínez and Delgadillo 2017). The species superficially resembles Stephaniella paraphyllina but differs from the latter in the plicate, subsymmetrical leaves which are mostly green (not colorless), absence of stolons, and the presence of foliose paraphyllia (more than 1 cell wide). Fertile plants are separated by the presence of a short marsupium (absent in Stephaniella).

Trichocoleaceae A family of three genera. Trichocoleaceae are recognized by (1) whitish-green, 1–3-pinnate plants with densely ciliate (hairy), (2–)4–8-lobed, succubous (rarely incubous) leaves; (2) underleaves large, bifid, densely ciliate; (3) rhizoids in tufts from underleaf bases; and (4) sporophyte in a fleshy perigynium with or without a small perianth at the top. Fulford (1963) placed the neotropical species of this family in Trichocolea Dumort., but Schuster (2000a, 2001b) showed that most of them belong in Leiomitra Spruce. The latter genus differs from Trichocolea in 1–2-pinnate branching (2–3-pinnate in Trichocolea), thick-walled stem cortex absent or 1-layered (thickwalled cortex present, 2–3-layered in Trichocolea), perianth present (absent in Trichocolea), and capsule spherical (ellipsoid in Trichocolea). Leiomitra Spruce Trichocolea p.p. (non typus) Plants small to large, whitish-green to pale brown, creeping, plants 1–2-pinnate, branches becoming as long as the main stem. Branches usually Frullania-type. Stems with a weakly differentiated cortex of thick-walled cells in 1 layer or thickwalled cortex lacking; paraphyllia present or absent. Leaves succubous, densely hairy, deeply divided into (2–)4–8 strongly ciliate segments, leaf insertion line oblique to sublongitudinal. Cells narrowly elongate, with uniformly thickened walls, cuticle striate-papillose; oil bodies small, homogeneous. Underleaves similar to the leaves but slightly smaller, transverse. Rhizoids lacking or very scarce, in bundles from underleaf bases. Gametoecia on long shoots, rarely on short ones. Sporophytes surrounded by a fleshy perigynium (coelocaule) with a short perianth

Trichocoleaceae

347

and calyptra at the tip. Setae thick. Capsules globose, wall 6–8-layered. Vegetative reproduction lacking. A genus of about ten species, five to six in Colombia and Ecuador. Literature: Fulford (1963 as Trichocolea), Schuster (2001b), Katagiri (2015a, 2016, in prep.) 1. Leaf lamina 1–3 cells high (measured from leaf base to the deepest sinus between the leaf lobes) or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf lamina (3–)4–8 cells high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Plants very small, maximally 1  mm wide. Leaves deeply 2–3-lobed (lobes densely ciliate). Plants rare . . . . . . Trichocolea filicaulis (see: Further record) 2. Plants 1–3 mm wide. Leaves deeply 4–8-lobed (lobes densely ciliate). Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaves 4–lobed, as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sprucei 3. Leaves 5–8-lobed, wider than long . . . . . . . . . . . . . . . . . . . . . . . . L. tomentosa 4. Stems without paraphyllia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. flaccida 4. Stems with paraphyllia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaves 4–6-lobed, dorsal leaf lobes much smaller than ventral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. paraphyllina 5. Leaves 6–8-lobed, dorsal leaf lobes not much smaller than ventral lobes. Plants only known from Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. robusta 1. Leiomitra flaccida Spruce Trichocolea flaccida (Spruce) J.B.Jack & Steph. Trichocolea herzogii Steph.

Fig. 35M

On bark, humic soil, and rock in montane forests and shrubby páramo, 600–4000  m. Colombia: Boy, Cau, Cho, Cun, Hui, Mag, Nar, Qui, Ris, Tol. Ecuador: Co, Na, Pi, Su, Tu, ZC. General distribution: tropical America. Leiomitra flaccida is a common and widespread neotropical species that is distinguished by the 3–6-cell-high leaf lamina (measured from the leaf base, near the middle of the leaf, to the deepest sinus between leaf segments) and the absence of stem paraphyllia. Leiomitra elliotii (Steph.) R.M.Schust. (= Trichocolea elliottii Steph.), a Caribbean species recorded a few times from Colombia (Gradstein and Uribe 2016a), is very similar to L. flaccida (see Fulford 1963) and may be a robust phenotype of the latter. Leiomitra flaccida may be confused with Lepidozia pinnaticruris, but the latter species has incubuous leaves (not succubous), less elongate leaf cells, and a smooth cuticle. 2. Leiomitra paraphyllina Spruce Trichocolea difficilis Steph. Trichocolea paraphyllina (Spruce) Steph.

Fig. 35J–L

On bark, humic soil, and rotten wood in upper montane cloud forest and páramo, 2100–4200  m. Colombia: Ant, Cau, Cho, Cun, Hui, Ris. Ecuador: Ca, Co (Burghardt 2019), Na (páramo de Oyacachi, Polylepis pauta forest, Benitez et al.

348

 

Keys and Descriptions

1302, HUTPL; ibid., Gradstein 12805, QCA), Tu (Spruce 1885). General distribution: Central America, West Indies, tropical Andes. Leiomitra paraphyllina is a characteristic, robust species of montane cloud forests that is recognized by (1) plants to 3  mm wide, irregularly 1–2-pinnate; (2) dorsal stem surface with numerous small, linear paraphyllia, the paraphyllia 5–8 cells long; (3) leaves 4–6-lobed, strongly asymmetrical, the dorsal lobes smaller than the ventral lobes. By its stem paraphyllia, this species can be readily separated from all other Leiomitra species except L. robusta; for differences with the latter species, see the key. 3. Leiomitra robusta (Steph.) R.M.Schust. Trichocolea robusta Steph. On bark, humic soil, and rotten wood in upper montane cloud forest and páramo, 2800–3600 m. Colombia: Ant, Boy, Cho, Cun, Qui, Ris. General distribution: only recorded from Colombia. Leiomitra robusta is morphologically close to L. paraphyllina; for differences see the key. 4. Leiomitra sprucei (Steph.) T.Katagiri Fig. 35P Trichocolea sprucei Steph. Trichocolea floccosa Herzog & Hatcher (fide T. Katagiri, pers. comm.) On tree trunks and rock in montane forests, (100–)1000–4000  m. Colombia: Cau, Cho, Cun, Mag, Qui, Ris. Ecuador: Az, Lo, Pi, ZC (Schäfer-Verwimp et al. 2013a). General distribution: scattered in tropical America. Leiomitra sprucei is a rather widespread neotropical species recognized by the delicate plants (ca. 1 mm wide) with leaves about as wide as long and deeply divided into mostly 4 ciliate lobes. The leaf lamina (measured from the leaf base, near the middle of the leaf, to the deepest sinus between leaf-segments) is only 1–3(−4) cells high. Leiomitra sprucei is similar to L. flaccida in leaf shape, but the leaves in the latter species are less deeply lobed and the leaf lamina is 4–6 cells high. Weak phenotypes of L. tomentosa with a very low lamina and indistinct lobes may be difficult to separate from L. sprucei. 5. Leiomitra tomentosa (Sw.) Lindb. Trichocolea tomentosa (Sw.) Gottsche Leiomitra sphagnoides Spruce Trichocolea allionii Steph. Trichocolea uleana Steph. On bark, humic soil, and rotten wood in montane forests and páramo, 600–4000 m. Colombia: Boy, Cau, Cho, Cun, Qui, Ris. Ecuador: Lo, MS, Na, Pa, Pi, Tu, ZC. General distribution: common and widespread in tropical America. Leiomitra tomentosa is the most common neotropical member of Trichocoleaceae. Characteristics are the rather robust, irregularly 1–2-pinnate plants (ca. 1.5  mm wide) with wider than long leaves (ca. 1.5–3× wider than long), 5–8 densely ciliate

Order Porellales

349

leaf lobes, and a low, (0–)1–4 cell high leaf lamina (measured from the leaf base, near the middle of the leaf, to the deepest sinus between leaf-segments). Weak phenotypes with leaves divided to near the base may be difficult to separate from L. sprucei. Further Record Trichocolea filicaulis Steph. (Fig. 35N, O) – Colombia: Cau, Cho, Cun, Hui, Mag, Nar, Qui, Ris (Gradstein and Uribe 2016a); to be expected from Ecuador. On bark in montane forests, 1300–3000 m. Trichocolea filicaulis is an uncommon neotropical species that is recognized by (1) plants very small, maximally 1 mm wide, irregularly 1–2-pinnate; (2) stem epidermis cells scarcely smaller than inner cells, multi-layered thick-walled cortex absent; and (3) leaves deeply 2–3-lobed, lamina 1–3 cells high. By the lack of a well defined thick-walled stem cortex, this species would fit in Leiomitra. However, its reproductive structures are unknown. The species is maintained in Trichocolea at the advice of Dr. T. Katagiri. Excluded Record Leiomitra argentea (Herzog) T.Katag. (Trichocolea argentea Herzog) – Colombia: Mag (Winkler 1976). This species is restricted to Brazil; the material from Colombia belongs to Lophocolea muricata (Katagiri 2015b).

Order Porellales Plants usually epiphytic. Branching lateral only, not ventral. Leaves incubous, originating from 2 or 3 initial cells; ventral lobule and sometimes a stylus present. Setae very short. Capsules usually rounded, wall 2–4(−6)-layered (mostly 2-layered). Spores usually multicellular and germination endosporic (occasionally unicellular and exosporic in Radula). Contains about 425 species in Colombia and Ecuador, in 52 genera and 5 families. Key to the families of Porellales from Colombia and Ecuador 1. Underleaves present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaves absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2. Lobule for most of its length attached to the lobe along a keel. Keel and lobule obliquely to widely spreading, away from stem. Plants variously colored but not reddish or purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeuneaceae 2. Lobule not for most its length attached to the lobe, keel lacking or short. Lobule parallel to the stem or obliquely spreading. Plants green, brown, reddish, or purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

350

 

Keys and Descriptions

3. Plants robust, 3–7 mm wide, green to brown. Underleaves undivided. Lobule plane, with rounded apex .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Porellaceae 3. Plants smaller, 0.5–2.5 mm wide, green to reddish or purple. Underleaves bifid (very rarely undivided in Frullania). Lobule sac-like or concave (occasionally plane, then with acute apex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Plants purely green, without any reddish pigmentation. Leaf margins toothed, rarely entire, apex. Lobules very small, rounded or flat, attached to the ventral margin of the lobe at some distance from the stem . . . . . . . . . . . . . Jubulaceae 4. Plants usually with reddish pigmentation, rarely purely green. Leaf margins entire, rarely toothed. Lobules rounded or elongate, usually very close to the stem (at some distance from stem in subg. Diastaloba, small reddish plants) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frullaniaceae 5. Stems rigid, epidermis of numerous rows of cells. Rhizoids in a bundle from the surface of the lobule. Lobules broadly attached to stem. Plants usually more than 1  mm wide. Oil bodies brown. Perianth narrowly cylindrical, without beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Radulaceae 5. Stems usually fragile, epidermis usually of only 5–6 rows of cells, rarely more. Rhizoids in bundles from the ventral stem surface. Lobules very narrowly attached to stem (by only 1–4 cells). Plants usually less than 1  mm wide, occasionally larger. Oil bodies colorless. Perianth shorter and broader, with a beak . . . . . . . . . . . . . . Lejeuneaceae (Cololejeunea, Myriocoleopsis)

Frullaniaceae A family with a single genus. Frullania Raddi Plants small to large, 1–20 cm long, 0.5–2.5 mm wide, reddish-green to reddishbrown or purplish or blackish, sometimes pure green, creeping, ascending or pendent, regularly or irregularly 1–3-pinnately branched. Stems rigid, made up of thick-walled cells. Branches Frullania-type, with a hemiphyll at the base; innovations normally lacking. Leaves incubous with a very short, subtransverse insertion, divided into a large dorsal lobe, a smaller ventral lobule, and a small stylus between lobule and stem. Leaf lobes usually ovate-orbicular, apex rounded to acute to acuminate, margins entire, rarely toothed. Leaf lobules almost free from the dorsal lobe, rounded to elongate, transformed into a sac or flattened, or both. Stylus usually linear and minute, rarely large, foliar. Cells isodiametrical to (more often) elongate, usually with trigones and intermediate thickenings, the trigones often confluent and cell walls often irregularly undulate-sinuose, cuticle smooth; oil bodies finely granular; ocelli sometimes present. Underleaves small or large, bifid, rarely undivided. Rhizoids in bundles from underleaf bases. Dioicous or autoicous, rarely paroicous. Androecia usually on a short, globose male branch with a few pairs of bracts and

Frullaniaceae

351

bracteoles. Gynoecia on short or elongate shoots, each with 2–5 archegonia. Perianths flattened or inflated, with 0–14 keels, the mouth contracted into a beak. Setae of numerous rows of cells. Foot of the sporophyte not penetrating into the stem. Capsules globose, wall 2-layered. Elaters attached to the capsule valves, arranged vertically inside the capsule, with 2–3 spirals. Spores large, multicellular, germination endosporic, spore surface with rosettes. Vegetative reproduction rare, by caducous or fragmenting leaves, exceptionally via discoid gemmae from the surface of leaf lobes (in F. gemmipara S.Hatt. & R.M.Schust.); reports of gemmae produced at leaf margins are usually erroneous and refer to regenerants (buds) sprouting from the leaves, a feature commonly seen in Frullania. A large, worldwide genus of at least 300 species, 54 in Colombia and Ecuador. The main features of Frullaniaceae are (1) plants usually pinnately branched, frequently reddish or purplish colored, branches Frullania-type, with a hemiphyll; (2) incubous leaves with undivided apex and entire margins (rarely toothed); (3) presence of a small ventral lobule, which is usually almost free from the dorsal lobe or connected along a keel parallel to the stem (connected along a keel oblique or perpendicular to the stem in Lejeuneaceae and Radulaceae) and usually involute, forming an inflated sac (lobule “saccate”), more rarely flat or concave (lobule “explanate” or “laminate”); (4) underleaves usually bifid, with rhizoids in a bundle at the base; (5) beaked perianths with 2–5 archegonia, innovations absent; (6) foot of the sporophyte not penetrating into the stem; (7) seta thick, of numerous rows of cells; and (8) elaters attached to the capsule valves. This treatment follows Gradstein and Uribe (2011) with updates. The species of Colombia and Ecuador are classified into six groups (cf. Hentschel et  al. 2015; Stotler and Crandall-Stotler 2017): Frullania subg. Chonanthelia Spruce (24 spp.), subg. Trachycolea Spruce (2 spp.), subg. Homotropantha Spruce (1 sp.), subg. Meteoriopsis Spruce sect. Intumescentes R.M.Schust. (17 spp.), subg. Meteoriopsis Spruce sect. Obtusilobae Verd. (1 sp.: F. kunzei), subg. Meteoriopsis sect. Meteoriopsis (6 spp.), and subg. Diastaloba Spruce (3 spp.). The latter subgenus is treated here in a traditional broad sense and consists of two unrelated clades, group Diastaloba I and group Diastaloba IV (Hentschel et  al. 2015; Söderström et  al. 2016). Group Diastaloba I contains F. subtilissima (= F. exilis, syn. nov.), the type of subg. Diastaloba Spruce, whereas the other two species in the region, F. lindenbergii and F. obcordata, belong to group Diastaloba IV. There has been some confusion about the lectotype of the genus Frullania, which was considered to be F. tamarisci (L.) Dumort. or F. dilatata (L.) Dumort. (Uribe and Gradstein 2003). Based on the latest version of the Code of Nomenclature (Turland et al. 2018), the selection of F. dilatata as the lectotype by Evans (1918) is inadmissable, and F. tamarisci must be considered the type (Stotler and CrandallStotler 2017). Since F. tamarisci is a member of subg. Thyopsiella Spruce – a group characterized by presence of ocelli (Hentschel et al. 2009) and absent in Colombia and Ecuador – it follows that the name subg. Frullania is the correct name for subg. Thyopsiella whereas the name subg. Trachycolea Spruce (type, F. dilatata) must be reinstated for the group called subg. Frullania by Hentschel et al. (2015).

352

 

Keys and Descriptions

Literature: Stotler (1969), Yuzawa (1991), Schuster (1992a), Uribe (2008), Gradstein and Uribe (2011), Hentschel et al. (2015), Atwood et al. (2018) General Key Note: characters of leaves, lobules, and underleaves should be studied on main stems, not on branches (unless mentioned otherwise). 1. Lobules distant from the stem, space between stem and lobule almost as wide as the width of the lobule. Plants small, creeping . . . . . . . . . . . . . . . . . . . . . . 2 1. Lobules very close to the stem (sometimes almost touching the stem), space between stem and lobule much narrower than the width of the lobule. Plants small or large, creeping or ascending or pendent . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaf lobes squarrose when moist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. ericoides (phenotype with explanate lobules distant from stem) 2. Leaf lobes not squarrose . . . . . . . . . . . . . . . . . . . Key 1 (subg. Diastaloba s.l.) 3. Lobules pendent, opening of the lobule positioned toward the stem or the apex of the plant. Plants of Amazonia (subg. Homotropantha). . . . . . . . F. nodulosa 3. Lobules upright, opening of the lobule positioned toward the base of the plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Saccate lobules with a short or long laminate (flattened) lower portion. Ventral surface of perianth with 2–6 keels or folds, smooth. Plants mostly monoicous, occasionally dioicous. . . . . . . . . . . . . . . . . . . . . . . Key 2 (subg. Chonanthelia) 4. Saccate lobules without laminate lower portion. Ventral surface of perianths with 0–1(−2) keels, smooth or rough by scales or tubercles. Plants mostly dioicous (monoicous in F. kunzei) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Saccate lobules 1–1.3× longer than wide. Perianth surface with scattered tubercles or scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 (subg. Trachycolea) 5. Saccate lobules 1.4–3× longer than wide. Perianth surface without tubercles or scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 (subg. Meteoriopsis) 6. Leaves squarrose when moist. Lobules mostly explanate, ca. 1/3–1/2× lobe length. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. ericoides 6. Leaves not squarrose when moist. Lobules mostly saccate, ca. 1/5–1/3× lobe length. Plants rare (Ecuador) . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. glomerata 7. Stem leaves convoluted around the stem when moist. Leaf base with two large auricles, one at the dorsal base and one at the ventral base, the auricles ± similar in size. Plants flaccid, longly pendent from branches in cloud forest and páramo, laxly branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key 3 (sect. Meteoriopsis) 7. Stem leaves not convoluted around the stem when moist, at least partially spreading. Leaf base without auricles or with a large auricle at the dorsal base and a much smaller auricle (or none) at the ventral base, not with two large auricles. Plants rigid, creeping or ascending, rarely pendent (in F. atrata and F. involuta), densely branched. . . . . . . . . . Key 4 (sect. Intumescentes and sect. Obtusilobae)

Frullaniaceae

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Key 1. Frullania subg. Diastaloba s.l. Hemiphyll: Leaf-like appendage at the ventral base of the branch (see Fig. 36C). Hemiphylls originate from an underleaf or a leaf and are undivided or bifid. 1. Leaf apex always rounded. Plants sparsely branched . . . . . . . . . . F. obcordata 1. Leaf apex at least in some leaves apiculate (especially on branches). Plants rather regularly (bi)pinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Hemiphyll bifid. Saccate lobules open (incised) in the lower half along the abaxial side. Leaf base frequently with a group of large, darker-colored cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. subtilissima 2. Hemiphyll undivided. Saccate lobules closed along the abaxial side. Leaf base without group of large, darker-colored cells . . . . . . . . . . . . . . . F. lindenbergii Key 2. Frullania subg. Chonanthelia Key adapted from Yuzawa (1991) and Atwood et al. (2018) 1. Lobules on the main stem with a long laminate (flattened) portion which extends downward beyond the ventral margin of the leaf lobe. Keel 0.35–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Lobules on the main stem with a short laminate portion, which does not extend beyond the ventral margin of the leaf lobe. Keel shorter, 0.1–0.3  mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 2. Margins of leaves and underleaves distinctly undulate. Lobules beaked (= distal end of the sac decurrent along the outer margin of the laminate part of the lobule) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Margins ± plane. Lobules with or without beak . . . . . . . . . . . . . . . . . . . . . . . 5 3. Underleaves undivided. Perianth 4-keeled. Plants from northern Peru, not yet known from Ecuador and Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. heinrichsii Gradst., Espinoza-Prieto & J.J.Atwood 3. Underleaves mostly bifid. Perianth pluripluricate (with more than 5 keels or small folds). Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . (F. obscura) 4 4. Laminate portion of the lobules entire. Plants widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. obscura var. obscura 4. Laminate portion of the lobules toothed. Plants only known from Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. obscura var. spiniloba 5. Laminate portion of the lobules toothed. Perianth always pluriplicate (with more than 5 keels or small folds). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Laminate portion of the lobules entire. Perianth 4(−6)-keeled or pluriplicate ............................................................ 7 6. Underleaf margins entire. Lobules beaked . . . . . . . . . . . . . . . . . . . . F. ringens 6. Underleaf margins toothed. Lobules without beak . . . . . . . . . . . F. blepharozia 7. Perianth 4(−6)-keeled. Lobule beaked (but beak sometimes folded, clearly visible only after careful preparation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Perianth clearly pluriplicate. Lobule with or without beak . . . . . . . . . . . . . . 11

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8. Laminate portion of lobules wider than the sac and with the outer margin strongly folded (just below the sac). Plants only known from Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. bogotensis 8. Laminate portion of lobules not wider than the sac, outer margin not strongly folded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Stylus large, more than 0.3 mm long, suborbicular. Plants rare F. confertiloba 9. Stylus small, less than 0.1 mm long, filiform. Plants very common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (F. rio-janeirensis) 10 10. Male branches short, with 3–6 pairs of bracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. rio-janeirensis var. rio-janeirensis 10. Male branches long, with 10–12 pairs of bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. rio-janeirensis var. megalostipa 11. Lobules distinctly beaked. Female bracts with short, blunt teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. tunguraguana 11. Lobules without beak. Female bracts with sharp or blunt teeth . . . . . . . . . . 12 12. Female bracts and bracteoles densey imbricate and incurved, forming a globose head. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. sphaerocephala 12. Female bracts and bracteoles spreading, not forming a globose head. Plants monoicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. ecklonii 13. Underleaves undivided (very large, orbicular). Plants of Venezuela and Bolivia, not in Colombia and Ecuador but expected there . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. holostipula S.Hattori & D.G.Griffin 13. Underleaves bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Leaf apex (sub)acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Leaf apex broadly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15. Underleaves of main stem distinctly auriculate. Appendage at the dorsal leaf base broadly rounded. Plants from the Andes of Ecuador and northern Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. haematocysta 15. Underleaves of main stem not auriculate. Appendage at the dorsal leaf base of the leaf lobe triangular, obtuse. Plants from the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. gradsteinii 16. Leaves squarrose when moist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Leaves not or hardly squarrose when moist . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Plants ca. 2 mm wide. Stylus large, rounded, leaf-like. Underleaves large, 3–4× stem width, bases auriculate. Perianth with 4 keels . . . . . . . . . . . . . . F. gibbosa 17. Plants smaller, 1–1.5 mm wide. Stylus minute, filiform. Underleaves smaller, 2–3× stem width, bases not auriculate. Perianth pluriplicate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. pluricarinata 18. Outer margin of the laminate portion of the lobule folded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. winteri var. vanderhammenii 18. Outer margin of the laminate portion of the lobule not folded . . . . . . . . . . . 19 19. Underleaves 1.5–3× wider than the stem, distant . . . . . . . . . . . . . . . . . . . . . 20 19. Underleaves larger, 3–5× wider than the stem, distant to imbricate . . . . . . . 24 20. Underleaves 2–3× longer than wide. Stylus 5–10 cells long. Plants very rare (Ecuador, ca. 3000 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. stenostipa 20. Underleaves less than 2× longer than wide. Stylus 3–6(−8) cells long. . . . . 21

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21. Lobules beaked (= distal end of the sac is decurrent along the outer margin of the laminate part of the lobule). Underleaves bifid to 1/3–1/2. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. cuencensis 21. Lobules not beaked. Underleaves less deeply bifid. Plants dioicous or autoicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Underleaves as long as wide or slightly longer than wide (maximally 1.2×). Leaves frequently somewhat squarrose, leaf lobes sometimes caducous. Perianth pluriplicate (or perianth unknown). Plants autoicous or dioicous, growing in rather dry habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22. Underleaves distinctly longer than wide (1.2–1.7×). Leaves not squarrose, leaf lobes not caducous. Perianth 4-keeled. Plants autoicous, growing in moist habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. tetraptera 23. Underleaves bifid to 1/4, slightly longer than wide, distant to contiguous. Perianth pluriplicate. Plants found in dry woodlands, roadside trees and parks, mostly in urban areas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. pluricarinata 23. Underleaves very shallowly bifid to 1/6–1/5, orbicular, distant. Perianth unknown. Plants very rare (only male plants known) . . . . . . . . . . . . F. decidua 24. Laminate portion of the lobules long, as long as the sac and reaching the ventral margin of the leaf lobe. Lobules beaked . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24. Laminate portion of the lobules much shorter than the sac, less than 0.4 mm long, not reaching the ventral margin of the leaf lobe. Lobules without beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25. Perianth 4-keeled. Stem underleaves with large auricles, the auricles approaching each other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. laxiflora 25. Perianth pluriplicate. Stem underleaves with small auricles, the auricles distant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. planifolia 26. Underleaves longer than wide, distant. Sac of the lobule inflated throughout, not flattened in the lower half. Plants dioicous, very rare (one nineteenth-century record from Colombia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. jelskii 26. Underleaves about as long as wide, imbricate. Sac of the lobule inflated in the upper half, flattened below. Plants autoicous, widespread . . . . . . . . . . . . . . . 27 27. Stylus minute, 3–5 cells long, filiform, 1 cell wide throughout. Plants growing in montane forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. dusenii 27. Stylus larger, more than 5 cells long, foliose in the lower half, the foliose part 2–8 cells wide. Plants growing in montane forest and páramo . . F. standaertii Key 3. Frullania subg. Meteoriopsis sect. Meteoriopsis Key adapted from Uribe (2008) 1. Leaf margins toothed. Plants rare (Colombia, Jamaica) . . . . . . . . F. dulimensis 1. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaf apex rounded to obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaf apex acute. apiculate to acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Underleaves ovate-oblong, 1.4–2 × longer than wide, not overlapping. Female bracts entire. Perianth distinctly exserted beyond the bracts. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. convoluta

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Keys and Descriptions

3. Underleaves suborbicular, 1–1.3  ×  longer than wide, overlapping. Female bracts toothed. Perianth not exserted beyond the bracts. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. grandifolia 4. Underleaf bases without conspicuous auricles. Female bracts toothed. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. peruviana 4. Underleaf bases with conspicuous auricles. Female bracts entire or weakly toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Underleaf margins recurved. Underleaf auricles large, more than 0.15 mm long. Median leaf cells longer than wide. Female bracts weakly toothed. Plants rare (high Andes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. phalangiflora 5. Underleaf margins plane. Underleaf auricles smaller, less than 0.1 mm long. Median leaf cells isodiametrical. Female bracts entire. Plants from the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. darwinii Key 4. Frullania subg. Meteoriopsis sect. Intumescentes and sect. Obtusilobae Note: Frullania kunzei belongs in sect. Obtusilobae, all other species in sect. Intumescentes (Söderström et al. 2016). Plants without perianths cannot always be identified with certainty. Hemiphyll: Leaf-like appendage at the ventral base of the branch (see Fig. 36C). Hemiphylls originate from an underleaf or a leaf and are undivided or bifid. Most species of sect. Intumescentes have an undivided hemiphyll. 1. Leaf apex very longly and finely acuminate, the acuminate portion to 1  mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. aculeata 1. Leaf apex not acuminate or less longly acuminate . . . . . . . . . . . . . . . . . . . . . 2 2. Dorsal margin of leaves toothed. Plants usually growing pendent . . . F. uleana 2. Leaf margins entire. Plants not pendent (except F. atrata, couplet 17) . . . . . . 3 3. Lobules on main stems ca. 1.5× longer than wide, upright or at an oblique angle with the stem. Plants small, 0.5–1.0 mm wide, prostrate, autoicous or dioicous. Subapical cells of leaf lobe ± isodiametrical, cell walls with trigones but without intermediate thickenings. Underleaves 1–2× stem width . . . . . . . F. kunzei 3. Lobules on main stems more elongate, 1.8–3× longer than wide, upright. Plants larger, prostrate, ascending or pendent, dioicous. Subapical cells of leaf lobe longer than wide, cell walls with trigones and intermediate thickenings (the intermediate thickenings sometimes confluent with the trigones). Underleaves usually more than 2× stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Ventral leaf base with 1–2 long and narrow appendages, one hanging down and the other standing up, parallel to the stem. . . . . . . . . . . . . . . . . . . . F. mirabilis 4. Ventral leaf base without long appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaf apex at least in part apiculate to acuminate . . . . . . . . . . . . . . . . . . . . . . . 6 5. Leaf apex mostly rounded to obtuse (rarely a few leaves subapiculate) . . . . 18 6. Underleaves reniform (much wider than long). Underleaf apex broadly truncate and with 2 small, subulate teeth . . . . . . . . . . . . . . . . . . . . . . . . F. bicornistipula

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6. Underleaves rounded, ovate to oblong, not reniform. Underleaf apex not broadly truncate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Underleaf apices finely acuminate. Hemiphyll longly acuminate, undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. setigera 7. Underleaf apices not finely acuminate. Hemiphyll rounded to acute, bifid or undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Underleaves with very long auricles, the auricles about ½ the length of the underleaf lamina (lamina measured from the middle of the insertion to the base of the leaf lobes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. lobatohastata 8. Underleaves without or with shorter auricles . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Leaf apex distinctly recurved in all leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Leaf apex plane, or slightly recurved in a few leaves . . . . . . . . . . . . . . . . . . 13 10. Stylus filiform, 1 cell wide (sometimes with a small appendage at the base in F. intumescens). Perianth (sub)terete or longly 3-keeled . . . . . . . . . . . . . . . . 11 10. Stylus not filiform, more than 1 cell wide at least in the lower half. Perianth subterete (slightly plicate only at the apex). . . . . . . . . . . . . . . . . . F. ambronnii 11. Perianth terete or with 3 very short, rounded keels at apex. Stylus without appendage. Plants of lowlands and montane, very common. . . . F. brasiliensis 11. Perianth sharply 3-keeled over most of its length. Stylus with or without foliose appendage at the base. Plants growing above 1000 m. . . . (F. intumescens) 12 12. Stylus with a foliose appendage at the base. Plants uncommon in Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. intumescens var. intumescens 12. Stylus without foliose appendage at the base. Plants common in Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. intumescens var. sabanetica 13. Underleaf margins strongly recurved. Leaf apex apiculate to acuminate . . . 14 13. Underleaf margins plane or slightly and very narrowly recurved. Leaf apex acute to shortly apiculate [when in doubt try both leads] . . . . . . . . . . . . . . . 15 14. Leaf apex distinctly acuminate, ending in a 1–2-cell-wide row of up to 10 cells. Female bracts and bracteoles toothed. Perianth 3-keeled . . . . . . . . . . F. pittieri 14. Leaf apex apiculate to shortly acuminate, ending in a 1–2-cell-wide row of 1–4 cells. Female bracts and bracteoles entire. Perianth subterete, slightly plicate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. macrocephala 15. Underleaves more than 3× as wide as the stem . . . . . . . . . . . . . . . . . . . . . . . 16 15. Underleaves (1.5–)2–3× as wide as the stem . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Underleaves longer than wide, distant. . . . . . . . . . . . . . . . . . . . . . F. meridana 16. Underleaves rounded or wider than long, ± imbricate . . . . . . . . . . . F. formosa 17. Plants less than 1 mm wide, dark reddish-brown to black, often pendent. All or some underleaves with slightly recurved margins. Leaf lobes 2–3(−4)× longer than the lobules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. atrata 17. Plants more robust, 1–2 mm wide, greenish-brown, stems ascending from the substrate, not pendent. Underleaf margins plane. Leaf lobes always more than 3× longer than the lobules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. beyrichiana 18. Underleaves with very long auricles, auricles to about ½ the length of the underleaf lamina (lamina measured from the middle of the insertion to the base of the leaf lobes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. lobatohastata

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Keys and Descriptions

1 8. Underleaves without or with short auricles . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19. Underleaves more than 3× as wide as the stem. Perianth terete . F. clandestina 19. Underleaves 1.5–3× as wide as the stem. Perianth 3-keeled. . . . . . . . . . . . . 20 20. Hemiphyll undivided. Underleaves 2–3× stem width, margins recurved. Margins of female bracteoles usually entire, rarely toothed . . . . F. montagnei 20. Hemiphyll bifid (to 1/4–1/2). Underleaves smaller, 1.5–2× stem width, margins plane to narrowly recurved. Margins of female bracteoles ciliate-laciniate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. paradoxa 1. Frullania aculeata Taylor Frullania acuminata Steph.

Fig. 36A, B

On bark, soil, and rock in moist, open evergreen woodlands, scrub and isolated trees in meadows and plantations, also in the canopy and at the margin of humid rainforests, 300–3000  m; widespread on the Galápagos Islands between 400 and 1000 m (Gradstein 2009). Ecuador: Az, Ch, Co, EO, Ga, Lo, Pi, LR; to be expected in Colombia. General distribution: Ecuador, Panama; further records need verification. Frullania aculeata (sect. Intumescentes) is a common Ecuadorian species that is immediately recognized by the very long and finely acuminate leaf tips, to 1 mm long (Uribe and Gradstein 2003). By the characteristic leaf shape, this species cannot be confused with any other member of the genus Frullania. Frullania aculeata is very abundant in the Galápagos Islands; in addition, the species has recently been recorded from Panama (Schäfer-Verwimp 2018). The frequently cited record from Peru (Stephani 1898–1924, type of F. acuminata; Menzel 1984) is erroneous; the “Peruvian” locality, Tambo de la Chonta, is in Ecuador (prov. El Oro). 2. Frullania ambronnii Steph. Frullania harpantha Herzog (fide Hentschel et al. 2015) On bark in montane rainforest, 2200–2400. Colombia: Cun (Tequendama, type of F. harpantha). General distribution: tropical Andes (Colombia, Bolivia) Frullania ambronnii Steph. (sect. Intumescentes) is a little-known Andean species that is only known from Bolivia (type) and the type of F. harpantha (synonym) from Colombia. The outstanding character of the species is the broad, foliose stylus, which is more than 1 cell wide over its whole length or in the lower half. A foliose stylus is not seen in any other species of sect. Intumescentes in the northern Andes (Stotler 1969). Except for the broad stylus, F. ambronnii is simlar to F. brasiliensis (e.g., in the recurved, apiculate leaf apices, the recurved underleaf margins, and the subterete perianth, which is weakly plicate near the apex only) and might be a variety of the latter. The relationship of the two taxa needs further study. The Colombian specimen of F. ambronnii (type of F. harpantha) stands out by the claw-shaped apices of the (toothed) female bracts and bracteoles. 3. Frullania atrata (Sw.) Mont. Frullania crinoidea Steph. Frullania lechleri Steph. Frullania subaculeata Spruce

Fig. 36C, D

Frullaniaceae

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Fig. 36 (A and B) Frullania aculeata. (A) Leaf. (B) Habit. (C and D) Frullania atrata. (C) Habit, showing a small, lanceolate hemiphyll (undivided) at the base of the branch. (D) Leaf. (E) Frullania beyrichiana, habit. (F) Frullania bicornistipula, habit. (G and H) Frullania bogotensis. (G) Leaf. (H) Habit. (I and J) Frullania brasiliensis. (I) Habit. (J) Gynoecium with perianth. (K and L) Frullania obcordata. (K) Habit. (L) Leaf. (M) Frullania cladestina, habit. (A–D from Uribe and Gradstein 2003; E, F, M redrawn from Stotler 1969; G, H redrawn from Yuzawa 1991; I, J from Gradstein and Uribe 2011; K, L from Gradstein and Ilkiu-Borges 2009). Scale bars: A, E–J, M = 0.5 mm; B = 1.25 mm; C = 400 μm; D, L = 100 μm; K = 250 μm

360

 

Keys and Descriptions

On bark of trees and shrubs in montane rainforests and lower páramo, usually in partially exposed sites, 600–3200(−4000) m. Colombia: Ant, Boy, Cas, Cau, Ces, Cun, Hui, Mag, Met, San, Tol. Ecuador: Az, Co, EO, Lo, MS, Tu, ZC. General distribution: tropical America. Frullania atrata (sect. Intumescentes) is a rather widespread neotropical species that is distinguished by the very slender habit (less than 1 mm wide), very dark, almost blackish plant color, small, sharply apiculate leaves, about 0.5–0.6 mm long and only little longer than wide, with flat apices, and underleaves less than 3× wider than the stem and with narrowly recurved or almost plane margins. The plant often grows pendent on branches of trees and shrubs. Frullania atrata has frequently been confused with members of sect. Meteoriopsis, especially with F. peruviana (e.g., by Spruce 1884), but the latter species are usually more elongate and more robust (more than 1 mm wide when leaves are flattened) and possess leaves with two large auricles. Frullania atrata, in contrast, has only one large auricle, at the dorsal leaf base; a much smaller auricle (or none at all) is present at the ventral base. The published records of F. atrata from Colombia and Ecuador, especially the older records, must be verified; some may be erroneous and belong to F. peruviana or other members of the sect. Meteoriopsis. Frullania atrata is morphologically close to F. uleana, but the latter species has toothed leaf margins. Frullania atrata may also be confused with other members of sect. Intumescentes, especially F. beyrichiana. For differences see under the latter species. 4. Frullania beyrichiana (Lehm. & Lindenb.) Lehm. & Lindenb. Frullania beyrichii Spruce Frullania breuteliana Gottsche (?) Frullania coalita Steph.

Fig. 36E

On bark of trees or terrestrial, 1800–2400 m. Colombia: Ant, Cun. Ecuador (new): Bo (East of Echeandía, 2400 m, Holm-Nielsen 18618, AAU, PC). General distribution: scattered in tropical America, mainly in Brazil. Frullania beyrichiana (sect. Intumescentes) is a rare species in the northern Andes. Characteristics are the flat leaves with apiculate apex and the flat, distant, orbicular to broadly ovate underleaves which are maximally 3× stem width. Frullania beyrichiana approaches F. formosa, but the underleaves in the latter species are much larger and usually imbricate. Frullania breuteliana, which has been reported from Colombia, is very similar to F. beyrichiana (Stotler 1969). Slight differences are seen in the leaf tip, which is shortly acuminate in F. breuteliana (acute or apiculate in F. beyrichiana), and the female bracts and bracteole, which are toothed in F. breuteliana (entire in F. beyrichiana). Possibly, these two species are conspecific (Gradstein and Uribe 2011). This needs study. 5. Frullania bicornistipula Spruce

Fig. 36F

On bark of trees and soil in montane rainforest, 1250–2300 m. Colombia: Cau, Hui, NSa, Ris. Ecuador: Ch, Pa, Pi, Tu, ZC.  General distribution: northern Andes, Guayana Highland, Costa Rica.

Frullaniaceae

361

Frullania bicornistipula (sect. Intumescentes) is a characteristic northern Andean-Costa Rican species that is readily distinguished by the longly mucronate leaf apex and the reniform, undivided underleaves with a broad truncate apex with 2 subulate teeth. The longly filiform stylus, 6–12 cells long, is another characteristic feature of F. bicornistipula. The species resembles F. pittieri in the longly mucronate leaf apex but is readily separated from the latter by the reniform underleaves. 6. Frullania blepharozia Spruce On bark, 1800–3300  m. Ecuador: Ch, Pi, ZC.  General distribution: only known from Ecuador. Frullania blepharozia (subg. Chonanthelia) is a rare Ecuadorian endemic near to F. ecklonii that is characterized by toothed underleaf margins, lobules with a very small sac and a large laminate portion and no beak, and pluriplicate perianths (Yuzawa 1991). 7. Frullania bogotensis Steph.

Fig. 36G, H

On logs, 700–2000 m. Colombia: Boy, Cun, San, Tol. General distribution: only known from Colombia. Frullania bogotensis (subg. Chonanthelia) is closely similar to F. rio-janeirensis but differs in the broader laminate portion of the lobule, which is broader than the sac (narrower than the sac in F. rio-janeirensis) and has a strongly folded margin (Yuzawa 1991). 8. Frullania brasiliensis Raddi Fig. 36I, J Frullania columbica Steph. Frullania cylindrica Steph. Frullania granatensis Gottsche (fide J. Uribe, in sched.) Frullania guadalupensis Steph. Type: Guadeloupe, l’Herminier s.n. (lectotype G-00120036!; isotype PC-0102248!) Frullania mucronata (Lehm. & Lindenb.) Lehm. & Lindenb. Frullania robusta Steph. Frullania spruceana Steph. Frullania tenuis Lehm. On bark of trees, logs, soil, and rock in lowland and montane rainforests; semideciduous and evergreen woodland, scrub, savannas, and meadows; also in the canopy of lowland rainforest and savanna but uncommon in lowlands, 100–3900 m; very common on the Galápagos Islands between 100 and 1200 m. Colombia: Ant, Boy, Cal, Cau, Ces, Cho, Cun, Hui, Mag, Met, Nar, Qui, Ris, San, Tol, Vau. Ecuador: Az, Co, EO, Ga, Lo, LR, MS, Na, Pa, Pi, Tu, ZC. General distribution: common and widespread in tropical America. Frullania brasiliensis (sect. Intumescentes) is the most common neotropical species of sect. Intumescentes. It is recognized by recurved apiculate leaf apices; a short, filiform stylus; underleaves bifid to 1/5–1/3; recurved underleaf margins; entire or toothed female bracts and bracteoles; and a terete perianth (sometimes weakly plicate near the apex). Frullania brasiliensis is quite similar to F.

362

 

Keys and Descriptions

intumescens, but the latter species has a sharply trigonous perianth and produces a small, foliose appendage at the base of the stylus in var. intumescens. Moreover, F. intumescens is restricted to montane forests, above 1000 m. Sterile or male plants of the two species without stylar appendage cannot be distinguished morphologically based on current knowledge. 9. Frullania clandestina (Nees & Mont.) Nees

Fig. 36M

On bark and rock, 1400–4500 m. Colombia: Cau (Stotler 1969). Ecuador: Pa (Stotler 1969), Tu (road Baños – Puyo, 1400 m, Holm-Nielsen 16317, AAU, PC). General distribution: tropical Andes (Colombia to Peru), Costa Rica. Frullania clandestina (sect. Intumescentes) is a rare Andean-Costa Rican species from high elevations. The species is characterized by the recurved and rounded (not apiculate) leaf apex; the large underleaves, more than 3× stem width and with flat or somewhat recurved underleaf margins; and the terete perianths (but perianth rare; seen in material from Peru: Sylvester s.n., PC). In its large underleaves, F. clandestina resembles F. formosa, but the leaf apex in the latter species is apiculate and the perianth 3-keeled. Frullania clandestina is most closely related to F. brasiliensis; both species share recurved leaf apices, large underleaves with somewhat recurved margins, and terete perianths. They only differ in the leaf apex, which is apiculate in F. brasiliensis and rounded in F. clandestina. The relationship of the two species should be analyzed with molecular methods; possibly F. clandestina is a variety of F. brasiliensis. Frullania clandestina was erroneously cited as a synonym of F. intumescens in León-Yánez et al. (2006). 10. Frullania confertiloba Steph. Frullania bomboizana Steph. On bark of trees and logs, 700–1700 m. Colombia: Boy, San. Ecuador: MS (Yuzawa 1991; type of F. bomboizana). General distribution: scattered in tropical America. Frullania confertiloba (subg. Chonanthelia) is a rare neotropical species that is closely similar to F. rio-janeirensis but differs in the large, rounded stylus (Yuzawa 1991). The species is known from only few localities and may have been overlooked. 11. Frullania convoluta Lindenb. & Hampe Frullania longipinna Steph.

Fig. 37A, B

Pendent from branches of trees and shrubs in upper montane cloud forest, 2000–3600 m, occurring at much lower elevation (ca. 1100 m) on the summit of volcán Alcedo on the Galápagos Islands. Colombia: Boy, Cho, Cun, Met, Nar, NSa, Put, Qui, Ris, San, Tol. Ecuador: Ca, Co, Ga, Na. General distribution: Central America, Dominican Republic, tropical Andes, Galápagos Islands, Guayana Highland. Frullania convoluta (sect. Meteoriopsis) is a common species of neotropical upper montane cloud forests that is readily distinguished by the long-pendent plants with narrowly rounded leaf apices and distant, ovate-oblong underleaves (Uribe

Fig. 37 (A and B) Frullania convoluta. (A) Habit. (B) Leaf. (C and D) Frullania cuencensis. (C) Leaf lobule. (D) Habit. (E) Frullania darwinii, habit. (F and G) Frullania decidua. (F) Leaf. (G) Habit. (H) Frullania dulimensis, habit. (I and J) Frullania dusenii. (I) Habit. (J) Leaf. (K) Frullania kunzei, habit (type of F. ecuadorensis). (L and M) Frullania ericoides. (L) Leaf. (M) Habit with gynoecium. (N and O) Frullania subtilissima. (N) Leaf. (O) Habit. (P and Q) Frullania gibbosa. (P) Habit with gynoecium. (Q) Leaf. (A, B, E, H redrawn from Uribe 2008; C, D, F, G, P, Q redrawn from Yuzawa 1991; K redrawn from Stotler 1969; rest from Gradstein et  al. 2001; Gradstein and Ilkiu-Borges 2009). Scale bars: A, B, I = 1 mm; C, F, L, O, Q = 250 μm; D, G, H, J, K, M, P = 0.5 mm; E = 1.25 mm; N = 100 μm

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Keys and Descriptions

2008). The species is closely similar to F. grandifolia; for differences see the key. Frullania convoluta has sometimes been confused with F. peruviana, but the latter species has an acute leaf apex. The shape of the leaf apex should be studied in flattened leaves. 12. Frullania cuencensis Taylor

Fig. 37C, D

On bark and logs in montane cloud forests and páramo, 1800–4100 m. Colombia: Cun, San. Ecuador: Az, Ch, Na, Pi, ZC. General distribution: tropical America. Frullania cuencensis (subg. Chonanthelia) is a common neotropical-montane species that is distinguished by (1) lobules beaked and with a short laminate portion; (2) underleaves bifid to 1/3–1/2; and (3) dioicous sexuality. The species has been confused with F. tetraptera, but the lobule in the latter species has no beak, the underleaves are more shallowly bifid (1/6–1/4) and somewhat longer than wide, and the plants are autoicous. According to A. Schäfer-Verwimp (in litt.), F. tetraptera is smaller than F. cuencensis and grows in more open, drier habitats. 13. Frullania darwinii Gradst. & Uribe

Fig. 37E

Pendent on trunks, branches, and rock, 250–650 m. Ecuador: Galápagos Islands (Floreana, Rábida, San Cristóbal, Santa Cruz). General distribution: only known from the Galápagos Islands where it is rather common. Frullania darwinii (sect. Meteoriopsis) is a striking Galápagos endemic recognized by longly pendent growth, pointed leaf apex, and auriculate underleaves with plane margins (Uribe 2008). The species may be confused with F. peruviana and F. phalangiflora; F. peruviana differs in strongly toothed female bracts (± entire in F. darwinii) and underleaf bases without auricules, while F. phalangiflora differs by recurved underleaf margins and globose perianths (± cylindrical in F. darwinii). 14. Frullania decidua Spruce Fig. 37F Type: Ecuador, Chimborazo, “Lucmas in valle Alausi,” Spruce s.n. (MANCH!) Frullania flammea Spruce, syn. nov. Type: Ecuador, Azuay, Cuenca, on trees, Jameson s.n., (MANCH!) On bark or rock, 2350–3800 m. Colombia: Boy (Yuzawa 1991). Ecuador: Az, Ch. General distribution: tropical Andes, southeastern Brazil. Frullania decidua (subg. Chonanthelia) is small Andean species distinguished by plane to slightly squarrose and somewhat caducous leaf lobes, lobules with a short laminate portion and without beak, and the small, orbicular underleaves that are very shallowly bifid (to 1/6). The species is known only from male (or sterile) plants. Frullania decidua closely resembles F. pluricarinata, but the latter species has more deeply bifid underleaves (to 1/4) that are often somewhat longer than wide and frequently have a tooth on the outer margins. Gradstein and Uribe (2016a) considered F. decidua a synonym of F. cuencensis, but the lack of a lobule beak and the very shallowly bifid underleaves of F. decidua separate this species from F. cuencensis.

Frullaniaceae

15. Frullania dulimensis Uribe

365

Fig. 37H

Pendent on isolated trees in open, upper montane cloud forest, with F. peruviana and F. convoluta, 3200 m. Colombia: Tol (Uribe 2006). General distribution: Colombia (type), Jamaica (Schäfer-Verwimp and van Melick 2016). Frullania dulimensis (sect. Meteoriopsis) is a rare neotropical species that is readily separated from other members of sect. Meteoriopsis by toothed leaves and underleaves. By its toothed leaves, the species resembles F. uleana, but the latter species differs in leaves with only one large auricle (at the dorsal leaf base), not two large auricles, and entire underleaves. Frullania dulimensis is known from two populations, a male population in Colombia and a female population in Jamaica. 16. Frullania dusenii Steph. Frullania transfalcicula L.Clark

Fig. 37I, J

On bark and rock in montane forest environments, 1700–3500  m. Colombia: Ces, Cun, Mag. Ecuador: Az, Ch, Co, Na, Pi, ZC. General distribution: tropical America. Frullania dusenii (subg. Chonanthelia) is a neotropical-montane species recognized by (1) lobules with a very short lamina, lobule sac with a constricted surface (= folded in the middle), lobule mouth somewhat narrowed (constricted); (2) stylus minute, filiform, 3–4 cells long; and (3) underleaves rather large, 3–4× wider than the stem, imbricate. The species is monoicous and frequently fertile. Frullania dusenii may be confused with F. standaertii, but the latter species has a larger, (partially) foliose stylus. 17. Frullania ecklonii (Spreng.) Gottsche, Lindenb. & Nees Frullania vimicola Spruce On bark, rock, and soil in montane forest areas, (500–)1800–3500  m; on the mainland above 1800  m but on the Galápagos Islands found at 500  m in moist Scalesia woodland. Colombia: Boy, Cun, Tol. Ecuador: Az, Ca, Ga, Pi, ZC. General distribution: pantropical. Frullania ecklonii (subg. Chonanthelia) is a widespread pantropical species that is distinguished by the pluriplicate perianth, the non-undulate margins of leaves and underleaves, the longly laminate portion of the lobules, the underleaves with large auricles, and, especially, the lack of a lobule beak. The species has sometimes been treated as a synonym of F. obscura (= F. arecae), but in the latter species, the leaf and underleaf margins are undulate, and the lobule is beaked (Yuzawa 1991). 18. Frullania ericoides (Nees) Mont. Fig. 37L, M Frullania allionii Steph., syn. nov. Type: Ecuador, Morona Santiago, Gualaquiza, “in arbore silvae supra rus. antiq. ‘hacienda Vega’,” 900 m, 13 Feb 1909, Allioni s.n., Bryotheca Levier 6443 (lectotype, fide Bonner (1965), G-00066795!, Icon. Ined. nr. 3403). Syntypes: ibid., “forêt Liuriapa, saxicola,” 900 m, 20 Aug 1909, Allioni s.n., Bryotheca Levier 6444 (G-00128070!); ibid., “ad truncos in silva Guayusa,” 20 Aug 1909, Allioni s.n., Bryotheca Levier 6565 (G-00128071!); ibid., Bomboiza, “ad saxum erraticum vulgo ‘La Mesa’

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Keys and Descriptions

prope Missionariorum domum,” 900  m, 23 Feb 1910, Allioni 281 (?) (G-00128072!) Frullania ocanniensis Steph., syn. nov. Type: Colombia, Antioquía, páramo de Ocaña, Pehlke s.n., “misit Irmscher nr. 12” (G-00069231!) Frullania squarrosa (Reinw., Blume & Nees) Dumort. On bark and rock in rather dry, open woodlands, savannas, and meadows, from sea level to 2200 m; widespread and common in the Galápagos Islands. Colombia: Ant, Boy, Cas, Cho, Cun, Qui, Ris, San, Tol. Ecuador: Ga, Lo, MS, Pa, Pi, Tu, ZC. Distribution: pantropical. The pantropical F. ericoides (subg. Frullania) is one of the most drought-tolerant epiphytic liverwort species and one of the very few liverworts occurring in arid coastal areas on the Galápagos Islands (Gradstein 2009). The species is readily distinguished by the brittle and more or less squarrose leaves. The plants are dioicous and rather small, 0.8–1.5  mm wide; the lobules are usually explanate and acuteelongate, rarely saccate (helmet-shaped); the stylus is very small lanceolate; the underleaves are imbricate, ca. 3× stem width, shortly bifid and frequently with 1 or a few blunt teeth on the outer margins; and the perianth is 3(−4)-keeled and covered by small scales, tubercles, or papillae. The explanate lobules are usually at some distance from the stem, like in members of subg. Diastaloba s.l. Frullania ericoides may be confused with F. gibbosa and F. glomerata. Frullania gibbosa (subg. Chonanthelia) shares with F. ericoides the squarrose leaves and occurrence in rather dry, open habitats but differs from the latter species in being autoicous, more robust (more than 2 mm wide), leaves not brittle, lobules usually saccate and with a short laminate portion, stylus large, leaf-like, and perianth surface smooth, 4-keeled. Frullania glomerata (subg. Frullania), which has been recorded from the surroundings of Baños, differs in plane leaves and lobules smaller and mostly saccate. Moreover, F. glomerata grows in more humid locations. The type material of F. ocanniensis is a mixture of F. ericoides and F. rio-janeirensis. Based on the protologue (description, Icones), the F. ericoides part is the type of F. ocanniensis. The record of F. ericoides from high elevation in Quito (SchäferVerwimp et al. 2013a) belongs to F. pluricarinata. 19. Frullania formosa Spruce On branches of trees in submontane rainforest and light scrub, 500–1400  m. Colombia: Ris (Gradstein and Uribe 2016a). Ecuador: MS, Na, Pa, Tu. General distribution: northern Andes (Colombia, Ecuador). Frullania formosa (sect. Intumescentes) is a rare northern Andean species that is distinguished by the flat, shortly apiculate leaf apex; the large, orbicular or wider than long, ± imbricate underleaves, which are more than 3× as wide as the stem; and the broadly 3-keeled perianth (Stotler 1969). By the large underleaves, F. formosa is similar to F. clandestina and F. meridana, but F. clandestina differs in the rounded leaf apex (not apiculate) and F. meridana in the longer than wide, distant underleaves. 20. Frullania gibbosa Nees

Fig. 37P, Q

Frullaniaceae

367

Frullania tolimana Steph. On bark and rock in rather dry, open places in lowland and montane areas, from sea level to 2300 m. Colombia: Ant, Boy, Cal, Cas, Cau, Ces, Cun, Mag, Ris, San, Tol. Ecuador: Ga. General distribution: widespread in tropical America. The unique characteristic of the neotropical F. gibbosa (subg. Chonanthelia) is the large, rounded, leaf-like stylus. The stylus is usually covered by the large underleaves and can only be observed by removal of the underleaves. Frullania gibbosa is a common species from rather open locations at low elevations and often grows together with F. ericoides. The two species can be confusingly similar; for differences between see under F. ericoides. Frullania gibbosa is common in Colombia but seems in Ecuador to be only known from the Galápagos Islands. 21. Frullania glomerata (Lehm. & Lindenb.) Nees & Mont. Frullania hirtiflora Spruce, syn. nov. Type: Ecuador, Baños, at the foot of Mt. Tungurahua, 1800 m, on trunk bases and rock, 1857, Spruce s.n. (MANCH!) Frullania squamulifera Spruce, syn. nov. Type: Ecuador, Tungurahua, Agoyán, 1857, Spruce s.n. (MANCH!) On bark and wet rock in lower montane forest areas, ca. 1500–1800 m. Ecuador: Tu (Spruce 1884 as F. squamulifera and F. hirtiflora). General distribution: scattered in tropical America. Frullania glomerata (subg. Frullania) is a rather widespread species that is surprisingly rare in the northern Andes where it has only been recorded from Baños, collected by Richard Spruce in 1857 and described as F. squamulifera and F. hirtiflora. The species resembles F. ericoides in the helmet-shaped lobules, dioicous sexuality, and 3-keeled perianths covered by small scales or hairly papillae but differs from the latter in rather plane leaves (not squarrose) and lobules mostly saccate, 3–5× shorter than leaf lobes (2–3× shorter than leaf lobes in F. ericoides). Moreover, F. glomerata seems to grow in moister habitats than F. ericoides. 22. Frullania gradsteinii Yuzawa, Mues & S.Hatt. Frullania mesophila Gradst., nom. inval.

Fig. 38A

Forming ascending tufts on bark in rather dry, semi-deciduous woodlands of the “transition zone” on the Galápagos Islands, 50–200 m. Ecuador: Ga (San Cristóbal, Santa Cruz). General distribution: only known from the Galápagos Islands. Frullania gradsteinii (subg. Chonanthelia), endemic to the Galápagos Islands, is one of the few species of subg. Chonanthelia with acute leaf lobes. The species is closely related to F. haematocysta from mainland Ecuador and northern Peru but differs from the latter in the absence of auricles at underleaf bases and the triangular-obtuse appendage at the dorsal leaf base. 23. Frullania grandifolia Steph.

Fig. 38B, C

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Keys and Descriptions

Fig. 38 (A) Frullania gradsteinii, habit. (B and C) Frullania grandifolia. (B) Leaf. (C) Habit. (D) Frullania involuta, habit. (E and F) Frullania kunzei. (E) Leaf. (F) Habit. (G) Frullania laxiflora, leaf. (H and I) Frullania lobatohastata. (H) Underleaf. (I) Habit. (J and K) Frullania macrocephala. (J) Gynoecium. (K) Habit. (L) Frullania meridana, habit. (M and N) Frullania nodulosa. (M) Leaf. (N) Habit. (O and P) Frullania obscura. (O) Lobule. (P) Habit. (A, G redrawn from Yuzawa 1991; B, C, L, O, P from Gradstein and Uribe 2011; D from Uribe and Gradstein 2003; H, I, J–K redrawn from Stotler 1969; rest from Gradstein et  al. 2001; Gradstein and Ilkiu-Borges 2009). Scale bars: A, G–J, M, N = 0.5 mm; B–D, K, L = 1 mm; E = 100 μm; F, O = 250 μm; P = 0.8 mm

Frullaniaceae

369

On bark in upper montane forest, 3400  m. Colombia: Tol (Uribe 2008). Ecuador: Co. General distribution: Mexico, Costa Rica, northern Andes (Colombia, Ecuador). Frullania grandifolia (sect. Meteoriopsis) is a rare high-montane species that is recognized by pendent growth, a rounded leaf apex, imbricate suborbicular underleaves, and toothed female bracts. The species is closely related to F. convoluta, but the underleaves in the latter species are not imbricate and distinctly longer than wide, and the margins of the female bracts are entire (Uribe 2008). 24. Frullania haematocysta Spruce On trees, ca. 1800 m. Ecuador: Ch (Pallatanga; Spruce 1885). General distribution: northern Andes (Ecuador, northern Peru). Frullania haematocysta (subg. Chonanthelia) is a very rare northern Andean species that is recognized by the acute leaf apex, lobules with a narrow laminate portion, auriculate underleaves, and a large, rounded auricle at the dorsal leaf base that extends down the stem (Yuzawa 1991). The species is not known from Colombia and has been collected in Ecuador only by Richard Spruce. 25. Frullania intumescens (Lehm. & Lindenb.) Lehm. & Lindenb. Frullania baptistae Gottsche Frullania cucullata Lindenb. & Gottsche Frullania longicollis Lindenb. & Gottsche Frullania pachoensis Steph. Frullania intumescens (sect. Intumescentes) is a common neotropical species that is distinguished by (1) leaf apices apiculate, recurved; (2) underleaf margins recurved; (3) stylus filiform; and (4) perianth sharply trigonous over most of its length, with a long beak. The species contains two varieties, var. intumescens with a foliose appendage at the base of the stylus and var. sabanetica without stylar appendage. The foliose stylar appendage of F. intumescens was first detected by Yuzawa and Koike (1989) in a study of the type material (Jamaica, Bancroft s.n., lectotype S-B27792!). They showed that F. intumescens as described by Stotler (1969) contains populations with and without stylar appendage and referred the plants without stylar appendage to F. closterantha Spruce, a species included in the synonymy of F. intumescens by Stotler (1969). Since F. intumescens and F. closterantha differ in only one character (presence/absence of a stylar appendage) and are sympatric, they are treated here as varieties. Frullania sabanetica Gottsche is an older name for F. closterantha and is used here for naming the plants without stylar appendage. Stotler (1969) provided a long list of synonyms of F. intumescens; a few of them are cited above. It remains unclear to which of the two varieties they belong. Also, the general distribution of the two varieties remains unclear. A full-scale revision of F. intumescens is necessary. 25a. Frullania intumescens (Lehm. & Lindenb.) Lehm. & Lindenb. var. intumescens

370

 

Keys and Descriptions

On rocky slope along road, 950  m. Ecuador: MS (Schäfer-Verwimp et  al. 2013a); to be expected in Colombia. General distribution: tropical America; distribution incompletely known. Frullania intumescens var. intumescens is recognized by recurved, apiculate leaf apices; underleaves with recurved margins and shortly auriculate bases; the filiform stylus with a large, foliose appendage at the base; and the sharply trigonous perianth. The variety has been recorded once from Ecuador and is apparently more rare in the northern Andes than the var. sabanetica. 25b. Frullania intumescens var. sabanetica (Gottsche) Gradst., comb. nov.1 Frullania sabanetica Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 170. 1864. Type: Colombia, Cundinamarca, Sabaneta, Lindig 1725 p.p. (isotype G-00115883!) Frullania closterantha Spruce (fide Yuzawa and Koike 1989) On bark and rock in montane rainforest areas and páramo, 1000–4000  m. Colombia (Gradstein and Uribe 2011): Ant, Boy, Cau, Ces, Cho, Cun, Hui, Mag, Qui, Ris, San, Tol, Val. Ecuador: Az (Herzog 1952), Ga (?), Pi (Herzog 1952), Tu, ZC. General distribution: tropical America; distribution incompletely known. Similar to F. intumescens var. intumescens except for the absence of a stylar appendage. Gradstein and Uribe (2011) screened the Colombian material of F. intumescens in herbarium COL and found that all specimens were referable to F. intumescens var. sabanetica. Several records have also become known from Ecuador. Apparently, var. sabanetica is the most common variety of F. intumescens in the northern Andes. Populations of F. intumescens var. sabanetica without perianths are morphologically indistinguishable from F. brasiliensis. 26. Frullania jelskii Loitl. Habitat unknown. Colombia: Cun (Apollinaire s.n.; Yuzawa 1991): General distribution: tropical Andes (Peru, Colombia). Frullania jelskii (subg. Chonanthelia) is a very rare Andean species that is characterized by lobules with a very short lamina and smooth surface (not plicate in the middle), large distant underleaves (3–4× stem width) which are somewhat longer than wide, and dioicous sexuality. The species is only known from the type locality in Peru and from Colombia (vicinity of Bogotá), where it was collected by Apollinaire in the nineteenth century (Yuzawa 1991); its continued existence needs confirmation. Frullania jelskii resembles F. dusenii and F. standaertii, but the latter two species are monoicous (and often fertile) and have lobules with a distinctly plicate surface. Frullania standaertii, moreover, has a large, foliose stylus. 27. Frullania kunzei (Lehm. & Lindenb.) Lehm. & Lindenb Figs. 37K, 38E, F Frullania ecuadorensis Steph., syn. nov. Type: Ecuador, Guayas, San José, Allioni s.n., Bryotheca. Levier 6551 (holotype G-00067443!) Frullania neesii Lindenb. Frullania parasitica sensu Mont. (non Lehm. & Lindenb.)

1  After this book went in press, Gradstein & Pócs (Nova Hedwigia 112) showed that the correct name is F. intumescens var. closterantha (Spruce) Gradst. & Pócs

Frullaniaceae

371

On bark in rather open, semi-deciduous and evergreen woodlands, rainforests, and isolated trees, occasionally on living leaves, soil, or rock, sometimes growing over larger bryophytes or foliose lichens, from sea level to 3000  m. Colombia: Ama, Boy (as F. ecuadorensis), Caq, Ris (as F. ecuadorensis), Vau (Campos et al. 2015). Ecuador: Ga, Gu, ZC.  General distribution: tropical and subtropical America. Frullania kunzei (sect. Obtusilobae) is one of the smallest neotropical species in the genus Frullania. It is readily distinguished by (1) plants heteroicous (autoicous or dioicous), dark purplish, prostrate, small, 0.5–1 mm wide; (2) leaves with rounded apex and weakly auriculate dorsal base; (3) hemiphyll undivided; (4) subapical leaf cells (sub)isodiametrical, the walls with trigones but without intermediate thickenings; (5) lobules moderately elongate, 1.3–1.7× longer than wide, sometimes positioned at an oblique angle to the stem (e.g., in the type of F. ecuadorensis); (6) underleaves distant to contiguous, ± flat, little wider than the stem (1.5–2.5×); and (7) female bracts entire, female bracteoles entire or toothed, perianth broadly 3-keeled. By its small size and prostrate growth, F. kunzei approaches F. obcordata and F. subtilissima but is separated from the latter two species by all (or part of) the lobules being positioned close to the stem (all lobules at considerable distance from the stem in F. obcordata and F. subtilissima). Frullania ecuadorensis is an epiphyllous phenotype of F. kunzei with lobules varying from upright and close to the stem to oblique and at some distance to the stem (the latter position as in subg. Diastaloba). The close similarity of F. ecuadorensis to F. kunzei (= F. neesii) was already noted by Stotler (1969). 28. Frullania laxiflora Spruce Fig. 38G Frullania laxiflora var. crossii Spruce, syn. nov. Type: Colombia, Cauca, Popayán, 1877, Cross 760c (MANCH!, c. spor.) On bark in montane rainforest areas, (450–)1500–3500  m; on the Galápagos Islands (Santa Cruz) in moist Scalesia woodland at 650–700 m. Colombia: Cau, Tol. Ecuador: Ch, Ga, Tu. General distribution: tropical Andes, Galápagos Islands, Costa Rica. Frullania laxiflora (subg. Chonanthelia) is an uncommon Andean species that is characterized by (1) leaves strongly convex; (2) lobule with a long laminate portion, which is about as long as the sac but does not extend beyond the ventral lobe margin; (3) lobule with a longly decurrent beak; and (4) underleaves broadly rounded, shallowly bifid and with large auricles (Yuzawa 1991). Frullania laxiflora may be confused with F. rio-janeirensis, but in the latter species, the laminate portion of the lobule extends beyond the ventral lobe margin (to be observed on leaves of the main stem) and the beak is not longly decurrent. 29. Frullania lindenbergii Lehm. On canopy branches of trees in disturbed montane forest and isolated trees in meadows, ca. 1900 m. Ecuador: ZC (Gradstein and Benitez 2017). General distribution: East Africa, Tristan da Cunha, Ecuador, Brazil.

372

 

Keys and Descriptions

Frullania lindenbergii (subg. Diastaloba s.l.) is a common African species that has been recorded from Brazil and the Dominican Republic (but see SchäferVerwimp et al. 2012) and was recently found in Ecuador (San Francisco Biological Reserve). The species is distinguished by (1) plants autoicous, ca. 1  mm wide, creeping, pale reddish-brown, densely (bi)pinnately branched with numerous small, microphyllous branches; (2) leaf apices flat, rounded on main stems and rounded to bluntly apiculate on branches; (3) leaf cells ± isodiametric, trigones distinct and not lined by a darkish wall, leaf base without group of large ocellus-like cells; (4) hemiphyll undivided; (5) lobules cylindrical, positioned at considerable distance from stem, abaxial side of the lobules closed; (6) stylus foliose, made up of many cells; (7) underleaves variable in size, 2–4× stem width, outer margins ± toothed; and (8) female bracts and bracteoles entire or toothed. Frullania lindenbergii superficially resembles F. subtilissima, but the latter species is readily separated from F. lindenbergii by the bifid hemiphyll, the partially open abaxial side of the lobule, and the presence of a group of large, darker-colored cells at the leaf base. Moreover, the stylus in F. subtilissima is usually linear, rarely foliose. 30. Frullania lobatohastata Steph. Frullania apollinarii Steph. Frullania villosa Steph.

Fig. 38H, I

On bark in montane forest and shrubby páramo, 1800–4000 m. Colombia: Boy, Cun, Qui, San. Ecuador: Az, Pi (Schäfer-Verwimp et al. 2013a). General distribution: northern Andes (Colombia, Ecuador). Frullania lobatohastata (sect. Intumescentes) is a characteristic northern Andean species that is distinguished by underleaves with very long auricules, almost 1/2 the length of the underleaf lamina (measured from the middle of the underleaf insertion line to base of the leaf lobes). The leaf apex is broadly rounded (occasionally shortly apiculate); the underleaves are large, bifid to 1/4–1/3 and with broadly recurved margins; and the perianth is sharply trigonous to subterete with a few weak folds above. Frullania lobatohastata somewhat resembles F. brasiliensis, but the leaf apex in F. brasiliensis is always apiculate and the underleaf auricles are much smaller. 31. Frullania macrocephala (Lehm. & Lindenb.) Lehm. & Lindenb. Fig. 38J, K Frullania bullatiflora Steph. Frullania mucronata auct. (non typus) Frullania repanda Gottsche (fide Hentschel et al. 2015) Frullania strobilantha Spruce Frullania tubiflora Spruce On bark of trees in montane forest, also on shaded, moist rock, 1000–2900 m. Colombia: Boy, Cun, Qui, Ris, San. Ecuador: Pa, MS, ZC. General distribution: tropical America. Frullania macrocephala (sect. Intumescentes) is closely similar to F. brasiliensis but differs from the latter in the flat leaf apex (strongly recurved in F. brasiliensis).

Frullaniaceae

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Both species are rather robust and have strongly recurved underleaf margins, terete perianths, and  ±  entire female bracts and bracteoles. The relationship of the two species should be analyzed with molecular methods; possibly F. macrocephala is a mere variety of F. brasiliensis. 32. Frullania meridana Steph.

Fig. 38L

On bark of trees in montane forest and páramo, 1500–4000 m. Colombia: Cun, Ris, Tol. Ecuador: Pi (Schäfer-Verwimp et  al. 2013a). General distribution: northern Andes. Frullania meridana (sect. Intumescentes) is a rare northern Andean species that is distinguished by the flat and shortly apiculate leaf apex and the large (more than 3× stem width), distant, longer than wide underleaves with plane margins. By the large underleaves, F. meridana resembles F. clandestina and F. formosa, but in the latter two species, the underleaves are imbricate and orbicular or wider than long. Moreover, F. clandestina has a rounded leaf apex. 33. Frullania mirabilis J.B.Jack & Steph. Frullania pendulostyla Steph. On bark of trees and soil in montane rainforests, 1200–2550 m. Colombia: Ant, Cau, Cho, Met, Nar, Val. Ecuador: MS, ZC.  General distribution: northern Andes, Guayana Highland, Costa Rica. Frullania mirabilis (sect. Intumescentes) is an unusual northern Andean-Costa Rican species that is recognized by the presence of 1–2 long appendices at the ventral base of the leaf, one standing up and the other one hanging down (Stotler 1969 as F. pendulostyla). The leaf apex in the F. mirabilis varies from rounded to apiculate. 34. Frullania montagnei Gottsche Frullania auritistipula Steph. Frullania gualaquizana Steph., syn. nov. Type: Ecuador, Morona Santiago, Gualaquiza, “in silvis,” Feb 1909, Allioni s.n., Bryotheca Levier 6480 (lectotype, fide Bonner (1965), G-00068154!, Icon. Ined. 4038). Syntype: ibid., Allioni s.n., Bryotheca Levier 6475 (G-00265392!) Frullania purpurascens Steph. On bark, 200–2250  m. Colombia: Mag, San, Tol, Vau. Ecuador: MS, Na, ZC. General distribution: tropical America. Frullania montagnei (sect. Intumescentes) is a little-known neotropical species that stands out by (1) plants 0.8–1.5 mm wide; (2) leaves convex, apices recurved or plane, rounded to obtuse (rarely a few leaves slightly apiculate); (3) stylus very short, filiform; (4) hemiphyll undivided; (4) subapical leaf cells longer than wide, leaf cell walls with trigones and intermediate thickenings; (5) underleaves distant to imbricate, 2–3× stem width, bifid to 1/4, margins recurved; and (6) perianth inflated, with three rounded keels in the upper half, female bracts and bracteoles usually entire, rarely somewhat toothed (Vaupés, Schultes & Cabrera 14296, CINC, PC). Frullania montagnei resembles F. paradoxa, but the latter species has a bifid hemiphyll (bifid to 1/4–1/2), smaller underleaves (1.5–2× stem width), strongly

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Keys and Descriptions

ciliate-laciniate female bract lobules and bracteoles, and a sharply 3-keeled perianth which is abruptly narrowed to a very long beak. Robust forms of F. kunzei approach F. montagnei but differ from the latter species in having shorter lobules, smaller underleaves, ± isodiametric apical leaf cells, and leaf cell walls without intermediate thickenings. 35. Frullania nodulosa (Reinw., Blume & Nees) Nees  Frullania replicata (Nees) Spruce

Fig. 38M, N

In the canopy of lowland rainforest and on isolated trees and shrubs in savannas, 100–250  m. Colombia: Ama, Vau. Ecuador: Na, Or. General distribution: pantropical. Frullania nodulosa (subg. Homotropantha) is a widespread pantropical species from low elevations that is readily separated from all other neotropical members of the genus Frullania by its pendent lobules, with the mouth up and the lobule apex turned to the base of the stem. The plants are robust and regularly (bi)pinnate, the leaf lobes are suborbicular with a broadly rounded apex, and the underleaves are large, imbricate, very shallowly bifid and with large basal auricles (Gradstein and Ilkiu-Borges 2009). The species is widespread in Amazonia but has been little collected in Colombia and Ecuador. 36. Frullania obcordata (Lehm. & Lindenb.) Lehm. & Lindenb. Frullania caroliniana Sull. Frullania compsotera Spruce Frullania gymnotis Mont. Frullania galapagona Ångstr. Frullania parasitica Lehm. & Lindenb. (non Mont.)

Fig. 36K, L

On bark in lowland and lower montane forest areas and scrub, growing tightly appressed to the substrate, from sea level to ca. 1000 m Colombia: Ama, Boy, Caq, Cas, Ces, Mag, Met, Qui, Ris, San, Tol, Vau. Ecuador: Ga, Pa. General distribution: tropical America. Frullania obcordata (subg. Diastaloba s.l.) is a common neotropical epiphyte from low elevations; records from above 1000 m need verification. In recent years the species was called F. caulisequa (Nees) Mont. following Yuzawa (1988), but according to G. Winter (pers. com.), F. caulisequa is a different species. Frullania obcordata is readily distinguished by (1) plants autoicous, small (0.9–1.4  mm wide), creeping, little branched, without microphyllous branches; (2) leaf apex rounded (never pointed); (3) leaf cells with rather small trigones and without group of large, colored cells at the leaf base; (4) hemiphyll undivided; (5) lobules positioned at some distance from the stem and closed on the adaxial side; (6) underleaves small, 2–3× stem width; and (7) female bracts entire or toothed. Plants with toothed bracts have been called F. gymnotis. Frullania obcordata resembles F. lindenbergii in the undivided hemiphyll, the absence of a group of large, ocellus-like cells at the leaf base and lobules not incised on the adaxial side, but differs from the latter species in the rounded leaf tips (never apiculate), sparse branching, absence of microphyllous branches, and small underleaves.

Frullaniaceae

375

37a. Frullania obscura (Sw.) Mont. var. obscuraFig. 38O, P Frullania arecae (Spreng.) Gottsche (fide Gradstein 2012b) Frullania campanensis Spruce Frullania hians (Lehm. & Lindenb.) Lehm. & Lindenb. On bark and rock in montane forest, (500–)1600–3400  m; on the Galápagos Islands in moist Scalesia woodlands at 500–750 m. Colombia: Boy, Cas, Cau, Ces, Cun, Hui, Mag, Qui, Ris, San, Tol. Ecuador: Az, Ch, Co, Ga, MS, Lo, Su, Tu, ZC. General distribution: pantropical. Frullania obscura is a common, tropical-montane species that is distinguished by the robust plants with undulate margins of leaves and underleaves, beaked lobules, and a pluriplicate perianth surrounded by a large and strongly toothed, flowerlike involucre. Frullania obscura may be confused with F. ecklonii; for differences see under the latter species. Two varieties have been recognized, var. obscura with entire laminate portion of the lobule and var. spiniloba characterized by the toothed laminate portion of the lobule. Frullania obscura var. obscura is widespread throughout the tropics, whereas var. spiniloba is only known from Ecuador. 37b. Frullania obscura var. spiniloba (Steph.) Hentschel & von Konrat Frullania spiniloba Steph. (Frullania arecae (Spreng.) Gottsche var. spiniloba (Steph.) Yuzawa) On bark, ca. 650–700 m. Ecuador: Ga, Ma. General distribution: Ecuador, Central America. Differs from Frullania obscura var. obscura in the toothed laminate portion of the lobule (entire in var. obscura). 38. Frullania paradoxa Lehm. & Lindenb. Fig. 39A Frullania flexicaulis Spruce (fide Hentschel et al. 2015) Frullania trollii Herzog, syn. nov. Type: Colombia, Bogotá, 3000–3200  m, Troll 2086 (JE!); the synonymy was already noted by R. Stotler (in sched.) but remained unpublished Frullania ulotricha Taylor On bark of trees and shrubs in upper montane forests, elfin forest and páramo, also on rock, 1500–4000 m. Colombia: Boy, Cau, Cun, Qui (páramo El Campanario), Ris, Val. Ecuador: Az, Ca, Ch, Ga (?), Lo, Na, Pi, Tu, ZC. General distribution: scattered in tropical America. Frullania paradoxa (sect. Intumescentes) is common in the northern Andes and is recognized by (1) plants rather small, leaf apex rounded to acute; (2) hemiphyll bifid to 1/4–1/2, the two segments often different in size; (3) underleaves distant to subimbricate, small, only 1.5–2× stem width, longer than wide, shallowly bifid (to 1/6–1/4) with plane or narrowly recurved margins; and (4) gynoecia terminal on the stem or on short branches, bract lobules and bracteoles ciliate-laciniate, bract lobe subentire to ciliate-laciniate, perianth sharply 3-keeled, abruptly narrowed to a long-linear beak (Stotler 1969). Frullania flexicaulis is a phenotype with rounded to obtuse leaf tips, gynoecia on short branches, and subentire female bract lobes.

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Keys and Descriptions

Fig. 39 (A) Frullania paradoxa, habit. (B) Frullania peruviana, habit. (C and D) Frullania pittieri. (C) Leaf. (D) Habit. (E) Frullania planifolia, habit. (F–H) Frullania pluricarinata. (F) Leaf. (G) Underleaf. (H) Perianth. (I and J) Frullania rio-janeirensis. (I) Leaf. (J) Habit. (K and L) Frullania setigera. (K) Underleaf. (L) Habit. (M–O) Frullania sphaerocephala. (M) Habit. (N) Underleaf, (O) Leaf. (P) Frullania standaertii, lobule. (Q and R) Frullania stenostipa. (Q) Lobule, (R) Underleaf. (S and T) Frullania tetraptera. (S) Lobule. (T) Underleaf. (U) Frullania winteri var. vanderhammenii, leaf. (A, C, D redrawn from Stotler 1969; B redrawn from Uribe 2008; E, F–H, M–U redrawn from Yuzawa 1991; I, J from Gradstein et al. 2001; K, L from Gradstein and Uribe 2011). Scale bars: A, C, D, I, J, N, O = 0.5 mm; B, E, H, L, M = 1 mm; F, G, P–U = 250 μm; K = 400 μm

Frullaniaceae

39. Frullania peruviana Gottsche Frullania atrata sensu Spruce (non typus) Frullania atrosanguinea Taylor Frullania haeckeriana Lindenb. Frullania mathanii Steph. Frullania remotifolia Steph.

377

Fig. 39B

On bark of tree branches in montane cloud forest areas, (450–)1600–4000 m; very common in moist evergreen woodlands and scrub on the Galápagos Islands at 450–1100 m. Colombia: Ant, Boy, Cal, Cho, Cun, Mag, Nar, Qui, Ris, Tol, Val. Ecuador: Az, Ca, Co, EO, Ga, Im, Lo, MS, Na, Pi, Tu, ZC. General distribution: Central America, tropical Andes, Galápagos Islands. Frullania peruviana (sect. Meteoriopsis) is a common neotropical montane cloud forest species that is distinguished by pendent growth, acute to short-acuminate leaf apices, underleaves without or with small auricles, and toothed female bracts (Uribe 2008). 40. Frullania phalangiflora Steph. Frullania bakeri Steph. On bark, 300–600  m. Ecuador: EO, Ga (Isabela, Santa Cruz; Uribe 2008). General distribution: tropical Andes (Venezuela, Bolivia), Galápagos Islands, Central America. Frullania phalangiflora (sect. Meteoriopsis) is a rare Andean species that resembles F. peruviana but differs in the underleaves with large auricles and the globose perianth (perianth elongate in F. peruviana; Uribe 2008). Frullania phalangiflora is also close to F. darwinii, but in the latter species, the underleaf auricles are smaller, the underleaf margins are not recurved, and the perianth is elongate. 41. Frullania pittieri Steph.

Fig. 38C, D

On bark of trees, from sea level to 3000  m. Colombia: Boy, Cau, Cho, Ris. Ecuador: ZC. General distribution: scattered in tropical America. Frullania pittieri is morphologically close to F. macrocephala, but the leaf tips in F. pittieri are more longly acuminate, ending in a row of up to 10 cells (row of 1–4 cells in F. macrocephala), the female bracts and bracteoles are toothed, and the perianth is 3-keeled (bracts and bracteoles entire and perianth subterete in F. macrocephala). 42. Frullania planifolia Steph. Frullania subalpina Spruce

Fig. 39E

On tree bases and rock on steep, cut road banks, 3000–3500 m. Ecuador: Pi (León-Yánez et al. 2006); not known from Colombia but expected there. General distribution: northern Andes (Ecuador, Venezuela), Costa Rica. Frullania planifolia (subg. Chonanthelia) is an uncommon northern Andean species from high elevations that is recognized by the very flat leaves; the rather large laminate portion of the lobule, which is about as long as the sac but does not extend

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Keys and Descriptions

beyond the ventral lobe margin; the longly decurrent beak of the lobule; and the pluriplicate perianth. 43. Frullania pluricarinata Gottsche

Fig. 39F–H

On tree trunks in dry woodlands and open areas, city parks, and along roads, occasionally on rock, often growing in urban areas, xerotolerant, 1700–3350  m. Colombia: Cun (type), Mag. Ecuador: Az, Pi. General distribution: a disjunct Latin American species, occurring at high elevation (above 1700 m) in Mexico and Guatemala and in the northern Andes and again at low elevation (below 800 m) in the Mediterranean region of central Chile (Gradstein and Cuvertino 2015). Frullania pluricarinata (subg. Chonanthelia) is a characteristic xerotolerant species of urban and open areas in the high Andes that has been little collected. The species is distinguished by (1) leaves ± squarrose when moist, sometimes caducous; (2) lobules with a small laminate portion, without beak; (3) underleaves distant, rather small, 2–3× stem width, slightly longer than wide, bifid to 1/4, outer margins sometimes with a tooth (in large underleaves), bases cuneate to rounded, not auriculate; and (4) perianth pluriplicate. The plants may be autoicous or dioicous. Plants without perianths resemble F. decidua and F. tetraptera, but the shape of the underleaves in the latter two species is different (see key). 44. Frullania ringens Spruce On bark of trees, 3400  m. Colombia: Cau (Cleef 493f, COL). Ecuador: Ch (Spruce 1884). General distribution: northern Andes (Colombia, Ecuador, Peru). Frullania ringens (subg. Chonanthelia) is a rare northern Andean species that is recognized by the lobules with a large, 2–3-toothed lamina and a longly decurrent beak; the large, rounded underleaves with two very large, ± overlapping auricles; and pluriplicate perianths (Yuzawa 1991). The species is close to F. ecklonii, but the latter species differs in having lobules with an entire lamina and without beak and by smaller underleaf auricles that are not overlapping. 45a. Frullania rio-janeirensis (Raddi) Ångstr. var. riojaneirensisFig. 39I, J Frullania arietina Taylor Frullania latilobula Herzog Frullania leptomitra Spruce Frullania tenera Spruce On bark and rock in open environments, from sea level to 3800  m; on the Galápagos Islands widespread and common in semi-deciduous and rather dry evergreen woodlands and scrub up to 1200 m. Colombia: Ant, Boy, Cal, Cas, Cau, Ces, Cho, Cun, Hui, Mag, Qui, Ris, San, Tol. Ecuador: Co, EO, Ga, Lo, MS, Or, Pa, Pi, Tu, ZC. General distribution: pantropical. Frullania rio-janeirensis (subg. Chonanthelia) is a common and variable pantropical species of open environments. It is one of the most commmon droughttolerant epiphytic liverworts in tropical America and has a wide elevational range. The species is distinguished by the long laminate portion of the lobule which is not wider than the sac (to be examined on stem leaves) and lacks teeth on the outer

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margin and by the distinct lobule beak and the 4-keeled perianth. The plants may be autoicous or paroicous; paroicous plants have been described as F. arietina. The populations of Colombia and Ecuador generally belong to var. rio-janeirensis characterized by short male branches, with 3–6 pairs of bracts (see also under F. riojaneirensis var. megalostipa). Frullania rio-janeirensis var. rio-janeirensis may be confused with F. laxiflora; for differences see under the latter species. The name of the species is usually spelled “F. riojaneirensis,” but the original spelling “rio-janeirensis” should be retained because Rio de Janeiro are separate words (ICN Art. 60.9). 45b. Frullania rio-janeirensis var. megalostipa (Spruce) Gradst., comb. nov. Frullania megalostipa Spruce, Trans Proc. Bot. Soc. Edinburgh 15: 15. 1884. Frullania amentacea Herzog On bark, 700–2000 m. Ecuador (Yuzawa 1991): MS, Pa. General distribution: Ecuador (type), Bolivia. Frullania rio-janeirensis var. megalostipa is a rare Andean taxon that differs from the typical variety in the long male branches, consisting of 10–12 pairs of bracts (Yuzawa 1991). In var. rio-janeirensis, the male branches consist of 3–6 pairs of bracts. 46. Frullania setigera Steph.

Fig. 39K, L

On bark of trees, soil, rock, and logs in montane forests, 1000–3600  m. Colombia: Ant, Boy, Cau, Cho, Hui, San, Val. Ecuador: Az, Lo, Na, Pi, ZC. General distribution: scattered in tropical America. Frullania setigera (sect. Intumescentes) is a little-known neotropical species that is recognized by the leaves with a flat, shortly acuminate apex; underleaves with apiculate to narrowly acuminate apices, recurved margins, and auriculate bases; the longly acuminate apex of the hemiphyll; entire female bracts and bracteoles; and a 3-keeled perianth (Stotler 1969). The species resembles F. beyrichiana, but in the latter species, the leaf apex is apiculate (not acuminate), the underleaf margins are more or less plane, and the apex of the hemiphyll is acute (not longly acuminate). 47. Frullania sphaerocephala Spruce

Fig. 39M–O

On bark of trees in upper montane cloud forest areas and shrubby páramo, ca. 2000–3900 m. Colombia: Cun, Met, Qui, Ris, Tol. Ecuador: Az, Lo. General distribution: tropical Andes. Frullania sphaerocephala (subg. Chonanthelia) is a characteristic high-Andean species that is readily distinguished by the strongly involute-imbricate, ball-shaped female involucres. By its ball-shaped female involucres, F. sphaerocephala cannot be confused with any other neotropical Frullania species. Further characters of the species include the lobules with a long lamina and without beak; the large trigones; the large auricles of leaves and underleaves; the minute, 1(−2)-celled stylus; and the pluriplicate perianth. Frullania sphaerocephala is related to F. ecklonii but differs from the latter species in the ball-shaped female involucres and dioicy (F. ecklonii is monoicous).

380

 

Keys and Descriptions

48. Frullania standaertii Steph. Fig. 39P Frullania albertii Steph. Frullania arsenii Steph. (“F. arnellii” in Gradstein and Uribe 2016a) On bark of trees and shrubs in upper montane forests and páramo, 2750–4000 m. Colombia: Boy, Cun, San. Ecuador: Pi (León-Yánez et al. 2006 as F. albertii). General distribution: tropical Andes; also in East Africa (Kenya). Frullania standaertii (subg. Chonanthelia) is identical to F. dusenii except for the larger, foliose stylus (Yuzawa 1991). The stylus in F. standaertii varies considerably in size, however, and ranges from filiform to partly foliose, 5–10 cells long, in plants from the Sierra Nevado del Cocuy identified by Yuzawa as F. albertii. This variation is seen on single shoots. The underleaves in the species are also more variable than previously described and vary from small, distant to large, imbricate, measuring ca. 3–5× stem width. Possibly, F. standaertii is a variety of F. dusenii. 49. Frullania stenostipa Spruce

Fig. 39Q, R

Unknown habitat, 3000  m. Ecuador: Pi (Yuzawa 1991). General distribution: Ecuador, Costa Rica. Frullania stenostipa (subg. Chonanthelia) is a very rare neotropical species that is recognized by narrowly elongate underleaves (2–3× as long as wide) and the 5–10-cell-long stylus (Yuzawa 1991). It is closely similar to F. tetraptera, but the underleaves in the latter species are less elongate and the stylus is shorter. Possibly, F. stenostipa is a mere variety of F. tetraptera. 50. Frullania subtilissima (Nees ex Mont.) Lindenb. (1845) Fig. 37N, O Frullania apiculata auct. (non typus) Frullania atrata (Sw.) Mont. var. [“β”] subtilissima Nees ex Mont. [“N. ab E. in litt.”). Type: French Guiana, “ad truncos Bixae Orellanae secùs flumen Kau om Guianâ, Maio 1838 lecta.  – Lepr. Coll. n. 254” (lectotype, designated here, PC-Mont.-0033300!). Although the herbarium label mentions “Leprieur 264,” the specimen in PC-Mont. cited here is selected as the lectotype as it is the only specimen of F. atrata β subtilissima in the Montagne herbarium. Frullania exilis Taylor (1846), syn. nov. Type: Guyana, Demarara, Apr 1842, unknown collector, ex herb. Greville (lectotype, designated here, FH-00290639!) Frullania confertiramea Steph. Frullania crenulifolia J.B.Jack & Steph., syn. nov. Type: Colombia, Antioquía, páramo de Sonson, 10.000 ft., 1872, Wallis s.n. (G!) Frullania paraensis Steph. Frullania taxodiocola R.M.Schust., syn. nov. Type: U.S.A., Florida, Monroe Co., Cypress swamp, 25 Dec 1958, Schuster 42241 (F n.v.); the type material cannot be found (L. Briscoe, pers. comm.) On bark in lowland and montane rainforests and scrub up to the páramo, growing tightly appressed to the substrate, from sea level to 3300 m. Colombia: Caq, Cho, Qui, Ris, San. Ecuador: Lo (Schäfer-Verwimp 21771, 31820, PC), MS, ZC. General distribution: tropical America, Africa (?).

Frullaniaceae

381

Frullania subtilissima (subg. Diastaloba s.l.) is a very small species that is distinguished by (1) plants 0.7–1 mm wide, prostrate, dark reddish-brown to purplish, regularly (bi)pinnate branched, without or with a few microphyllous branches; (2) leaf apex varying from rounded to apiculate; (3) leaf base with a group of large, often darker-colored, ocellus-like cells; (4) lobules cylindrical, distant from the stem, open (incised) in the lower half on the adaxial side; (5) hemiphyll bifid; (6) stylus linear, rarely foliose; (7) underleaves small, 2–3× stem width, entire; and (8) plants autoicous, female bracts and bracteoles entire or toothed. The leaf cells in F. subtilissima are isodiametrical to somewhat elongate, and the cell walls are often quite sinuose and deeply reddish-purple in plants from high elevation. The trigones in these high-elevation plants are often conspicuously hyaline-whitish and lined by a darkish pigmented wall. Based on the detailed description and illustration in Schuster (1992a), F. taxodiocola is a synonym of F. subtilissima; the type material of F. taxodiocola has not been located. The lectotype of Frullania subtilissima is a mixture of two species, one belonging to subg. Diastaloba and the other to subg. Meteoriopsis sect. Intumescentes. Since the leaves of F. subtilissima are described in the protologue as obtuse to mucronate (Gottsche, Lindenb. & Nees, 1844–1847, p. 443), the Diastaloba element must be considered the genuine F. subtilissima, as already proposed by A.  Schäfer-Verwimp (in sched.). Gradstein and Hekking (1989) selected the Intumescentes element as the type, but this is not in agreement with the protologue since the leaf apex in this plant, which belongs to F. montagnei, is rounded. Frullania subtilissima, as lectotypified here, is identical to F. exilis and widespread in tropical America. The species has also been called “Frullania apiculata” (e.g., Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009; Gradstein and Uribe 2011), but the latter species, described from Asia, is a more robust plant, with larger underleaves and all leaves sharply apiculate (Gradstein 2013b). The relationship of F. subtilissima and F. apiculata (Reinw., Blume & Nees) Nees deserves study. 51. Frullania tetraptera Nees & Mont. Frullania semiconnata Lindenb. & Gottsche

Fig. 39S, T

On bark of trees in montane forest areas and páramo in rather open, moist habitats, (1000–)2000–4000 m. Colombia: Cun, Met, Ris. Ecuador: Az, Pi. General distribution: tropical America. Frullania tetraptera (subg. Chonanthelia) is a widespread neotropical species that is distinguished by the small plants with a short laminate portion of the lobule, absence of a lobule beak, shallowly bifid underleaves (bifid to 1/6–1/4) that are longer than wide and 1.5–2.5× wider than the stem, 4-keeled perianths, and autoicous sexuality (Yuzawa 1991). The species has been confused with F. cuencensis, but the lobules in the latter species are beaked and the underleaves are broader and more deeply bifid (to 1/3–1/2). Moreover, F. cuencensis is dioicous. Frullania tetraptera also somewhat resembles F. pluricarinata, but the latter species has subsquarrose and somewhat caducous leaves and a pluriplicate perianth.

382

 

Keys and Descriptions

52. Frullania tunguraguana L.Clark & Frye Frullania brachyclada Spruce, nom. illeg. On bark of shrubs and on soil in rather mesic montane forest areas including parks, also in Polylepis pauta forest, up to the páramo, 500–4000 m. Colombia: San. Ecuador: MS (Schäfer-Verwimp et al. 2013a), Pi (Mojanda, León-Yánez s.n., QCA), Tu. General distribution: tropical Andes, Mexico. Frullania tunguraguana (subg. Chonanthelia) is a rare Andean species characterized by pluriplicate perianths; female involucre with short, obtuse teeth; almost plane leaf and underleaf margins (not clearly undulate); beaked lobules with a long, entire laminate portion; and autoicous sexuality. Frullania tunguraguana is close to F. obscura and has sometimes been considered a synonym of the latter, but F. obscura differs in having clearly undulate leaf and underleaf margins, a sharply toothedciliate female involucre (obtusely toothed in F. tunguraguana), and underleaves with a narrower sinus (Yuzawa 1991). 53. Frullania uleana Steph. Frullania involuta Steph.

Fig. 38D

Forming thread-like, blackish garlands in summit bush, 450–650 m. Ecuador: Ga (Pinta, Pinzón; Gradstein 2009). General distribution: scattered in tropical America, mainly in the West Indies. Frullania uleana (sect. Intumescentes) is a Caribbean species that has been recorded from the Galápagos Islands but not from the mainland of Ecuador and Colombia. Uribe and Gradstein et al. (2003) described the species under the name F. involuta, but the name F. uleana is older and has priority. Frullania uleana is readily distinguished by the very slender habit; pendent growth; toothed leaf margins; dorsal leaf base with a large auricle and ventral leaf base without or with a much smaller auricle; distant, deeply bifid, flat underleaves, which are hardly wider than the stem; and cuneate or slightly rounded underleaf bases. The species resembles F. atrata and F. dulimensis; for differences see under the latter species. 54. Frullania winteri Steph. var. vanderhammenii (Haarbrink) Yuzawa Frullania vanderhammenii Haarbrink On bark of trees in montane rainforest, 1700–2550 m. Colombia: Boy, Qui, Ris, San. Ecuador: ZC.  General distribution: northern Andes (Colombia, Ecuador), Mexico. Frullania winteri (subg. Chonanthelia) is a rather rare northern Andean-Mexican species that is recognized by the folded margin of the (short) laminate portion of the lobule. Yuzawa (1991) recognized two varieties: var. winteri with a relatively large stylus (10–13 cells long, 5–10 cells wide) and var. vanderhammenii with a smaller stylus (5–8 cells long, 1–2 cells wide). The plants from the northern Andes belong to var. vanderhammenii.

Jubulaceae

383

Further Records Frullania crispiloba Steph. – Colombia: Cho (Vasco et al. 2002). Frullania moritziana Lindenb. & Gottsche – Colombia: Qui (Orrego and Uribe 2004), Ris (Wolf 1993). Excluded Records Frullania armata Herzog & L.Clark  – Colombia: Mag (Winkler 1976). The material was probably misidentified. Frullania serrata Gottsche  – Ecuador: ZC (Reserva Biólogica San Francisco; Nöske et al. 2003). The material probably belongs to F. lindenbergii. Frullania tamarisci (L.) Dumort.  – Ecuador: Ga (Hooker 1847; Evans 1902). Holarctic species; the material from the Galápagos Islands was probably misidentified. Frullania vaginata (Sw.) Dumort. – Ecuador: Ga (Hooker 1847; Evans 1902). Asiatic species; the material from the Galápagos Islands was probably misidentified.

Jubulaceae A small family of three genera, one genus in Colombia and Ecuador. Jubula Dumort. Plants glossy deep green, creeping, regularly pinnate. Branches Frullania-type or Lejeunea-type; innovations normally present. Leaves incubous with a long, J-shaped insertion, divided into a dorsal lobe and a small ventral lobule; leaf lobe ovate, apex mucronate to long-piliferous, margins sharply toothed or entire; leaf lobule small, rounded and saccate (sometimes ± flat), attached to the ventral margin of the lobe at some distance from the stem; stylus absent. Cells thin-walled, with minute trigones; oil bodies finely granular. Underleaves bifid, with longly decurrent bases and deeply arched insertion. Autoicous. Androecia on tiny, elongate branches. Gynoecia on elongate shoots, with 1–2 Radula-type innovations and 2–3 archegonia. Perianths flattened, with 3 sharp keels (2 lateral, 1 ventral), perianth mouth contracted into a beak. Setae of only 20 rows of cells: 4 inner rows and 16 outer rows. Spore surface without rosettes. A widespread genus (ca. 5 spp.), one species or subspecies in Colombia and Ecuador. For recognition of Jubula, see under the species.

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Keys and Descriptions

Literature: Guerke (1978) 1. Jubula bogotensis Steph. Fig. 63A–C Jubula hutchinsiae (Hook.) Dumort. subsp. bogotensis (Steph.) Verd. Jubula pennsylvanica (Steph.) A.Evans subsp. bogotensis (Steph.) Guercke On bases of trees, rock, and humic soil in the shaded understory of submontane and montane rainforest, also on rock in rivers, 600–3000 m. Colombia: Cun, Hui, Mag, Qui, San. Ecuador: MS, Pa, Tu. General distribution: scattered in the neotropics, known from Mexico, Central America, the West Indies, and the tropical Andes. Jubula bogotensis is an uncommon neotropical species that is recognized by the green to greenish-brown plants without any reddish pigmentation, laxly pinnate branching, entire to sharply toothed leaf margins, a longly mucronate leaf apex, and a small, rounded or flat lobule which is attached to the ventral margin of the lobe at some distance from the stem. The underleaves are bifid, 2–3× stem width, with entire or toothed margins, cuneate and longly decurrent bases, and a deeply arched insertion. The plants are autoicous and frequently fertile. Jubula bogotensis is very similar to Frullania, but the latter genus is frequently reddish to purplish and occurs in open, sunny habitats. The green color of J. bogotensis coincides with its occurrence in shaded habitats in the forest understory. The species is close to J. pennsylvanica from North America and J. hutchinsiae from Eurasia and has been considered a subspecies of either of them (e.g., Guerke 1978; Pätsch et al. 2010). However, all studies on J. bogotensis were based on a very limited number of specimens. A detailed study of the characters and relationships of the species based on a broad sampling is necessary. As shown by Katagiri et al. (2010), detailed morphological study of Jubula species may reveal new and little-known species characters. Pending more detailed study, J. bogotensis is provisionally maintained here at species level.

Lejeuneaceae Plants green, yellowish, brown, black, or whitish, never reddish; creeping to ascending or pendent, pinnate, forked or irregularly branched. Branches usually Lejeuneatype, sometimes Frullania-type; innovations of the Radula-type; stolons lacking. Stems with or without hyalodermis. Leaves incubous, divided into a large dorsal lobe and a small ventral lobule, the lobule broadly attached to the dorsal lobe along a keel; stylus reduced. Cells with homogeneous or granular oil bodies; ocelli sometimes present. Underleaves undivided or bifid, rarely lacking. Rhizoids in bundles from underleaf bases. Gametoecia on leading shoots or on short branches. Androecia usually with 1–2 antheridia per bract, antheridium globose, on a long 1-seriate stalk. Gynoecia with one archegonium. Sporophytes surrounded by a perianth; mouth of the perianth usually contracted into a short beak. Sporophyte foot reduced, not penetrating the stem. Setae very short, thin, of 16 or 20 rows of cells, rarely thicker. Capsules globose, wall 2-layered. Elaters attached to the capsule valves, arranged

Lejeuneaceae

385

vertically inside capsule, spirals well developed or reduced. Spores large, germination endosporic. Vegetative reproduction by multicellular gemmae, caducous leaves or leaf lobes, caducous branches or fragmentation. The largest family of the liverworts with more than a thousand species in about 70 genera worldwide, ca. 330 species in 48 genera in Colombia and Ecuador. The unique morphological character of Lejeuneaceae is the presence of only one archegonium in the gynoecium. Further important family characters are (1) plants without reddish pigmentation; (2) vegetative branches mainly Lejeunea-type, innovations Radula-type; (3) leaves incubous, divided in a large dorsal lobe and a smaller ventral lobule which is attached to the lobe along a long keel; (4) leaf developing by means of 3 initial cells, one producing the lobe (indicated by a hyaline papilla at the dorsal leaf base), the second producing the lobule (indicated by a hyaline papilla at the lobule apex), and the third producing the stylus, which is normally reduced to a hyaline papilla at the base of the lobule (stylus well developed in a few spp. of Cololejeunea); (5) underleaves attached to the stem by 2–4 (or more) U-shaped cells (“superior central cells”; visible in longitudinal and cross section of stem); (6) rhizoids in bundles from the bases of underleaves; the bundles may develop into a large, adhesive disc in epiphyllous plants; (7) sporophyte in a beaked perianth; (8) foot of sporophyte short, not penetrating into the stem; (9) capsule wall 2-layered; (10) elaters attached to capsule valves; and (11) spore germination endosporic, within the capsule. Characters 4 and 7–11 are shared with Frullaniaceae and Jubulaceae. Lejeuneaceae are subdivided into two subfamilies, Ptychanthoideae and Lejeuneoideae, based on sporophyte characters (see below). Lejeuneoideae are further subdivided into three tribes (Gradstein 2013a).  lassification of the genera of Lejeuneaceae of Colombia C and Ecuador Subfamily Ptychanthoideae: Acrolejeunea, Archilejeunea, Bryopteris, Caudalejeunea, Dibrachiella, Frullanoides, Fulfordianthus, Lopholejeunea, Marchesinia, Schiffneriolejeunea, Spruceanthus, Thysananthus, Verdoornianthus Subfamily Lejeuneoideae Tribe Brachiolejeuneeae: Acanthocoleus, Blepharolejeunea, Brachiolejeunea, Dicranolejeunea, Lindigianthus, Neurolejeunea, Odontolejeunea, Stictolejeunea Tribe Symbiezidieae: Symbiezidium Tribe Lejeuneeae: Allorgella, Anoplolejeunea, Bromeliophila, Ceratolejeunea, Cheilolejeunea, Cololejeunea, Cyclolejeunea, Colura, Diplasiolejeunea, Drepanolejeunea, Harpalejeunea, Lejeunea, Lepidolejeunea, Leptolejeunea, Luteolejeunea, Metalejeunea, Microlejeunea, Myriocoleopsis, Pictolejeunea, Prionolejeunea, Pycnolejeunea, Rectolejeunea, Reinerantha, Schusterolejeunea, Vitalianthus, Xylolejeunea

386

 

Keys and Descriptions

Synoptic key to the subfamilies and tribes of Lejeuneaceae 1. Sporophyte “ptychanthoid”: spores isodiametric; upper ends of elaters attached to valve surface and margins, spiral bands well developed, brown; inner wall of capsule valves brownish, covered by a fenestrate thickening layer; capsule dehiscence “explosive.” Seta with 16 or more outer rows of cells. Underleaves undivided. Ventral merophyte 4 or more cells wide. Ocelli absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subfamily Ptychanthoideae 1. Sporophyte “lejeuneoid”: spores elongate-rectangular; upper ends of elaters attached to valve margins only, spiral bands usually rudimentary, pale-colored; inner wall of capsule valves pale, lacking a thickening layer (rarely with a thickening layer: in Stictolejeunea, Neurolejeunea, and Pycnolejeunea); capsule dehiscence gentle, not “explosive.” Seta with 12–16 outer rows of cells. Underleaves bifid, undivided or absent. Ventral merophyte 2–4 cells wide, rarely wider. Ocelli present or absent . . . . . . . . . . subfamily Lejeuneoideae 2 2. Seta with 12 outer rows of cells. Underleaves mostly bifid, sometimes undivided or lacking. Vegetative branches Lejeunea-type only . . tribe Lejeuneeae 2. Seta with 16 outer rows of cells. Underleaves undivided. Vegetative branches Lejeunea-type and Frullania-type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Sporophyte foot made up of 4–5 transverse rings of cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tribe Brachiolejeuneeae 3. Sporophyte foot made up of only 3 transverse rings of cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tribe Symbiezidieae  rtificial key to the genera of Lejeuneaceae of Colombia A and Ecuador 1. Leaves highly specialized, distal part forming an inflated sac (extension of the lobule) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colura 1. Leaves less specialized, distal part not forming an inflated sac . . . . . . . . . . . . 2 2. Underleaves lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Underleaves present (sometimes very small) . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Plants large, 3–5  mm wide, rheophilous, growing pendent on small shrubs along blackwater rivers on the eastern slopes of the Ecuadorian Andes. Stems very robust, ventral merophyte 10–14 cells wide. Leaves crispate-undulate. Branching Radula-type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colura irrorata 3. Plants different, stems less robust . . . . . . . Cololejeunea (and Myriocoleopsis) 4. Underleaves undivided to weakly emarginate. . . . . . . . . . . . . . . . . . . . . Key 1 4. Underleaves bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key 2 Key 1. Genera of Lejeuneaceae with undivided underleaves 1. Leaf margins toothed, at least near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 2. Ventral merophyte 4 or more cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Ventral merophyte 2(−3) cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Width of the ventral merophyte = number of epidermis cells across the ventral stem surface, in the area between where the underleaves are attached (but not in

Lejeuneaceae

387

the direct neighborhood of the underleaf base, because there the number of epidermis cells is variable) 3. Leaf cells with evenly thickened walls, trigones lacking. Lobules with a very long (5–10 cells), curved tooth. Underleaves toothed, emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fulfordianthus 3. Leaf cells with trigones, walls not evenly thickened. Lobules with or without a short tooth (less than five cells long) or with several teeth. Underleaves entire or toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Trigones cordate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Trigones not cordate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5. Underleaves toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Underleaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Plants pinnate or dichotomous, branches Frullania-type. Innovations lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bryopteris 6. Plants irregularly branched, branches Lejeunea-type. Innovations present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thysananthus amazonicus 7. Lobules with 7–9 teeth. Underleaf apex ± rounded, base auriculate. Perianth 8–10-keeled, with innovations. . . . . . . . . . . . . . . . . Frullanoides laciniatiflora 7. Lobules with 1–3 teeth. Underleaf apex emarginate, base not auriculate. Perianth 3-keeled, without innovations . . . . . . . . Caudalejeunea lehmanniana 8. Underleaf insertion line ± straight. Plants 1.5–2 mm wide. Perianth with 2 ventral keels, innovations lacking . . . . . . . . . . . . . . . . . . Lopholejeunea nigricans 8. Underleaf insertion line deeply arched. Plants usually more than 2 mm wide. Perianth without ventral keels, innovations present or absent. . . . . . . . . . . . . 9 9. Apical portion of leaf broadly recurved. Ventral merophyte 4 cells wide, epidermal cells thin-walled. Plants glossy brown . . . . . . . Lindigianthus cipaconeus 9. Leaf ± plane. Ventral merophyte more than 4 cells wide, epidermal cells thickwalled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Lobules plane, with (1–)2–4 distinct teeth. Perianth terminal on main stem or elongate branches, with 2 long innovations. . . . . . . . . . . . . . . . . . Marchesinia 10. Lobules strongly inflated-rounded, small (hidden behind the underleaves), ± without teeth. Perianth on a very short branch, with 1 short innovation or without innovation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symbiezidium dentatum 11. Underleaves 6–10× stem width, at apex short-bifid or notched. Perianth terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeunea sulphurea 11. Underleaves smaller, apex undivided (emarginate in Harpalejeunea). Perianth with keels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Plants pale green. Disciform gemmae usually produced on dorsal leaf margins. Underleaf apex truncate . . . . . . . . . . . . . . . . . . . . . Cyclolejeunea convexistipa 12. Plants brownish green. Disciform gemmae lacking . . . . . . . . . . . . . . . . . . . . 13 13. Leaves with (5–)7–25 teeth. Underleaves toothed or entire. Plants very commonly growing on living leaves, less common on bark . . . . . . Odontolejeunea 13. Leaves with 1–5 teeth. Underleaves entire. Plants growing on bark or rock, rarely on living leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

388

 

Keys and Descriptions

14. Branches predominantly Frullania-type. Female bracteole toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dicranolejeunea axillaris 14. Branches predominantly Lejeunea-type. Female bracteole entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthocoleus 15. Underleaves toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 15. Underleaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Leaves with ocelli (scattered, in a row or 1–2 ocelli near leaf base) . . . . . . . 17 16. Leaves without ocelli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 17. Ocelli in a row or 1–2 near leaf base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Ocelli scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 18. Row of ocelli 7–22 cells long. Ventral merophyte 4–6 cells wide. Lobule tooth 3–5 cells long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neurolejeunea 18. Row of ocelli 1–6 cells long. Ventral merophyte 2(−3) cells wide. Lobule tooth 1 cell long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19. Lobule fully inflated, bottle-shaped, with a long, falcate tooth. Perianth sharply 5-keeled, the keels smooth or toothed, never extending into horns. Plants usually green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harpalejeunea 19. Lobule inflated in the lower half, flattened above, rectangular, with a short, blunt tooth. Perianth 4-keeled, the keels smooth and extended into inflated horns. Plants usually brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratolejeunea 20. Ventral merophyte 4 or more cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20. Ventral merophyte 2–3 cells wide (see note in couplet 2) . . . . . . . . . . . . . . . 22 21. Underleaf apex undivided. Frullania-type branches present. Leaf apex rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stictolejeunea 21. Underleaf apex emarginate. Frullania-type branches lacking. Leaf apex acute, rarely rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidolejeunea eluta 22. Leaf cells with large trigones. Ocelli conspicuous, usually larger than green leaf cells. Epidermal cells thick-walled. Lobules curved downward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Luteolejeunea herzogii 22. Leaf cells with minute trigones. Ocelli inconspicuous, equal in size to or smaller than other leaf cells. Epidermal cells thin-walled. Lobules straight or somewhat curved upward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidolejeunea sullivantii 23. Ventral merophyte 4 or more cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. Ventral merophyte 2 cells wide (see note in couplet 2) . . . . . . . . . . . . . . . . 47 24. Trigones ± cordate. Leaves convolute when dry (rarely spreading) . . . . . . . 25 24. Trigones not cordate. Leaves mostly spreading when dry (rarely convolute) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 25. Underleaf apex emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25. Underleaf apex rounded or truncate, not emarginate . . . . . . . . . . . . . . . . . . . 27 26. Plants dark green to brown to blackish, rather robust, 2–3.5 mm wide, leaves when dry strongly convolute. Stems rigid, without hyalodermis, ventral merophyte 8 cells wide. Perianth with innovations. . . . . . Thysananthus amazonicus 26. Plants light green to pale yellowish-brown, smaller, 1.5–2  mm wide, leaves usually not convolute. Stems flaccid, with a distinct hyalodermis, ventral merophyte 4 cells wide. Perianth without innovations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caudalejeunea lehmanniana

Lejeuneaceae

389

2 7. Lobules with 3–10 teeth (the teeth sometimes inflexed) . . . . . . . . . . . . . . . . 28 27. Lobules with 0–2 teeth (occasionally 3 teeth: Thysananthus innovans) . . . . 30 28. Plants blackish in older stem portions. Perianth with 5–10 keels, with innovations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frullanoides 28. Plants not blackish. Perianths with 3–10 keels, innovations present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 29. Underleaf insertion line slightly curved, underleaf base plane. Perianth with 5–10 keels, innovations lacking. Flagelliform branches (producing caducous leaves) frequently present. Plants of lowlands (0–800 m). . . . . . Acrolejeunea 29. Underleaf insertion line deeply arched, underleaf bases folded. Perianth with 3(−4) keels, innovations present. Flagelliform branches lacking. Plants montane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachiolejeunea 30. Ventral epidermis cells not or scarcely larger than medulla cells (stem cross section). Medulla cells thick-walled. Plants brown green to blackish-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 30. Ventral epidermis cells distinctly larger than medulla cells. Medulla cells thinwalled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 31. Leaf apex acute. Female bracts and bracteole toothed. Plants of Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thysananthus amazonicus 31. Leaf apex rounded. Female bracts and bracteole without teeth. . . . . . . . . . . 32 32. Plants turning olive to reddish-brown. Innovations lacking. Female bract apices acute-acuminate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schiffneriolejeunea 32. Plants turning dark brown to black. Innovations present. Female bract apices rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thysananthus 33. Lobules mostly reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthocoleus 33. Lobules well developed, never reduced, with 2 teeth . . . . . . . . . . . . . . . . . . 34 34. First lobule tooth short, incurved and blunt, second tooth long, pointing outward, sharp. Lobule truncate. Leaves not squarrose when moist. Plants of upper montane (above 2000 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blepharolejeunea 34. Lobule teeth equal or the first tooth larger than the second. Lobule oblique or truncate. Leaves squarrose when moist. Plants of lowlands and montane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachiolejeunea 35. Median leaf cells distinctly elongate, ca. 2× longer than wide. (Plants robust. Ventral merophyte more than 10 cells wide. Lobules often reduced. Perianth with 5–8 keels) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spruceanthus theobromae 35. Median leaf cells isodiametrical or slightly elongate . . . . . . . . . . . . . . . . . . 36 36. Lobules with 2–4 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 36. Lobules with 0–1 tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 37. Leaf apex strongly and broadly recurved, acute. Epidermal cells thin-walled. Plants glossy brown, growing above 2000 m . . . . . . Lindigianthus cipaconeus 37. Leaf apex ± plane, rounded to acute-acuminate to apiculate. Epidermal cells thick-walled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38. Insertion line of underleaves deeply arched (more than 100 μm deep). Ventral merophyte 6–12 cells wide. Plants more than 2 mm wide, often black. Perianth flat, without ventral keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marchesinia

390

 

Keys and Descriptions

38. Insertion of underleaves straight or shallowly curved. Ventral merophyte 4 cells wide. Plants smaller, never black. Perianth with 2 ventral keels. . . . . . . . . . 39 39. Well-developed lobules with a large second tooth (at the lobule apex) and a much smaller first tooth (between apex and the end of the keel). Stem epidermis cells distinctly larger than medulla cells. Oil bodies 1–2(−3) per leaf cell, very large, filling up the cell lumen. Plants pale green to whitish-green to gray. Androecia on short-specialized branches, male bracteoles reduced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea (sect. Leucolejeunea) 39. Second lobule tooth not larger than the first tooth. Stem epidermis cells not or little larger than medulla cells. Oil bodies more than 3 per leaf cell, rather small, not filling up the cell lumen. Plants pale green to brown. Androecia on elongate shoots, male bracteoles not reduced . . . . . . . . . . . . . . . . . . . . . . . . Dibrachiella 40. Underleaf insertion line deeply arched, over 100 μm deep . . . . . . . . . . . . . . 41 40. Underleaf insertion line straight or curved . . . . . . . . . . . . . . . . . . . . . . . . . . 42 41. Leaf cells with small ± radiate trigones, intermediate thickenings usually present; oil bodies smooth, ± homogeneous. Lobules 1/10–1/4(−2/5)× leaf length, strongly swollen, free margin deeply inflexed. Perianth on a very short branch (appearing lateral on the stem), keels ciliate-laciniate. Plants of lowlands and montane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symbiezidium 41. Trigones usually large, not radiate, intermediate thickening absent; oil bodies coarsely granular. Lobules 1/4–1/2× leaf length. Perianth on an elongate shoot, keels smooth or lacking. Plants montane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea (sect. Omphalanthus) 42. Epidermis cells distinctly larger than medulla cells (stem cross section) . . . 43 42. Epidermis cells not or little larger than medulla cells (when in doubt try both leads) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 43. Plants pale green to whitish-green or grayish. Oil bodies 1–2(−3) per leaf cell, very large, coarsely segmented. Perianth keels smooth, innovations present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea (sect. Leucolejeunea) 43. Plants dark green to black or brown. Oil bodies 10–40 per cell, very small, homogeneous. Perianth keels ciliate-laciniate, innovations lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lopholejeunea 44. Midleaf cells 10–20 μm in diameter, trigones confluent, cell walls almost evenly thickened. Lobules flask-shaped, often darker than the leaf lobe, with a long, curved tooth made up of elongate cells. Perianth 3-keeled, the lateral keels expanded into auricles. Plants greenish-brown to blackish. . . . Neurolejeunea 44. Midleaf cells larger, trigones not confluent. Lobules not flask-shaped, not darker than the leaf lobe, tooth short or long. Perianth 4–5-keeled, the lateral keels not expanded into auricles. Plants green, brown or black . . . . . . . . . . 45 45. Innovations lacking. Free margin of lobule incurved, at least near apex. Leaves when moist ± squarrose. Plants of Amazonia, in rainforest canopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verdoornianthus 45. Innovations present. Free margin plane. Leaves when moist not squarrose. Widespread, common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

Lejeuneaceae

391

46. Lobules never reduced. Underleaves (3–)4–7× stem width, imbricate. Innovations pycnolejeuneoid (first appendage an underleaf). Plants dioicous (rarely paroicous), yellow-brown to dark brown. . . . . . . . . . . . . Archilejeunea 46. At least some lobules reduced. Underleaves 2–4× stem width, distant to subimbricate. Innovations lejeuneoid (first appendage a lateral leaf). Plants autoicous, pale green to greenish-brown to black. . . . . . . . . . . . . . . . . . . . . . Dibrachiella 47. Lobules strongly inrolled, ball-shaped, free margin inrolled 2–3× . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoplolejeunea conferta 47. Lobules not ball-shaped, free margin plane or only weakly incurved . . . . . . 48 48. Frullania-type branches present (sometimes only few). Lobules when well developed with 2 teeth. Plants greenish-brown . . . . . . . . . . . . . . . . . . . . . . . 14 48. Frullania-type branches lacking. Lobules with 1 tooth. Plants pale green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 49. Plants tiny, 0.4–0.7  mm wide. Leaf lobes suberect to obliquely spreading. Lobule tooth long-acute, consisting of a strongly elongate cell (at least 4× as long as wide). Underleaves very small, 1.5–2× stem width, orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea holostipa 49. Plants larger. Leaves wide-spreading. Lobule tooth blunt or acute, short. Underleaves larger. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeunea Key 2. Genera of Lejeuneaceae with bifid underleaves 1. Leaves highly specialized, upper part forming an inflated sac . . . . . . . Colura 1. Leaves less specialized, upper part not forming an inflated sac . . . . . . . . . . . 2 2. Plants large, 3–5  mm wide, rheophilous, growing pendent on small shrubs along blackwater rivers on the eastern slopes of the Ecuadorian Andes. Stems very robust, ventral merophyte 10–14 cells wide. Branching Radula-type. Leaves crispate-undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colura irrorata 2. Plants different, stems less robust, ventral merophyte less than 10 cells wide, branching Lejeunea-type (except innovations) . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Underleaf lobes diverging (spreading outward) . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Underleaf lobes not diverging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 4. One underleaf to each leaf, the underleaves usually imbricate. Leaf apex rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplasiolejeunea 4. One underleaf to each pair of leaves, the underleaves usually distant. Leaf apex acute-acuminate to rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Attachment of leaf lobe and lobule to the stem very short, by only 1–2 cells, leaf insertion transverse. Plants very rare, epiphyllous . . . . . . Reinerantha foliicola 5. Attachment longer, leaf insertion J-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Underleaf lobes 1 cell wide at least in the upper half, apex acute . . . . . . . . . . 7 6. Underleaf lobes broader, apex blunt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 7. Leaf apex rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Leaf apex acute or acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Lobule tooth short and blunt, not falcate. Ocellus near leaf base very large, 2.5–4× longer than adjacent leaf cells. Gynoecium without innovation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptolejeunea

392

 

Keys and Descriptions

8. Lobule tooth elongate, falcate. Ocellus near leaf base smaller, maximally 2× longer than adjacent leaf cells. Gynoecium with innovation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanolejeunea polyrhiza 9. Ocellus near leaf base very large, 2.5–4× longer than surrounding leaf cells. Lobule tooth short and blunt, not falcate. Gynoecium without innovation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptolejeunea 9. Ocellus at leaf base smaller, 1–2× longer than surrounding leaf cells, or lacking. Lobule with a long, falcate tooth (rarely short and blunt). Gynoecium usually with innovation (rarely without) . . . . . . . . . . . . . . . . . . . . . . Drepanolejeunea 10. Leaf lobes with an unbroken row of 4–8 ocelli, ocelli usually glossy yellowishbrown. Plants of Amazonian rainforest canopies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea urubuensis 10. Leaf lobes with a broken of unbroken row of (1–)2–3 ocelli in the lower half, the ocelli colorless or grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Ocelli in a broken row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanolejeunea 11. Ocelli in an unbroken row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Innovations pycnolejeuneoid. Cells on dorsal leaf surface mammillose, thickwalled, with a broad, lens-shaped papilla. Lobule with or without pre-apical tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanolejeunea 12. Innovations lejeuneoid. Cells on dorsal leaf surface smooth, thin-walled (rarely thick-walled and with a conical papilla, H. scabra). Lobule without pre-apical tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harpalejeunea 13. Underleaves very large, at least 6× wider than the stem . . . . . . . . . . . . . . . . 14 13. Underleaves smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Leaf cells uniformly thin-walled or with very small trigones, not mammillose. Lobules often reduced. Plants glistening pale green, green or whitish (when whitish, stems often long and slender, ± pendent). Oil bodies small, numerous per cell, finely granular or homogeneous. Hyaline papilla proximal to the largest lobule tooth (= first tooth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeunea 14. Leaf cells with conspicuous trigones, often mammillose. Lobules not reduced, well developed and conspicuously inflated. Plants dull pale green to olive green. Oil bodies usually large, few per cell, sausage-shaped, granular. Hyaline papilla distal to the largest lobule tooth (= second tooth) . . . . . Cheilolejeunea 15. Lobules when well developed very large, 3/5–4/5 of lobe length. Leaves usually suberect, almost parallel to the stem. Plants minute, 0.2–0.6 mm wide . . . . 16 15. Lobules smaller. Leaves spreading. Plants small or large . . . . . . . . . . . . . . . 17 16. Plants monoicous. Leaf base without ocelli. Innovations pycnolejeuneoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metalejeunea cucullata 16. Plants dioicous. Leaf base usually with one or more ocelli. Innovations lejeuneoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microlejeunea 17. Ocelli present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Ocelli lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 18. Ocelli present in leaves and underleaves . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18. Ocelli present in leaves only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 19. Ocelli in dried plants reddish or brown. Leaf cells often papillose. Caducous leaves absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pictolejeunea

Lejeuneaceae

393

19. Ocelli colorless or gray, rarely brownish (in Rectolejeunea halinae). Leaf cells smooth. Caducous leaves present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Caducous leaves present, much smaller than ordinary leaves, produced on flagelliform branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rectolejeunea 20. Caducous leaves absent or present, when present not smaller than ordinary leaves, not produced on flagelliform branches . . . . . . . . . . . . . . Lepidolejeunea 21. Underleaf lobes broadly rounded. (Ocelli 2–3 in an unbroken row in the lower half of the leaf) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 21. Underleaf lobes (sub)acute to acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Leaves deeply convex-falcate, dorsal leaf margin with several very large teeth (3–7 cells long), ventral leaf margin entire. Plants of Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanolejeunea palmifolia 22. Leaves plane or convex, margins entire or with smaller teeth. . . . . . . . . . . . 23 23. Leaves with a very large ocellus at the base, the basal ocellus 2.5–4× larger than adjacent leaf cells. Underleaf lobes subulate (1–2 cells wide), inserted at the outer edges of the underleaf. Underleaf margins bordered by 6 large cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptolejeunea 23. Basal ocellus absent or smaller, maximally 2× larger than adjacent leaf cells. Underleaf lobes wider, not inserted at outer edges of underleaf. Underleaf margins not bordered by 6 large cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 24. Plants brown or brownish-green. Walls of leaf cells ± pale brown, trigones radiate (or lacking). Branch bases sometimes with a huge, strongly inflated lobule (“utricle”). Perianth usually with inflated horns (horns sometimes absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratolejeunea 24. Plants green or whitish (rarely brown). Walls of leaf cells colorless, trigones not radiate. Utricles lacking. Perianth without inflated horns . . . . . . . . . . . . . . . 25 25. Leaf lobes with a row of 3–8 ocelli. Plants of Amazonia . . . . . . . . . . . . . . . 26 25. Leaf lobes not with a row of 3–8 ocelli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 26. Vitta 3 cells long, ± brown. Plants very small, less than 0.7 mm wide, leaves oblong, obliquely speading . . . . . . . . . . . . . . . . . . . . . Vitalianthus aphanellus 26. Vitta 4–8 cells long, glossy yellowish-brown. Plants larger, leaves ovate, widely spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea urubuensis 27. Underleaf margins denticulate, crenulate or subentire. Gemmae frequently present. Perianth flattened, 2-keeled, the keels expanded above into short auricles. Plants growing on living leaves, occasionally on bark or rock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyclolejeunea 27. Underleaf margins entire (occasionally with a small tooth). Gemmae lacking (ligulate gemmae present in Otigoniolejeunea, couplet 32). Perianth 3–5-keeled in the upper half, the lateral keels expanded or not. Plants growing on bark or rotten wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28. Caducous leaves present, very small, much smaller than oridinary leaves, produced on flagelliform branches. Leaf cells very small, 14–18 μm long in the middle of the lobe, walls thin or uniformly thickened, without trigones. Lobule small, usually with a long, falcate tooth. (Plants less than 1 mm wide; leaves wide-spreading) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rectolejeunea

394

 

Keys and Descriptions

28. Caducous leaves present or absent, when present not smaller than ordinary leaves. Leaf cells larger. Lobule tooth short, not falcate . . . . . . . . . . . . . . . . 29 29. Leaves suberect or obliquely spreading. Plants very small, 0.2–0.5(−0.7) mm wide. Ocelli grayish or brown . . . . . . . . . . . . . . . . . . . . . . . . . . Microlejeunea 29. Leaves wide-spreading. Plants more than 0.5  mm wide. Ocelli colorless or grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 30. Leaf lobe insertion very short, maximally 1/5× lobe width. Lobule attached to stem by only 2(−3) cells. (Lobule strongly swollen, ball-shaped, without tooth. Underleaves very small, 1.5–2× stem width). Plants of Amazonian lowland rainforest, not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplolejeunea amazonica Ilk.-Borg. & Gradst. 30. Leaf insertion longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 31. Leaf base with 2 transverse rows of larger, elongate cells, the upper, suprabasal row consisting of 2–5 adjacent ocelli. Underleaves distant. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xylolejeunea crenata 31. Leaf base not with 2 transverse rows of elongate cells, ocelli adjacent or scattered, 1–20 per cell. Underleaves distant or imbricate . . . . . . . . . . . . . . . . . 32 32. Leaf cells thin-walled, without or with very small trigones. Underleaves distant. Perianth apex with flat, sometimes branched lacinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Otigoniolejeunea huctumalcensis 32. Leaf cells with conspicuous trigones. Underleaves usually imbricate (rarely distant). Perianth apex without lacinia . . . . . . . . . . . . . . . . . . . . Pycnolejeunea 33. Leaf margins at least in part toothed by conically projecting cells with thickwalled tips. Dorsal leaf surface papillose or smooth. . . . . . . . . . . . . . . . . . . 34 33. Leaf margins entire or crenate or  ±  toothed, but never toothed by conically projecting cells with thick-walled tips. Dorsal leaf surface smooth . . . . . . . 36 34. Leaf base with 2 transverse rows of larger, elongate cells, the upper row made up of ocelli (but ocelli sometimes not well visible in dried material) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xylolejeunea 34. Leaf base not with 2 transverse rows of larger, elongate cells . . . . . . . . . . . . 35 35. Dorsal leaf surface spinose. Perianth inflated, with 5 keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeunea (sect. Echinocolea) 35. Dorsal leaf surface smooth or papillose, not spinose. Perianth flattened, with 2 keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prionolejeunea 36. Vegetative reproduction by means of multicellular, rounded or elongate gemmae from leaf margins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 36. No vegetative reproduction by means of multicellular gemmae . . . . . . . . . . 39 37. Gemmae orbicular, with rhizoids. Cuticle of leaf cells papillose. Innovations pycnolejeuneoid. Perianth flattened, with 2 keels . . . . . . . . . . . . Cyclolejeunea 37. Gemmae elongate, without rhizoids. Cuticle of leaf cells smooth. Innovations absent or lejeunoid. Perianth flattened or inflated, with 2–5 keels . . . . . . . . 38 38. Perianth with 2 keels. Leaf margins crenate or denticulate. . . Prionolejeunea 38. Perianth with 5 keels. Leaf margins entire or crenulate . . . . . . . . . . . . Lejeunea 39. Leaves with a very short insertion, attached to the stem by only 2–3 cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 39. Leaves with a longer insertion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

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40. Lobules flat, obliquely spreading. Underleaves longer than wide, not wider than the stem. Plants growing on rock and rotten wood along Amazonian rivers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schusterolejeunea inundata 40. Lobules inflated, wide-spreading. Underleaves wider than long, 2–3× stem width. Plants growing on living leaves in montane forest areas, rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reinerantha foliicola 41. Leaves conspicuously elongate, (1.2–)2–4× longer than wide. . . . . . . . . . . . 42 41. Leaves less elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 42. Leaf apex acute-acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanolejeunea 42. Leaf apex rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 43. Plants growing in leaf axils of bromeliads, often submerged. Midleaf cells in elongate leaves 35–70  μm long, 1.5–4× longer than wide. Perianth inflated, 5-keeled, keels not expanded into auricles . . . . . . . . . . . Bromeliophila helenae 43. Plants growing on growing on living leaves in lowland rainforest. Midleaf cells smaller, 20–30 μm long, subisodiametrical. Perianth flattened, 2-keeled, keels expanded into large auricles . . . . . . . . . . . . . . . . . . . . . . . . . Alorgella schnellii 44. Walls of leaf cells ± pale brown, trigones radiate (or lacking). Branch bases sometimes with a huge, strongly inflated lobule (utricle). Perianth usually with horns. Plants brown or brownish-green. . . . . . . . . . . . . . . . . . . Ceratolejeunea 44. Walls of leaf cells colorless, trigones not radiate. Utricles lacking. Perianth without horns. Plants green or whitish, rarely greenish-brown. . . . . . . . . . . 45 45. Trigones large or small. Lobules strongly inflated, usually not reduced (sometimes reduced in C. aneogyna). Leaf cells mammillose or not. Oil bodies large (more than 1/2 the cell lumen in length), coarsely segmented. Hyaline papilla distal to the largest lobule tooth (or between 2 small, closely associated teeth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilolejeunea 45. Trigones lacking or very small. Lobules inflated or not, frequently reduced. Leaf cells never mammillose. Oil bodies smaller (less than 1/4 the cell lumen in length), finely segmented or smooth. Hyaline papilla proximal to the largest lobule tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeunea Acanthocoleus R.M.Schust. Plants 0.8–2  mm wide, 1–4  cm long, dull green to brown, creeping. Branches Lejeunea-type, occasionally (robust vegetative shoots) Frullania-type. Stems with a hyalodermis; ventral merophyte 2(−4) cells wide. Leaf lobes wide-spreading, when dry ± convolute, apex rounded to acuminate, margins entire or toothed. Cells slightly longer than wide, trigones cordate; oil bodies finely granular. Lobules 1/4–2/3 leaf length, often reduced, with (1–)2 teeth, second tooth smaller. Underleaves undivided, margins entire, bases decurrent, insertion line shallowly curved to arched. Male bracts just below the gynoecium or on separate branches, with epistatic lobules. Gynoecia with (1–)2 pycnolejeuneoid innovations, female bracteoles entire. Perianths with (0–)2–5 keels, usually winged and ciliate-laciniate near apex, occasionally smooth. Sporophytes lejeuneoid. Vegetative reproduction not observed. A pantropical genus of eight species (Kruijt 1987), one species in Colombia and Ecuador. The main features of Acanthocoleus are (1) creeping, dull brownish-green

396

 

Keys and Descriptions

plants with a 2(−4)-cell-wide ventral merophyte; (2) leaf cells with small, cordate trigones; (3) lobules with 2 teeth; (4) undivided underleaves; and (5) gynoecia with usually 2 pycnolejeuneoid innovations and 2(−5)-keeled perianths with wingedlaciniate keels. Acanthocoleus is close to Dicranolejeunea and may be a subgenus of the latter; for differences see under Dicranolejeunea. Literature: Kruijt (1987), Gradstein et al. (1994) 1. Acanthocoleus aberrans (Lindenb. & Gottsche) Kruijt var. laevis Gradst. Fig. 40A Dicranolejeunea loxensis (Gottsche) Steph. Dicranolejeunea renauldii Steph. On bark and rock in rather dry woodlands or on isolated trees, 700–2600  m. Colombia: Ant, Boy, Cho, Cun, Hui, Mag, Met, NSa, Ris, San, Tol. Ecuador: Ga, Lo. General distribution: tropical America, Africa. For characters of the widespread Afro-American A. aberrans, see under the genus Acanthocoleus. Gradstein et al. (1994) recognized two varieties, var. aberrans characterized by a 4–5-keeled perianth with 2 sharp keels on the ventral surface and A. aberrans var. laevis Gradst. characterized by a 2(−3)-keeled perianth with a smooth ventral surface. The plants from Colombia and Ecuador belong to var. laevis. Acrolejeunea (Spruce) Steph. Plants 1–2 mm wide, yellow-green to yellow-brown, creeping. Branches Lejeuneatype, occasionally Frullania-type. Stems with a hyalodermis; ventral merophyte 4–6 cells wide. Leaf lobes wide-spreading and often ± squarrose, when dry strongly convolute, apex rounded, margins entire. Cells slightly longer than wide, trigones cordate; oil bodies homogeneous; ocelli lacking. Lobules 2/5–1/2 leaf length, never reduced, with (2–)3–8 teeth. Underleaves undivided, margins entire, insertion line shallowly curved. Male bracts rather similar to leaves, with epistatic lobules. Gynoecia without innovations. Perianths with (5–)8–10 smooth keels. Sporophytes ptychanthoid. Vegetative reproduction sometimes by caducous leaves produced on upright flagelliform shoots. A pantropical genus of 21 species (Wang et al. 2014), 2 in Colombia and Ecuador. The neotropical species of Acrolejeunea are recognized by (1) glossy green to yellowish-brown plants on bark of trees in exposed and rather dry lowland environments; (2) leaves densely imbricate and strongly convolute when dry; (3) lobules with (2–)3–8 teeth; (4) leaf cells with cordate trigones and numerous small, homogeneous oil bodies; (5) undivided underleaves; (6) gynoecia with pluriplicate perianths and no innovations; and (7) vegetative reproduction by small caducous leaves produced on tiny, upright flagelliform shoots. The strong convolution of the leaves seen in Acrolejeunea and many other genera of Ptychanthoideae is considered an adaption to drought, as most genera showing this character occur in rather dry, exposed environments, e.g., in the outer canopy of the rainforest (Gradstein 1994).

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Fig. 40 (A) Acanthocoleus aberrans, habit with gynoecia. (B–D) Acrolejeunea torulosa. (B) Habit with gynoecium and sporophyte. (C) Leaf. (D) Gynoecium. (E) Acrolejeunea emergens, leaf. (F–H) Allorgella schnellii. (F) Habit with gynoecium and androecia. (G) Habit. (H) Portion of shoot with underleaf. (I and J) Anoplolejeunea conferta. (I) Habit with gynoecium. (J) Leaf. (From Gradstein and Ilkiu-Borges 2009; Gradstein and Costa 2003; Gradstein et al. 2001)

398

 

Keys and Descriptions

Literature: Gradstein (1975, 1994), Wang et al. (2014) 1. Lobule of stem leaves with 5–8 teeth, distalmost tooth of the lobule longer than the other teeth and frequently falcate. Caducous leaves minute, much smaller than branch leaves, with a long, subapical rhizoid . . . . . . . . . . . . . A. torulosa 1. Lobule of stem leaves with 2–3(−4) teeth, distalmost tooth of the lobule not longer than the other teeth and never falcate. Caducous leaves larger, resembling branch leaves, without subapical rhizoid . . . . . . . . . . . . . . . A. emergens 1. Acrolejeunea emergens (Mitt.) Steph.

Fig. 40E

On bark in the canopy of lowland rainforest, in orchards, and on trunks of isolated trees, 150–220  m. Colombia: Cho, NSa. Ecuador (new): Or (Tiputini Biodiversity Station, on branch in outer canopy of huge Ceiba tree, Gradstein, Deleg & Bader 12812, HUTPL). General distribution: tropical America, Africa, India. Acrolejeunea emergens is a widespread, almost pantropical species that is closely related to A. torulosa but differs in the somewhat broader lobule with a semicircular, somewhat wavy free margin and only 2–3(−4) teeth, and in the less modified caducous leaves, resembling small branch leaves, without subapical rhizoids. Moreover, the keel in A. emergens usually forms an almost straight line with the ventral margin of the leaf lobe, whereas in A. torulosa the keel stands at an angle of 140–160° with the ventral margin. Occasionally the two species grow mixed. 2. Acrolejeunea torulosa (Lehm. & Lindenb.) Schiffn.

Fig. 40B–D

On bark in the canopy of lowland and submontane rainforests, in orchards, and on trunks of isolated trees, from sea level to 1000 m. Colombia: Ama, Boy, Cas, Cho, Guv, Met, Nar, Ris, San, Vau. Ecuador (new): Or (Yasuni National Park, near Yasuni Station, in the canopy of lowland rainforest in valley, 228  m, 2019, M. Berdugo et al. 18VC, QCA). General distribution: tropical South America. The main characters of A. torulosa, a common lowland species of northern tropical South America, are the lobules with 5–8 teeth and the minute caducous leaves with the lobule about as large as the lobe and with a long subapical rhizoid. The first, distalmost tooth on the lobule of stem leaves is longer than the other teeth and frequently falcate, and the free margin of the lobule is straight or slightly curved. For differences with A. emergens, see under the latter species. Allorgella Tixier Otolejeunea subg. Allorgella (Tixier) Grolle Plants small, creeping, 1–2 cm long, ca. 1–1.5 mm wide, yellowish-green when dry. Stems with hyalodermis; ventral merophyte 2 cells wide. Leaf lobes obliquely to widely spreading, oblong, ca. 2× longer than wide, apex rounded, margins entire or slightly denticulate, plane, dorsal margin curved, ventral margin straight. Midleaf cells 20–30 μm long, subisodiametrical, thin-walled, trigones small, cuticle smooth; oil bodies granular; ocelli lacking. Lobules reduced to a fold, 1/5 of leaf length, narrow-rectangular, tooth reduced, hyaline papilla on the inner side of the reduced

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tooth. Underleaves small, distant, 1.5–2.5× stem width, bifid, tips acute, margins entire or with a blunt tooth, insertion line straight. Autoicous. Androecial bracteoles restricted to the base of the spike. Gynoecia on a very short branch without innovations. Perianths ± flattened with 2 broad lateral keels expanded above into a large auricle. Sporophytes lejeuneoid. Vegetative reproduction not observed. A mainly paleotropical genus of about ten species, one species in the neotropics. Alorgella was traditionally considered a subgenus of Otolejeunea Grolle & Tixier, but molecular analysis has shown that Allorgella is a separate genus (Bechteler et al. 2016). For main features of Allorgella, see under A. schnellii. Literature: Grolle and Reiner-Drehwald (2000 as Otolejeunea), Bechteler et al. (2016) 1. Allorgella schnellii Tixier Otolejeunea schnellii (Tixier) R.L.Zhu & M.L.So

Fig. 40F–H

On living leaves in lowland rainforest, ca. 100 m. Colombia: Cho (Benavides and Sastre-de Jesús 2011); to be expected in Amazonas. General distribution: Brazil (Amazonas), Chocó. Allorgella schnellii is a characteristic epiphyllous species of the Brazilian Amazon that has been recorded once from the Chocó. The species is distinguished by (1) oblong leaves (ca. 2× longer than wide); (2) thin-walled, isodiametric leaf cells; (3) absence of ocelli; (4) lobules narrowly rectangular; (5) underleaves very small, scarcely wider than the stem, bifid; (6) gynoecia on a short branch, without innovations; and (7) perianth large, flattened, 2-keeled, keels expanded above into a large auricle. Anoplolejeunea (Spruce) Schiffn. Plants 1–1.5 mm wide, to 1–2 cm long, pale whitish-green to pale brown, creeping. Branches often microphyllous. Stems with hyalodermis; ventral merophyte 2 cells wide. Leaf lobes wide-spreading, convex, apex rounded, margins entire. Cells with small, concave-sided trigones; oil bodies finely granular; ocelli usually absent (occasionally present, 1–3 at leaf base). Lobules ca. 1/3–2/5 leaf length, highly inflated with strongly inrolled margin, occasionally reduced, tooth obscure, hyaline papilla distal. Underleaves undivided, broadly obovate-obcuneate, apex rounded to truncate, insertion line slightly curved. Gynoecia on elongate shoots or short branches, with 1(−2) pycnolejeuneoid innovations. Perianths sharply 5-keeled above. Sporophytes lejeuneoid. Vegetative reproduction not observed. A neotropical genus with a single species; for the main features of Anoplolejeunea, see under the species. Literature: Evans (1908), Gradstein et al. (2001) 1. Anoplolejeunea conferta (Spreng.) A.Evans Lejeunea herpestica Spruce

Fig. 40I, J

Sun epiphyte on twigs, branches of trees, and shrubs in montane forests and scrub in open habitats, in dense forest mainly in the canopy, also on tree trunks and

400

 

Keys and Descriptions

occasionally epiphyllous, from sea level to 3700  m. Colombia: Ama, Ant, Boy, Caq, Cau, Ces, Cho, Cun, Hui, Mag, Put, Qui, Ris, San, Tol, Vau. Ecuador: Az, Lo, MS, Na, Pi, ZC. General distribution: tropical America. Anoplolejeunea conferta is a widespread neotropical species with a remarkably large elevational range. It is readily recognized by the small, pale green to whitishgreen plants with convex leaves, undivided underleaves (obovate-obcuneate in shape) and ball-shaped lobules with a strongly inrolled free margin. The frequent presence of microphyllous branches is also characteristic of this species. Archilejeunea (Spruce) Steph. Plants 1–3.5 mm wide, glossy brownish-green, becoming brown when dry, creeping or ascending. Branches Lejeunea-type, occasionally Frullania-type. Stems simple, without hyalodermis, epidermis cells similar to medullary cells, all cell walls colorless; ventral merophyte 4–6(−8) cells wide. Leaf lobes wide-spreading, when dry not appressed or convolute, apex rounded, margins entire. Cells (sub)isodiametric, trigones simple-triangular to radiate; oil bodies coarsely granular; ocelli lacking. Lobules shortly rectangular, 1/3–1/2 leaf length, never reduced, with 0–1 teeth, free margin usually plane. Underleaves undivided, usually imbricate, with entire margins and straight or weakly curved insertion line. Dioicous, rarely paroicous (A. badia). Gynoecia with 1–2 pycnolejeuneoid innovations. Perianths (2–)4–5 keeled, the keels smooth or slightly toothed above. Sporophytes ptychanthoid. Vegetative reproduction by regeneration from leaf cells. An Afro-American genus of six species (Shi and Zhu 2015), five species in Colombia and Ecuador. Archilejeunea was traditionally split into two subgenera, subg. Archilejeunea and subg. Dibrachiella (Spruce) Schiffn. (e.g., Gradstein 1994). The latter subgenus was recently raised to generic level on molecular grounds (Shi et  al. 2015). The main features of the genus Archilejeunea, separating it from Dibrachiella, are (1) pycnolejeuneoid innovations (lejeuneoid in Dibrachiella); (2) lobules never reduced (frequently reduced in Dibrachiella), with 0–1 tooth; (3) glossy brownish-green plant color; (4) large imbricate underleaves; and (5) mostly dioicous (autoicous in Dibrachiella). Further characteristics that are also seen in Dibrachiella include the rather uniform stems without enlarged epidermal cells, the isodiametric leaf cells with simple-triangular to radiate trigones, and the dry plants with wide-spreading leaves (not appressed or convolute). Literature: Shi and Zhu (2015) 1. Leaf apex pointed, obtuse to acute. Underleaf apex emarginate . . A. nebeliana 1. Leaf apex broadly rounded. Underleaf apex rounded . . . . . . . . . . . . . . . . . . . 2 2. Ventral margin of the leaf lobe ± auriculate at the junction with the keel. Underleaf margins undulate, rarely plane . . . . . . . . . . . . . . . . A. crispistipula 2. Ventral margin not auriculate at the junction with the keel. Underleaf margins never undulate, usually recurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Plants creeping, 1–2 mm wide. Leaves mostly suborbicular, 1–1.3× longer than wide. Angle between ventral margin of the leaf lobe and the keel usually rather sharp, less than 150°. Plants very common in Amazonia . . . . . . . A. juliformis

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3. Plants ascending from the substrate, more robust, 2–3.5  mm wide. Leaves ovate-oblong, more than 1.3× longer than wide. Angle between ventral margin and keel wide, usually more than 150° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Plants paroicous (male bracts just below the gynoecium), ca. 2  mm wide. Leaves less than 2 mm long. Lobules strongly swollen, lobule tooth long, the uniseriate tip 2–5 cells long. Gynoecia always with 1 innovation. . . . A. badia 4. Plants dioicous, 2.5–3.5 mm wide. Leaves more than 2 mm long. Lobules rather flat, lobule tooth lacking or short, with a 1–2 cell long uniseriate tip. Oldest gynoecium with 2 innovations, younger ones with 1 innovation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (A. ludoviciana) 5 5. Lobule apex with a distinct tooth. Plants of the Chocó region and eastern Amazonia. . . . . . . . . . . . . . . . . . . . . . . . . . . A. ludoviciana subsp. ludoviciana 5. Lobule apex blunt, without tooth. Plants widespread in Amazonia, not in the Chocó region . . . . . . . . . . . . . . . . . . . . . . . . A. ludoviciana subsp. porelloides 1. Archilejeunea badia (Spruce) Steph. Lejeunea recurvans Spruce

Fig. 41D

Ascending to pendent on small branches in the outer canopy of Amazonian rainforest and on twigs in scrub and savanna, 200–250 m. Colombia: Ama (Campos et al. 2014), Vau (Schultes & Cabrera 17272, CINC, PC). General distribution: Amazonia. Archilejeunea badia is a rare twig epiphyte of Amazonian rainforests and of shrubs in savanna. The species is rather similar to the common A. juliformis but differs in the more elongate leaves (more than 1.3× longer than wide), strongly swollen lobules with a rather long and often curved tooth, and, especially, paroicous sexuality. The plants are usually copiously fertile, but the androecia, which are located just below the gynoecia, are rather inconspicuous and are easily overlooked. 2. Archilejeunea crispistipula (Spruce) Steph. Lejeunea crispistipula Spruce

Fig. 41B

On bark of trees in the understory of periodically inundated rainforests and upland rainforests, 100–350  m. Colombia: Ama, Caq, Cas, Mag, Met, Put, Vau (Campos et  al. 2015); not recorded from Ecuador but expected there. General distribution: Amazonia, Guyana. Archilejeunea crispistipula is a characteristic Amazonian species that is recognized by (1) plants prostrate, rather small, 1–1.4 mm wide; (2) underleaf margins undulate, rarely plane; (3) ventral margin of the leaf lobe forming a sharp angle (ca. 90°) with the keel and more or less auriculate at the junction with the keel; and (4) perianth with 2 lateral keels and 2 sharp ventral keels and sometimes with one or two additional small folds, the keels irregularly crenate-dentate. 3. Archilejeunea juliformis (Nees) Gradst. Archilejeunea fuscescens (Lehm.) Fulford Archilejeunea spruceana Steph. Marchesinia extensa (Steph.) Steph.

Fig. 41C

Fig. 41 (A) Archilejeunea nebeliana, habit with gynoecium. (B) Archilejeunea crispistipula, habit. (C) Archilejeunea juliformis, leaf and lobule apex. (D) Archilejeunea badia, leaf and lobule apex. (E) Archilejeunea ludoviciana subsp. porelloides, leaf and lobule apex. (F) Blepharolejeunea saccata subsp. harlingii, habit. (G) Blepharolejeunea incongrua, apical free margin of lobule, showing enlarged margin cells. (H) Blepharolejeunea securifolia, apical free margin of lobule. (I and J) Brachiolejeunea laxifolia. (I) Habit with gynoecia. (J) Leaf. (K) Brachiolejeunea leiboldiana, gynoecium. (L) Brachiolejeunea phyllorhiza, perianth. (M and N) Bromeliophila helenae. (M) Gynoecium, showing stalked perianth. (N) Habit. (A from Gradstein and Schäfer-Verwimp 2012; B, D, E from Gradstein and Buskes 1985; F–H redrawn from van Slageren and Kruijt 1985; L from Kruijt and Gradstein 1986; rest from Gradstein 1994; Gradstein et al. 2001)

Lejeuneaceae

403

On bark of trees in lowland rainforest, occasionally on rotten logs, 100–500 m. Colombia: Ama, Caq, Cas, Mag, Met, Put, Vau. Ecuador: Na, Or. General distribution: common and widespread in the Amazon and Orinoco basins and in the Guianas, furthermore on Trinidad, Cocos Island (Costa Rica) and in E Brazil (Pernambuco, Bahia). Archilejeunea juliformis is one of the most common epiphytic bryophytes in the Amazon Basin (e.g., Gradstein 1994; Ilkiu-Borges 2000; Gradstein and Costa 2003; Mota de Oliveira and ter Steege 2013; Campos et al. 2015; Sierra et al. 2018; all as A. fuscescens). The species is distinguished by the dioicous, glossy brownish-green, prostrate plants with suborbicular leaves, large imbricate underleaves with usually recurved margins (margins not undulate), and large lobules which are never reduced and possess a short or long tooth (1–5 cells long); occasionally there is an indication of a second, rudimentary tooth. The species has usually been called A. fuscescens, but according to Söderström et al. (2016), A. juliformis is the correct name. 4a. Archilejeunea ludoviciana (De Not.) P.Geissler & Gradst. subsp. ludoviciana Archilejeunea porelloides (Spruce) Steph. subsp. chocoensis Gradst., nom. inval. Leucolejeunea ecuadorensis Steph. Marchesinia longistipula Steph. (Homalolejeunea longistipula Steph. nom. inval., in sched.) On bark of trees in open places in lowland and submontane rainforest and scrub, from sea level to 1100  m. Colombia: Cho, Met (Gradstein and Geissler 1994). Ecuador: MS, Na, ZC (Schäfer-Verwimp et al. 2013a). General distribution: Chocó biogeographic region and at the foothills of the northern Andes. Archilejeunea ludoviciana is close to A. juliformis but is a more robust plant (2.5–3.5 mm wide) with projecting to pendent growth, more elongate leaves, and a flat lobule. Two subspecies are recognized (Gradstein and Geissler 1994), subsp. ludoviciana with a sharp lobule tooth and subsp. porelloides without lobule tooth. Both occur in Colombia and Ecuador, but their general distribution is different. Subspecies ludoviciana is distributed throughout the Chocó biogeographic region and additionally occurs in the eastern foothills of the northern Andes. Subspecies porelloides, on the other hand, is widely distributed in inner Amazonia but does not occur in the Chocó biogeographic region and is rare in the eastern foothills of the Andes. The types of Leucolejeunea ecuadorensis and Marchesinia longistipula (Gualaquiza, leg. M. Allioni) are from the province of Morona Santiago, not Azuay as mentioned on the labels (e.g., Gradstein and Geissler 1994, 1997). 4b. Archilejeunea ludoviciana subsp. porelloides (Spruce) Gradst. Archilejeunea porelloides (Spruce) Steph.

Fig. 41E

On twigs and branches and on small trunks in lowland rainforest and scrub, 100–450 m. Colombia: Ama, Hui, Put, Vau. Ecuador: MS (Gradstein and Geissler 1994). General distribution: inner Amazonia (Venezuela to Peru).

404

 

Keys and Descriptions

Recognized by the robust, Porella-like plants with ascending to pendent growth, elongate leaves and large, flat lobules with an obtuse apex, without any trace of a tooth. For differences with the typical subspecies, see under subsp. ludoviciana. 5. Archilejeunea nebeliana Gradst. & Schäf.-Verw.

Fig. 41A

On bark of thin trunks and branches of shrubs in the understory of submontane rainforest, 1000–1075 m. Colombia: Put (Gradstein et al. 2018c). Ecuador: ZC (Gradstein and Schäfer-Verwimp 2012). General distribution: northern Andes (Colombia, Ecuador). Archilejeunea nebeliana is a rare, submontane species known from only two localities in the eastern foothills of the northern Andes. The species is recognized by (1) leaf apex pointed; (2) lobule cells very thin-walled, without trigones, and almost colorless (very different from leaf lobe cells); (3) cell walls of the stem epidermis with a darker-colored middle-lamella; (4) underleaf apex emarginate; and (5) female bracts acuminate, 2.5–3× longer than wide. In other species of Archilejeunea, the apex of leaves and underleaves is broadly rounded, the lobule cells are similar to those of the lobe, the middle-lamella of the cell walls of the stem epidermis is not darker-colored, and the female bracts are less elongate, with a rounded or obtuse apex (Gradstein and Schäfer-Verwimp 2012). Blepharolejeunea S.W.Arnell Plants 1–2  mm wide, glossy green to brown or blackish, creeping or ascending, often forked. Branches Frullania-type or Lejeunea-type. Stems with a weak hyalodermis; ventral merophyte 2–4 cells wide. Leaf lobes wide-spreading and often convex, when dry convolute, apex rounded or apiculate, margins entire or toothed. Cells subisodiametric, trigones simple-triangular or cordate; oil bodies finely granular or homogeneous; ocelli lacking. Lobules ca. 1/2 leaf length, never reduced, truncate, with 2 teeth, second tooth larger than the first. Underleaves undivided, margins entire, insertion line straight or arched. Male bracts just below the gynoecium, with epistatic lobules. Gynoecia with 2 pycnolejeuneoid innovations. Perianths with 3(−5) keels, smooth or ciliate-laciniate near apex. Sporophytes lejeuneoid. Vegetative reproduction not observed. A New World genus of five species, three in Colombia and Ecuador, in two subgenera, subg. Blepharolejeunea and subg. Oreolejeunea R.M.Schust. (van Slageren and Kruijt 1985, Gradstein 1994). Blepharolejeunea is close to Brachiolejeunea and Dicranolejeunea but differs in the truncate lobule with a short and blunt first tooth (sometimes inconspicuous) and a long and sharp second tooth. The leaf margins are sharply toothed (B. saccata) or entire. Literature: van Slageren and Kruijt (1985), Gradstein et al. (1994) 1. Leaf margins ± toothed. Underleaves distant, small, 1–2× stem width. Oil bodies homogeneous. Plants dioicous, usually sterile, below 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (B. saccata) 2

Lejeuneaceae

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1. Leaf margins fully entire. Underleaves contiguous to imbricate, larger, 3–5× stem width. Oil bodies finely granular. Plants paroicous, usually fertile, growing above 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaf margins with 6–8 long teeth, the teeth up to 5 cells long. Plants of the northern Andes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. saccata subsp. harlingii 2. Leaf margins with (0–)1–4 small teeth, the teeth 1–2 cells long. Plants of the West Indies and Central America, not known in the Andes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. saccata subsp. saccata 3. Apical margin of lobule (= margin between second tooth and keel) bordered by large cells, larger than inner lobule cells. Perianth obcuneate, lateral keels rather sharp and narrow (rarely broadly rounded) . . . . . . . . . . . . . . . . . B. incongrua 3. Apical margin of lobule (= margin between second tooth and keel) not bordered by large cells. Perianth oblong, lateral keels broadly rounded . . . . . . . .  B. securifolia 1. Blepharolejeunea incongrua (Lindenb. & Gottsche) van Slageren & Kruijt Fig. 41G Brachiolejeunea nitidiuscula (Gottsche) Schiffn. On bark of trees and shrubs and rock in open high-montane environments, at forest edges, in scrub, and in páramo, 1800–4100 m. Colombia: Ant, Ara, Boy, Cas, Cun, Met, Ris, Tol. Ecuador: Ca, Co, Pi, ZC.  General distribution: Central America, tropical South America. Blepharolejeunea incongrua (subg. Oreolejeunea) is a characteristic sun epiphyte of high elevations in the Andes, forming glossy reddish-brown mats on bark or rock in open environments. The species is distinguished by the apical lobule margin bordered by large cells (larger than adjacent lobule cells) and perianths obcuneate, brownish, with ± sharp lateral keels (rarely broadly rounded) and a broadly rounded ventral keel. The lateral keels may be smooth or ciliate. 2. Blepharolejeunea saccata (Steph.) van Slageren & Kruijt subsp. harlingii (S.W.Arnell) Gradst., comb. nov.Fig. 41F Blepharolejeunea harlingii S.W.Arnell, Svensk Bot. Tidskr. 56: 335. 1962 On bark in wet lower montane rainforest and scrub, 1200–1850 m. Colombia: Ant. Ecuador: Na, Pa, Tu. General distribution: northern Andes. Blepharolejeunea saccata (subg. Blepharolejeunea) differs from the other species in the genus in the very small, distant underleaves, which are only 1–2× wider than the stem, and occurrence at much lower elevation. The plants are very fragile, the leaves are strongly falcate, the leaf apex is apiculate to acuminate and plane or recurved, and the leaf margins have 6–8 large teeth (up to 5 cells long) in the plants from the northern Andes (Ecuador, Colombia). Based on the long marginal teeth, the northern Andean plants are placed in a separate subspecies, subsp. harlingii. In the subsp. saccata, which occurs in the West Indies, Central America, and the Guayana Highland, the leaf margins are entire or with a few small, 1–2-cell-long teeth (van Slageren and Kruijt 1985).

406

 

Keys and Descriptions

3. Blepharolejeunea securifolia (Steph.) R.M.Schust. Fig. 41H Brachiolejeunea securifolia Steph. (Lejeunea securifolia Spruce, nom. illeg.) Brachiolejeunea asplundii Herzog Brachiolejeunea hans-meyeri Steph. On branches, twigs, and rock in open high-montane environments, 2450–4100 m. Colombia: Boy, Cas, Cun, Mag, Met, Ris. Ecuador: Ca, Ch, Im, MS, Na, Pi. General distribution: Central America, tropical South America. Blepharolejeunea securifolia (subg. Oreolejeunea) is very close to B. incongrua but differs from the latter in the rounded perianth keels and the unbordered lobule apical margin. Some populations of B. securifolia may be difficult to separate from B. incongrua. Blepharolejeunea securifolia resembles Brachiolejeunea laxifolia in the perianth with rounded keels (and with 2 innovations) but differs from the latter species in the truncate lobule with 2 teeth (3–4 in B. laxifolia), the underleaves with a plane apex (recurved in B. laxifolia), and the stem without a dorsal ridge. Brachiolejeunea (Spruce) Schiffn. Plants 1–3 mm wide, to 6 cm long, green or brown, creeping or ascending, often forked. Branches Frullania-type or Lejeunea-type. Stems with a hyalodermis; ventral merophyte 4–8 cells wide. Leaf lobes wide-spreading and often squarrose, when dry convolute, apex rounded to short-acuminate, margins entire. Cells slightly longer than wide, trigones cordate; oil bodies homogeneous; ocelli lacking. Lobules 1/3–1/2 leaf length, never reduced, with (1–)2–4(−10) teeth. Underleaves undivided, margins entire, bases decurrent, insertion line deeply arched. Plants paroicous or autoicous. Male bracts just below the gynoecium or on a separate branch, with epistatic lobules. Gynoecia with 1–2 pycnolejeuneoid innovations. Perianths with 3(−4) keels in the upper half, the keels usually broadly rounded and smooth, occasionally ciliate-laciniate and ± winged near apex. Sporophytes lejeuneoid. Vegetative reproduction not observed. A New World genus of six species, three in Colombia and Ecuador, in two sections, sect. Brachiolejeunea and sect. Phyllorhizae Gradst. (van Slageren 1985; Gradstein 1994). The main features of Brachiolejeunea are the greenish-brown plants with predominantly Frullania-type branches, ± squarrose leaves with 2–4(−10)-toothed lobules, cordate trigones, homogeneous oil bodies, gynoecia with winged bracts, and 3-keeled perianths subtended by (1–)2 innovations. The perianth keels are usually broadly rounded and short, being limited to the upper half of the perianth. Literature: van Slageren (1985), Gradstein et al. (1994) 1. Lobule of stem leaves with 3–4 teeth. Perianth smooth . . . . . . . . . . B. laxifolia 1. Lobule with 2 teeth (in some leaves occasionally with a small third tooth). Perianth smooth or ciliate-toothed above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Perianth ciliate-toothed above. Dorsal stem surface smooth, without ridge. Plants autoicous. Androecia on very short branches, conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. phyllorhiza

Lejeuneaceae

407

2. Perianth smooth. Dorsal stem surface with a 1 cell high longitudinal ridge. Plants paroicous (or dioicous). Androecia on main stem or long branches, inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. leiboldiana 1. Brachiolejeunea laxifolia (Taylor) Schiffn. Thysananthus mexicanus Taylor

Fig. 41I, J

On bark and rock, occasionally on living leaves, in moist, rather open montane forest and scrub, 1500–3800  m. Colombia: Ant, Boy, Cas, Cau, Ces, Cun, Hui, Mag, Met, NSa, Ris, Qui, San, Tol. Ecuador: Az, Ca, Ch, Na, Pi, Tu, ZC. General distribution: tropical America. Brachiolejeunea laxifolia (sect. Brachiolejeunea) is a widespread neotropicalmontane species that is distinguished by the 3–4-dentate lobules and by the perianths with 3 smooth, rounded keels and paired innovations. Further characteristics of the species are the recurved underleaf apices and the presence of a long, longitudinal, 2–3 cell high outgrowth (ridge) on the dorsal surface of the stem. The ridge is readily observed in stem cross section. 2. Brachiolejeunea leiboldiana (Gottsche & Lindenb.) Schiffn.

Fig. 41K

On bark in rather dry, open woodlands, and on roadside trees, 1500–1700  m. Colombia: Cal, Cas. Ecuador: Tu. General distribution: tropical America. Brachiolejeunea leiboldiana (sect. Brachiolejeunea) closely resembles B. laxifolia, but the lobules have only 2 (rarely 3) teeth and the innovations are usually single instead of paired. Moreover, the longitudinal ridge on the dorsal surface of the stem is only 1 cell high in B. leiboldiana (2–3 cells high in B. laxifolia). 3. Brachiolejeunea phyllorhiza (Nees) Kruijt & Gradst.

Fig. 41L

On bark and rock in rather dry, open woodlands, plantations, and on roadside trees, 300–2500  m. Colombia: Cho, Cun, Ris, San, Tol. Ecuador: Pi Tu, ZC. General distribution: tropical America, Africa. Brachiolejeunea phyllorhiza (sect. Phyllorhizae) is a widespread but uncommon Afro-American species that is recognized by the densely imbricate and somewhat squarrose leaves; the large underleaves with recurved margins; the broad and somewhat truncate lobules with 2 small, widely spaced teeth; the short male branches; and, especially, the ciliate-toothed perianth keels. Bromeliophila R.M.Schust. Plants 1.5–2 mm wide, pale yellow-green, turning dark brownish to black when dry, creeping or floating in water. Stems with hyalodermis; ventral merophyte 2 cells wide. Leaf lobes laxly inserted and wide-spreading, with a long insertion line (11–16 cells long), ovate to linear-lanceolate, apex rounded or acuminate, plane or recurved, margins ± sinuate to crenulate. Cells in narrowly elongate leaves 50–70 × 15–40 μm, ca. 1.5–4× as long as wide, in broad-ovate leaves shorter, completely thin-walled, without trigones, cuticles smooth; oil bodies unknown; ocelli lacking. Lobules very short, with long insertion line (9–10 cells), with (1–)2(−3)

408

 

Keys and Descriptions

short, blunt teeth, hyaline papilla proximal to the first tooth. Underleaves rather small, bifid, tips rounded or obtuse. Autoicous, with numerous short male branches and some female branches side by side. Male branches linear, with 3–10(−20) pairs of bracts. Female branches short or long, without or with 1(−2) sterile lejeuneoid innovations. Perianths inflated, 5-keeled, the keels undulate and broadly 2-winged, perianth base usually stalked. Sporophytes lejeuneoid. Vegetative reproduction unknown. A neotropical genus of two species, one in Colombia and Ecuador. For principal morphological characters of Bromeliophila, see under the species. The genus is unique among liverworts by its exclusive occurrence on the leaves of tank bromeliads but shares the same habitat with the moss Philophyllum tenuifolium (Mitt.) Broth. Literature: Schuster (1994), Gradstein (1997) 1. Bromeliophila helenae Gradst.

Fig. 41M, N

Semi-aquatic in leaf axils of tank bromeliads growing on the inner surface of the bromeliad leaves, frequently submerged, in montane environments above 1000 m, in Colombia found at 3200  m. Colombia: Ant (Benavides and Callejas 2004). General distribution: Colombia, Venezuela (Guayana Highland), Lesser Antilles (Dominica, Martinique). Bromeliophila helenae is an ususual neotropical liverwort that grows in the leaf axils of tank bromeliads. The species resembles members of the genus Lejeunea but can be separated from the latter by the conspicuously elongate, oblong leaves (but sometimes shorter, ovate) and the elongate, fully thin-walled leaf cells, without any trace of trigones. The leaf apex is rounded, and the perianths are frequently stalked and furnished with five double-winged keels. The species has been little collected and may have been overlooked. For photographs of the habit and habitat of B. helenae, see Lavocat Bernard (2018). Bryopteris (Nees) Lindenb. Bryolejeunea (Spruce) Lacout. Plants robust, 3–20 cm long, 2–4 mm wide, dull green to brown, erect from a rhizome-like shoot, (bi)pinnate or forked. Vegetative branches usually Frullaniatype, sexual branches Lejeunea-type. Stems with a dark brown cortex of thickwalled cells in 3–5 layers surrounding a colorless, thin-walled medulla, epidermis cells ca. 2× larger than subepidermis cells; ventral merophyte more than 10 cells wide. Leaf lobes wide-spreading, when dry usually convolute, apex pointed, margins toothed or entire. Cells longer than wide, trigones cordate, intermediate thickenings 0–3 per wall; oil bodies homogeneous; ocelli lacking. Lobules small, to 1/4 leaf length, never reduced, with 0–4 teeth. Underleaves undivided, quadrate to spathulate, apex sharply toothed, insertion line straight. Gametoecia on very short branches. Gynoecia without innovations. Perianths with 3 smooth keels. Sporophytes ptychanthoid. Vegetative reproduction by caducous branchlets.

Lejeuneaceae

409

A small Afro-American genus of three extant species (Hartmann et  al. 2006), two in Colombia and Ecuador. Bryopteris is readily recognized by the large, featherlike or forked habit of the plants, the Frullania-type branches, the toothed leaves and underleaves (leaves sometimes entire), the elongate leaf cells with well-defined cordate trigones, and the gynoecia on short-specialized branches lacking innovations and with a sharply 3-keeled perianth. Growth of the plants is dendroid, with a creeping rhizome and erect or pendent fronds. Bryopteris may be confused with Fulfordianthus which is similar in habit. The latter genus, however, has Lejeuneatype instead of Frullania-type branches, smaller leaf cells with uniformly thickened walls (lacking trigones) and without oil bodies, and gynoecia on elongate shoots with 2 innovations. Literature: Gradstein et al. (1994), Hartmann et al. (2006) 1. Plants dichotomous. Lobule with several large teeth. Leaf margin entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. diffusa 1. Plants regularly or irregularly pinnate. Lobule without teeth. Leaf margin usually toothed (rarely entire) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. filicina 1. Bryopteris diffusa (Sw.) Nees

Fig. 42C

On branches of trees and shrubs in rather open, submontane rainforests and woodlands, also on logs, 600–1250  m. Colombia: Qui, Ris. Ecuador: MS, ZC (Benitez and Gradstein 2011). General distribution: tropical America. Bryopteris diffusa is readily separated from B. filicina by dichotomous branching and lobules with several large, multicellular teeth. The species occurs at rather low elevations throughout the neotropics and is much less common than B. filicina. 2. Bryopteris filicina (Sw.) Nees Bryopteris fruticulosa Taylor Bryopteris galapagana Gottsche Bryopteris liebmanniana Lindenb. & Gottsche Bryopteris longispica (Spruce) Steph. Bryopteris tenuicaulis Taylor Bryopteris trinitensis (Lehm. & Lindenb.) Lehm. & Lindenb Thysananthus olivaceus S.W.Arnell (fide Sukkharak 2015)

Fig. 42A, B

Common on bark and logs in montane rainforests, cloud forests, and scrub, occasionally on moist rock, (250–)500–2150(−3000) m. Colombia: Boy, Cas, Cau, Ces, Cho, Cun, Hui, Mag, Met, Nar, Put, Qui, Ris, San, Tol, Val, Vau. Ecuador: Bo, Ch, EO (250–400 m), Es, Ga (very common), Gu, Lo, LR, MS, Na, Pa, Tu, ZC. General distribution: widespread in tropical America. Bryopteris filicina is very common, robust neotropical species that is readily distinguished by its feather-like, dendroid habit, with long, pinnately branched stems arising from a short, rhizome-like creeping shoot. The leaves and underleaves are usually toothed. A similar, feather-like habit is seen in Fulfordianthus pterobryoides, but the latter species has Lejeunea-type vegetative branches (Frullania-type in B. filicina), deeply emarginate underleaf apices (rounded to truncate in

Fig. 42 (A and B) Bryopteris filicina. (A) Habit with gynoecium. (B) Leaf. (C) Bryopteris diffusa, leaf. (D and E) Caudalejeunea lehmanniana. (D) Habit with gynoecium. (E) Leaf. (F) Ceratolejeunea brevinervis, habit. (G) Ceratolejeunea ceratantha, habit. (H) Ceratolejeunea coarina, leaf. (I) Ceratolejeunea confusa, leaf. (J and K) Ceratolejeunea cornuta. (J) Habit with gynoecium. (K) Portion of shoot, showing utricle at branch base. (L) Ceratolejeunea cubensis, stem with leaf. (M) Ceratolejeunea desciscens, leaf and underleaf. (N) Ceratolejeunea fallax, portion of stem with underleaves. (O) Ceratolejeunea filaria, portion of stem with underleaf. (P) Ceratolejeunea globulifera, portion of stem with leaf. (Q) Ceratolejeunea grandiloba, habit. (R) Ceratolejeunea guianensis, portion of stem with leaf. (S) Ceratolejeunea laetefusca, habit showing shoot partially denuded by caducous leaf lobes. (F, I, N–R redrawn from Dauphin 2003; G, M, S from Campos et al. 2014; rest from Gradstein et al. 2001; Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009)

Lejeuneaceae

411

B. filicina), lobules with a long falcate tooth (tooth lacking in B. filicina), and very small leaf cells without trigones and without oil bodies (cells with trigones and oil bodies in B. filicina). Moreover, the gynoecia in Fulfordianthus have 2 innovations (innovations lacking in B. filicina). Caudalejeunea Schiffn. Plants small- to medium-sized, 1.5–2.5  mm wide, green to brown, creeping to ascending. Branches Lejeunea-type. Stems with a hyalodermis; ventral merophyte 4 cells wide. Leaf lobes obliquely spreading and somewhat falcate, when dry spreading or appressed to stem, apex obtuse, margins entire or toothed. Cells longer than wide, trigones cordate, intermediate thickenings 1–2 per wall; oil bodies homogeneous; ocelli lacking. Lobules 1/3–2/5 leaf length, sometimes reduced, with (1–)2–3 teeth. Underleaves emarginate, margins entire, bases decurrent, insertion line curved. Gynoecia without innovations. Perianths with 3(−5) sharp keels, keels smooth or dentate. Sporophytes ptychanthoid. Vegetative reproduction by disciform gemmae from dorsal leaf surfaces. A pantropical genus (ca. 13 spp.), one species in the neotropics. The main feature of Caudalejeunea is asexual reproduction by means of disciform gemmae. The gemmae are produced on specialized gemmiparous leaves, on projecting vegetative branches. For further features of the genus, see under the species. Caudalejeunea is the only genus in the subfamily Ptychanthoideae that is frequently epiphyllous. Large adhesive rhizoid discs, produced at underleaf bases, serve as holdfasts for the epiphyllous plants. These discs are an adaptation to growth on living leaves and are lacking in corticolous plants. Literature: Gradstein et al. (1994) 1. Caudalejeunea lehmanniana (Gottsche) A.Evans Caudalejeunea harpaphylla (Spruce) Schiffn. Caudalejeunea harpaphylla fo. longiloba Spruce

Fig. 42D, E

On living leaves, twigs, and branches in lowland rainforests, usually in rather open habitats, often in secondary forests and scrub, 100–500 m. Colombia: Ant, Cho, Hui, San. Ecuador: Or (Schäfer-Verwimp et al. 2013a), Pa (Spruce 1884 as L. harpaphylla). General distribution: tropical America reaching northward to Florida, West Africa, India (Dey and Singh 2009). The elongate, obliquely spreading and somewhat falcate leaves, emarginate underleaves, elongate lobules with 2 teeth, and the sharply trigonous perianths without innovations are characteristic. Caudalejeunea lehmanniana has often been misidentified and seems to be uncommon in Colombia and Ecuador. The record from Boyacá (Gradstein and Hekking 1979) is Cyclolejeunea convexistipa; that from the Galápagos Islands (Clark 1953) is Dicranolejeunea axillaris (Gradstein 1994).

412

 

Keys and Descriptions

Ceratolejeunea (Spruce) J.B.Jack & Steph. Plants small- to medium-sized, 1–5  cm long, 0.5–2  mm wide, glossy greenishbrown to dark brown, creeping to ascending or pendent. Stems mostly with a hyalodermis; ventral merophyte 2 cells wide. Leaf lobes wide-spreading, apex rounded to acute, margins entire or toothed. Cells with elongate, triradiate trigones or uniformly thick-walled, the walls pale brown with darker brown middle lamellae, cuticle smooth, rarely elevated-papillose; oil bodies small, finely granular; ocelli present or absent, when present 1–10 near the base or in the lower half of the leaf, sometimes in a short row, rarely present in the underleaves (C. grandiloba). Lobules 1/6–1/3(−1/2) leaf length, sometimes reduced, inflated, ovate to globose, at the base of branches sometimes much enlarged, forming utricles, tooth 1-celled, the cell short and obtuse or long and sharp, hyaline papilla proximal. Underleaves bifid or undivided, small or large, insertion line shallowly curved to deeply arched. Gynoecia on long or short branches, with 1–2 pycnolejeuneoid innovations. Perianths normally with 4 keels extended above into narrow, horn-like or bulbous projections. Sporophytes lejeuneoid. Vegetative reproduction by caducous leaf lobes. A pantropical genus (ca. 37 spp.), the majority of the species in the neotropics, 18 species in Colombia and Ecuador. The neotropical species were monographed by Dauphin (2003) who recognized two subgenera, subg. Ceratolejeunea (= subg. Ceratophora R.M.Schust.) and subg. Caduciloba R.M.Schust. (= subg. Ceratolejeunea) (for nomenclature of the two subgenera, see Söderström et  al. 2016). Scheben et al. (2016), however, found that subg. Ceratophora is nested in subg. Ceratolejeunea. The main features of Ceratolejeunea are (1) plants glossy greenish-brown to brown, stem usually with a thin-walled epidermis (hyalodermis) and a thick-walled medulla, sometimes epidermis thick-walled; (2) cell walls of leaves pale brownish, with a darker pigmented middle lamella and triradiate trigones or with a continuous sheath of thickening; (3) leaf lobes usually with ocelli and with entire to toothed margins; (4) frequent presence at branch bases of “utricles” (= huge, inflated lobules with reduced lobes); and (5) perianth keels with a short-bulbous or narrow-tubular, horn-like projection (occasionally absent). Literature: Dauphin (2003), Pócs (2011), Scheben et al. (2016) 1. Underleaves undivided or very shortly bifid . . . . . . . . . (subg. Ceratophora) 2 1. Underleaves deeply bifid . . . . . . . . . . . . . . . . . . . . . . (subg. Ceratolejeunea) 4 2. Dorsal leaf margin toothed in the upper half or over its whole length. Ocelli (1–)2 per leaf, in a short unbroken row above the leaf base . . . . C. globulifera 2. Dorsal leaf margin entire or with a few small teeth near apex. Ocelli 2–20 per leaf, scattered or in an unbroken row of 2–6 ocelli . . . . . . . . . . . . . . . . . . . . . 3 3. Ocelli scattered and sometimes in a short row. Leaf apex rounded to acute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grandiloba 3. Ocelli in a row, never scattered. Leaf apex rounded (never acute) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. desciscens 4. Ocelli in a row in the leaf lobe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Ocelli single or clustered near leaf base, not in a row, or absent. . . . . . . . . . . 9

Lejeuneaceae

413

5. Ocelli in a broken row (“seriate”), row interrupted by normal green leaf cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Ocelli in an unbroken row (“moniliate”). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Underleaves small, distant, 2(−3)× stem width, base cuneate, insertion line straight or slightly curved. Leaf apex usually rounded . . . . . . . . C. ceratantha 6. Underleaves larger, contiguous to imbricate, 3–6× stem width, base cordateauriculate, insertion line arched. Leaf apex usually acute. . . . . . . . . . C. fallax 7. Leaf apex acuminate, upper leaf margin with long teeth, the teeth 1–7 cells long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pungens 7. Leaf apex rounded, upper leaf margin entire or with short teeth, the teeth 1(−2) cells long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Leaf apex plane. Leaves asymmetrical, dorsal margin strongly arched, ventral margin straight. Underleaves ± as long as wide. Ocelli in 1–2 rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rubiginosa 8. Leaf apex usually broadly recurved. Leaves ± symmetrical. Underleaves wider than long. Ocelli in 1 row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brevinervis 9. Underleaves at least in part more than 3× stem width . . . . . . . . . . . . . . . . . . 10 9. Underleaves 1.5–2.5(−3)× stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 10. Underleaf lobes acuminate, ending in a row of 2–4 cells. Leaf apex usually acute. Ocelli 0–1 at leaf base. Plants very rare (Colombia) . . . . . . C. oxygonia 10. Underleaf lobes acute or obtuse, ending in a row of 1(−2) cells. Leaf apex rounded, obtuse or subacute. Ocelli (0–)1–7 at leaf base. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Flagelliferous branches present. Underleaf bases on main stems auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. filaria 11. Flagelliferous branches absent. Underleaf bases not or scarcely auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Perianth with horns. Leaves toothed or entire. Lobules frequently dimorphic, small ovoid or larger spherical (spherical lobules not to be confused with the huge utricles at branch bases, which have reduced leaf lobes). Ocelli (0–)2–7 per leaf. Plants very common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cornuta 12. Perianth without horns. Leaves entire. Lobules never dimorphic, all lobules small ovoid. Ocelli 1–2 per leaf. Plants rare . . . . . . . . . . . . . . . . . . . C. confusa 13. Plants producing caducous leaf lobes. Plants dioicous (rarely monoicous). Perianth without horns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13. Caducous leaf lobes not produced. Plants monoicous or dioicous. Perianth with or without horns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Lobules 1/2–2/3× leaf length, flask-shaped: with a conspicuously elongate and curved-narrowed upper portion. Ventral margin forming a sharp angle with the keel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. patentissima 14. Lobules smaller, 1/5–1/3× leaf length, not flask-shaped. Ventral margin forming a wide angle with the keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laetefusca 15. Leaves conspicuously asymmetrical, dorsal margin strongly arched, ventral margin much less arched to almost straight. Dorsal leaf margin entire or toothed throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

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Keys and Descriptions

15. Leaves almost symmetrical. Dorsal leaf margin entire or toothed only at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Teeth on leaf margins large, (1–)2–5 cells long. Plants dioicous, on bark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spinosa 16. Teeth smaller, 1(−2) cells long, or absent. Plants monoicous, usually on living leaves, occasionally on bark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. coarina 17. Leaf apex mostly recurved. Microphyllous branches frequently present. Perianth without horns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. guianensis 17. Leaf apex ± plane. Microphyllous branches absent. Perianth with horns. . . 18 18. Spherical lobules lacking. Ocelli often in pairs. Oil bodies finely granular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cubensis 18. At least a few leaves with spherical lobules (not to be confused with utricles at branch bases). Ocelli not in pairs. Oil bodies smooth, homogeneous or divided into 2–5 coarse segments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cornuta 1. Ceratolejeunea brevinervis (Spruce) A.Evans

Fig. 42F

On bark of trees in montane rainforest, 1800–2150  m. Ecuador: ZC (Nöske et al. 2003). General distribution: West Indies, Ecuador. Ceratolejeunea brevinervis (subg. Ceratolejeunea) is an uncommon Caribbean species that is distinguished by the symmetrical leaf lobes with broadly recurved, rounded apices, subentire margins, and an unbroken row of 1–6 ocelli in the lower half of the leaf. The species is morphologically close to C. rubiginosa, but the leaf lobes in the latter species are conspicuously asymmetrical (dorsal margin arched, ventral margin straight) and the leaf apex is plane. 2. Ceratolejeunea ceratantha (Nees & Mont.) Schiffn.

Fig. 42G

On branches of trees in the canopy of Amazonian rainforest, 115–190  m. Colombia: Ama, Vau (Campos et al. 2014). General distribution: scattered in tropical America. Ceratolejeunea ceratantha (subg. Ceratolejeunea) is an uncommon neotropical lowland rainforest species that is recognized by leaf lobes with rounded apices and 1–3 ocelli in a broken row in the lower half of the leaf. The underleaves are small, distant, 2(−3)× stem width, with cuneate bases. 3. Ceratolejeunea coarina (Gottsche) Schiffn. Ceratolejeunea diversicornua (Prantl) Sydow Ceratolejeunea monticola (Spruce) Steph. Ceratolejeunea plumula (Spruce) Steph.

Fig. 42H

Common on living leaves in lowland rainforest, also bark and rotten wood, 100–450  m. Colombia: Ama, Caq, Cho, NSa, San, Vau. Ecuador: Es, Na, Or (Schäfer-Verwimp et  al. 2013a), Pa (Lorocachi, Jaramillo et  al. 31132, PC). General distribution: tropical America, tropical Africa. Ceratolejeunea coarina (subg. Ceratolejeunea) is a common Afro-American epiphyll (occasionally on bark) of lowland rainforests that is distinguished by (1) plants pale green to dark brown, autoicous, frequently fertile; (2) leaf lobes

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conspicuously asymmetrical (dorsal margin arched, ventral margin straight), dorsal margin usually toothed, with small teeth (1–2 cells long); (3) 1–4(−7) large ocelli in a group above the leaf base; (4) lobules ovate to spherical to flask-shaped, 0.2–0.4× leaf length; (5) underleaves distant, ca. 2× stem width, (sub)orbicular, with cuneate bases; and (6) perianths usually with four erect horns, occasionally horns poorly developed or absent (Jaramillo 31132). Scheben et al. (2016) accepted C. diversicornua from Africa (considered a synonym of C. coarina; Gradstein 2013b) as a good species but failed to study C. coarina. 4. Ceratolejeunea confusa R.M.Schust.

Fig. 42I

On bark of trees in Amazonian rainforest, 115–190  m. Colombia: Ama, Vau (Campos et al. 2014). General distribution: tropical America. Ceratolejeunea confusa (subg. Ceratolejeunea) is a little-known Amazonian species that is recognized by entire leaves; large, imbricate underleaves (3–5× stem width) with cuneate to subauriculate bases; leaves with a broadly rounded apex and entire margins; 1–2 ocelli near leaf base; small ovoid lobules; and, especially, perianths without horns. Plants without perianth are very difficult to distinguish from the common C. cornuta; for differences see under C. cornuta. A molecular analysis of the two taxa is recommended. 5. Ceratolejeunea cornuta (Lindenb.) Steph. Ceratolejeunea andicola Steph. Ceratolejeunea calabariensis Steph. Ceratolejeunea hylophila (Spruce) Steph. Ceratolejeunea litoralis Herzog Ceratolejeunea maritima (Spruce) Steph. Ceratolejeunea megalophysa (Spruce) Steph. Ceratolejeunea microrhegma (Spruce) Steph. Ceratolejeunea valida A.Evans Ceratolejeunea variabilis (Lindenb.) Pears.

Fig. 42J, K

Common on bark of trees and shrubs, sometimes on rock and living leaves, often in open environments, from sea level to 2550 m. Colombia: Ama, Ant, Boy, Caq, Cas, Cho, Cun, Mag, Met, Nar, Put, Ris, San, Val, Vau. Ecuador: Az, Ch, Es, Ga, LR, MS, Na, Or, Pa, Pi, ZC. General distribution: probably pantropical. Ceratolejeunea cornuta (subg. Ceratolejeunea) is a common and variable, widespread species that is distinguished by dimorphic lobules, which are either smallovate, ca. 1/5–1/4× leaf length, or swollen-spherical and larger, 1/3–1/2× leaf length, and furthermore by enormously variable underleaves, varying from narrowly ovate, 2–3× stem width to broadly reniform, 4–6× stem width. The variation in lobule shape and underleaf shape is seen on single stems and separates C. cornuta from the closely similar C. cubensis, which lacks this variation and has uniformly small lobules and small underleaves, 1.5–2.5× wider than the stem. A further difference between the two species is seen in the oil bodies, being homogeneous in C. cornuta and finely granular in C. cubensis (Dauphin 2003). However, oil bodies have been

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Keys and Descriptions

seen in only a few specimens, and this difference therefore needs confirmation. Ceratolejeunea cornuta is very similar to C. confusa, and sterile plants may be difficult to separate. The latter species, however, lacks the variation in underleaf and lobule shape seen in C. cornuta, underleaves of C. confusa being always large, 3–5× stem width, and lobules invariable small ovoid. Moreover, leaves of C. confusa are never toothed, and ocelli are absent or few, 1–2 (up to 7 per leaf in C. cornuta). Plants of C. confusa with perianths are readily separated from C. cornuta by the lack of perianth horns. Scheben et  al. (2016) resolved Ceratolejeunea cornuta in a well-supported molecular clade together with several paleotropical species (C. calabariensis, C. belangeriana [Gottsche] Steph., C. mascarena Steph.). Earlier, Pócs (2011) had found that C. calabariensis was a synonym of C. cornuta, whereas C. belangeriana differed from C. cornuta by the lack of spherical lobules and the presence of a longexserted perianth and was maintained as a separate species. Possibly, C. belangeriana may be treated as a subspecies or variety of C. cornuta. 6. Ceratolejeunea cubensis (Mont.) Schiffn. Ceratolejeunea amazonica Steph. Ceratolejeunea commixta (Spruce) Steph. Ceratolejeunea granatensis Steph. Ceratolejeunea parisii Steph. Ceratolejeunea tenuicornuta Steph.

Fig. 42L

On bark, rock, and rotten wood, from sea level to 1500 m. Colombia: Ama, Boy, Caq, Cho, Hui, Put, Ris, San, Val, Vau. Ecuador: Es, EO, MS, Or, Pa. General distribution: tropical America and subtropical America. Ceratolejeunea cubensis (subg. Ceratolejeunea) is a widespread neotropical species that is very common in Amazonian rainforests. The species stands out by small plant size, entire or slightly toothed leaf lobes, ocelli mostly in pairs, and small underleaves (1.5–2.5× stem width). The species is morphologically close to C. cornuta but differs from the latter in the lack of spherical lobules and consistently small underleaves. According to Dauphin (2003), C. cubensis and C. cornuta also differ in oil bodies (homogeneous in C. cornuta, finely granular in C. cubensis), but this needs confirmation based on more observations. 7. Ceratolejeunea desciscens (Sande Lac.) Schiffn. Ceratolejeunea ecuadorensis S.W.Arnell

Fig. 42M

On bark in lowland and submontane rainforest, 100–1100 m. Colombia: Caq, Put, Vau (Campos et  al. 2015). Ecuador: Pa. General distribution: northern Andes, Amazonia, Guianas, Panama. Ceratolejeunea desciscens (subg. Ceratophora) is a characteristic species of Amazonian and Guianan rainforests that is recognized by (1) plants glossy brown; (2) ventral merophyte 2 cells wide; (3) leaf apex rounded, margins entire; (4) ocelli 2–6 in a row in the lower half of the leaf; (5) lobules large, inflated, ca. 1/2× lobe length, with 1 tooth; (6) underleaves undivided, without ocelli; and (7) gynoecia usually with (0–)1 pycnolejeuneoid innovation, perianths with 4 bulbous horns.

Lejeuneaceae

8. Ceratolejeunea fallax (Lehm. & Lindenb.) Bonner Ceratolejeunea brasiliensis (Gottsche) Steph. Ceratolejeunea corniculata Steph.

417

Fig. 42N

On bark and rock in lowland and montane rainforest, 100–2000 m. Colombia: Cho, Ris. Ecuador: ES, MS, Pa. General distribution: tropical America. Ceratolejeunea fallax (subg. Ceratolejeunea) is a rather widespread neotropical species that is distinguished by the large, (sub)imbricate underleaves, 3–6× stem width, with cordate-auriculate bases and an arched insertion line. Further characters include (1) plants brown to blackish; (2) leaf lobes with acute and frequently recurved apex and toothed to entire margins; (3) ocelli few in the lower half of the leaf, isolated or contiguous, frequently in a small row; and (4) lobules small. 9. Ceratolejeunea filaria (Taylor) Steph. Ceratolejeunea fastigiata (Spruce) Steph. Ceratolejeunea sacculosa Herzog

Fig. 42O

On bark in montane forests, scrub, and isolated trees, rarely in lowland rainforest, 250–2400  m; on the Galápagos Islands forming large festoons on twigs and branches of Miconia robinsoniana at the forest line, ca. 600  m. Colombia: Ant, Cau, Ces, Cho, Ris, Val, Vau. Ecuador: Ch, EO, Es, Ga, LR, MS, ZC. General distribution: tropical America. Ceratolejeunea filaria (subg. Ceratolejeunea) resembles C. fallax in the large underleaves with cordate-auriculate bases but differs in ocelli only 0–1(−2) per leaf (never in a row) and the frequent presence of short flagelliform branches (absent in C. fallax). The species is closely related to C. cornuta (Scheben et al. 2016), but the underleaf bases in the latter species are not or only slightly auriculate, the lobules in C. cornuta vary in shape from ovate to spherical, and ocelli are usually more numerous per leaf. 10. Ceratolejeunea globulifera Herzog

Fig. 42P

On bark, 1400–1500 m. Colombia: Cau (type), Qui. General distribution: tropical Andes, Costa Rica. Ceratolejeunea globulifera (subg. Ceratophora) is a rare Andean species that is close to C. desciscens but differs in toothed dorsal leaf margins, fewer ocelli per leaf (only 1–2), and smaller underleaves, 1–2× stem width (2–4× stem width in C. desciscens). Moreover, C. globulifera occurs at higher elevation (1400–1500 m). 11. Ceratolejeunea grandiloba J.B.Jack & Steph. Fig. 42Q Ceratolejeunea szyszylowiczii (Loitl.) Steph., syn. nov. Type: Peru, Tambillo, Jelski 527 (lectotype, designated here, W; isotype, G-00121836!) On bark of trees and shrubs and on living leaves in rather open high-montane environments, 1400–3000  m. Colombia: Ant (type), Ris. Ecuador: Az, ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical Andes, Guayana Highland; a separate subspecies, C. grandiloba subsp. inflata (Mizut.) Gradst., on Java (Gradstein 2013c).

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Keys and Descriptions

Ceratolejeunea grandiloba (subg. Ceratophora) is a characteristic Andean species that is close to C. desciscens but differs in (1) ocelli scattered in leaf lobes, rarely in a row (in a row only in C. desciscens), ocasionally ocelli also present in underleaves, and (2) leaf tips plane or recurved and varying from rounded to acute (always rounded in C. desciscens). Moreover, C. grandiloba occurs at higher elevation. Plants with shortly bifid, reniform underleaves belong to C. panamensis I. Fossatti et al. (Phytotaxa 459: 248. 2020). 12. Ceratolejeunea guianensis (Nees & Mont.) Steph.

Fig. 42T

On bark in lowland rainforest and scrub, 30–250 m. Colombia: Ama, Caq, Cho, Guv, Vau. General distribution: northern South America, Cuba, Florida (?). Ceratolejeunea guianensis (subg. Ceratolejeunea) is a rare neotropical lowland rainforest species that is close to C. cubensis but differs in perianths without horns, leaf apices usually recurved, ocelli in a group of 2–6 ocelli near the leaf base, and the frequent presence of microphyllous branches. The leaf apices in C. guianensis are rounded or acute, the leaf margins are entire or with a few small teeth at the apex, and the underleaves are small, 1.5–2.5× stem width, with sharply acute and sometimes connivent lobes and cuneate to slightly rounded bases. 13. Ceratolejeunea laetefusca (Austin) R.M.Schust. Ceratolejeunea caducifolia (Spruce) Steph.

Fig. 42S

On bark and rock in open, lowland and submontane environments, from sea level to 1300(−2500) m. Colombia: Ama, Put (Campos et al. 2014). Ecuador: Es, LR, Or, ZC. General distribution: southeastern USA, tropical America. Ceratolejeunea laetefusca (subg. Ceratolejeunea) is a widespread neotropical species that is recognized by the dioicous plants with entire leaf margins, no utricles, small underleaves, perianth keels without distinct horns, and occasional presence of caducous leaf lobes. The species mostly occurs at low elevations (Dauphin 2003). 14. Ceratolejeunea oxygonia Steph.

Fig. 43A, B

On bark in river valley, ca. 1200 m. Colombia: Boy (Dauphin 2003). General distribution: Cuba (type), Colombia. Ceratolejeunea oxygonia (subg. Ceratolejeunea) is a very rare species that is known from one collection from Cuba (type) and one from Boyacá. It is distinguished by (1) plants reddish-brown, dioicous; (2) leaf lobes with acute tips, crenulate margins and 0–1 ocelli near leaf base; (3) lobules small, hidden behind the large underleaves; and (4) underleaves imbricate, 4–6× stem width, lobes conspicuously acuminate, terminating in a row of 2–4 cells, margins sinuose, bases cordate-auriculate, insertion line arched. The species approaches C. filaria, but the leaf tips in the latter species are rounded to subacute, the ocelli are more numerous (to 7 per leaf), and the underleaf lobes are acute, not acuminate.

Fig. 43 (A and B) Ceratolejeunea oxygonia. (A) HabIT. (B) Underleaf apex. (C) Ceratolejeunea patentissima, habit. (D and E) Ceratolejeunea pungens. (D) Leaf. (E) Tooth on leaf margin. (F) Ceratolejeunea rubiginosa, leaf. (G) Ceratolejeunea spinosa, leaf. (H) Cheilolejeunea acutangula, habit with gynoecium. (I) Cheilolejeunea adnata, habit in ventral view. (J) Cheilolejeunea aneogyna, portion of stem with underleaf and lobule. (K) Cheilolejeunea asperrima, leaf. (L) Cheilolejeunea beyrichii, portion of stem with underleaves. (M) Cheilolejeunea trifaria var. clausa, habit. (N) Cheilolejeunea comans, underleaves. (O and P) Cheilolejeunea filiformis. (O) Habit. (P) Leaf. (Q) Cheilolejeunea holostipa, leaf. (R) Cheilolejeunea inflexa, habit. (S) Cheilolejeunea jackii, habit. (T) Cheilolejeunea laevicalyx, habit with gynoecium. (A–G redrawn from Dauphin 2003; J from Campos et al. 2014; K redrawn from Stephani 1916; 12 redrawn from Schuster 1987; L redrawn from Reiner-Drehwald 2006; N redrawn from Reiner-Drehwald 1998; T redrawn from Zhu 2006; rest from Gradstein et al. 2001; Gradstein and Ilkiu-Borges 2009)

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Keys and Descriptions

15. Ceratolejeunea patentissima (Hampe & Gottsche) A.Evans Ceratolejeunea wallisii Steph. Harpalejeunea longilobula Herzog

Fig. 43C

On bark of trees and shrubs, rotten logs, and soil in rather open, montane environments, 450–3050 m. Colombia: Hui, Ris. Ecuador: Lo, Ma, Tu, ZC. General distribution: scattered in tropical America. Ceratolejeunea patentissima (subg. Ceratolejeunea) is a very distinct, neotropical species recognized by the peculiar, flask-shaped lobules (ca. 1/2× leaf length) and the abundance of caducous leaf lobes. Perianth horns are absent or very low in C. patentissima. 16. Ceratolejeunea pungens Steph.

Fig. 43D, E

Pendent on bark of trees in wet submontane rainforest, (200–)500–900  m. Colombia: Cho (Dauphin 2003). Ecuador: Ch, Es, LR, MS, Pa (type). General distribution: northern Andes. Ceratolejeunea pungens (subg. Ceratolejeunea) is a robust, pendent northern Andean species with strongly toothed leaf lobes (teeth to 7 cells long), a long-acuminate leaf apex, and 2–6 ocelli in an unbroken row in the lower half. The lobules are very small, and the underleaves are 2–4× stem width, with a weakly cordateauriculate base. 17. Ceratolejeunea rubiginosa Steph.

Fig. 43F

On bark and living leaves in lowland and submontane rainforests, from sea level to 1000 m. Colombia: Cho (Uribe and Gradstein 1998). General distribution: West Indies, Florida, Costa Rica, Colombia. Ceratolejeunea rubiginosa (subg. Ceratolejeunea) is an uncommon Caribbean species that is morphologically close to C. brevinervis; for differences see under the latter species. 18. Ceratolejeunea spinosa (Gottsche) Steph. Ceratolejeunea platyceras (Spruce) Steph.

Fig. 43G

On bark of trees in wet lowland and montane rainforest and cloud forest, from sea level to 1600 m. Colombia: Cau, Cho, Nar, Ris, Val. Ecuador: Ch, EO, Es, Pa. General distribution: tropical America. Ceratolejeunea spinosa (subg. Ceratolejeunea) is a characteristic neotropical rainforest species (mostly montane) that is recognized by (1) plants often reddishbrown in color, dioicous; (2) leaf lobes conspicuously asymmetrical (dorsal margin arched, ventral margin straight), with strongly toothed margins, the teeth 2–5 cells long; and (3) underleaves rather small, longly ovate (2–3× stem width), lobes acuteacuminate, margins subentire, bases cuneate to rounded to subauriculate. The record of C. spinosa from the Galápagos Islands (Clark 1953) is probably a misidentification.

Lejeuneaceae

421

Cheilolejeunea (Spruce) Steph. Aureolejeunea R.M.Schust., Cyrtolejeunea A.Evans, Cystolejeunea A.Evans, Euosmolejeunea (Spruce) Steph., Leucolejeunea A.Evans, Omphalanthus Lindenb. & Nees, Omphalolejeunea (Spruce) Lacout., Peltolejeunea (Spruce) Schiffn., Potamolejeunea (Spruce) Lacout., Strepsilejeunea (Spruce) Schiffn., Trachylejeunea (Spruce) Steph. Plants small to large, 0.6–8 cm long, 0.4–2.5 mm wide, usually dull pale green to olive green to glossy brown, sometimes whitish, gray or yellowish-green, creeping to ascending, occasionally pendent. Stems usually with or without enlarged epidermis cells, cells ± thick-walled; ventral merophyte 2–12 cells wide. Leaf lobes suberect to widely spreading, convolute or spreading when dry, convex, apex rounded to acute to acuminate plane or recurved, margins entire or sinuate by mammillose or tuberculate cells, rarely toothed. Cells plane or mammillose and sometimes with a broad, lenticular papilla, trigones small to large, intermediate thickenings scarce or lacking; oil bodies usually large, sometimes filling the cell lumen, 1–5 per cell, sausage-shaped, coarsely granular (small and finely granular in C. holostipa); ocelli mostly lacking. Lobules 1/10–1/2(−3/4) leaf length, usually not reduced, ovate to globose or rectangular, strongly inflated with plane or incurved free margin, usually with one tooth, 1–6 cells long (= second tooth; the first tooth is reduced) and a distal hyaline papilla; occasionally with 2 closely associated teeth and the hyaline papilla positioned between the teeth (in former genus Trachylejeunea). Underleaves small or large, bifid or undivided, margins entire, insertion line straight to deeply arched. Gynoecia on long or short shoots, with 1(−2) lejeuneoid or pycnolejeuneoid innovations, ocasionally without innovations. Perianths with 3–5(−10) smooth keels or flattened and with only 2 keels or terete, with or without beak. Sporophytes lejeuneoid. Vegetative reproduction rare, by caducous leaf lobes. A large pantropical genus (ca. 125 spp.), 30 species in Colombia and Ecuador, in at least 8 sections (Ye et al. 2015). The genus Cheilolejeunea is broadly defined here following Ye et al. (2015) and others, who showed based on molecular evidence that several holostipous genera (Aureolejeunea, Leucolejeunea, Cyrtolejeunea, Cystolejeunea, Omphalanthus) are nested in Cheilolejeunea. The main features of Cheilolejeunea are (1) stems with or without hyalodermis, ventral merophyte 2–12 cells wide; (2) leaf margins usually entire; (3) leaf cells with small or large trigones, mostly without intermediate thickenings, walls without brownish pigmentation; (4) oil bodies large, few per cell, coarsely granular (rarely finely granular: sect. Cyrtolejeunea); (5) ocelli usually absent (present in C. urubuensis and C. vittata); (6) lobules inflated, rarely reduced, with distal hyaline papilla (first tooth normally reduced); (7) underleaves undivided or bifid, when bifid with upright lobes; and (8) vegetative reproduction rare, by caducous leaf lobes. Literature: Schuster (1980a, 1987 as Aureolejeunea), Reiner-Drehwald (1998), Ye et al. (2015), Bastos (2017), Bastos and Gradstein (2020) 1. Underleaves undivided. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaves bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

422

 

Keys and Descriptions

2. Plants very small, ca. 0.4–0.7  mm wide, ventral merophyte 2 cells wide. Underleaves small, distant, insertion line straight. Leaf lobes suberect to obliquely spreading. Lobule tooth consisting of 1 strongly elongate cell (at least 4× as long as wide) . . . . . . . . . . . . . . . . . . . C. holostipa (sect. Cyrtolejeunea) 2. Plants larger, more than 1  mm wide, ventral merophyte 4–12 cells wide. Underleaves large, contigsuous to imbricate, insertion line arched. Leaf lobes widely spreading. Lobule tooth consisting of 1, or several, short cells. . . . . . 3 3. Plants creeping, pale colored. Lobule with a tooth (1–5 cells long). Ventral merophyte 4 cells wide . . . . . . . . . . . . . . . . . . . . . . . . (sect. Leucolejeunea) 4 3. Plants creeping, ascending to pendent, pale colored to brown. Lobule without distinct tooth. Ventral merophyte (2–)4–12 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (sect. Omphalanthus) 5 4. Ventral margin of leaf lobe strongly revolute . . . . . . . . . . . . . . C. xanthocarpa 4. Ventral margin of leaf lobe ± plane . . . . . . . . . . . . . . . . . . . . . . . . . C. unciloba 5. Lobules maximally 1/3× lobe length, subquadrate-trapezoid. Plants dioicous, yellowish-green to pale brown, never dark brown (former genus Omphalanthus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Lobules 2/5–1/2× lobe length, rectangular. Plants autoicous, yellowish-green to dark brown (former genus Aureolejeunea) . . . . . . . . . . . . . . . . . . . . . . . . . . 10 6. Underleaves orbicular or wider than long. Leaf apex rounded. Perianth usually terete and fully smooth, rarely keeled or angled . . . . . . . . . . . . (C. filiformis) 7 6. Underleaves clearly longer than wide. Leaf apex obtuse to acuminate. Perianth 5-keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7. Perianth terete. Plants very common. . . . . . . . . . . . C. filiformis var. filiformis 7. Perianth obtusely angled or sharply keeled, not terete. Plants rare. . . . . . . . . 8 8. Perianth obtusely 4-angled . . . . . . . . . . . . . . . . . . C. filiformis var. platycoleus 8. Perianth sharply 5-keeled . . . . . . . . . . . . . . . . . . . . . . . C. filiformis var. wallisii 9. Leaf apex acuminate. Underleaves undivided to short-bifid . . . . . . . . C. jackii 9. Leaf apex obtuse. Underleaves undivided . . . . . . . . . . . . . . . . . . . . . . C. ovalis 10. Dorsal surface of leaf cells mammillose-papillose, the cells distinctly elevated and the outer wall thickened by a tuberculiform papilla . . . . . . . . . C. aurifera 10. Dorsal surface of leaf cells smooth or mammillose, not papillose . . . . . . . . 11 11. Perianth with 2–5 narrow folds on the ventral surface and 1–3 narrow folds on the dorsal surface. Plants orange-brown to dark brown, ascending to pendent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. paramicola 11. Perianth sharply 5-keeled, or with 3–4 broadly rounded keels in the upper half (rarely without keels). Plants pale-colored to dark brown, creeping to ascending, not pendent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Ventral merophyte 4–6 cells wide. Perianth with 3–4 low, rounded keels in the upper half, keels sometimes lacking. Plants of montane forest and subpáramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tonduzana 12. Ventral merophyte 2 cells wide. Perianth sharply 5-keeled. Plants of páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. quinquecarinata 13. Leaf lobes with a conspicuous, unbroken row of 4–8 ocelli, the ocelli usually yellowish-brown. Plants of Amazonian rainforest canopies . . . . C. urubuensis

Lejeuneaceae

423

1 3. Leaf lobes without ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Leaf margins toothed. (Lobule keel with large papillae. Underleaves small, distant). Plants from Panama (on tree in secondary forest); not known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . C. norisiae G.Dauphin & Gradst. 14. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15. Underleaves shallowly bifid to maximally 1/5(−1/4) of underleaf length, large, 4–7× stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15. Underleaves more deeply bifid [when in doubt try both leads] . . . . . . . . . . . 21 16. Leaf apex acute-apiculate to acuminate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Leaf apex rounded to obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Underleaves 2–3× longer than wide, bases cuneate. Leaves ovate-lanceolate, apex acuminate, plane. Plants from submontane forests in western Ecuador and the Galapagos Islands, below 1500 m . . . . . . . . . . . . . . . . . . . . . . . . . C. jackii 17. Underleaves wider than long, bases rounded. Leaves broadly ovate, apex acuteapiculate, recurved. Plants from upper montane forest and páramo, above 2500 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. asperrima 18. Ventral merophyte 4–7 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18. Ventral merophyte 2 cells wide . . . . . . . . . . . . . . . . . . . . . . . C. fragrantissima 19. Lobule apex conspicuously curved downward, reaching the ventral margin of the lobe (or even passing beyond the ventral margin). Lobule apical margin (= margin between apex and ventral lobe margin) very short. Hyaline papilla proximal to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . Pycnolejeunea decurviloba 19. Lobule apex not conspicuously curved downward. Lobule apical margin long. Hyaline papilla distal to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Underleaf apex strongly recurved. Flagelliform branches usually present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. revoluta 20. Underleaf apex plane. Flagelliform branches absent. . . . . . . . . . . . C. beyrichii 21. Lobules globose, strongly swollen, free margin strongly involute. Perianths flat, with only 2 keels, without beak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lineata 21. Lobules not globose, swollen or not, free margin plane or incurved. Perianths with 4–5 smooth keels and with a beak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Lobule decurved, apex almost touching the ventral margin of the lobe. Lobule apex with 2 tightly associated teeth. Plants from Amazonia. . . . C. neblinensis 22. Lobule not decurved, apex not almost touching the ventral lobe margin. Lobule apex usually with 1 tooth (with 2 tightly associated teeth in C. aneogyna, couplets 29, 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23. At least some leaf apices obtuse or acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. All leaf apices broadly rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 24. Keel with large, 7–14 μm high, tubercle-like papillae. . . . . . . . . . . . . . . . . . 25 24. Keel without papilla or with low, to maximally 3  μm high, lens-shaped papillae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25. Leaf apex rounded to obtuse to apiculate, plane or recurved. Underleaves small, 1.5–2.5× stem width, bases cuneate, insertion straight . . . . . . . . . . . C. ornata 25. Leaf apex sharply acute-acuminate, recurved. Underleaves larger, bases rounded, insertion arched. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. inflexa

424

 

Keys and Descriptions

26. Ventral merophyte 4–8 cells wide. Lobules very small, ca. 1/6–1/5× leaf length. Gynoecia without innovations. Plants from Venezuela, not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . C. valenciae (Steph.) X.L.He 26. Ventral merophyte 2(–4) cells wide. Lobules small or large. Gynoecia with innovation(s) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27. Underleaf bases (sub)auriculate. Plants growing below 2000 m . . . . C. comans 27. Underleaf bases cuneate to rounded, not auriculate . . . . . . . . . . . . . . . . . . . . 28 28. Underleaves 1.5–2.5× stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 28. Underleaves 2.5–5× stem width [when in doubt try both leads]. . . . . . . . . . 30 29. Leaf cells mammillose and with a low lens-shaped papilla on the outer, bulging wall; trigones well developed. Lobules never reduced, apex with 1 tooth. Microphyllous branches and caducous leaf lobes lacking. Plants autoicous, mostly montane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lobulata 29. Leaf cells not mammillose, without lens-shaped papilla; trigones lacking or small. Lobules sometimes reduced, apex with 2 very small, paired teeth (teeth very closely side by side and sometimes overlapping). Plant sometimes with numerous microphyllous branches or with caducous leaf lobes. Plants dioicous, in lowlands (100–200 m). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aneogyna 30. Leaf cells with small, distant trigones. Dorsal leaf surface mammillose but not papillose, outer cell wall thin. Perianth without keels, or with 3–4 low and  broadly rounded keels. Innovations lejeuneoid. Plants growing above 2500 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laevicalyx 30. Leaf cells with large, contiguous to confluent trigones. Dorsal leaf surface mammillose-papillose, outer cell wall thickened. Perianth sharply 5-keeled. Innovations lejeuneoid or pycnolejeuneoid. . . . . . . . . . . . . . . . . . . . . . . . . . 31 31. Innovations lejeuneoid. Apices of underleaf lobes rounded to obtuse. Leaves usually strongly convex. Plants growing at high elevation (2150–4500 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. choachina 31. Innovations pycnolejeuneoid. Apices of underleaf lobes subacute. Leaves weakly convex. Plants growing below 2500 m . . . . . . . . . . . . . . C. acutangula 32. Lobule apex with a pair of short, blunt teeth (to be carefully studied, the 2 teeth positioned very closely side by side and often overlapping each other). Plants frequently with caducous microphyllous branches, in Amazonian rainforests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aneogyna 32. Lobule apex with only 1 tooth, not with a small pair of teeth. Plants without caducous microphyllous branches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 33. Plants producing caducous leaf lobes (dioicous). Leaf cells usually smooth, ± thin-walled, with small trigones (mammillose and with large trigones in C. clausa). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33. Plants not producing caducous leaf lobes (autoicous or dioicous). Leaf cells smooth or mammillose, with small or large trigones . . . . . . . . . . . . . . . . . . . 36 34. Underleaves imbricate, 4–6× stem width, orbicular. Lobule with a short, blunt tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. trifaria var. clausa 34. Underleaves distant, 1.5–3× stem width. Lobule with a long and sharp tooth (or short and blunt: C. aneogyna, couplet 32). . . . . . . . . . . . . . . . . . . . . . . . . . . 35

Lejeuneaceae

425

35. Ventral merophyte on main stems 4–6 cells wide. Caducous leaf lobes produced on upright flagelliform shoots with densely imbricate underleaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. schiavoneana 35. Ventral merophyte on main stems 2 cells wide. Caducous leaf lobes produced on ordinary leafy shoots with distant underleaves, upright flagelliform shoots lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. adnata 36. Leaves suberect to obliquely spreading. Underleaves very small, less than 2× stem width, distant. Leaf cells very thin-walled, trigones minute. Innovations pycnolejeuneoid. Plants very small, to 0.6  mm wide, autoicous, found in Brazil,  the West Indies and Costa Rica; not yet known from Colombia and Ecuador . . . . . . . . . C. discoidea (Lehm. & Lindenb.) R.M.Schust. & Kachroo 36. Leaves widely spreading. Underleaves larger, 2–6× stem width, distant to imbricate. Trigones small or large. Innovations lejeuneoid (rarely pycnolejeuneoid, C. intertexta). Plants larger, autoicous or dioicous. . . . . . . . . . . . . . . 37 37. Underleaves 2–3.5× stem width, distant to contiguous. Underleaf bases cuneate to slightly rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 37. Underleaves larger, (3.5–)4–6× stem width, (sub)imbricate. Underleaf bases broadly rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (C. trifaria) 39 38. Plants autoicous or paroicous, always fertile, very rare, found once in Colombia. Innovations pynolejeuneoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. intertexta 38. Plants dioicous, sterile or fertile, very common in lowland rainforests, less common in montane forests. Innovations lejeuneoid . . . . . . . . . . . C. rigidula 39. Underleaves usually wider than long, 1.0–1.5× wider than long, rarely orbicular. Plants autoicous, usually fertile, very common. . . . C. trifaria var. trifaria 39. Underleaves orbicular. Plants dioicous, mostly sterile, not common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. trifaria var. clausa 1. Cheilolejeunea acutangula (Nees) Grolle Fig. 43H Jungermannia acutangula Nees (Strepsilejeunea acutangula [Nees] Schiffn.). Type: Brazil, Minas Gerais, Martius s.n., c. gyn. (lectotype, designated here, STR-Nees!; isolectotype PC-Mont.-MB1668!). Syntype: Brazil, Rio de Janeiro, Serra dos Orgaos, Mandiocca, Martius s.n. (isosyntype M, fide Grolle 1979) Harpalejeunea puelensis Steph. Lejeunea kunthiana Lindenb. (Strepsilejeunea kunthiana [Lindenb.] Schiffn.). Type: Rio de Janeiro, Gaudichaud s.n. (isotypes PC-0103866!, PC-0103867!, PC-0103868!) On tree trunks and on branches of trees and shrubs in rather open, montane environments, also on rock and living leaves, 900–2100 m. Colombia: Ant, Boy (Bastos 2017), Hui, Mag, Ris. Ecuador: EO, ZC.  General distribution: scattered in tropical America. Cheilolejeunea acutangula (sect. Strepsilejeunea) is an uncommon neotropicalmontane species that is recognized by (1) plants autoicous, 0.6–0.9 mm wide, ventral merophyte 2 cells wide; (2) leaves slightly convex, apex acute to short-acuminate, terminating in a row of 1–2 cells, plane or recurved; (3) leaf cells with large, swollen, confluent trigones (cell lumen angular to somewhat stellate), dorsal outer wall

426

 

Keys and Descriptions

usually mammillose and thickened, with a low, lenticular papilla; (4) lobules ovate to short-rectangular, 1/3–2/5× lobe length, fully inflated, tooth long, acute, 1-celled, keel smooth or crenate by mammillose cells, forming a sharp angle with the ventral margin of the leaf lobe; (5) underleaves ovate-orbicular, 2.5–4× stem width, bifid to 2/5 with a wide V-shaped sinus, apices obtuse to subacute, bases cuneate to somewhat rounded (not auriculate); (6) androecia with bracteoles limited to the base; and (7) gynoecia with 1–2 pycnolejeuneoid innovations, perianth sharply 5-keeled, with a beak. Cheilolejeunea acutangula is morphologically close to C. choachina and C. erostrata R.M.Schust., but the latter two species grow at higher elevation (usually above 2500 m); moreover, C. choachina differs in having lejeuneoid innovations and more strongly convex leaves, while C. erostrata (Venezuela; see Bastos 2016) has larger underleaves (4–5× stem width), larger lobules with a reduced tooth, and 3-keeled perianths without a beak. Cheilolejeunea acutangula is also close to C. lobulata, but the latter species is a more delicate plant with smaller underleaves and obtuse leaf tips. Cheilolejeunea acutangula was described by Nees based on two collections from Brazil, one from Minas Gerais, and one from Rio de Janeiro. Only the syntype from Minas Gerais is kept in the Nees herbarium (STR), which is selected here as the lectotype. Bastos (2017) suggested that Grolle (1979) had selected both syntypes as a single lectotype but Grolle did not propose a lectotype; moreover, Grolle did not study the original material in the Nees herbarium. 2. Cheilolejeunea adnata (Lehm.) Grolle Cheilolejeunea decidua (Spruce) A.Evans

Fig. 43I

On bark of trees in lowland and lower montane rainforests, from sea level to 1800 m. Colombia: Ama, Cho, Hui. Ecuador: Ch, Ga (?), MS, Or, Pa. General distribution: tropical America, extending northward to Florida. Cheilolejeunea adnata (sect. Cheilolejeunea) is a widespread neotropical species that is distinguished by (1) plants dioicous, small, 0.7–1 mm wide, pale green, stems fragile, ventral merophyte 2 cells wide; (2) leaves caducous, widely spreading to somewhat falcate, ovate, apex broadly rounded, margins often with short rhizoids; (3) leaf cells with very small trigones; (4) lobules 1/4–1/3× leaf length, never reduced, bottle-shaped (conspicuously constricted toward the mouth), with a long and sharp tooth; (5) underleaves distant, small, scarcely wider than the stem (1–2× stem width), bifid, orbicular to obovate, bases cuneate, insertion line straight; (6) androecia with bracteoles limited to the base; and (7) gynoecia without or with 1 short innovation, perianth somewhat flattened, 3-keeled with a broad ventral keel, lateral keels somewhat expanded beyond the apex, beak short, recessed. The record of C. adnata from the Galápagos Islands (Clark 1953; Arnell 1962) probably refers to Lejeunea deplanata, a very common species in the Galápagos (Gradstein 2009 as Rectolejeunea maxonii).

Lejeuneaceae

3. Cheilolejeunea aneogyna (Spruce) A.Evans  Trachylejeunea aneogyna (Spruce) Grolle Cheilolejeunea assurgens (Spruce) Steph. Potamolejeunea riparia (Spruce) Steph. Pycnolejeunea surinamensis Steph. Rectolejeunea heteroclada (Spruce) Steph.

427

Fig. 43J

On bark in the canopy of lowland rainforests; in scrub, gardens, and plantations; in rather open environments, 100–250 m. Colombia: Ama, Caq, Put, Vau (Campos et  al. 2014). Ecuador: Az (Schäfer-Verwimp et  al. 2013a). General distribution: widespread in Amazonia, scattered elsewhere in tropical America. Cheilolejeunea aneogyna (sect. Trachylejeunea) is a common Amazonian species that resembles C. rigidula but differs from the latter in the frequent presence of caducous leaves or caducous microphyllous branches, thin-walled leaf cells without or with very small trigones, lobules sometimes reduced (never reduced in C. rigidula), and lobule apex with a pair of 2 very small, closely associated teeth. The paired teeth are sometimes difficult to observe because the lobule apex is often incurved and the teeth may be overlapping each other. The underleaves in C. aneogyna are small (1.5–2.5× stem width), and the tips of the underleaf lobes are usually blunt. Cheilolejeunea assurgens (Spruce) Steph. was accepted as a separate species close to C. aneogyna by Reiner-Drehwald and Grolle (2012) but is placed here in the synonymy of C. aneogyna following Bastos (2012). 4. Cheilolejeunea asperrima (Steph.) Grolle Taxilejeunea asperrima Steph.

Fig. 43K

On trunks and branches in upper montane forest and páramo, 2500–3350  m. Colombia: Boy (J. Gil, pers. comm.), Mag (Grolle 1985); to be expected in Ecuador. General distribution: tropical Andes (Venezuela, Colombia, Bolivia). Cheilolejeunea asperrima (sect. Strepsilejeunea) is a rare high-Andean species that is distinguished by (1) plants dioicous, robust, ventral merophyte 4–6 cells wide; (2) leaves broadly ovate, apex acute to apiculate, recurved; (3) leaf cells mammillose-papillose, with huge trigones; (4) lobules ca. 1/3× leaf length, with a sharp tooth; and (5) underleaves shallowly bifid (to 1/10–1/5), broad, ca. 5× stem width, apices rounded, bases broadly rounded (but not auriculate) and ± folded, insertion line deeply arched (Bastos 2017). In general habit C. asperrima approaches the members of sect. Omphalanthus. 5. Cheilolejeunea aurifera (R.M.Schust.) W.Ye, R.L.Zhu & Gradst. Aureolejeunea aurifera R.M.Schust. On shrubs in páramo, ca. 3500–4160 m. Colombia: Ara (Cocuy, Cleef 10373, U), Cun (Sumapaz, Cleef 1697, 1700, U). General distribution: northern Andes, Costa Rica, Dominican Republic. Cheilolejeunea aurifera (sect. Omphalanthus) is a northern Andean-Costa Rican species that is closely similar to C. quinquecarinata but differs in having mammillose leaf cells crowned by a low lenticular papilla (papilla lacking in C. quinquecarinata) and a 4-cell-wide ventral merophyte (2 cells wide in C. quinquecarinata).

428

6. Cheilolejeunea beyrichii (Lindenb.) M.E.Reiner Euosmolejeunea beyrichii (Lindenb.) Steph. Cheilolejeunea rupestris C.J.Bastos & Gradst.

 

Keys and Descriptions

Fig. 43L

On bark and rock in montane forest areas, 1000–2500 m. Colombia: Boy, Hui. Ecuador: Az (Reiner-Drehwald 2006), ZC. General distribution: Guatemala, northern Andes (Venezuela to Ecuador), Guayana Highland, Brazil (Bastos 2017). Cheilolejeunea beyrichii (sect. Euosmolejeunea?) is an uncommon neotropical species that is distinguished by large, reniform underleaves (4–6× stem width) with a shallow notch at the apex; leaves with recurved, obtuse apices; small lobules with a short tooth; and stems with a 4–6-cell-wide ventral merophyte. The species is very similar to C. fragrantissima and the two may be easily confused; for differences see under C. fragrantissima. 7. Cheilolejeunea choachina (Gottsche) Gradst. Strepsilejeunea choachina (Gottsche) Steph. On shrubs and soil in upper montane forest and páramo, 2150–4500  m. Colombia: Ara, Boy, Cal, Cun, Ris, Tol. Ecuador: Lo, Pi, ZC. General distribution: northern Andes. Cheilolejeunea choachina (sect. Strepsilejeunea) is a characteristic northern Andean species that is recognized by (1) plants usually monoicous, ventral merophyte 2 cells wide; (2) leaves usually strongly convex, apex recurved, obtuse to acute; (3) leaf cells with large, often confluent trigones; (4) underleaves 3–5× stem width, bifid, ± obovate, tips rounded to obtuse; and (5) androecia with bracteoles limited to the base, gynoecia with 1–2 lejeuneoid innovations, perianth sharply 5-keeled. The species is morphologically close to C. acutangula and C. laevicalyx, but C. acutangula has pycnolejeuneoid innovations and occurs at lower elevation, while C. laevicalyx has more thin-walled leaf cells with small, distant trigones, leaves only slightly convex, and perianths terete or with 3–4 low, broadly rounded keels. Cheilolejeunea invaginata R.M.Schust. from Venezuela is very similar to C. choachina and may be a synonym; for a description of the species, see Bastos (2016, 2017). 8. Cheilolejeunea comans (Spruce) R.M.Schust.  Fig. 43N Lejeunea comans Spruce. Type: Venezuela, San Carlos del Río Negro, “in arborum ramis,” Spruce s.n. (lectotype, designated here, MANCH-cc19029!; isolectotype JE-H2370, fide Grolle 1988) On tree trunks and branches in lowland and montane rainforests, from sea level to 2000  m. Colombia: Boy, San (Campos and Uribe 2006). Ecuador: Ch, Tu, ZC. General distribution: tropical South America, Dominican Republic. Cheilolejeunea comans (sect. Strepsilejeunea) is an uncommon neotropical species that is distinguished by (1) plants monoicous or dioicous, ventral merophyte 2 cells wide; (2) leaf apex broad, rounded to short-apiculate by 1 cell, plane; (3) leaf cells with large, swollen trigones (but cell lumen oval, not angular), smooth or slightly convex, outer wall thin; (4) lobules small, 1/5–1/4× lobe length, fully

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inflated, with a short blunt tooth, keel smooth, ± straight, forming a wide angle with the ventral margin of the leaf lobe; (5) underleaves ovate-orbicular, 3–4× stem width, bifid to 1/3 with a rather narrow V-shaped sinus, apices obtuse to acute by 1 cell, insertion line arched, bases (sub)auriculate (auricles ca. 50 μm high); and (6) androecia with bracteoles limited to the base, gynoecia with 1–2 lejeuneoid innovations, perianth sharply 5-keeled. Because the type material of C. comans is purely male, the innovation type cannot be determined on the type specimen. Cheilolejeunea comans may be confused with C. acutangula, but the latter species differs in sharper leaf apices terminating in a row of 1–2 cells, mammillose leaf cells with thickened outer wall, lobules with a long and sharp tooth, underleaf bases without auricles, and innovations pycnolejeuneoid. 9a. Cheilolejeunea filiformis (Sw.) W.Ye, R.L.Zhu & Gradst. var. filiformis Fig. 43O, P Omphalanthus filiformis (Sw.) Nees (Jungermannia filiformis Sw.). Type: Jamaica, Swartz s.n. (lectotype, designated here, S-B21099!) Omphalanthus filiformis var. laxus Gottsche Omphalanthus geminiflorus (Nees) Gottsche Omphalanthus grandistipulus Steph. Creeping or loosely pendent on bark, rock, and soil in submontane and montane forests and páramo, (100–)800–3900 m; in the Galápagos Islands very common in the moist zone, 300–1000 m. Colombia: Ant, Ara, Boy, Cas, Ces, Cho, Cun, Hui, Mag, Met, Nar, Put, Qui, Ris, San, Tol. Ecuador: Ca, EO, Ga, Lo, LR, MS, Na, Pa, Pi, Tu, ZC. General distribution: common throughout the mountains of tropical America. Cheilolejeunea filiformis var. filiformis (sect. Omphalanthus) is a very common neotropical taxon that is readily recognized by (1) plants dioicous, ascending to pendent, growing rather loosely, pale yellow to pale brown, shoots long and little branched; (2) stems rigid, of thick-walled cells, ventral merophyte 4–6 cells wide; (3) leaves obliquely spreading, ovate-orbicular, apex broadly rounded; (4) leaf cells with large swollen trigones, intermediate thickenings absent; (5) lobules shortly trapezioid, 1/4–1/3× leaf length, without tooth; (6) underleaves undivided, large, 4–6× stem width, rounded to reniform; (7) perianth terete, with 1 lejeuneoid innovation. Based on perianth shape, two further varieties have been recognized in C. filiformis, C. filiformis var. platycoleus with an obtusely four-angled perianth and var. wallisii with a sharply 5-keeled perianth. These two varieties are rare and largely restricted to the northern Andes (Schäfer-Verwimp et  al. 2013a; Gradstein and Uribe 2016a). 9b. Cheilolejeunea filiformis var. platycoleus (Herzog) W.Ye, R.L.Zhu & Gradst. Omphalanthus platycoleus Herzog (Omphalanthus filiformis var. platycoleus [Herzog] Gradst.). Type: Colombia, Cauca, San Antonio, 1922, Pennell & Killip 7301 (JE!)

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Keys and Descriptions

On branches and twigs of shrubs, often growing pendent, ca. 1500–3000  m. Colombia: Cau. Ris. Ecuador: ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical Andes (Colombia, Ecuador, Bolivia), Costa Rica. A rare Andean-Costa Rican taxon, characterized by the obtusely four-angled perianth. Cheilolejeunea filiformis var. platycoleus can be confused with C. tonduzana, but the latter species is autoicous (C. filiformis is dioicous) and the leaf lobule in C. tonduzana is more elongate, 2/5–1/2× lobe length, rectangular. 9c. Cheilolejeunea filiformis var. wallisii (J.B.Jack & Steph.) W.Ye, R.L.Zhu & Gradst. Omphalanthus wallisii J.B.Jack & Steph. (Omphalanthus filiformis var. wallisii [J.B.Jack & Steph.] Gradst.). Type: Colombia, Antioquía, páramo de Sonson, 1872, Wallis s.n. (G-00061044!) On branches of trees and shrubs, ca. 1900–3000 m. Colombia: Ant. Ecuador: ZC (Nöske et  al. 2003). General distribution: northern Andes (Venezuela, Colombia, Ecuador), Mexico. A rare Andean-Mexican taxon, characterized by the sharply 5-keeled perianth. By the shape of the perianth, C. filiformis var. wallisii may be confused with C. ovalis and C. quinquecarinata. Cheilolejeunea ovalis differs in longer than wide underleaves (as long as wide or wider than long in C. filiformis) and the obtuse to subacute leaf apex (rounded in C. filiformis), whereas C. quinquecarinata differs in autoicous sexuality (C. filiformis is dioicous), ventral merophyte only 2 cells wide, and leaf lobule more elongate, 2/5–1/2× lobe length, rectangular. 10. Cheilolejeunea fragrantissima (Spruce) R.M.Schust. Euosmolejeunea fragrantissima (Spruce) Steph. Euosmolejeunea fragrantissima var. columbica Herzog On bark of trees, 1500–2500 m. Colombia: Ant (Benavides and Gutiérrez 2011), San (Herzog 1955). General distribution: northern South America, Dominican Republic. Cheilolejeunea fragrantissima (sect.?) is a rare neotropical species that is distinguished by (1) plants autoicous, ventral merophyte 2 cells wide; (2) leaf apex plane, rounded; (3) leaf cells with large, confluent trigones and a smooth cuticle; (4) lobules very small, 1/5–1/6× leaf length, with a long and sharp tooth; and (5) underleaves very broad, reniform, 4–6× stem width, very shallowly bifid, to maximally 1/5 of underleaf length, bases broadly rounded. Cheilolejeunea fragrantissima is closely similar to C. beyrichii, but the leaf apex in the latter species is recurved and obtuse, the lobule tooth is short and blunt, and the ventral merophyte is 4–6 cells wide (Reiner-Drehwald 2006). 11. Cheilolejeunea holostipa (Spruce) Grolle & R.L.Zhu Cyrtolejeunea holostipa (Spruce) A.Evans

Fig. 43Q

On bark of trees in the understory and canopy of lowland and montane rainforests, from sea level to 3400 m. Colombia: Ama, Caq, Cau, Cho, Cun, Mag, Put, Ris, Val. Ecuador: Az, Lo, LR, MS, Na, Or, Su, ZC. General distribution: scattered in tropical America.

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Cheilolejeunea holostipa (sect. Cyrtolejeunea) is a widespread neotropical species that is readily recognized by the very small plants (0.4–0.7  mm wide) with suberect to obliquely spreading leaves, zig-zag stems, and small, undivided, orbicular underleaves. The lobules are 1/2× leaf length, ovoid, with a long-falcate tooth, and the leaf cells have small trigones and finely granular (Jungermannia-type) oil bodies. 12. Cheilolejeunea inflexa (Hampe) Grolle Trachylejeunea inflexa (Hampe) Steph.

Fig. 43R

On bark in montane rainforest and cloud forest, 1300–2550 m. Colombia: Ant, Cho (Gil 3358, UPTC), Mag, Qui, Ris. Ecuador: ZC.  General distribution: scattered in tropical America. Cheilolejeunea inflexa (sect. Strepsilejeunea) is a characteristic neotropicalmontane species that is readily distinguished by (1) leaf apex sharply acute-acuminate, recurved; (2) cells on dorsal leaf surface and keel mammillose and with a huge, rounded papillae, the papilla about half the diameter of the cell; and (3) underleaves large, (sub)imbricate, 3–4× stem width, base cordate. 13. Cheilolejeunea intertexta (Lindenb.) Steph. Cheilolejeunea compacta (Steph.) M.E.Reiner On tree trunks, frequently in rather open situations, in woodlands, along roads, in urban areas, etc., from sea level to 2100 m. Colombia: Hui (Palermo, road to El Carmen, 2100 m, Schuster 88-1327, det. C. J. Bastos, F). General distribution: pantropical, scattered in tropical America. Cheilolejeunea intertexta (sect. Euosmolejeunea) has recently been discovered among the Colombian collections made by R. M. Schuster (Bastos 2017). The species has been little recorded in the neotropics and has mainly been reported from the Mata Atlântica of Brazil. It has also been found on a roadside tree in Panama City, near sea level. In Colombia and Ecuador, the species has probably been overlooked. Cheilolejeunea intertexta closely resembles C. rigidula but differs in autoicous or paroicous sex distribution (antheridia on small separate branches or on the female shoot just below the perianth), pycnolejeuneoid innovations, and possibly the occasional presence of microphyllous branches. The species shares autoicy and pycnolejeuneoid innovations with C. discoidea (= C. myriantha), but the latter species is a smaller, minute plant with very small underleaves and with suberect to obliquely spreading leaves (see key). Cheilolejeunea discoidea is not known from Colombia and Ecuador but may be expected there. 14. Cheilolejeunea jackii (Prantl) W.Ye, R.L.Zhu & Gradst. Fig. 43S Omphalanthus jackii (Prantl) Gradst. (Peltolejeunea jackii Prantl). Type: Ecuador (“Peru”), Manabí, Wallis s.n. (lectotype, designated here, G-09613!) Pendent from tree branches and shrubs in submontane rainforest, cloud forest, and scrub, often forming dense, pale green to yellowish-green festoons, 200–1200 m. Ecuador: Bo, Cañ, EO, Ga (Pinzón, Pinta, Rábida), LR, Ma, Pi; to be expected in Colombia. General distribution: Bolivia, Ecuador, Nicaragua; the record from Peru is erroneous (see type, above).

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Keys and Descriptions

Cheilolejeunea jackii (sect. Omphalanthus) is a very distinct submontane species that is restricted in Ecuador to the western foothills of the Andes and the Galápagos Islands. A somewhat similar distribution, but extending further eastward and to higher elevations, is seen in Frullania aculeata. Cheilolejeunea jackii is readily distinguished by (1) plants dioicous (but male and female plants sometimes growing mixed), long pendent, pale green to yellowish-green; (2) leaves ovate-lanceolate, obliquely spreading, apex acuminate; and (3) underleaves oblong (2–3× longer than wide), apex short-bifid or undivided, bases cuneate, insertion line deeply arched. The gynoecia and androecia are similar to those of C. ovalis. A very copious collection from the cloud forest of Reserva Buenaventura in southwestern Ecuador (Nov 2018, Benitez et al. s.n., HUTPL) stands out by somewhat mammillose leaf cells with dorsally thickened walls and underleaves with undivided to minutely bifid tips. 15. Cheilolejeunea laevicalyx (J.B.Jack & Steph.) Grolle Strepsilejeunea laevicalyx J.B.Jack & Steph.

Fig. 43T

On bark and rock in upper montane cloud forest and páramo, 2500–3600  m. Colombia: Ant (type), Cun. Ecuador: Lo (Tapichalaca). General distribution: tropical Andes (Colombia, Ecuador, Bolivia). Cheilolejeunea laevicalyx (sect.?) is a characteristic high-Andean species that is characterized by (1) plants autoicous, ventral merophyte 2 cells wide; (2) leaves with smooth surface and rounded to acute, broadly recurved apex; (3) leaf cells with small trigones; (4) underleaves 3–5× stem width, longer than wide, bifid, apices rounded; and (5) gynoecia with lejeuneoid innovations, perianth terete or slightly keeled (Zhu 2006). Cheilolejeunea laevicalyx is somewhat similar to C. choachina; for differences see under the latter species. 16. Cheilolejeunea lineata (Lehm. & Lindenb.) Schiffn.  Cystolejeunea lineata (Lehm. & Lindenb.) A.Evans

Fig. 44A, B

On bark of trees in wet lowland rainforest and cloud forests, from sea level to 950 m. Colombia: Cho, Cau (Schuster 88-1621c, F), Val. Ecuador: ZC (Arnell, 1962). General distribution: West Indies, northern South America, southeastern Brazil. Cheilolejeunea lineata (sect. Cheilolejeunea) is a Caribbean species that is readily recognized by the ball-shaped lobules with a strongly involute free margin, bulging leaf cells with rather large trigones, and flattened perianths without beak. The plants are 1.5–2.5 mm wide; vegetative reproduction is by caducous leaf lobes from

Cheilolejeunea trifaria var. trifaria, portion of stem with underleaves. (M) Cheilolejeunea unciloba, portion of stem with leaf. (N) Cheilolejeunea urubuensis, portion of stem with leaf and underleaf. (O and P) Cheilolejeunea xanthocarpa. (O) Leaf. (P) Lobule apex. (Q) Cololejeunea antillana, habit. (R) Cololejeunea azorica, leaf. (S and T) Cololejeunea bekkerae. (S) Leaf. (T) Lobule. (U) Cololejeunea bischleriana, leaf. (V and W) Cololejeunea camillii. (V) Leaf. (W) Gemma. (X) Cololejeunea cardiocarpa, portion of stem with leaf. (Y) Cololejeunea cingens, lobule. (C, N from Campos et al. 2014; J from Gradstein and Reiner-Drehwald 2017b; R, V–Y redrawn from Pócs et al. 2014; U redrawn from Tixier 1980; rest from Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009)

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Fig. 44 (A and B) Cheilolejeunea lineata. (A) Habit. (B) Lobule, showing inflexed free margin. (C) Cheilolejeunea neblinensis, lobule. (D) Cheilolejeunea lobulata, portion of stem with leaf. (E) Cheilolejeunea ornata, portion of stem with leaf and underleaf. (F) Cheilolejeunea ovalis, habit. (G) Cheilolejeunea paramicola, perianth. (H) Cheilolejeunea revoluta, portion of stem with underleaf. (I) Cheilolejeunea rigidula, portion of stem with underleaf. (J) Cheilolejeunea schiavoneana, habit showing caducous-leaved flagelliform shoot. (K) Cheilolejeunea tonduzana, portion of stem with leaf. (L)

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Keys and Descriptions

normal shoots. In the field the species stands out by its rather loose and somewhat pendent growth and dull pale green color. The lobules of C. lineata somewhat resemble those of Anoplolejeunea conferta, but the latter species is a smaller plant (1–1.5  mm wide) with undivided underleaves, small trigones, and 5-keeled perianths with innovations. 17. Cheilolejeunea lobulata (Lindenb.) Gradst. & C.J.Bastos Cheilolejeunea oncophylla (Ångstr.) Grolle & M.E.Reiner Harpalejeunea verrucosa Herzog Trachylejeunea dominicensis Steph.

Fig. 44D

On bark of trees and shrubs and on rock in montane rainforests and páramo, also in the canopy of Amazonian lowland forest (Campos et  al. 2015), 200–4500  m. Colombia: Ama, Ant, Cas (Gradstein 12712, PC), Put, Ris, Vau. Ecuador: Lo, Na, Tu, ZC. General distribution: tropical America. Cheilolejeunea lobulata (sect. Strepsilejeunea) is a common and variable neotropical species with a remarkably wide elevational amplitude, ranging from lowland rainforest to páramo. The species was usually called C. oncophylla, but the name C. lobulata has priority (Bastos and Gradstein submitted). Grolle (1979) treated C. lobulata (= Strepsilejeunea lobulata [Lindenb.] Steph.) as a synonym of C. acutangula, but the leaves in the type of C. lobulata are obtuse and the underleaves only 2× wider than stem, like in C. oncophylla. The species is characterized by (1) plants autoicous, small, 0.4–0.9 mm wide, ventral merophyte 2 cells wide; (2) leaves convex, apex mostly obtuse, sometimes rounded or bluntly apiculate (tip consisting of one cell), plane or recurved; (3) leaf cells with one large oil body and with large swollen trigones, dorsal cell wall convex-mammillose and thickened, with a low and broad, lens-shaped papilla; (4) lobule tooth long and acute, lobule cells mammillose and with a low and broad, lens-shaped papilla, keel crenate; (5) underleaves small, 1.5–2.5× stem width, bifid to 1/4–1/3, with a wide V-shaped sinus, apices obtuse, bases cuneate, insertion line straight; (6) gynoecia with 1 pycnolejeuneoid innovation, perianth sharply 5-keeled, keels smooth; (7) vegetative reproduction by caducous leaf lobes and cladia (Grolle and Reiner-Drehwald 1997 as C. oncophylla). Cheilolejeunea lobulata resembles the rare C. ornata, but the latter species is dioicous and has large, rounded, tubercle-like papillae on keel, lobule, and dorsal leaf surface. The species is also close to C. acutangula, but the underleaves in the latter species are larger, 3–4× stem width, with a distinctly curved insertion line, and the leaf apices are more sharply pointed, acute-acuminate, terminating in a row of 1–2 cells. 18. Cheilolejeunea neblinensis Ilk.-Borg. & Gradst.

Fig. 43C

On bark of trees in Amazonian lowland rainforest, 100–250 m. Colombia: Ama, Caq, Put, Vau (Campos et  al. 2014) General distribution: upper Amazonia (Brazil, Colombia, Venezuela). Cheilolejeunea neblinensis (sect. Trachylejeunea) is an Amazonian species that is distinguished by (1) leaf apex rounded to obtuse to subacute (sometimes purely

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rounded), usually recurved; (2) leaf cells with conspicuous trigones, cuticle smooth or with a low lens-shaped papilla; (3) lobules 1/3–1/2× leaf length, decurved, lobule apex almost reaching the ventral leaf margin (like in Pycnolejeunea decurviloba and Stictolejeunea squamata), with 2 tightly associated teeth, hyaline papilla between the 2 teeth, keel crenate, sometimes bordered by a row of enlarged cells; (4) underleaves 2–4× stem width, bifid to ca. 1/3–2/3, outer margins entire to sinuose to obtusely toothed; and (5) innovations present or absent, pycnolejeuneoid. Cheilolejeunea neblinensis is readily recognized by the decurved lobule with twinned teeth at the apex. A further characteristic feature, seen in the type specimen from Cerro de la Neblina, is the enlarged cells along the lobule keel (Ilkiu-Borges and Gradstein 2008). However, in populations from the Colombian Amazon, the cells along the keel are usually not enlarged. 19. Cheilolejeunea ornata C.J.Bastos

Fig. 44E

On bark of tree in lower montane rainforest, 1890 m. Ecuador: ZC (Reserva Biológica San Francisco; Gradstein and Benitez 2017). General distribution: eastern Brazil (Bahia), southern Ecuador. Cheilolejeunea ornata (sect. Strepsilejeunea) is a rare species from eastern Brazil that has been found once in Ecuador (A. Schäfer-Verwimp, JE). The species is habitually similar to C. lobulata but is distinguished by the large, 7–10 μm high, tubercle-like papillae (one per cell) on the cells of leaf lobes, lobules, and keel (Bastos 2011). In C. lobulata the papillae are low, lens-shaped, 2–3  μm high. Moreover, C. ornata is dioicous, whereas C. lobulata is autoicous. The Ecuadorian specimen slightly differs from the type of C. ornata in the obtuse to subacute leaf apex (acute to apiculate in the type) and is in this respect more similar to C. grosseoleosa Bastos & Schäf.-Verw., a Brazilian species characterized by the presence of one huge, coarsely granular oil body in leaf cells (Bastos and Schäfer-Verwimp 2017). Cheilolejeunea ornata, in contrast, has several smaller oil bodies per cell. The oil bodies of the Ecuadorian plants are unknown. 20. Cheilolejeunea ovalis (Lindenb. & Gottsche) Ye et al. Fig. 44F Omphalanthus ovalis (Lindenb. & Gottsche) Gradst. (Lejeunea ovalis Lindenb. & Gottsche). Type: Venezuela, Mérida, Moritz s.n. (lectotype, designated by R. Grolle [in sched.], G-19432!) Peltolejeunea rotundistipula Steph. Pendent on trunks and branches in moist evergreen woodlands, in gardens, and in disturbed lowland and montane rainforest, rarely on soil, 100–3000 m; very common in the Galápagos Islands. Colombia: Cas, Mag, Nat, Val. Ecuador: Az, Ch, EO, Ga, Lo, LR, Na, Pi, Tu, ZC (pass El Tiro, 3000  m, Burghardt 7104, QCA). General distribution: scattered in tropical America. Cheilolejeunea ovalis (sect. Omphalanthus) is a rather widespread neotropical species that is closely related to C. filiformis but differs in the longer than wide underleaves (as long as wide or wider than long in C. filiformis) and the obtuse to subacute leaf apex (rounded in C. filiformis). Moreover, the perianth in C. ovalis is 5-keeled (mostly eplicate in C. filiformis; 5-keeled in the rare C. filiformis var.

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Keys and Descriptions

wallisii). Cheilolejeunea ovalis and C. filiformis are very common in the Galápagos Islands and may grow mixed there. 21. Cheilolejeunea paramicola (Herzog) W.Ye, R.L.Zhu & Gradst. Fig. 44G Aureolejeunea paramicola (Herzog) R.M.Schust. (Brachiolejeunea parami­ cola Herzog) Aureolejeunea paramoensis R.M.Schust. On bark of trees and shrubs in upper montane forest and páramo, 2600–4000 m. Colombia: Can, Cun, Qui (El Campanario, Gradstein et al. s.n., HUQ), Ris, San. Ecuador: Az, Ca, Pi, ZC (páramo of Saraguro and Yacuambi, Benitez & Gradstein s.n., HUTPL). General distribution: northern Andes. Cheilolejeunea paramicola (sect. Omphalanthus) is a characteristic northern Andean páramo species that is readily recognized by the brown to glossy orangebrown, creeping to ascending to pendent, autoicous plants with undivided underleaves, large rectangular lobules, large swollen trigones, gynoecia with 2 innovations, and perianths broadly 3-keeled and pluriplicate, with 4–8 narrow folds: 3–5 on the ventral perianth surface (mostly on the broad ventral keel) and 1–3 on the dorsal suface. 22. Cheilolejeunea quinquecarinata (R.M.Schust.) W.Ye, R.L.Zhu & Gradst. Aureolejeunea quinquecarinata R.M.Schust. On branches and twigs of shrubs in upper montane dwarfforest and páramo, 3000–4100 m. Colombia: Cas, Cun, Met, Ris. Ecuador: Lo, ZC. General distribution: northern Andes, Guayana Highland, Dominican Republic. Cheilolejeunea quinquecarinata (sect. Omphalanthus) is an uncommon northern Andean páramo species that is distinguished by (1) plants autoicous, glossy brown; (2) ventral merophyte 2 cells wide; (3) leaves convex, apex rounded, broadly recurved; (4) leaf cells mammillose (but without papilla), with large swollen trigones, intermediate thickenings absent; (5) lobules rectangular, 2/5–1/2× lobe length, tooth 2–3 cells long; (6) underleaves undivided, large (4–5× stem width), orbicular or somewhat wider than long, margin often narrowly recurved, bases broadly rounded, insertion line deeply arched; and (7) perianth sharply 5-keeled, with 1–2 lejeuneoid innovations. Cheilolejeunea quinquecarinata is closely related to C. aurifera and C. tonduzana; for differences see the key. 23. Cheilolejeunea revoluta (Herzog) Gradst. & Grolle  Pycnolejeunea revoluta Herzog

Fig. 44H

On rock and bark in shaded, humid environments, 700–2650 m. Colombia: San (type). Ecuador: ZC (Nöske et  al. 2003). General distribution: southeastern Brazil, northern Andes, Costa Rica. Cheilolejeunea revoluta (sect.?) is a rare neotropical species that is recognized by (1) plants dioicous, stems robust, with a 6–7-cell-wide ventral merophyte; (2) leaves with a rounded apex; (3) lobules small, flask-shaped, with a short tooth; and

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(4) underleaves large (5–6× stem width), reniform, shallowly bifid, with broadly recurved apical margin. 24. Cheilolejeunea rigidula (Mont.) R.M.Schust. Fig. 44T Cheilolejeunea duriuscula (Nees) Schiffn. (Euosmolejeunea duriuscula [Nees] Steph.) Cheilolejeunea serpentina (Mitt.) Mizut. Euosmolejeunea neglecta Steph. Common on bark of trees in the canopy and the understory of lowland rainforests, in scrub, plantations, on isolated trees, etc., also on rock, less common in montane rainforest, from sea level to 2800  m. Colombia: Ama, Ant, Caq, Ces, Cho, Hui, Mag, Nar, Put, Qui, Ris, Tol, Vau. Ecuador: EO, Es, Ga, Lo, MS, Na, Or. General distribution: pantropical. Cheilolejeunea rigidula (sect. Euosmolejeunea) is one of the most common epiphytic liverworts in Amazonian rainforests (Campos et  al. 2015). The species is distinguished by (1) plants dioicous (rarely autoicous), rather small, ca. 0.7–1.2 mm wide, dull grayish-green; (2) leaves wide-spreading, flat, short-ovate, apex rounded; (3) leaf cells with small or large trigones, intermediate thickenings lacking; (4) lobules ovoid, never reduced, 1/4–1/3× leaf length, with a short or long, blunt tooth; (5) underleaves 2–3.5× stem width, bifid, bases cuneate to slightly rounded, insertion line curved; and (6) gynoecia with 1–2 lejeuneoid innovations. Robust phenotypes of C. rigidula with slightly rounded underleaf bases resemble C. polyantha A.Evans, which might be a variety of C. rigidula. Such robust plants may be confused with C. trifaria, especially C. trifaria var. clausa. However, the underleaves in C. trifaria are 4–6× stem width and mostly imbricate, while those of C. rigidula are maximally 3.5× stem width and usually distant. Small phenotypes of C. rigidula may be confused with C. discoidea and C. intertexta, but the latter two species are monoicous and have pycnolejeuneoid innovations. For further differences see the key and under C. intertexta. Cheilolejeunea rigidula may also be confused with C. aneogyna; for differences see under C. aneogyna. 25. Cheilolejeunea schiavoneana M.E.Reiner & Gradst. 

Fig. 44J

On canopy branches in submontane rainforest on the Pacific side of the Western Cordillera of Colombia, 1200 m. Colombia: Ris (Gradstein and Reiner-Drehwald 2017b). General distribution: only known from the type from Colombia. Cheilolejeunea schiavoneana (sect. Cheilolejeunea) is a rare Colombian endemic that is characterized by (1) plants dioicous (?), rather robust, 1.8–2.4 mm wide, with numerous microphyllous branches, stems rigid, ventral merophyte on main stems 5–6 cells wide; (2) leaf lobes with rounded apex and entire margins apex rounded; (3) leaf cells 25–40 × 20–30 μm in midleaf, conspicuously smaller toward the margin, walls with triradiate trigones and intermediate thickening; (4) lobules shortly flask-shaped, 1/6–1/4× leaf length, fully inflated, with a long and sharp tooth, lobule cells much smaller than leaf lobe cells; (5) underleaves distant, 2.5–3× stem width, bifid to 1/2, margins entire; and (6) caducous leaf lobes produced on short upright, flagelliform shoots with densely imbricate underleaves, caducous leaf lobes smaller than normal leaves, margins with short rhizoids.

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Keys and Descriptions

Cheilolejeunea schiavoneana approaches C. adnata, but the latter species differs in smaller plant size, smaller size of leaves and underleaves, thinner stems (of 7 rows of cortical cells surrounding 10 medullary rows), ventral merophyte only 2 cells wide, absence of microphyllous branches, and caducous leaf lobes unmodified, produced on normal leafy shoots. 26. Cheilolejeunea tonduzana (Steph.) W.Ye, R.L.Zhu & Gradst. Aureolejeunea tonduzana (Steph.) Gradst. Aureolejeunea fulva R.M.Schust.

Fig. 44K

On branches and twigs in montane rainforest and scrub, 1800–3100  m. Colombia: Ris (Wolf 1993), San (Virolin, R. M. Schuster 88-1069, det. C. J. Bastos, F). Ecuador: Lo, ZC (Nöske et  al. 2003). General distribution: Mexico to Bolivia, Southeastern Brazil, Lesser Antilles. Cheilolejeunea tonduzana (sect. Omphalanthus) is a rather widespread neotropical species that resembles C. filiformis but differs in having longer, rectangular lobules (2/5–1/2× leaf length), reniform underleaves, and autoicous sexuality. The ventral merophyte is 4–6 cells wide, and the perianth in C. tonduzana is 3–4-keeled to almost terete. The plants grow prostrate to ascending. 27a. Cheilolejeunea trifaria (Reinw., Blume & Nees) Mizut. var. trifariaFig. 44L Cheilolejeunea azureomontana C.J.Bastos & Schäf.-Verw. (fide Bastos and Gradstein submitted) On rock, soil, or bark of trees, in shaded or open places, along roads, etc., from sea level to 2200  m. Colombia: Ama, Caq, Nar, Put, Qui, Ris, San (Cuta 394, UPTC), Vau. Ecuador: Az, EO, Ga, MS, Or, ZC.  General distribution: pantropical. Cheilolejeunea trifaria var. trifaria (sect. Euosmolejeunea) is a very common pantropical taxon that is readily recognized by (1) plants autoicous, dull green to pale yellowish-brown, ventral merophyte 2 cells wide; (2) leaves with broadly rounded apex and small, ovoid lobules; (3) leaf cells usually mammillose and with conspicuous trigones; (4) lobules ca. 1/4–1/3 of leaf length, never reduced, ovoid, with a short, blunt tooth; and (5) underleaves (sub)imbricate, broadly ovate to suborbicular, usually wider than long (rarely as wide as long), large, ca. 4–6× stem width, bifid to 1/5–1/3, with rounded to auriculate bases. The plants are frequently fertile and the gynoecia have 1–2 lejeuneoid innovations. Cheilolejeunea trifaria is closely similar to C. rigidula, but the latter species has smaller underleaves (2–3.5× stem width) with cuneate to slightly rounded bases and is usually dioicous. 27b. Cheilolejeunea trifaria var. clausa (Nees & Mont.) C.J.Bastos & Gradst. Fig. 43M Cheilolejeunea clausa (Nees & Mont.) R.M.Schust. Euosmolejeunea laxiuscula (Spruce) Steph. Euosmolejeunea opaca (Gottsche) Steph. Lejeunea nemoralis Gottsche, syn. nov. Type: Colombia, Cundinamarca, Bogotá, near Aserradero, 2500  m, Lindig 1735 p.p. (isotype G-00282340!, c. gyn.)

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On bark of trees in lowland and montane rainforest, also on rock, 70–2500 m. Colombia: Ama, Cun, Put (Campos et al. 2014 as C. clausa). Ecuador: Ga (?), Es, MS, Or, ZC. General distribution: scattered in tropical and subtropical America. Cheilolejeunea trifaria var. clausa differs from var. trifaria in being dioicous. In addition, the underleaves of var. clausa are rounded, whereas in var. trifaria they are mostly wider than long, rarely rounded. The existence of autoicous C. trifaria with rounded underleaves (described as C. azureomontana) indicates that C. clausa must be a variety of C. trifaria, not a separate species (Bastos and Gradstein submitted). The records of C. trifaria var. clausa from the northern Andes should be verified; some may be misidentifications. The record from the Galápagos Islands (Weber 1975 as C. clausa) probably refers to var. trifaria. 28. Cheilolejeunea unciloba (Lindenb.) Malombe Leucolejeunea unciloba (Lindenb.) A.Evans

Fig. 44M

On bark and rock in rather open evergreen and semi-evergreen forest and scrub and on isolated trees and shrubs, especially in coastal environments, from sea level to 3100  m. Colombia: Cho, Cun, Mag. Ecuador: Lo, ZC.  General distribution: tropical and subtropical America, Africa. Cheilolejeunea unciloba (sect. Leucolejeunea) is a widespread Afro-American species that is rare in the northern Andes. The species stands out by its pale whitishgreen to greenish-brown color, appressed growth, large undivided underleaves, 4-cell-wide ventral merophyte, lobules with 1–4 cell long, straight or curved tooth, and plane ventral margin of leaf lobe. The leaf cells are plane and thin-walled on the dorsal surface (Schuster 1980a as Leucolejeunea unciloba). 29. Cheilolejeunea urubuensis (C.E.Zartman & I.L.Ackerman) R.L.Zhu & Y.M.WeiFig. 44N Vitalianthus urubuensis C.E.Zartman & I.L.Ackerman On twigs in the outer canopy of Amazonian lowland rainforest, 100–220  m. Colombia: Ama, Caq, Put (Campos et al. 2014). General distribution: Amazonia (Brazil, Colombia). Cheilolejeunea urubuensis (sect.?) is a characteristic canopy species of Amazonian rainforest that is easily distinguished by the leaf lobes with a row of 4–8 yellowish to yellow-brown ocelli. The leaf apex is rounded; the lobules are very large, 1/2–2/3× lobe length, with long, curved, multicellular tooth; and the underleaves are deeply bifid (to more than 1/2), with slightly diverging, blunt lobes, cuneate bases, and a straight insertion line. The plants are autoicous and frequently fertile, the gynoecia have 1–2 pycnolejeuneoid innovations, and the perianth is obconical with a broad, truncate apex and 4 smooth keels. 30. Cheilolejeunea xanthocarpa (Lehm. & Lindenb.) Malombe Leucolejeunea xanthocarpa (Lehm. & Lindenb.) A.Evans

Fig. 44O, P

On bark and rock in evergreen and semi-evergreen montane forest and scrub, also on isolated trees and shrubs, 450–3500 m. Colombia: Boy, Cas, Cun, Mag, San, Tol. Ecuador: EO, Ga, Lo, ZC. General distribution: pantropical.

440

 

Keys and Descriptions

Cheilolejeunea xanthocarpa (sect. Leucolejeunea) is a common pantropical species that is readily recognized by the strongly revolute ventral margin of the leaf lobe, undivided underleaves, 4-cell-wide ventral merophyte, and the large, rectangular lobule with a distinct, 1–5-cell-long tooth. The leaf cells are mammillose and usually thick-walled on the dorsal surface (Schuster 1980a as Leucolejeunea xanthocarpa). Excluded Records Cheilolejeunea curvatiloba (Herzog) Grolle – Colombia: Val (Herzog 1955 as Harpalejeunea curvatiloba Herzog). Not a good species (R.L. Zhu, pers. comm.). Cheilolejeunea discoidea (Lehm. & Lindenb.) R.M.Schust. & Kachroo (= C. myriantha (Nees & Mont.) Schiffn.) – Colombia: Cho (Vasco et al. 2002); the specimen was misidentified and belongs to C. rigidula. Cheilolejeunea asperiflora (Spruce) Gradst. & Ilk.-Borg. – Colombia: Ant, montane cloud forest, ca. 1800  m (Benavides and Gutiérrez 2011 as Trachylejeunea asperiflora [Spruce] Schiffn.). This is a rare Amazonian lowland species, endemic to the upper Rio Negro region of Brazil and Venezuela (Bastos 2017); its occurrence in montane cloud forest is highly unlikely. Cololejeunea (Spruce) Steph. Aphanolejeunea A.Evans Plants very small- to medium-sized, 0.25–1.5  mm wide, pale or dull green, creeping. Branches Lejeunea-type (with collar) or Aphanolejeunea-type (without collar). Stems usually zig-zag and with a hyalodermis, in cross section made up of 1 medullary cell surrounded by 5–6 epidermis cells (rarely more, C. madothecoides, C. stotleriana); ventral merophyte 1(−6) cells wide. Leaf attached to stem by only few cells, leaf insertion very short, transverse. Leaf lobes distant or subimbricate, spreading, ovate-obovate to narrowly lanceolate, sometimes reduced; leaf surface smooth, papillose or mammillose by conically elevated cells, leaf apex rounded to acute-acuminate, margins entire to denticulate, sometimes with a hyaline border of translucent cells without chloroplasts. Cells thin-walled with or without small trigones, sometimes sigmoid, cuticle smooth, striped or punctulate; oil bodies botryoidal or finely granular or smooth; ocelli lacking or present as a 1–2-cell-wide vitta. Lobules small or large, flat, ligulate or saccate, inflated, sometimes reduced, apex with (0–)1–2 teeth, hyaline papilla proximal, distal, or ental. Underleaves absent, only rhizoids in small bundles at the ventral side of the stem, usually near each leaf base (except in C. stotleriana). Androecia variable, bracteoles lacking, bracts with hypostatic lobules. Gynoecia with 0–2 innovations, bracteoles lacking. Perianths inflated or flattened, 0–5-keeled, perianth surface smooth or papillose. Sporophytes lejeuneoid. Vegetative reproduction by multicellular, disciform gemmae from leaf surfaces or from leaf margins.

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A large pantropical and warm-temperate genus of about 300 species, especially in tropical Asia, 32 species in Colombia and Ecuador, in 8 subgenera, subg. Aphanolejeunea (A.Evans) Pócs, subg. Chlorocolea R.M.Schust., subg. Chlorolejeunea Benedix, subg. Cololejeunea, subg. Diaphanae R.M.Schust., subg. Leptocolea (Spruce) Schiffn., subg. Pedinolejeunea Mizut., and subg. Taeniolejeunea (Zwickel) Benedix (Pócs et al. 2014). The subgeneric classification was not supported by DNA sequences, however, and one subgenus, C. subg. Protocolea R.M.Schust., has been transferred to Myriocoleopsis (Yu et al. 2013, 2014). Most of the species grow on living leaves or twigs and are very small and fragile. The genus Cololejeunea is readily recognized by the very short, transverse leaf insertion, the absence of underleaves, and the very thin stems consisting of only 1 row of medullary cells and a limited number of epidermal rows (usually 5–6; occasionally more). Very short leaf insertion, absence of underleaves, and stems with only 1 row of medullary cells are also seen in Myricoleopsis. The latter genus differs in having upright leafy shoots arising from a creeping stoloniform stem, without hyalodermis. Rhizoids in Cololejeunea develop in a “pendular” fashion: instead of one rhizoid bundle to each leaf pair (as is usual in Lejeuneaceae), each leaf is associated with a rhizoid bundle, indicating the potential presence of one underleaf to each leaf. The pendular leaf segmentation seen in Cololejeunea and the thin stems with only one row of medullary cells are characteristic of juvenile leafy liverworts; their retention in the adult Cololejeunea plants is an example of paedomorphosis or neoteny (“primary neoteny”; Gradstein et al. 2003). The neoteny in Cololejeunea is interpreted as an adaptation to the short-lived, epiphyllous or ramicolous habitat of the plants, enhancing rapid completion of the life cycle. Literature: Pócs (2009a), Pócs et al. (2014) 1. Lobules often reduced to a long and narrow, flat fold or strip (ca. 0.5× leaf length) along the ventral leaf margin, sometimes rather inconspicuous. Leaves sometimes squarrose. Perianths very numerous, ± stalked. Plants on bark or rock, rarely on leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . genus Myriocoleopsis 1. Lobules not reduced to a long and narrow fold or strip along the ventral leaf margin. Leaves not squarrose. Perianth not stalked. Pland usually on leaves or twigs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (genus Cololejeunea) 2 Here also keys out Myriocoleopsis minutissima subsp. minutissima, couplet 39 2. Leaf lobe cells sigmoid (narrowly elongate and with undulate walls). Stems flattened, ribbon-like. Plants growing very tighly fixed to the surface of living leaves in lowland rainforest. . . . . . . . . . . . . . . . . . . . . . . . . . . C. linopteroides 2. Leaf lobe cells not sigmoid. Stems not ribbon-like. . . . . . . . . . . . . . . . . . . . . 3 3. Plants robust, 2–3.5 mm wide. Branches with numerous, clustered gynoecia. Stems rigid, epidermis composed of 9–12 rows of cells, ventral merophyte 3–5 cells wide. Plants on periodically inundated river banks in southern Ecuador, epiphyllous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stotleriana 3. Plants smaller. Branches not with clustered gynoecia. Stems fragile and very thin, epidermis composed of 5(−6) rows of cells, ventral merophyte 1(−2) cells wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

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Keys and Descriptions

4. Most leaves reduced to a small, linear band of 2–4 cell rows. . . . . . . . . . . . . . 5 4. Most leaves ovate to oblong to lanceolate, not reduced to a small linear band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 5. Leaf margins toothed. Cells on the dorsal leaf surface conically elevated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. gracilis 5. Leaf margins entire or crenulate. Dorsal leaf surface smooth or with a few, ­scattered, conically elevated cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Leaves and discoid gemmae often extended at apex into a long, ribbon-like gemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. minuscula 6. Leaves and discoid gemmae not extended into a long, ribbon-like gemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. papillosa 7. Lobules mostly reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Lobules mostly well developed [when in doubt try both leads] . . . . . . . . . . 13 8. Dorsal leaf surface, margins and keel rough due to concally elevated cells crowned a papilla. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Dorsal leaf surface smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 9. All branches Lejeunea-type (with collar). Distal free margin of well-developed lobules formed by 3–4 large cells. Lobule surface smooth. Plants of southern Ecuador, rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. kuciana 9. Branches Lejeunea-type or Aphanolejeunea-type (with or without collar). Distal free margin of well-developed lobules not formed by 3–4 large cells. Lobule surface smooth or papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Surface of well-developed lobules papillose; lobule tooth reduced. Plants not known with certainty from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. contractiloba A.Evans 10. Surface of well-developed lobules smooth; lobule tooth well developed, 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Well-developed lobule almost as large as the leaf lobe (more than 1/2 of lobe surface). Plants of northern and southern subtropical America; not known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. jamesii (Austin) M.E.Reiner & Pócs (= C. clavatopapillata Steph.) 11. Well-developed lobule much smaller than the lobe (less than 1/2 of lobe surface) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. papilliloba 12. Plants minute, less than 0.5  mm wide, very rare (Ecuador). Leaves without gemmae. Perianth apex with 5 wing-like auricles. . . . . . . . . . . . C. pterocolea 12. Plants larger. Leaves with gemmae on margins. Perianth apex without winglike auricles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. camillii 13. Lobule almost as long as the lobe, ca. 0.9× lobe length. Apex of leaf lobe broadly rounded, deflexed, partly covering the lobule apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. subsphaeroidea 13. Lobules shorter, 0.3–0.8× lobe length. Apex of leaf lobe obtuse to acute, plane. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Branches Lejeunea-type or Aphanolejeunea-type (with or without collar at the base). Plants very small, leaves distant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

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14. Branches exclusively Lejeunea-type (always with a collar at the base). Plants small or large, leaves distant or imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 15. Leaves strongly contracted at the junction of lobe and lobule, with 2 lobe cells protruding at the junction. (Leaves ovate-oblong, surface, margin and keel smooth, lobules ca. 1/2× lobe length, first lobule tooth 2 cells long, straight or incurved) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. winkleri 15. Leaves not contracted at the junction of lobe and lobule, lobe cells not protruding at the junction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Lobule cells sigmoid (narrowly elongate and with sinuate walls). Apex of leaf lobe rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. crenata 16. Lobule cells not sigmoid (narrowly elongate and with sinuate walls). Apex of leaf lobe rounded to acute-acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. First lobule tooth straight, sometimes reduced. Apex of leaf lobe rounded to subacute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. First lobule tooth falcate (but sometimes incurved and almost invisible). Apex of leaf lobe acute-acuminate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 18. First lobule tooth 1-celled or reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18. First lobule tooth 2 cells long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 19. Leaves lanceolate, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. antillana 19. Leaves broadly ovate, papillose. Not known with certainty from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. contractiloba 20. Lobules less than 1/2× lobe length. Leaf apex acute. Gemmae angular . . . . 21 20. Lobules more than 1/2× lobe length. Leaf apex rounded to obtuse. Gemmae rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21. Leaves on stems and branches very different in shape: stems with broadly ovate-rhomboid leaves, branches with narrowly ovate-lanceolate leaves (like in C. camillii, but with well-developed lobules). Lobule teeth separated by a broad, 2-cell-wide gap. Plants of Costa Rica, Venezuela and Bolivia; to be expected in Colombia and Ecuador. . . . . . . . . . . . C. yelitzae Pócs & Bernecker 21. Leaf shape of stems and branches uniform, ovate-lanceolate. Lobule teeth separated by a narrow, 1-cell-wide gap . . . . . . . . . . . . . . . . . . . . . . . . . C. diaphana 22. Leaves smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. azorica 22. Leaves papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. papillosa 23. Well-developed leaves 2–3× longer than wide. Lobule 1/2–3/4× lobe length. Dorsal leaf surface usually with sharply conical mammillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. microscopica 23. Well-developed leaves 4–6× longer than wide. Lobule 1/3–1/2× lobe length. Dorsal leaf surface smooth or ± papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 24. Keel crenate. First lobule tooth 1 cell long . . . . . . . . . . . . . . . . . . . C. sicifolia 24. Keel smooth. First lobule tooth 2 cells long . . . . . . . . . . . . . . . . . . C. cingens 25. All cells on dorsal leaf surface, margins and keel strongly conically (or fingerlike) projecting and crowned by a papilla. Plants minute, ca. 0.3 mm wide, rare (southern Ecuador). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. kuciana 25. Cells on dorsal leaf surface and keel not strongly conically (or finger-like) ­projecting. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

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2 6. Leaf apex with one or more finger-like projecting hyaline cells . . . . . . . . . . 27 26. Leaf apex without finger-like projecting hyaline cells . . . . . . . . . . . . . . . . . . 30 27. Dorsal leaf margin with a narrow, 1-cell-wide hyaline border . . . . . . . . . . . 28 27. Dorsal leaf margin without hyaline border . . . . . . . . . . . . . . . . . . . . . . . . . . 29 28. Lobules large, at least 1/3 of lobe length, inflated. Cuticle smooth . . . . . . . . . . 28. Lobules smaller, less than 1/4 lobe length, inflated or flat. Cuticle smooth or finely striate-papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. verwimpii 29. Leaf apex with one projecting hyaline cell. Stylus 3–8 cells long, linear. Plants rare (Ecuador, Panama) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ecuadoriensis 29. Leaf apex with several projecting hyaline cells. Stylus 1 cell long or lacking. Plants common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cardiocarpa 30. Leaf lobes with a white margin of large, rectangular, hyaline cells . . . . . . . . 31 30. Leaf lobes without white margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 31. Lobules all or in part flat, ± erect, ligulate to triangular-subquadrate . . . . . . 32 31. Lobules inflated-saccate, wide-spreading, ovate . . . . . . . . . . . . . . . . . . . . . . 35 32. Cuticle finely papillose, leaf cells opaque . . . . . . . . . . . . . . . . C. surinamensis 32. Cuticle smooth, leaf cells transparent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 33. Lobules narrowly ligulate, distal base of lobule (at the distal end of the keel) adnate to the surface of the leaf lobe across 1–3 cells. . . . . . . . . . . C. bekkerae 33. Lobules triangular to broadly ligulate, distal base of lobule not adnate to leaf lobe surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 34. Ventral merophyte 2–4 cells wide. Lobules constantly flat (usually ligulate) and without teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. latilobula 34. Ventral merophyte 1 cell wide. Lobules flat or saccate on the same plant, with 1–2 teeth or a small lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. planissima 35. Ventral merophyte 2–3 cells wide. Dorsal leaf base very broad, widely arching across and beyond the stem. Lobules with 2 unequal teeth, the second tooth shorter and obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vitaliana 35. Ventral merophyte 1 cells wide. Dorsal leaf base narrow, arching across but not beyond the stem. Lobules with 2 ± equal, acute teeth. . . . . . . . C. bischleriana 36. Base of leaf lobes with a group of enlarged cells (6–18). Leaf cells uni-papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. platyneura 36. Base of leaf lobes without enlarged cells. Leaf cells smooth or papillose . . . . . 37 37. Leaf apex narrowly obtuse to acute. Leaves narrowly oblong, 2–3 longer than wide, margins entire. Leaf cells smooth. Lobules with 1 tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hildebrandii 37. Leaf apex broadly rounded. Leaves ovate-oblong, 1–1.5(−2)× longer than wide, margins denticulate, occasionally entire. Leaf cells usually papillose or smooth. Lobules usually with 2 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38. Leaf margins denticulate. Leaf cells uni-papillose. Distal end of the lobule not adnate to the leaf lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. obliqua 38. Leaf margins entire. Leaf cells smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39. Leaf lobes ovate-oblong. Lobule less than 1/2 the length of the leaf lobe. Distal end of the lobule (at the end of the keel) adnate to the leaf lobe across 2–3 cells. Perianth terete, without keels. Plants on living leaves in wet lowland rainforest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. planiuscula

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39. Leaf lobes broadly ovate-orbicular. Lobule more than 1/2 the length of the leaf lobe. Distal end of the lobule not adnate to the leaf lobe. Perianth 5-keeled. Plants on bark and leaves in dry forests, parks and gardens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myriocoleopsis minutissima subsp. minutissima 1. Cololejeunea antillana Pócs Aphanolejeunea longifolia Jovet-Ast

Fig. 44Q

On living leaves in lowland and lower montane rainforest, from sea level to 1700 m. Colombia: Cho, San (Campos and Uribe 2006). Ecuador: Na (SchäferVerwimp et  al. 2013a). General distribution: West Indies, Central America, northern South America. Cololejeunea antillana (subg. Diaphanae) is a Caribbean species distinguished by lanceolate leaf lobes (4–5× longer than wide) with a subacute apex and entire margins and lobules with a reduced tooth. The species resembles C. sicifolia in leaf shape, but the lobule tooth in C. sicifolia is well developed (2 cells long, falcate) and the leaf cells are frequently papillose. 2. Cololejeunea azorica V.Allorge & Jovet-Ast Aphanolejeunea azorica (V.Allorge & Jovet-Ast) Pócs & Bernecker

Fig. 44R

On fern fronds in upper montane cloud forest, 2180–2880 m. Ecuador: Lo, ZC (Schäfer-Verwimp et  al. 2013a). General distribution: tropical America, Macaronesia. Cololejeunea azorica (subg. Diaphanae) is a widespread neotropical species that closely resembles C. diaphana but differs by the characters given in the key. 3. Cololejeunea bekkerae Tixier [“bekkeri”]Fig. 44S, T On living leaves in lowland rainforest, 230 m. Ecuador: Or (Schäfer-Verwimp et al. 2013a). General distribution: Guianas, Ecuador. Cololejeunea bekkerae (subg. Pedinolejeunea) is a rare South American lowland rainforest species that is recognized by the suborbicular leaves with a wide hyaline border and flat, ligulate lobules that are standing upright nearly parallel to the stem and adnate at the tip of the keel to the surface of the leaf lobe (along 2–3 cells). The keel is short (2–3 cells long), the proximal base of the ventral margin of the leaf lobe is usually arched, and the angle between keel and ventral margin of the leaf lobe is sharp. Cololejeunea bekkerae is related to C. planissima, but in the latter species, the lobule is not adnate to the lobe surface and the angle between keel and ventral leaf margin is very wide, forming an almost straight line. 4. Cololejeunea bischleriana Tixier

Fig. 44U

On living leaves in montane rainforest, 1900–2500  m. Colombia: Cun (type; Tixier 1980). Ecuador: ZC (Parolly et  al. 2004). General distribution: Colombia, Ecuador, Brazil. Cololejeunea bischleriana (subg. Pedinolejeunea) is a little-known South American species that is recognized by leaf lobes with a white border, dorsal leaf base not arching beyond the stem, lobules inflated-saccate, and ventral merophyte 1

446

 

Keys and Descriptions

cell wide. Cololejeunea bischleriana resembles C. vitaliana, but the ventral merophyte in the latter species is 2–3 cells wide and the dorsal leaf bases are arching widely across and beyond the stem. 5. Cololejeunea camillii (Lehm.) A.Evans Aphanolejeunea camillii (Lehm.) R.M.Schust. Aphanolejeunea ensifolia (Spruce) Herzog [Spruce] Steph.) Aphanolejeunea liliputana R.M.Schust. Aphanolejeunea misiones M.E.Reiner

Fig. 44V, W (Cololejeunea

ensifolia

On living leaves, occasionally on bark and rotten wood, in lowland and montane rainforests, from sea level to 2400  m. Colombia: Mag (Gradstein et  al. 2016). Ecuador: EO, Es, MS, Or, ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical America. Cololejeunea camillii (subg. Diaphanae) is a widespread neotropical species that is distinguished by the obliquely spreading, broadly ovate-rhomboid to ovatelanceolate leaves with an obtuse to acute apex and reduced lobules. The leaf surface is smooth, the leaf cells are 2–3× longer than wide and thin-walled, and the leaf margin cells are subquadrate. The record of C. camillii from the Chocó (Winkler 1970 as C. ensifolia) was erroneous (Gradstein et al. 2016). Cololejeunea lanceolata E.W.Jones from West Africa is very similar to C. camillii and may be a synonym. 6. Cololejeunea cardiocarpa (Mont.) A.Evans

Fig. 44X

Usually on living leaves, also on twigs and rock, in rainforests, rather dry woodlands, scrub, plantations, and gardens, from sea level to 1825 m. Colombia: Ama, Caq, Put, Vau (Campos et al. 2014). Ecuador: Az, ES, Ga, Or, ZC. General distribution: pantropical. Cololejeunea cardiocarpa (subg. Pedinolejeunea) is a widespread pantropical species that is readily recognized by the ovate-oblong leaves with a group of fingerlike hyaline cells at the apex. The leaf margins are otherwise entire and without hyaline border, and the lobules are well developed, 20–50 cells wide, ovate, inflated, ca. 1/3–1/2× lobe length, with isodiametric cells and 2 teeth. The Amazonian C. schusteri Pócs resembles C. cardiocarpa but differs in its asymmetrical, subfalcate leaves with a narrower lobule, only 12–14 elongate cells across. Its occurrence in lowland areas of Colombia and Ecuador may be expected. 7. Cololejeunea cingens (Herzog) Bernecker & Pócs Aphanolejeunea cingens Herzog

Fig. 44Y

On living leaves in lowland and montane rainforest, 250–2000 m. Ecuador: Ga (?), Ma, MS, ZC. General distribution: scattered in tropical America. Cololejeunea cingens (subg. Aphanolejeunea) is an uncommon neotropical species that is morphologically close to C. sicifolia but differs in the lobules with a smooth keel and a 2-celled, longly falcate tooth. In C. sicifolia the keel is crenate and the tooth is 1-celled (Pócs et al. 2014). The leaves in C. cingens are ovate to

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447

lanceolate and falcate, and the lobules when well developed are up to 2/3 of leaf length. 8. Cololejeunea crenata (A.Evans) Pócs Aphanolejeunea crenata A.Evans

Fig. 45A

On living leaves in montane rainforest, ca. 2800 m. Ecuador: Lo (Cajanuma, Norris et al. 91876, det. T. Pócs, EGR). General distribution: Greater Antilles, Central America, Colombia. Cololejeunea crenata (subg. Diaphanae) is morphologically close to C. diaphana but differs from the latter species in the broader, ovate-elliptical to spathulate leaves with a rounded apex, the lobule tooth 1–2 cells long and directed sideward, not forward, and by the sigmoid lobule cells (cells elongate, with undulate walls) (Pócs et al. 2014). 9. Cololejeunea diaphana A.Evans Fig. 45B Aphanolejeunea truncatifolia Horik. (Cololejeunea truncatifolia [Horik.] Pócs) Cololejeunea ensifolia (Spruce) Herzog var. pygmaea Herzog On living leaves and bark of trees in lowland and montane rainforests, occasionally on rock, 180–3100 m. Colombia: Ama, Vau (Campos et al. 2014). Ecuador: Lo, MS, ZC (Schäfer-Verwimp et al. 2013a). General distribution: pantropical. Cololejeunea diaphana (subg. Diaphanae) is a widespread pantropical species that is recognized by the ovate-lanceolate leaf lobes (2–3× longer than wide) with an obtuse apex and crenate margins due to conically elevated cells. The lobules are 1/3–1/2× lobe length and the first lobule tooth is straight and 2 cells long. Cololejeunea diaphana is closely related to C. azorica, C. crenata, C. microscopica, C. sicifolia, and C. yelitzae. Cololejeunea azorica differs in shorter leaf lobes (< 2× longer than wide) and longer lobules (> 1/2× lobe length); C. crenata in broader, ovate-elliptical leaves with a rounded apex and sigmoid lobule cells; C. microscopica in conically elevated leaf cells and a falcate lobule tooth; and C. sicifolia in strongly elongate leaves (4–6× longer than wide) and a falcate lobule tooth. For differences with C. yelitzae, see the key. 10. Cololejeunea ecuadoriensis Pócs

Fig. 45C

On living leaves and twigs of shrubs, 500–1950  m. Ecuador: Es, Pi (type). General distribution: Ecuador, Panama. Cololejeunea ecuadoriensis (subg. Pedinolejeunea) is a rare neotropical species that is distinguished by the ovate-oblong, falcate leaves without white margin and with an obtuse apex tipped by an elongate, papilla-like hyaline cell. The plants are ca. 1–1.5 mm wide; the lobules are large, 2/5–1/2× leaf length; the first lobule tooth is 2 cells long; and the stylus is well developed, 3–8 cells long (Pócs 2002). 11. Cololejeunea gracilis (Jovet-Ast) Pócs Aphanolejeunea gracilis Jovet-Ast Aphanolejeunea heterophylla R.M.Schust.

Fig. 45D

On living leaves, also on bark, 200–2050 m. Colombia: Boy, Cho, Hui, Put, Ris. Ecuador: Es, Ga, MS, Or, ZC. General distribution: pantropical.

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Keys and Descriptions

Fig. 45 (A) Cololejeunea crenata, leaf. (B) Cololejeunea diaphana, portion of stem with leaves. (C) Cololejeunea ecuadoriensis, portion of stem with leaf. (D) Cololejeunea gracilis, habit. (E) Cololejeunea hildebrandii, portion of stem with leaf. (F) Cololejeunea linopteroides, leaf. (G) Cololejeunea microscopica, leaf. (H) Cololejeunea minuscula, habit. (I and J) Cololejeunea obliqua. (I) Leaf cells, dorsal view. (J) Lobule apex. (K) Cololejeunea papilliloba, leaf. (L) Cololejeunea papillosa, habit. (M) Cololejeunea planissima, lobule. (N) Cololejeunea planiuscula, leaf.

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Cololejeunea gracilis (subg. Aphanolejeunea) is a very small and common epiphyll (less than 0.4 mm wide) with mostly reduced, linear leaves with toothed margins. The species is widely distributed and closely similar to C. minuscula and C. papillosa; for differences see the key. 12. Cololejeunea hildebrandii (Austin) Steph.

Fig. 45E

On living leaves in Pacific coastal rainforest, 100 m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). General distribution: pantropical. Cololejeunea hildebrandii (subg. Leptocolea) is a widespread pantropical species that is readily recognized by the characters given in the key. The species seems to be rare in Colombia and Ecuador and has only been recorded from the everwet lowland rainforests of the Chocó. 13. Cololejeunea kuciana Pócs & Schäf.-Verw. On filmy ferns in montane rainforest, ca. 1900  m. Ecuador: ZC (Pócs and Schäfer-Verwimp 2012). General distribution: only known from the type from southern Ecuador. Cololejeunea kuciana (subg. Cololejeunea) is a rare Ecuadorian endemic that is characterized by the minute plants (ca. 0.3 mm wide) with acute, ovate-lanceolate leaves with strongly serrate-papillose margins and surface and large lobules (ca. 1/2 of leaf length) with a smooth surface, 2 small obtuse teeth, and distal free margin formed by 3–4 large cells. Cololejeunea kuciana is closely similar to C. calcarea (Lib.) Steph., the type of the genus Cololejeunea, but differs from the latter in the lack of a stylus (stylus well developed in C. calcarea). It also resembles the papillose members of subg. Diaphanae (C. contractiloba A.Evans, C. jamesii (Austin) M.E.Reiner & Pócs, C. papilliloba) but differs from them by the absence of Aphanolejeunea-type branches and the distal free margin of the lobule with 3–4 large cells. 14. Cololejeunea latilobula (Herzog) Tixier On living leaves in lowland rainforest, 450  m. Ecuador: Na (Jatun Sacha, R.  Lücking s.n., det. T.  Pócs, EGR). General distribution: pantropical; in the neotropics only known from Ecuador and Bolivia (Pócs et al. 2014).

(O) Cololejeunea platyneura, leaf, showing vitta. (P) Cololejeunea pterocolea, habit with gynoecium. (Q and R) Cololejeunea sicifolia. (Q) Leaf. (R) Lobule. (S) Cololejeunea stotleriana, habit, showing numerous small male branches. (T) Cololejeunea subcardiocarpa, leaf. (U) Cololejeunea subsphaeroidea, leaf. (V) Cololejeunea surinamensis, portion of leaf showing hyaline border. (W and X) Cololejeunea verwimpii. (W) Lobule. (X) Leaf. (Y) Cololejeunea vitaliana, portion of stem with leaf. (Z) Cololejeunea winkleri, habit. (A, D, E, I, J, M, U, Y redrawn from Pócs et al. 2014; C redrawn from Pócs 2002; N redrawn from Tixier 1991; P redrawn from Herzog 1952; S from Gradstein et al. 2011; T redrawn from Tixier 1980; rest from Gradstein and Ilkiu-Borges 2009)

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Keys and Descriptions

Cololejeunea latilobula (subg. Pedinolejeunea) is widespread in the paleotropics but apparently rare in the neotropics. It is closely similar to C. planissima but differs in the 2–5-cell-wide ventral merophyte (1 cell wide in C. planissima) and in the constantly flat lobules without teeth. 15. Cololejeunea linopteroides H.Rob.

Fig. 45F

On living leaves in lowland rainforest, ca. 450 m. Ecuador: Na (Jatun Sacha, Schäfer-Verwimp & Nebel 31728, det. T. Pócs, EGR); to be expected in Colombia. General distribution: Central America, Guianas, Ecuador. Cololejeunea linopteroides (subg. Chlorocolea) is a very characteristic lowland rainforest species that seems to have no close relatives in Ecuador and Colombia. The plants grow very tightly fixed to the substrate; the stems are flattened, ribbonlike with a broad, 2–5-cell-wide ventral merophyte; the leaves are suborbicular, with highly reduced lobules consisting of only one cell; and the leaf cells are smooth and strongly sigmoid (cells elongate, walls sinuate). In its suborbicular leaves with highly reduced lobules, the species somewhat resembles C. tamasii Schäf.-Verw. from Panama, but the stems in the latter species are not flattened and the leaf cells in C. tamasii are not sigmoid. 16. Cololejeunea microscopica (Taylor) Schiffn. Aphanolejeunea exigua A.Evans Aphanolejeunea microscopica (Taylor) A.Evans (incl. vars.) Aphanolejeunea verrucosa Jovet-Ast Cololejeunea megalandra (Spruce) Steph.

Fig. 45G

On living leaves, bark, and rotten wood in lowland and montane rainforest, Polylepis forest, and páramo, from sea level to 3800 m. Colombia: Ama, Boy, Caq, Cas (Gradstein 17210 p.p. among Drepanolejeunea inchoata, PC), Cho, Mag, Put, Vau (Campos et al. 2015 as C. contractiloba). Ecuador: Im, Ba, Or, Pa, Pi, Su, ZC.  General distribution: tropical and subtropical America, Africa, western Europe. Cololejeunea microscopica (subg. Aphanolejeunea) is a common, widespread species with a remarkably wide elevational range. It is the only Cololejeunea species that occurs in páramo. Habitually (under the handlens) it resembles a small Microlejeunea, but the leaves in C. microscopica are more distant. The main characters of the species are (1) branches without collar at the base; (2) leaves longer than wide, apex acute-acuminate; (3) cells on the dorsal leaf surface usually conically mammillose; and (4) lobules large, 1/2–3/4× lobe length, with a falcate first lobule tooth. Several varieties have been described based on the degree of mamillosity of the leaves (Pócs 2003), but these are part of a variation continuum and are no longer recognized (T. Pócs, pers. comm.). Cololejeunea sintenisii (Steph.) Pócs (= Aphanolejeunea ephemeroides R.M.Schust., Cololejeunea ephemeroides [R.M.Schust.] Stotler & Crand.-Stotl.), which has been recorded a few times from Colombia and Ecuador, is very similar to C. microscopica, but reduced leaves are more common in C. sintenisii and the welldeveloped lobules are smaller, 0.3–0.5× leaf length.

Lejeuneaceae

17. Cololejeunea minuscula Pócs Aphanolejeunea minuta R.M.Schust.

451

Fig. 45H

On living leaves in (sub)montane rainforest, rarely on bark, 950–1500  m. Colombia: Ris (Gradstein and Uribe 2016a). Ecuador: EO, ZC (Schäfer-Verwimp et al. 2013a). General distribution: scattered in tropical and subtropical America. Cololejeunea minuscula (subg. Aphanolejeunea) is an uncommon, very small epiphyll (less than 0.4  mm wide) with stems without medullary cells and with mostly reduced, linear leaves with entire or crenulate margins. The species is closely similar to C. gracilis and C. papillosa; for differences see the key. 18. Cololejeunea obliqua (Nees & Mont.) Schiffn. Cololejeunea scabrifolia Steph.

Fig. 45I, J

On living leaves and bark of trees in lowland and lower montane rainforest, from sea level to 2000 m. Colombia: Ces, Cho. Ecuador: Es, MS, Na, Or, ZC. General distribution: pantropical. Cololejeunea obliqua (subg. Leptocolea) is a common and widespread rainforest species that is distinguished by ovate-oblong leaves with a broadly rounded apex, denticulate margins, unipapillose leaf cells, and fully or partially inflated lobules (1/3–1/2× lobe length) with 1–2 straight teeth. 19. Cololejeunea papilliloba (Steph.) Steph. Aphanolejeunea papilliloba (Steph.) Pócs & Bernecker Aphanolejeunea kunertiana Steph. Aphanolejeunea pustulosa (Jovet-Ast) Bernecker & Pócs

Fig. 45K

On living leaves in montane rainforest, 1900 m. Colombia: Cho (Winkler 1970 as A. kunertiana). Ecuador: ZC (Parolly et  al. 2004). General distribution: scattered in tropical America; also in Africa (Bioko). Cololejeunea papilliloba (subg. Diaphanae) is an uncommon species that is recognized by the rounded leaf apex; the conically elevated leaf cells crowned by a large, rounded papilla; and the frequently reduced lobules. The lobule surface is smooth and the first tooth of the lobule is long and straight, 2-celled. Cololejeunea papilliloba is morphologically close to C. contractiloba A.Evans, a species not known with certainty from Colombia and Ecuador (reports from the Colombian Amazon are C. microscopica), but C. contractiloba differs from C. papilliloba in the papillose lobule surface and the reduced first lobule tooth. 20. Cololejeunea papillosa (K.I.Goebel) Mizut.  Aphanolejeunea angustissima Steph. Aphanolejeunea cyathiphylla Herzog

Fig. 45L

On living leaves in lowland and montane rainforest and scrub, 200–2850  m. Ecuador: Es, EO, LR, Lo, Or, Su, ZC; not known from Colombia but expected there. General distribution: tropical America, Southeast Asia; to be expected in tropical Africa. Cololejeunea papillosa (subg. Aphanolejeunea) is a widespread. possibly pantropical species that is distinguished by the very small, epiphyllous plants (less than

452

 

Keys and Descriptions

0.4 mm wide) with mostly reduced, linear leaves with entire or crenulate margins. The species is closely similar to C. gracilis and C. minuscula; for differences see the key. 21. Cololejeunea planissima (Mitt.) Abeyw.  Cololejeunea lanciloba Steph. Cololejeunea manaosensis (Herzog) Herzog Cololejeunea planifolia (A.Evans) R.M.Schust. Cololejeunea subscariosa (Spruce) Pócs (?) Cololejeunea tonkinensis Steph.

Fig. 45M

On living leaves, bark, and dead wood in lowland and montane rainforest, from sea level to 2000 m. Colombia: Cho (Winkler 1970 as C. planifolia). Ecuador: Es, ZC (Schäfer-Verwimp et al. 2013a as C. manaosensis and C. planifolia). General distribution: pantropical. Cololejeunea planissima (subg. Pedinolejeunea) is a widespread and variable, pantropical species that is recognized by the white-bordered leaves with a smooth cuticle and with a flat, ligulate to broadly triangular lobule, which is frequently ± erect and parallel to the stem (Pócs et  al. 2014). A few wide-spreading, inflated lobules may sometimes be present as well. The species is rare in Colombia and Ecuador, where it has been recorded three times. Cololejeunea lanciloba, which is often accepted as a good species, is included here in C. planissima. Cololejeunea subscariosa, a species very similar to C. planissima (see Pócs et al. 2014), may be a further synonym. 22. Cololejeunea planiuscula Tixier

Fig. 45N

On living leaves in Pacific coastal rainforest, 100 m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). General distribution: Mexico, Dominican Republic, Colombia, Brazil. Cololejeunea planiuscula (subg. Leptocolea) is a rare epiphyll that resembles C. obliqua in habit and leaf shape; for differences see the key. 23. Cololejeunea platyneura (Spruce) A.Evans

Fig. 45O

On living leaves in Amazonian and Pacific coastal rainforests, 100–250  m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). Ecuador: Or (SchäferVerwimp et al. 2013a). General distribution: pantropical; in tropical America only known from Amazonia and the Chocó. Cololejeunea platyneura (subg. Cololejeunea) is a characteristic epiphyll of lowland rainforests that is readily recognized by the presence of a group of enlarged cells (6–18) at the leaf base. The plants are rather small, to 1 mm wide, the leaves are ovate-falcate with entire margins and a rounded apex, and the leaf cells are unipapillose on the dorsal surface. 24. Cololejeunea pterocolea Herzog  Aphanolejeunea pterocolea (Herzog) R.M.Schust.

Fig. 45P

On living leaves in submontane rainforest, 700 m. Ecuador: MS (type; Herzog 1952). General distribution: only known from the type from Ecuador.

Lejeuneaceae

453

Cololejeunea pterocolea (subg. Aphanolejeunea) is a very rare epiphyll that is hitherto only known from one locality in the eastern foothills of the Ecuadorian Andes. The plants are very small, 0.5 mm wide; the leaves are lanceolate (ca. 3× longer than wide) with an obtuse apex, thin-walled cells, and reduced lobules; and the perianths have 5 wing-like, flat horns. Cololejeunea pterocolea is morphologically close to C. camillii but differs from the latter species in the horned perianth, smaller plant size, and narrower leaves. In its horned perianth, C. pterocolea is similar to C. cornuta E.W.Jones from tropical Africa. The latter species is a more robust plant with less elongate leaves. 25. Cololejeunea sicifolia (A.Evans) Pócs & Bernecker [“sicaefolia”]Fig. 45Q, R Aphanolejeunea sicifolia A.Evans On living leaves, rotten wood, and rock, 200–2700 m. Colombia: Val (Herzog 1955). Ecuador: Ga (Gradstein 2009), ZC (Schäfer-Verwimp et  al. 2013a). General distribution: tropical America. Cololejeunea sicifolia (subg. Aphanolejeunea) is a common and characteristic, mostly montane species recognized by narrowly lanceolate leaf lobes (3–4× longer than wide) with an acuminate apex and lobules less than 1/2× leaf length, with a crenate keel and a falcate tooth, the tooth 1–2 cells long. Rudimentary leaves are rather scarce. The plants from Zamora Chinchipe belong to C. sicifolia subsp. jamaicensis (R.M.Schust.) Bernecker & Pócs, characterized by acutely papillose dorsal leaf sufaces. Cololejeunea sicifolia resembles C. cingens; for differences see under the latter species. 26. Cololejeunea stotleriana Gradst., Ilk-Borg. & Vanderp. Schusterolejeunea andina Gradst. & Ilk.-Borg., nom. inval.

Fig. 45S

Rheophyte on periodically inundated living leaves of treelets, shrubs, and ferns along river in wet lower montane rainforest area, 900–1000 m. Ecuador: ZC (Río Nangaritza; Gradstein et al. 2011). General distribution: only known from the type locality in southeastern Ecuador. Cololejeunea stotleriana (subg.?) is a robust rheophilous epiphyll that is only known from the lower Río Nangaritza (Ecuador), where it occurs abundantly on the leaves of periodically inundated shrubs and ferns (Gradstein et al. 2011). The species was considered to be a member of subg. Chlorolejeunea, a small group of rheophilous species with robust stems and clustered gynoecia, but molecular analysis showed that subg. Chlorolejeunea is polyphyletic (Yu et al. 2013). Cololejeunea stotleriana is distinguished by (1) plants 2–3.5  mm wide, densely pinnate, with numerous short male and female branches, the female branches with clustered gynoecia; (2) stems rigid, without hyalodermis, with 9–12 rows of epidermal cells, ventral merophyte 3–5 cells wide; (3) leaves wide-spreading, oblong, with rounded tips and smooth, thin-walled cells; (4) lobules flat, ± triangular; and (5) rhizoid discs large, swollen, not developed at each leaf base. By its distinguishing characters, C. stotleriana cannot be confused with any other neotropical species of Cololejeunea.

454

27. Cololejeunea subcardiocarpa Tixier Cololejeunea yanoanae Tixier

 

Keys and Descriptions

Fig. 45T

On living leaves, 0–2000 m. Ecuador: ZC (Parolly et al. 2004). General distribution: scattered in tropical America. Cololejeunea subcardiocarpa (subg. Pedinolejeunea) is a poorly defined neotropical species distinguished by the leaves with a narrow, white or hyaline border (1 cell wide) and with finger-like hyaline cells at the apex. The lobules are relatively large, 1/3–1/2× lobe length and inflated, and the leaf cells are smooth. The species is quite similar to C. verwimpii, but the lobules in the latter species are smaller. Possibly the two species are conspecific. 28. Cololejeunea subsphaeroidea (R.M.Schust.) Pócs Aphanolejeunea subsphaeroidea R.M.Schust.

Fig. 45U

On living leaves of terrestrial bromeliad in upper montane rainforest, 2890 m. Ecuador: Lo (Schäfer-Verwimp et al. 2013a). General distribution: scattered in tropical America. Cololejeunea subsphaeroidea (subg. Aphanolejeunea) is a rare neotropical species recognized by the minute, subglobose leaves with lobules almost as long as the lobes and with a decurved, rounded leaf apex covering the mouth of the lobules. 29. Cololejeunea surinamensis Tixier

Fig. 45T

On bark and living leaves in lowland rainforest, 100–200  m. Colombia: Caq (Benavides 2004), Cho. General distribution: Amazonia, Guianas, Chocó. Cololejeunea surinamensis (subg. Pedinolejeunea) is an Amazonian rainforest species distinguished by leaves with a white border and an entire margin, inflated lobules with 2 conspicuous teeth, and a 1-cell-wide ventral merophyte. The species is most closely similar to C. submarginata Tixier, a widespread neotropical species which has not yet been recorded from Colombia and Ecuador, but in the latter species, the leaf margin is crenulate and the lobule has only 1 tooth. Cololejeunea surinamensis also resembles C. vitaliana, but the latter species has a 2–4-cell-wide ventral merophyte and a broad dorsal leaf base arching widely across and beyond the stem. 30. Cololejeunea verwimpii Tixier

Fig. 45W, X

On living leaves in submontane and montane rainforests, 650–2000  m. Colombia: Qui (Orrego and Uribe 2004). Ecuador: MS, ZC. General distribution: scattered in tropical America. Cololejeunea verwimpii (subg. Pedinolejeunea) is an uncommon neotropical epiphyll recognized by leaves with a narrow, white or hyaline border (1 cell wide) and with finger-like hyaline cells at the apex. The lobules are small, less than 1/4 of lobe length, inflated or flat, and the leaf cells are smooth to finely striate-papillose. The species resembles C. subcardiocarpa, but the lobules in the latter species are larger, usually more than 1/3× lobe length, and always inflated.

Lejeuneaceae

31. Cololejeunea vitaliana Tixier

455

Fig. 45Y

On living leaves in montane rainforest, 2020 m. Ecuador: ZC (Schäfer-Verwimp et  al. 2013a). General distribution: Mexico, Costa Rica, southern Ecuador, southeastern Brazil. Cololejeunea vitaliana (subg. Pedinolejeunea) is a neotropical epiphyll distinguished by (1) leaves with a white border and a wide dorsal base, which is widely arching across and beyond the stem; (2) lobules inflated and with 2 conspicuous teeth; and (3) ventral merophyte 2–3 cells wide. The species is closely similar to C. surinamensis; for differences see the key. 32. Cololejeunea winkleri (M.I.Morales & A.Lücking) Pócs

Fig. 45Z

On living leaves in lowland and submontane rainforest, 200–650 m. Ecuador: MS, Na, Or (Schäfer-Verwimp et al. 2013a). General distribution: Amazonia, Costa Rica. Cololejeunea winkleri (subg. Aphanolejeunea) is a tiny Amazonian species (0.3–0.4 mm wide) with ovate-oblong leaves which are strongly contracted at the junction of lobe and lobule; 2 lobe cells are protruding at the junction. The leaf cells are smooth and the lobules are large, ca. 1/2× lobe length, with a 2-celled, straight or slightly incurved first tooth. Note that the correct spelling of the epithet is “winkleri” (not “winklerii”) (see Turland 2013, pp. 91–93).

Further Records Cololejeunea conchifolia (Gottsche) Gradst. – Colombia: Cun (Gottsche 1864 as Lejeunea conchifolia Gottsche); the identity of the species is unclear (Pócs et al. 2014). Cololejeunea cremersii Tixier (= Cololejeunea fefeana Tixier) – Colombia: Cho (Vasco et  al. 2002 as C. fefeana). A rare species from French Guiana and Costa Rica; the identification should be checked.

Excluded Record Cololejeunea contractiloba A.Evans  – Colombia: Ama, Caq, Put, Vau (Campos et al. 2015); the material belongs to C. microscopica. Colura (Dumort.) Dumort. Colurolejeunea (Spruce) Steph., Macrocolura R.M.Schust., Myriocolea Spruce Plants 0.5–2 mm wide (to 5 mm in C. irrorata), pale green to yellow green to dark green, becoming brownish when dry, creeping. Stems with or without

456

 

Keys and Descriptions

hyalodermis, usually of 10–11 rows of cells (to 250 rows of cells in C. irrorata); ventral merophyte 2(−14) cells wide. Branching Lejeunea-type and Radula-type. Leaf attached to stem by only few cells, insertion very short, transverse. Leaves usually upright, longer than wide, in the upper half usually with a closed sac formed by the lobule, sac occasionally absent, margins entire or toothed. Cells of the lobe isodiametric-hexagonal to elongate, smooth or mammillose-papillose, usually with small trigones; oil bodies homogeneous or finely granular; ocelli lacking. Lobules longer than the lobe, cylindrical, from a narrow base usually widening above into a closed sac, the sac rounded, conical or cylindrical, sometimes with a long, linear beak, the opening at the base closed by a small valve. Underleaves present near the base of each leaf, distant or overlapping, bifid, lobes often strongly diverging, margins entire, insertion line almost straight. Gynoecia on long or short shoots, with 1–2 innovations. Perianths with 3–5 smooth or horned keels. Sporophytes lejeuneoid. Vegetative reproduction by multicellular, disciform gemmae. A large pantropical genus (ca. 70 spp.), 12 species in Colombia and Ecuador. Colura is readily recognized by the peculiar leaves with a very short insertion and an apical sac formed by the lobule. The sac varies greatly in shape and size and is sometimes lacking (e.g., in C. irrorata); it has a complicated aperture mechanism made up of a valve and a hinge. Based on valve morphology, Colura is subdivided into several sections (JovetAst 1953). One of these sections, sect. Lingua Jovet-Ast with one species in tropical America (C. sagittistipula), has been raised to generic level, Macrocolura R.M.Schust., characterized by large valves without hinge and asymmetrically decurrent underleaves (Schuster 1994; Zhu and Grolle 2002). Macrocolura was not strongly supported in a molecular analysis (Heinrichs et al. 2013b), however, and is maintained here in Colura. The underleaves in Colura are produced in a pendular fashion like in Diplasiolejeunea, each leaf being associated with an underleaf. This is a neotenic feature and interpreted as an adaptation to survival of the species to the short-lived, epiphyllous or ramicolous habitats (see also under Cololejeunea). Literature: Jovet-Ast (1953), Zhu and Grolle (2002 as Macrocolura), Heinrichs et al. (2013b) 1. Plants robust, 3–5 mm wide, growing upright or pendent along rivers. Leaves without apical sac, lobule usually absent (present on creeping shoots). Upright stems rigid, made up of 150–250 rows of cells, ventral merophyte 10–14 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. irrorata 1. Plants smaller, 0.5–1.5  mm wide. Leaves with apical sac and lobule. Stems fragile, made up of 10–11 rows of cells, ventral merophyte 1–2 cell wide. . . 2 2. Sac above with a linear, horn-like or beak-like prolongation . . . . . . . . . . . . . 3 2. Sac without prolongation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 3. Prolongation very short, maximally 1/5 of the length of the sac, often less (resembling a short beak). Plants growing in páramo . . . . . C. ornithocephala 3. Prolongation longer, 1/3–3/4 of the length of the whole leaf (including prolongation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Lejeuneaceae

457

4. Prolongation shorter than the rest of the leaf, sac rather abruptly narrowed to the prolongation. Plants growing above 2000 m . . . . . . . . . . . . C. calyptrifolia 4. Prolongation longer than the rest of the leaf, sac very gradually narrowed to the prolongation. Plants growing from sea level to the páramo . . . . . C. tenuicornis 5. Underleaves imbricate, emarginate or short-bifid, to 1/4 of underleaf length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Underleaves distant, more deeply bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Underleaves with a large triangular tooth near the base. Sac about 1.5× larger than the rest of the leaf. Plants growing below 2000 m, on bark. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sagittistipula 6. Underleaf without tooth near the base. Sac 2–3× larger than the rest of the leaf. Plants growing above 4000 m, in bogs . . . . . . . . . . . . . . . . . . . . . . . C. stotleri 7. Plants of upper montane cloud forest and páramo, 2500–4000 m. Perianth with 5 short keels or horns. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Plants of lowland and lower montane rainforest, below 2000  m. Perianth 3-keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Sac 1.5–2×wider than the rest of the leaf. Surface of sac smooth. Apex of sac broadly rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. naumanii 8. Sac scarcely wider than the rest of the leaf. Surface of sac crenulate due to mammillose-papillose cells. Apex of sac obtuse to acute (sometimes terminating in a short beak) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ornithocephala 9. Surface of sac crenate due to mammillose cells, especially near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Surface of sac smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 10. Sac oblong, 1.5–2× longer than wide, apex broadly rounded. Valve firmly attached at the base, not easily detached . . . . . . . . . . . . . . . . . C. greig-smithii 10. Sac narrowly conical-triangular, ca. 3–4× longer than wide, apex obtuse to acute. Valve loosely attached at the base, easily detached . . . . . . . . . . . . . . . 11 11. Valve with 15–25 median cells, of which the two basal ones are quadrate to narrowly rectangular in shape. Perianth 3-keeled. Plants of lowland rainforest (below 1000 m), on bark or living leaves . . . . . . . . . . . . . . . . . . . . C. tortifolia 11. Valve with ca. 10 median cells, of which the two basal ones are triangular in shape. Perianth with 5 acute horns. Plants of Peru, in montane rainforest (ca. 2000  m), epiphyllous (Pócs 2019a), to be expected in Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ochyrana Pócs 12. Apex of sac narrowly obtuse. Valve with 1 median basal cell (valve to be detached for observation of this character) . . . . . . . . . . . . . . . . . . . . . . . C. ulei 12. Apex of sac broadly rounded. Valve with 2 median basal cells . . . . . . . . . . . 13 13. Sac 1–1.5× longer than wide. Underleaf lobes 6–9 cells wide at base. Plants only known from the Galápagos Islands . . . . . . . . . . . . . . . . . . . . . . . C. andoi 13. Sac 2–4× longer than wide. Underleaf lobes 4 cells wide at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cylindrica 1. Colura andoi Gradst. & Jovet-Ast

Fig. 46A

On bark of tree near the forest line on the Galápagos Islands, 600–650  m. Ecuador: Ga (Santa Cruz; Gradstein 1986). General distribution: only known from the Galápagos Islands.

458

 

Keys and Descriptions

Colura andoi is a rare Galápagos endemic that is closely similar to C. cylindrica but differs in the shorter, ovate-subcylindrical sac (1.5–2× longer than wide) and the broader underleaf lobes, which are 6–9 cells wide at base (4 cells wide at base in C. cylindrica). 2. Colura calyptrifolia (Hook.) Dumort.

Fig. 46B

On branches and twigs of shrubs and on living leaves in upper montane rainforest and páramo, 2300–3750  m. Colombia: Cas, Ris (Gradstein et  al. 2018c). Ecuador: Ca, Lo, Na, Pi (Schäfer-Verwimp et al. 2013a). General distribution: oceanic and cool-temperate regions of South America, northern Andes, West Indies, tropical Africa, western Europe. Colura calyptrifolia, type of the genus Colura and a widespread species, is rare in the Andes where it occurs at high elevation, mostly above 3000 m. It is readily distinguished by the sac with a rather short prolongation, ca. 1/3–2/5× the length of the leaf. The species is close to the pantropical C. tenuicornis, but in the latter species, the prolongation of the sac is longer, at least half the length of the whole leaf, and the transition between sac and prolongation is more gradual. Moreover, C. tenuicornis occurs at lower elevation. 3. Colura cylindrica Herzog

Fig. 46C

Sun epiphyte on bark, twigs, and living leaves in lowland and (sub)montane rainforests, woodlands, and plantations, 100–650(−1800) m. Colombia: Cho, Hui, Vau. Ecuador: Ga, MS.  General distribution: scattered in tropical America; also in tropical Africa (Tanzania). Colura cylindrica is a widespread Afro-American species that is characterized by the smooth, cylindrical sac (2–4× longer than wide), which is not narrowed to apex. 4. Colura greig-smithii Jovet-Ast

Fig. 46D

On living leaves, bark, and rotten wood in lowland rainforest, 100–650  m. Colombia: Ama, Caq, Cho, Put, Vau (Campos et al. 2015). Ecuador: MS, Na, Or (Schäfer-Verwimp et  al. 2013a). General distribution: scattered in tropical America. Colura greig-smithii is an uncommon neotropical species recognized by the oblong sac with rounded apex and a crenulate surface due to mammillose cells. The valve in this species is firmly fixed at the base and not easily detached. Colura greigsmithii resembles C. cylindrical, but in the latter species, the surface of the sac is smooth and the valve is loosely attached and easily detached. 5. Colura irrorata (Spruce) Heinrichs et al. Myriocolea irrorata Spruce

Fig. 46J

Rheophilous epiphyte growing on periodically inundated subshrubs, especially Cuphaea bombonasae, in river beds with large granite boulders, 1200–1700  m. Ecuador: Tu (Río Topo, type), ZC (Río Nangaritza, Gradstein and Benitez 2014a; Río Machinaza, Perez 11329, PC, QCA); to be expected in Colombia. General distribution: only known from the Ecuadorian Andes.

Fig. 46 (A) Colura andoi, leaf. (B) Colura calyptrifolia, leaf. (C) Colura cylindrica, leaf. (D) Colura greig-smithii, leaf. (E) Colura naumannii, leaf. (F) Colura ornithocephala, leaf. (G) Colura sagittistipula, leaf. (H) Colura tortifolia, leaf. (I) Colura ulei, leaf. (J) Colura irrorata, portion of stem with leaves. (K and L) Cyclolejeunea accedens. (K) Habit. (L) Leaf with gemmae. (M and N) Cyclolejeunea chitonia. (M) Habit. (N) Leaf with gemma. (O) Cyclolejeunea convexistipa. (P and Q) Cyclolejeunea foliorum. (P) Habit with gynoecium. (Q) Leaf. (R–T) Cyclolejeunea glimeana. (R) Habit. (S) Portion of stem with underleaf and lobule. (T) Gemma. (U) Cyclolejeunea luteola, habit. (V) Cyclolejeunea peruviana, habit. (A from Gradstein 1986; B redrawn from Smith 1990; E, I redrawn from Jovet-Ast 1953; F redrawn from Herzog 1952; K, L redrawn from BerneckerLücking 1998; P, Q redrawn from Herzog 1931; R–T from Gradstein and Reiner-Drehwald 2017a; rest from Gradstein et al. 2001; Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009)

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Keys and Descriptions

Colura irrorata is a striking Ecuadorian endemic that is morphologically very different from other members of Colura and from Lejeuneaceae in general. The species is known from blackwater rivers in the Ecuadorian Andes (Río Topo, Río Machinaza, Río Nangaritza) characterized by large granite boulders, very nutrientpoor water, and strongly fluctuating water levels. At the type locality (Río Topo), the species is endangered due to a hydroelectric project. Colura irrorata is distinguished by (1) plants very robust and superfertile, to 5 cm long and 5 mm wide, standing away perpendicularly from the substrate or pendent, with a short creeping rhizomelike shoot, fertile shoots with strongly undulate-crispate leaves and numerous shortsexual branches arising from almost every leaf axil, each branch giving rise to dense clusters of gynoecia and androecia; (2) stems rigid, very thick, ventral merophyte 10–14 cells wide, rhizoids absent; (3) branching Radula-type (without collar), rarely Lejeunea-type; (4) leaves 2–4 mm long, imbricate, suborbicular, undulate-crispate, transversely inserted along a long insertion line, lobule absent (present on leaves of creeping shoots); (5) underleaves minute, bifid, or absent; and (6) plants autoicous or paroicous, antheridia 2–7 per bract (Spruce 1884 as Myriocolea irrorata; Gradstein et al. 2004 as M. irrorata). In spite of its unusual morphology, this species is clustered with members of Colura in a molecular-phylogenetic analysis (Heinrichs et al. 2013b). The distribution and habitat of C. irrorata resemble that of Lejeunea topoensis. Rivers in the Andes with a similar ecology should be searched for additional localities of these unusual rheophytes (see Gradstein and Benitez 2014a, Fig. 1). 6. Colura naumannii (Schiffn.) Steph. Colura patagonica Jovet-Ast

Fig. 46E

On bark of trees and shrubs in upper montane forest and páramo, 2550–3850 m. Colombia: Boy, Ris. Ecuador: Ca, Co, Lo, Pi, ZC. General distribution: Costa Rica to Tierra del Fuego. Colura naumannii is a characteristic Andean-Patagonian disjunct recognized by the large, suborbicular sac, distinctly wider than the rest of the leaf and with a broadly rounded apex. The underleaves are deeply bifid (to 1/2–3/4), the leaf cells have conspicuous trigones, and the perianth has 5 low, bulbous horns. 7. Colura ornithocephala Herzog Colura kilimanjarica Jovet-Ast & Pócs

Fig. 46F

On tree trunks and on twigs of shrubs in (sub)páramo, 2950–4020 m. Colombia: Cun, Ris, Tol. Ecuador: Az, Ca, Co, Na, Pi. General distribution: northern Andes (Venezuela to Ecuador), East Africa. Colura ornithocephala is characteristic of trees and shrubs near the forest line in the northern Andes. The record from low elevation on the Galápagos Islands (Clark 1953) should be erroneous (León-Yánez et al. 2006). The species is recognized by the sac about as wide as the rest of the leaf, with an obtuse or acute apex and a crenulate surface due to conically projecting, mammillose cells crowned by a papilla. Sometimes the sac terminates in a short beak (hence its name “ornithocephala”). The perianth has 5 strongly crenate keels, each terminating in a short horn.

Lejeuneaceae

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Colura ornithocephala is most closely related to C. naumannii, which grows in the same habitat, but the sac in C. naumannii is smooth, distinctly wider than the rest of the leaf and with a broadly rounded apex. 8. Colura stotleri J.C.Benavides & Uribe In Distichia bogs in superpáramo, 4400  m. Colombia: Boy (Benavides et  al. 2020). General distribution: only known from Colombia. This new species grows up to 3-cm-high, yellow-golden tufts on Distichia cushions (J. C. Benavides, pers. comm.) and is characterized by the huge, orbicular sac, which is much larger than the rest of the leaf, and the ligulate underleaves. By its distinctive characters, C. stotleri approaches C. itatyana Steph. from Serra do Itatiaia, Brazil. 9. Colura sagittistipula (Spruce) Steph. Macrocolura sagittistipula (Spruce) R.M.Schust. Colura clavigera Jovet-Ast Colura lyrata Steph.

Fig. 46G

On bark of trees in lowland and lower montane rainforests and elfin forest, from sea level to 1800  m. Colombia: Ama, Caq, Put, Ris, Vau (Campos et  al. 2015). Ecuador: Na, Or, ZC. General distribution: scattered in tropical America. Colura sagittistipula is an uncommon neotropical species that is readily recognized by its peculiar, large, imbricate underleaves, which are shallowly bifid (to 1/4); possess a triangular, lobe-like tooth on the outer margins, near the base (hence the name “sagittistipula”); and are longly decurrent on one side, in an alternate, leftright fashion (Zhu and Grolle 2002). The leaves stand upward and are narrowly elongate, to 2 mm long and ca. 5× longer than wide; the sac is bulbous-ellipsoid, about 1/2× leaf length, with a smooth or mammillose surface and a rounded apex; and the valve is very large, lingulate, with a whitish border and without hinge. The perianth has 5 inflated, smooth keels. 10. Colura tenuicornis (A.Evans) Steph. Sun epiphyte on bark and living leaves in lowland and montane rainforests, from sea level to 2900 m. Colombia: Ant, Caq, Cun, Put, Ris, San, Vau. Ecuador: Az, Ca, EO, Ga, Lo, LR, Pi, ZC. General distribution: pantropical. This widespread species is recognized by the long-linear, horn-like prolongation of the sac, measuring more than half the length of the entire leaf (including prolongation). Colura tenuicornis is closely related to C. calyptrifolia and may be confused with the latter species, but the prolongation of the sac is shorter in C. calyptrifolia, measuring less than half the length of the whole leaf. 11. Colura tortifolia (Nees & Mont.) Trevis.

Fig. 46H

On bark and living leaves in lowland and submontane rainforests, preference for rather open environments, from sea level to 1060 m. Colombia: Cho (Vasco et al. 2002). Ecuador: EO (Reserva Buenaventura, on twigs, Benitez et  al. 1312, HUTPL), Es, MS, Pa, ZC. General distribution: widespread in tropical America.

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Keys and Descriptions

Colura tortifolia is a rather common neotropical species that is recognized by the narrowly elongate, conical-triangular sac (3–4× as long as wide) with a crenate surface, especially near the tip. 12. Colura ulei Jovet-Ast

Fig. 46I

On bark and living leaves in wet lowland rainforest, ca. 100 m. Colombia: Cho, Val. General distribution: scattered in tropical America. Colura ulei is an uncommon neotropical species that is similar to C. tortifolia in having a narrowly elongate, conical-triangular sac (3–4× as long as wide), but the surface of the sac is smooth (crenate in C. tortifolia). Possibly, C. ulei is merely a variety of C. tortifolia.

Excluded Record Colura herzogii Jovet-Ast – Colombia: Cho (Vasco et al. 2002). Malesian species; the Colombian specimen is presumably misidentified. Cyclolejeunea A.Evans Plants small, 0.8–2 mm wide, pale green to brownish, sometimes blackish in herbarium, creeping and with upright gemmiparous shoots. Stems with hyalodermis, ventral merophyte 2 cells wide. Leaf lobes wide-spreading, apex rounded or apiculate, margins finely crenulate to denticulate, dorsal lobe surface smooth or elevated by large papillae (one per cell). Cells convex, with trigones and intermediate thickenings, cuticle smooth or with a large papilla; oil bodies small, finely granular, usually 2–4 per cell; ocelli (0–)1–3 above the lobe base, side by side. Lobules ca. 1/3 leaf length, ovate-inflated, free margin incurved, with a short, 1-celled tooth, hyaline papilla proximal. Underleaves bifid or undivided, margins entire or toothed, insertion line curved. Androecial bracteoles usually restricted to the base of the spike, sometimes present throughout (C. convexistipa). Gynoecia on short or long branches, with 1 pycnolejeuneoid innovation. Perianths flattened-pyriform, somewhat inflated ventrally and with 2 broad lateral keels expanded above into short auricles. Sporophytes lejeuneoid. Vegetative reproduction by means of multicellular, disciform or elongate gemmae produced on upright, gemmiparous shoots. A neotropical genus of seven species, all of them in Colombia and Ecuador. The species have been placed in four subgenera, subg. Chocolejeunea M.E.Reiner & Gradst. (C. glimeana), subg. Cyclolejeunea (C. chitonia, C. convexistipa, C. foliorum, C. peruviana), Hyalolejeunea Grolle (C. accedens), and subg. Nephrolejeunea Grolle (C. luteola). The subgeneric classification has been supported by DNA (Heinrichs et  al. 2014a). Cyclolejeunea is one of the most common neotropical bryophyte genera inhabiting living leaves. None of the species of Cyclolejeunea is an obligate epiphyll, however; all of them occur also on bark. Characteristic of

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Cyclolejeunea is the presence of a few ocelli above the leaf base (absent in C. accedens); the flattened, 2-keeled perianth with keels expanded above into low auricles; pycnolejeuneoid innovations; and vegetative reproduction by means of large, disciform or elongate, leaf-borne gemmae with rhizoids. The leaf margins in Cyclolejeunea are usually toothed (rarely entire). Cyclolejeunea closely resembles Prionolejeunea in the toothed leaf margins and the 2-keeled perianth with keels expanded above, but Prionolejeunea lacks ocelli, innovations are lejeunoid or absent, and gemmae in Prionolejeunea lack rhizoids. The two genera are classified in a separate subtribe, Lejeuneaceae subtribe Cyclolejeuneinae (Gradstein 2013a). Literature: Grolle (1984b), Bernecker-Lücking (1998), Gradstein and IlkiuBorges (2009), Heinrichs et al. (2014a), Gradstein and Reiner-Drehwald (2017a) Note: Characters of leaves and underleaves are to be studied on creeping stems, not on upright gemmiparous stems. 1. Underleaves undivided. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. convexistipa 1. Underleaves bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Underleaves of creeping shoots very shallowly bifid (1/6–1/5), lobes rounded or apiculate, margins entire. Gemmae wider than long (reniform) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. chitonia 2. Underleaves of creeping shoots more deeply divided, lobes acute, margins entire or toothed. Gemmae rounded or longer than wide . . . . . . . . . . . . . . . . 3 3. Plants 1.5–3 mm wide. Margins of leaves and underleaves of creeping shoots sharply toothed, the teeth 1–4 cells long. Lobule tooth ± reduced, obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. peruviana 3. Plants smaller, 0.5–1.5(−1.7) mm wide. Margins of leaves and underleaves of creeping shoots entire or crenulate-denticulate (like in Prionolejeunea), the teeth less than 1 cell long. Lobule tooth reduced or well developed. . . . . . . . 4 4. Cells of leaf margin and dorsal leaf surface mammillose and with a broad papilla. Leaves without ocelli. Gemmae orbicular . . . . . . . . . . . . C. accedens 4. Cells of leaf margin and dorsal leaf surface not mammillose, without papilla. Leaves with (1–)2–6 ocelli near leaf base. Gemmae orbicular or elongate. . . 5 5. Lobule tooth well developed, acute. Gemmae oblong to ligulate, with rhizoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. luteola 5. Lobule tooth small and obtuse or absent. Gemmae (sub)orbicular, with rhizoids, or ligulate without rhizoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Leaf margins ± toothed. Lobule with a small, obtuse tooth. Innovations present. Gemmae orbicular and with marginal rhizoids, produced on sharply toothed branch leaves. Androecia produced on short-specialized branches without vegetative leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. foliorum 6. Leaf margins entire or crenulate. Lobule without tooth. Innovations lacking. Gemmae ligulate, without rhizoids, produced on old, decaying shoots. Androecia produced on elongate branches with vegetative leaves, underleaves mostly restricted to the base of the male shoot . . . . . . . . . . . . . . . C. glimeana

464

1. Cyclolejeunea accedens (Gottsche) A.Evans Cyclolejeunea affixa (Taylor) Steph. Cyclolejeunea grandistipula Steph. Cyclolejeunea papillata Steph. Prionolejeunea leptocardia (Spruce) Steph. Prionolejeunea luxurians Steph.

 

Keys and Descriptions

Fig. 46K, L

On living leaves and shrubs in lowland and submontane rainforests, sometimes growing periodically submerged, from sea level to 1500 m. Colombia: Cho, Hui, Ris, Val. Ecuador: Ch, LR, Pa, ZC. General distribution: tropical America. Cyclolejeunea accedens (subg. Hyalolejeunea) is readily recognized by the mammillose leaf cells with a broad dorsal papilla. Moreover, it is the only species in the genus without ocelli. 2. Cyclolejeunea chitonia (Taylor) A.Evans Cyclolejeunea ecuadorensis Steph. (?)

Fig. 46M, N

On living leaves and twigs in lowland rainforest, from sea level to 300  m. Colombia: Cho, Vau. Ecuador: EO (Reserva Buenaventura, ca. 300 m, on canopy branch, Benitez et al. 1313, HUTPL), Ma (Stephani 1898–1924 as C. ecuadorensis), Pa (Arnell 1962 as C. ecuadorensis). General distribution: scattered in tropical America. Cyclolejeunea chitonia (subg. Cyclolejeunea) is an uncommon neotropical species. The plants are rather robust plant (1.5–3 mm wide) with broad and very shallowly bifid underleaves (bifid to 1/6–1/4 of underleaf length) and reniform gemmae. Cyclolejeunea ecuadorensis, described from Manabí and characterized by shallowly bifid underleaves and  ±  denticulate leaf margins, may be a synonym of C. chitonia. 3. Cyclolejeunea convexistipa (Lehm. & Lindenb.) A.Evans  Fig. 46O Cyclolejeunea subintegerrima Herzog Cyclolejeunea triocellata Herzog Cyclolejeunea truncatulata Spruce Lejeunea stachyclada Spruce Lejeunea surinamensis (Mont.) Gottsche, Lindenb. & Nees, nom. illeg. On living leaves, twigs, bark, and rock in lowland and lower montane rainforest, 70–1800 m. Colombia: Cau, Cho, Qui, Ris, San, Val. Ecuador: EO, Es, LR, MS, Na, Pa. General distribution: widespread in tropical America. Cyclolejeunea convexistipa (subg. Cyclolejeunea) is a common neotropical species that is readily recognized by the undivided underleaves with a truncate apex. The plants are relatively small, 0.5–1.5 mm, pale green to brownish, the leaves are conspicuously asymmetrical (dorsal margin arched, ventral margin straight), and the leaf margins vary from subentire on creeping shoots to strongly toothed on ascending shoots. The underleaves are appressed to the stem on creeping shoots but are standing upward, perpendicular to the stem, on upright shoots.

Lejeuneaceae

4. Cyclolejeunea foliorum (Nees) Grolle Cyclolejeunea angulata Herzog

465

Fig. 46P, Q

On living leaves and bark in lowland and submontane rainforest, from sea level to 1000 m. Colombia: Cho, Hui. Ecuador: Na, Or, ZC (Schäfer-Verwimp et al. 2013a). General distribution: Amazonia, Guianas, Chocó biogeographic region (Dauphin et al. 2015). Cyclolejeunea foliorum (subg. Cyclolejeunea) is an uncommon neotropical species that is morphologically similar to C. peruviana but differs in the smaller plant size (0.5–1.5 mm wide) and stem leaves and underleaves with finely denticulate leaf margins, like in Prionolejeunea (margins not sharply toothed). In addition, the underleaves in C. foliorum are very small, hardly wider than the stem. 5. Cyclolejeunea glimeana M.E.Reiner & Gradst.

Fig. 46R, S

On palm trunks in everwet lowland rainforest along the Pacific coast, 30  m. Colombia: Cho (Gradstein and Reiner-Drehwald 2017a). General distribution: only known from the type locality fom the Chocó. Cyclolejeunea glimeana (subg. Chocolejeunea) is a striking Colombian endemic that is distinguished by (1) plants very transparent and pale-colored, growing in dense ± colorless mats on smooth palm trunks; (2) margins of leaves and underleaves entire or sligthly crenulate; (3) leaf lobes with 2–3 large, suprabasal ocelli; (4) lobules bluntly angled at the apex, without tooth; and (5) gynoecia on short branches without innovation. The species lacks disciform gemmae but occasionally produces elongate, ribbon-like gemmae without rhizoids on old, decaying leaves (Gradstein and Reiner-Drehwald 2017a). Such elongate gemmae are also seen in C. luteola, Lejeunea, and Prionolejeunea, but the gemmae of C. luteola produce rhizoids, whereas Lejeunea and Prionolejeunea differ in ocelli absent and innovations lejeuneoid (or absent). 6. Cyclolejeunea luteola (Spruce) Grolle Cyclolejeunea angulistipa (Steph.) A.Evans

Fig. 46U

On tree trunks, rock, and rotten wood in lowland and montane rainforests, occasionally on living leaves, from sea level to 2000 m. Colombia: Ama, Caq, Mag, Put, Vau (Campos et al. 2015). Ecuador: ZC (Parolly et al. 2004). General distribution: throughout tropical America. Cyclolejeunea luteola (subg. Nephrolejeunea) is a common neotropical species recognized by elongate (oblong to narrowly ligulate) gemmae (with rhizoids) produced on the margins of toothed branch leaves. The plants are relatively small (0.5–1.5 mm wide), the leaf margins are strongly crenulate, the lobule has a sharp tooth, and the underleaves are small, bifid, with subentire margins. Winkler (1976) misidentified a specimen of Cyclolejeunea luteola from Magdalena as Trachylejeunea didrichsenii Steph. The latter species, described from Brazil, is a synonym of Lejeunea acanthogona s.l. (Bastos and Gradstein 2020). The listing of T. didrichsenii as a synonym of C. luteola in the Catalogue of the Plants and Lichens of Colombia (Bernal et al. 2016) is an error.

466

7. Cyclolejeunea peruviana (Lehm. & Lindenb.) A.Evans Odontolejeunea peruviana (Lindenb.) Steph.

 

Keys and Descriptions

Fig. 46V

On living leaves, occasionally on bark, in lowland and montane rainforests, 30–2700 m. Colombia: Boy, Cau, Cho, Hui, Vau. Ecuador: EO, LR, MS, Or, Na, Pa, Pi, Tu, ZC. General distribution: widespread in tropical America. Cyclolejeunea peruviana (subg. Cyclolejeunea) is one of the most common species in the genus. It resembles C. chitonia in the robust habit (1.5–3 mm) and rather strongly toothed leaves, but the underleaves are more deeply bifid (1/4–1/2 of underleaf length), with sharply toothed margins, and the gemmae are orbicular (wider than long in C. chitonia).

Further Record Cyclolejeunea ocellata S.Winkl., nom. inval. – Colombia: Cho (Winkler 1970). The material should be checked. Dibrachiella (Spruce) X.Q.Shi, R.L.Zhu & Gradst. Archilejeunea subg. Dibrachiella (Spruce) Schiffn. Autoicous. Plants small to large, 1–3.5 mm wide, pale green to greenish-brown to black, creeping or ascending. Branches Lejeunea-type, occasionally Frullaniatype; flagelliform branches absent. Stems without enlarged epidermis, all cell walls colorless, ± thickened, subepidermis lacking; ventral merophyte 4–6(−8) cells wide. Leaf lobes wide-spreading, when dry not appressed or convolute, apex rounded, margins entire. Cells (sub)isodiametric, trigones simple-triangular to radiate; oil bodies coarsely granular; ocelli lacking. Lobules 1/3–1/2 leaf length, often reduced, with 1–2 teeth, free margin usually plane. Underleaves undivided, distant to subimbricate, with entire margins and straight or weakly curved insertion line. Gynoecia with 1–2 lejeuneoid innovations. Perianths with 4–5 keels, the keels smooth or slightly toothed above. Vegetative reproduction by regeneration from leaf cells. An Afro-American genus of nine species, three in Colombia and Ecuador. Dibrachiella was long considered a subgenus of Archilejeunea but was raised to generic rank on molecular grounds (Shi et al. 2015). The genus differs morphologically from Archilejeunea in lejeuneoid innovations (pycnolejeuneoid in Archilejeunea) and frequently reduced lobules (not reduced in Archilejeunea). The neotropical species of Dibrachiella differ furthermore from Archilejeunea by distant underleaves (imbricate in Archilejeunea) and autoicy (Archilejeunea is dioicous or paroicous). Dibrachiella is closely related to Spruceanthus, a mainly Asiatic genus with one species in Ecuador (Shi et  al. 2015), but the latter genus has more robust stems (ventral merophyte up to 16 cells wide) with a distinct subepidermis, flagelliform branches at stem bases, and homogeneous oil bodies.

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Literature: Gradstein and Buskes (1985 as as Archilejeunea subg. Dibrachiella), Gradstein et al. (1994 as A. subg. Dibrachiella), Shi et al. (2015) 1. Lobule with 2 teeth. Plants growing in exposed, rather dry habitats, glossy light green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. auberiana 1. Lobule with 0–1 tooth. Plants growing in sheltered, wet habitats in the rainforest understory, green to brown to blackish. . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Lobule short-rectangular with truncate apex, tooth sharp. Ventral margin of the leaf lobe forming a rather sharp angle (ca. 120°) with the keel. Underleaves reniform. Plants of the Chocó . . . . . . . . . . . . . . . . . . . . . . . . . . D. bischleriana 2. Lobule ovate with oblique apex, tooth blunt or lacking. Ventral margin forming an almost straight line with the keel. Underleaves (sub)orbicular. Plants widespread. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. parviflora 1. Dibrachiella auberiana (Mont.) X.Q.Shi, R.L.Zhu & Gradst. Archilejeunea auberiana (Mont.) A.Evans Archilejeunea florentissima (Spruce) Herzog

Fig. 47A

On bark of trees and shrubs in rather open environments, in evergreen and semideciduous rainforests, 0–300 m. Colombia: Cho (Gradstein et al. 2018c). Ecuador: Es, Gu, LR. General distribution: scattered in tropical America; in Colombia and Ecuador restricted to the Pacific coastal region. Dibrachiella auberiana is distinguished by the small, pale green plants with ovate leaves, undivided underleaves, longly rectangular, 2-toothed lobules that are sometimes reduced, isodiametric leaf cells with simple-triangular trigones, stems without enlarged epidermis, and a 5-keeled perianth subtended by 1–2 lejeuneoid innovations. Dibrachiella auberiana is close to D. bischleriana, a species endemic to the Chocó of Colombia, but the latter species has shorter, rectangular lobules with only 1 tooth and reniform underleaves. 2. Dibrachiella bischleriana (Gradst.) X.Q.Shi, R.L.Zhu & Gradst. Archilejeunea bischleriana Gradst. On twigs and cyperaceous leaves in the understory of wet lowland rainforest, 50–80 m. Colombia: Cho (Gradstein 1994). General distribution: only known from the Chocó. Dibrachiella bischleriana is a rare Chocó endemic that is recognized by the shortly rectangular lobules with a truncate apex and a sharp tooth and distant reniform underleaves. 3. Dibrachiella parviflora (Nees) X.Q.Shi, R.L.Zhu & Gradst. Archilejeunea parviflora (Nees) Steph. Archilejeunea cruegeri (Lindenb.) Schiffn. Archilejeunea leprieurii (Mont.) Schiffn. Archilejeunea viridissima (Lindenb.) A.Evans

Fig. 47B

On tree bases and roots in the understory of lowland and submontane rainforests, rarely on living leaves, from sea level to 1500 m. Colombia: Ama, Caq, Cau, Ces,

Fig. 47 (A) Dibrachiella auberiana, leaf. (B) Dibrachiella parviflora, habit with gynoecia and sporophyte. (C and D) Dicranolejeunea axillaris. (C) Median cells. (D) Habit with gynoecium. (E) Diplasiolejeunea alata, habit. (F) Diplasiolejeunea brunnea, leaf. (G) Diplasiolejeunea cavifolia, leaf. (H) Diplasiolejeunea involuta subsp. andicola, habit. (I) Diplasiolejeunea johnsonii, leaf and underleaf. (J) Diplasiolejeunea lanceolata, habit. (K) Diplasiolejeunea paukertii, leaf and underleaf. (L and M) Diplasiolejeunea pellucida. (L) Leaf. (M) Habit with gynoecium. (E redrawn from Schuster 1970; H redrawn from Winkler 1976; I redrawn from Reyes 1982; J redrawn from Grolle 1992; rest from Gradstein et  al. 2001, Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009). Scale bars: A, F, G, L = 250 μm; B, H, K, M = 500 μm; C = 25 μm; D = 1000 μm; E = 400 μm; I = 200 μm; J = 350 μm

Lejeuneaceae

469

Cho, Met, NSa, Put, Qui, Ris, Vau. Ecuador: LR, Or. General distribution: throughout tropical America. Dibrachiella parviflora is a common, dark green to brown or blackish species of tree bases in neotropical rainforests. Characteristics are the wide-spreading leaves (both wet and dry) with rounded apices; isodiametric leaf cells with small, simpletriangular trigones; small ovate lobules that are frequently reduced; and distant, suborbicular (to broadly ovate) underleaves with an almost straight insertion line. A molecular analysis of D. parviflora has shown that the species is polyphyletic and consists of more than one, genetically distant race (Shi et al. 2015). The correlation of the genetic variation with morphology has not yet been studied. Dicranolejeunea (Spruce) Schiffn. Paroicous. Plants rather long and narrow, to 7 cm long, 1–2 mm wide, glossy light green to yellow-brown, creeping, ascending or pendent, often forked. Branches Frullania-type, rarely Lejeunea-type. Stems with a hyalodermis, epidermis cells mammillose, strongly bulging outward; ventral merophyte only 2 cells wide. Leaf lobes obliquely spreading and convex, when dry convolute, apex acute-acuminate, margins entire or weakly toothed at apex by 1–3(−5) teeth. Cells slightly longer than wide, trigones cordate or rounded; oil bodies finely granular; ocelli lacking. Lobules ca. 1/4–1/3 leaf length, occasionally reduced, with 2 teeth, the second tooth smaller than the first one. Underleaves ovate-orbicular, undivided, margins entire, bases decurrent, insertion line arched. Male bracts just below the gynoecium, with epistatic lobules. Gynoecia with (1–)2 pycnolejeuneoid innovations, male bracteoles toothed. Perianths 3-keeled with a low, broad ventral keel, ciliate-laciniate near apex. Sporophytes lejeuneoid. Vegetative reproduction not observed. A neotropical genus with a single species. Dicranolejeunea is readily recognized by the brownish-green, laxly forked plants with obliquely spreading leaves with a ± toothed, acute-acuminate apex, small, 2-toothed lobules, widely spaced orbicular underleaves, 2-celled ventral merophyte, and flattened ciliate-laciniate perianths with 2 long, repeatedly fertile innovations. The genus is closely related to Acanthocoleus; the two groups are separated by growth habitat, stem anatomy, and characters of the gynoecia. Dicranolejeunea is usually more robust than Acanthocoleus, with stems ascending from the substrate and often freely pendent and branches always of the Frullania-type. In contrast, species of Acanthocoleus are typically creeping plants with branches mostly of the Lejeunea-type. Furthermore, in Dicranolejeunea the stem epidermis cells are very large and strongly bulging outward, and the walls of the outer medullary cells are conspicuously thickened, forming a subepidermis and contrasting with the thin walls of the rest of the medullary cells. In Acanthocoleus, the epidermis cells are smaller and never bulging outward, and the outer medullary cells are scarcely thicker-walled than the inner medullary cells. Finally, female bracts and bracteoles are toothed in Dicranolejeunea while being mostly entire in Acanthocoleus.

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Keys and Descriptions

Literature: Kruijt (1987), Gradstein et al. (1994) 1. Dicranolejeunea axillaris (Nees & Mont.) Schiffn.  Fig. 47C, D Dicranolejeunea ciliatiflora (Spruce) Steph. Dicranolejeunea hypoacantha (Spruce) Steph. Dicranolejeunea pulcherrima Steph. Dicranolejeunea tridentata S.Winkl. Peltolejeunea acuminata Herzog (?) Type: Ecuador, Los Rios, Samama, Hacienda La Clementina, 600 m, Harling 532 p.p., 481 p.p. (JE; not found) Peltolejeunea galapagona Steph. On bark and rock in open, montane environments (rarely in lowlands), in forest canopies and at forest margins, on roadside trees and shrubs, in pastures, etc., a common invader of disturbed habitats, occasionally on living leaves, (50–) 500–3200 m. Colombia: Ant, Boy, Cal, Cho, Cun, Hui, Mag, Qui, Ris, San Tol. Ecuador: EO, Es, Ca, Ch, Ga, MS, Na, Pa, Tu, ZC. General distribution: widespread in tropical America but records from Brazil doubtful (to be verified). For characters of D. axillaris, see under the genus. Based on the original description and illustration, Peltolejeunea acuminata – described from Ecuador based on sterile material – is probably a synonym of D. axillaris; the type could not be found in JE. León-Yánez et al. (2006) treated P. acuminata as a synonym of Omphalanthus jackii, but the latter species differs by strongly elongate underleaves. Diplasiolejeunea (Spruce) Schiffn. Plants small- to medium-sized, 1–3 cm long, 0.7–3(−3.5) mm wide, pale green to pale brown, creeping, often closely appressed to the substrate. Stems of thickwalled, pale cells (7 epidermis cells, 3 medullary cells), epidermis not or slightly enlarged, hyalodermis lacking; ventral merophyte 2 cells wide. Leaf attached to stem by only few cells, insertion very short. Leaf lobes wide-spreading, convex, apex rounded, margins entire, sometimes hyaline-bordered. Cells isodiametric-hexagonal, smooth, walls thin or uniformly thickened, occasionally with trigones; oil bodies small, usually coarsely granular-botryoidal; ocelli frequently present, scattered in leaf lobes, sometimes lacking. Lobules usually large, 1/3–3/4 leaf length, inflated, apex with 2 short or long teeth, the second tooth usually smaller than the first one, hyaline papilla at the inner base of the first tooth. Underleaves one to each lateral leaf, often overlapping, deeply bifid with diverging lobes and a broad, V-shaped sinus, insertion line curved. Androecia on short-speciliazed branches, bracts with hypostatic lobules, bracteoles limited to the base of the male spike or present throughout. Gynoecia on long or short shoots, with 1 pycnolejeuneoid innovation. Perianths with 4–5 sharp, smooth keels, rarely without keels. Sporophytes lejeuneoid. Vegetative reproduction frequently by multicellular, disciform gemmae from leaf surfaces. A pantropical genus of about 50 species, 18  in Colombia and Ecuador. Diplasiolejeunea is readily recognized by the very short leaf insertion and the deeply bifid underleaves, which are present one to each leaf (instead of one per leaf

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pair) and with widely diverging lobes. Confusion is only possible with species of Colura with a reduced sac. The species typically grow tightly appressed on living leaves (epiphyllous) or smooth bark of twigs (ramicolous) in the understory of the forest or in rather open environments. Ocelli may be present or absent in leaf lobes; their presence/absence does not serve to subdivide the genus into sections as proposed by Schuster (1970) (see Dong et al. 2012). However, an interesting correlation with habitat is seen in the species of Colombia and Ecuador: all shade epiphytes (D. brunnea, D. cavifolia, D. johnsonii, D. pellucida, D. pocsii) have ocelli, while most sun epiphytes (D. alata, D. buckii, D. columbica, D. involuta, D. lanceolata, D. papillionacea, D. pauckertii, D. pluridentata, D. replicata) lack ocelli. Literature: Reyes (1982), Schäfer-Verwimp (2004), Dong et al. (2012) 1. Leaves ovate-lanceolate, ca. 3× longer than wide, apex acute-acuminate. Underleaves very small, not wider than the stem . . . . . . . . . . . . D. lanceolata 1. Leaves orbicular to ovate-oblong, 1–2× longer than wide, apex rounded. Underleaves wider than the stem. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leaves with a border of hyaline cells. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Leaves without border . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Second lobule tooth 3–4 cells long; first tooth T-shaped. Underleaf lobes terminating in a row of 2–3 cells. Plants rare. . . . . . . . . . . . . . . . . . . . . . . . D. pocsii 3. Second lobule tooth reduced or 1–2 cells long; first tooth finger-like or T-shaped. Underleaf lobes terminating in a row of 1–2 cells. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. pellucida 4. Underleaves on main stems small, 1.5–2× as wide as stem, lobes 2–4 cells wide  at the base. Perianth with a long stalk at the base, sharply and deeply 5-keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. alata 4. Underleaves on main stems larger, 2.5–5× as wide as stem, lobes usually more than 4 cells wide at the base. Perianth without long stalk. . . . . . . . . . . . . . . . 5 5. Lobules with more than 2 teeth (study carefully, teeth often inflexed) . . . . . . 6 5. Lobules with (1–)2 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 6. Lobule teeth about equal in length, first tooth incurved, 3–4 cells long and 1 cell wide (except at the base). Underleaf bases rounded, apices acute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. pluridentata 6. Lobule teeth very unequal in length, first tooth straight, 4–7 cells long and 2 cells wide, much longer than the other teeth. Underleaf bases auriculate, apices rounded or acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Underleaves imbricate, lobes broadly triangular, 16–28 cells wide at the base, apices broadly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. papillionacea 7. Underleaves distant, lobes narrower, oblong, 18–15 cells wide at the base, apices acute. Plants from Costa Rica and Venezuela; not yet known from Colombia and Ecuador. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. eggersii Pócs 8. Ventral margin of the leaf lobe revolute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Ventral margin of the leaf lobe plane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 9. Lobules huge, almost as long as the lobe, more than 1/2× lobe width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. involuta subsp. andicola

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Keys and Descriptions

9. Lobules less than 1/2× lobe width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Lobules 3–4× as long as wide. Perianth without beak, terete. Plants rare ­(subpáramo of southern Ecuador) . . . . . . . . . . . . . . . . . . . . . . . . . . D. erostrata 10. Lobules 1.5–2.5× as long as wide. Perianth with a beak, 5-keeled or terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Free margin of lobule plane. Underleaf lobes rounded . . . . . . . . . D. johnsonii 11. Free margin of lobule incurved. Underleaf lobes subacute to shortly acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Ocelli present. Underleaf lobes terminating in a row of 2–4 cells. Perianth terete, with a huge beak (ca. 1/4 of perianth width). Plants very rare (páramo of southern Ecuador) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. grandirostrata 12. Ocelli absent. Underleaf lobes terminating in a row of 1–2 cells. Perianth 5-keeled, beak smaller. Plants common in the northern Andes. . D. pauckertii 13. First tooth of lobule 2 cells wide from the base to near apex, finger-like. Leaves without ocelli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13. First tooth of lobule not 2 cells wide from base to near apex, finger-like or T-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Underleaf lobes ca. 15–22 cells wide at base . . . . . . . . . . . . . . . D. columbica 14. Underleaf lobes 4–8 cells wide at base . . . . . . . . . . . . . . . . . . . . . D. replicata 15. Leaf lobes without ocelli. Lobules ca. 3× as long as wide, its first tooth T-shaped. On twigs in the outer canopy of Amazonian rainforest. . . . . . . . . . . . D. buckii 15. Leaf lobes with ocelli. Lobules 1–2× as long as wide, its first tooth finger-like or T-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Apex of underleaf lobes rounded or obtuse. First lobule tooth finger-like, 3–15 cells long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. unidentata 16. Apex of underleaf lobes acute (acute to obtuse in the rare D. mayaykuensis). First lobule tooth T-shaped or finger-like, 1–10 cells long . . . . . . . . . . . . . . 17 17. Free margin of lobule flat. At least some lobules with a T-shaped first tooth, second tooth straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cavifolia 17. Free margin of lobule inflexed. Lobules never with a T-shaped first tooth, second tooth inflexed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18. Old stem leaves squarrose. Underleaf lobes acute to obtuse. Plants rare (southern Ecuador). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. mayaykuensis 18. All stem leaves appressed. Underleaf lobes acute. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. brunnea 1. Diplasiolejeunea alata Jovet-Ast

Fig. 47E

Sun epiphyte on branches and twigs of shrubs and isolated trees in open, montane environments, occasionally on living leaves, 1700–3050  m. Colombia: Ant (Benavides and Gutiérrez 2011 “cf.”), San (El Hato, Cuta 1006, UPTC; Virolin, Schuster 88-1069a, F, fide C.J. Bastos). Ecuador: Lo, Na, Pi, ZC. General distribution: scattered in tropical America. Diplasiolejeunea alata is a delicate, uncommon neotropical species with plants measuring barely more than 1 mm wide. The leaves lack ocelli, the lobules are large and swollen, and the underleaves are small, 1.5–2× stem width, with narrowly lanceolate lobes (2–4 cells wide at base). The perianth of D. alata is longly exerted

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beyond the bracts due to the presence of a long basal stalk, and the perianth is sharply and deeply 5-keeled. 2. Diplasiolejeunea brunnea Steph. Diplasiolejeunea bidentiloba Steph. (fide R. Grolle, in sched.) Diplasiolejeunea galloana Jovet-Ast Diplasiolejeunea lanciloba Tixier

Fig. 47F

On living leaves and bark in lowland and lower montane rainforests, plantations, and gardens, from sea level to 2500  m. Colombia: Caq, Cho, Hui, Put, San. Ecuador: Az, Es, EO, LR, MS, Na, Or, Pa, Pi, Su, ZC. General distribution: widespread in tropical America. Diplasiolejeunea brunnea is a common neotropical species that is distinguished by ocellate leaves without hyaline margin; a large lobule with a conspicuous, erect, finger-like first tooth and the second tooth well developed but often incurved and inconspicuous; and deeply bifid underleaves with widely diverging, lanceolate lobes with acute tips. 3. Diplasiolejeunea buckii Grolle Sun epiphyte on twigs in the outer canopy of Amazonian rainforest, 120–220 m. Colombia: Caq, Vau (Campos et  al. 2014). General distribution: northern Amazonia. Diplasiolejeunea buckii is an Amazonian canopy species that has been very little collected and has probably been overlooked due to neglect of the forest canopy. The species is readily recognized by the leaves without ocelli and with strongly swollen, narrowly elongate lobules with a T-shaped first tooth and a long second tooth. The underleaves are small, 2–3× stem width, deeply bifid with narrowly lanceolate lobes. 4. Diplasiolejeunea cavifolia Steph. Diplasiolejeunea brachyclada A.Evans

Fig. 47G

On living leaves and bark in lowland and montane rainforests, from sea level to 2800 m. Colombia: Ant (cf.), Boy, Cho, Hui, Put, Vau. Ecuador: Es, MS, Na, Pi, Tu, ZC. General distribution: pantropical. Diplasiolejeunea cavifolia is a common, pantropical species that is characterized by ocellate leaves with a 1–2-toothed lobule and deeply bifid underleaves with widely diverging, narrowly lanceolate lobes with acute tips. The species is rather similar to D. brunnea; for differences see the key. 5. Diplasiolejeunea columbica Tixier Sun epiphyte on twigs in montane scrub and open, boggy vegetation, 2300–2500 m. Colombia: Cun, Hui (Schäfer-Verwimp 2004). General distribution: only known from Colombia. Diplasiolejeunea columbica is a rare Colombian species known from only two localities. The species is distinguished by leaves without ocelli, large lobules with 1–2 teeth, a 2-cell-wide first tooth and a plane free margin, and very broad underleaf lobes, 15–22 cells wide at base. By the broad underleaf lobes, D. columbica

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Keys and Descriptions

resembles D. papillionacea, but the lobules in the latter species have 3 teeth and an incurved free margin. 6. Diplasiolejeunea erostrata Schäf.-Verw. On shrubs and small trees in subpáramo, in rather open environments, 2600–3050  m. Ecuador: Lo, ZC (Schäfer-Verwimp 2004). General distribution: only known from southern Ecuador. Diplasiolejeunea erostrata is a rare Ecuadorian endemic that is recognized by ocellate leaves, narrowly elongate lobules with an incurved free margin and 2 small teeth, acute underleaf lobes, and a terete, beakless perianth. 7. Diplasiolejeunea grandirostrata Schäf.-Verw. On twigs of shrubs in páramo, 3000 m. Ecuador: ZC (Schäfer-Verwimp 2004). General distribution: only known from the type from southern Ecuador. Diplasiolejeunea grandirostrata is only known from the páramo of San Francisco Biological Reserve (Ecuador) and somewhat resembles D. erostrata but differs in the less elongate lobules, acuminate underleaf lobes terminating in a row of 2–4 cells, and perianths with a huge beak (ca. 1/4 of perianth width). 8. Diplasiolejeunea involuta S.Winkl. subsp. andicola Pócs

Fig. 47H

Sun epiphyte on shrubs in open, upper montane rainforest areas, scrub, and páramo, 2150–3200 m. Ecuador: Lo, Pi, ZC; to be expected in Colombia. General distribution: northern Andes (Venezuela, Ecuador) Diplasiolejeunea involuta is a very distinct neotropical species that is easily recognized by the huge lobules which are more than 1/2× the width of the leaf lobes, greatly swollen and with strongly involute free margin. The plants from the northern Andes belong to the subspecies andicola, characterized by underleaf lobes 12–18 cells wide at the base (22–30 cells wide in subsp. involuta) (León et al. 1998). 9. Diplasiolejeunea johnsonii A.Evans

Fig. 47I

On bark and living leaves in the understory of montane rainforest, 1900  m. Ecuador: ZC (Nöske et  al. 2003). General distribution: Greater Antilles, Central America, Ecuador. Diplasiolejeunea johnsonii is an uncommon neotropical species with ocellate leaves with a conspicuously recurved ventral margin; large lobules with a plane free margin and a short, finger-like first tooth; and plump underleaves with rounded lobes. 10. Diplasiolejeunea lanceolata Grolle

Fig. 47J

Sun epiphyte on bark and living leaves at margins of submontane rainforest, 500–960 m. Ecuador: Es (type), EO (Grolle 1992; Schäfer-Verwimp et al. 2013a). General distribution: only known from the foothills of the Andes in western Ecuador. Diplasiolejeunea lanceolata is a rare Ecuadorian endemic that is immediately recognized by the lanceolate leaves, 3–4× longer than wide, with acute to shortacuminate tips and no ocelli, and tiny, bifid underleaves, which are not wider than

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the stem and with upright, narrowly lanceolate lobes. In other neotropical Diplasiolejeunea species, the leaves are 1–2× longer than wide, with broadly rounded tips, and the underleaves are wider than the stem (2–8×) and with widely diverging lobes. 11. Diplasiolejeunea mayaykuensis Schäf.-Verw. & Heinrichs Sun epiphyte on isolated shrub in cultivated land, 850 m. Ecuador: ZC (type; Schäfer-Verwimp et al. 2013b). General distribution: only known from the foothills of the Cordillera del Condor, southern Ecuador. Diplasiolejeunea mayaykuensis is very close to D. brunnea but differs in the squarrose older stem leaves, the lobules with a strongly involute free margin and 2 well-developed teeth, and the underleaf lobes with acute to obtuse apices. In D. brunnea the leaves are not squarrose and the underleaf lobes are acute (not obtuse). 12. Diplasiolejeunea papillionacea R.M.Schust. Sun epiphyte on branches and twigs of shrubs and bamboo in open situations, 1850–3500 m. Colombia: Ris. Ecuador: ZC (Schäfer-Verwimp 2004). General distribution: northern Andes. Diplasiolejeunea papillionacea is a characteristic northern Andean species that is recognized by the large underleaves (6–8× stem width) with broadly rounded apices and the lobules with 3–4 teeth and a long and straight first tooth (4–7 cells long). In its lobule with several teeth, the species approaches D. pluridentata and D. eggersii (Pócs 2009b); for differences see the key. 13. Diplasiolejeunea pauckertii (Nees) Steph. Diplasiolejeunea reflexiloba (Gottsche) Steph.

Fig. 47K

On branches and twigs of trees and shrubs and on living leaves in upper montane rainforest and páramo, (1500–)2000–4100 m. Colombia: Ant, Boy, Cau, Cun, Hui, Ris. Ecuador: Az, Ca, Lo, Na, Tu, ZC. General distribution: scattered in the mountains of tropical America. Diplasiolejeunea pauckertii is a characteristic species of neotropical upper montane forests and páramo, recognized by (1) ventral margin of the leaf lobe revolute; (2) lobule less than 1/2× leaf width, ovate, free margin incurved and with 1–2 teeth; and (3) underleaf lobes subacute, terminating in a single cell. The species may be confused with D. involuta and D. replicata which occur in the same habitat, but D. involuta has a much larger lobule, almost as broad as the leaf lobe, and D. replicata has a plane ventral leaf margin. 14. Diplasiolejeunea pellucida (Spreng.) Schiffn. Fig. 47L, M Diplasiolejeunea caribea Tixier, syn. nov. Type: Guadeloupe, Palmiste, bois Joseph, 620 m, 20 Dec 1960, Le Gallo s.n. (PC-0015330!) On living leaves in lowland and lower montane rainforests, plantations and gardens, 70–2500 m. Colombia: Ama, Boy, Cau, Cho, San. Ecuador: Az, Es, MS, Na, Pa, Pi, ZC. General distribution: widespread in tropical America.

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Keys and Descriptions

Diplasiolejeunea pellucida is a very common and variable neotropical species that is recognized by leaves with a hyaline border and many ocelli, lobules 1–2-toothed, and underleaves deeply bifid, with widely diverging lobes which are 4–8 cells wide at base and terminate in a row of 1–2 cells. The first lobule tooth is straight or T-shaped, and the second tooth is straight or incurved, or absent. Plants with a T-shaped lobule tooth were described as D. caribea and have been recorded from southern Ecuador (Schäfer-Verwimp 2004). The latter species is included here in D. pellucida based on the study of Dong et al. (2012) who found that D. caribea is nested in D. pellucida. Diplasiolejeunea evansii Tixier and D. intermis Tixier may be further synonyms of D. pellucida (T. Pócs, per. comm.). 15. Diplasiolejeunea pluridentata Schäf.-Verw.

Fig. 48A

On branches and twigs of shrubs in open locations in primary and secondary montane rainforests and subpáramo, 2150–3150 m. Colombia: Boy (Gradstein and Uribe 2016a). Ecuador: ZC (Nöske et al. 2003). General distribution: Costa Rica, northern Andes (Ecuador, Colombia). Diplasiolejeunea pluridentata is an uncommon northern Andean-Costa Rican species recognized by lobules with 4–6 teeth and underleaves with rounded bases and acute lobes. Pluridentate lobules (with more than 2 teeth) are also seen in D. papillionacea and D. eggersii, but in the latter two species, the first tooth of the lobule is much larger than the other teeth and the underleaf bases are auriculate. Moreover, the tips of the underleaf lobes in D. papillionacea are rounded, not acute (Pócs 2009b). 16. Diplasiolejeunea pocsii Reyes Montoya On living leaves in shrubby vegetation, 2000–2800 m. Ecuador: ZC (SchäferVerwimp 2004). General distribution: West Indies (Cuba, Dominica), southern Ecuador. A rare Caribbean species recognized by ocellate leaves with a hyaline border. The species is very close to D. pellucida; for differences see the key. 17. Diplasiolejeunea replicata (Spruce) Steph. On branches and twigs of trees and shrubs and on living leaves in rather open montane forest areas and páramo, 1200–4150  m. Colombia: Cas (Gradstein 12707a, PC), Tol. Ecuador: Az, Ca, Cañ, Lo, Na, Pa, Pi, Tu, ZC. General distribution: scattered in the mountains of tropical America. Diplasiolejeunea replicata is a rather common neotropical-montane species characterized by plane ventral margin of the leaf lobe, lobule with incurved free margin and 1–2 teeth, and subacute underleaf lobes terminating in 1–2 cells in a row. The species may be confused with D. pauckertii which occurs in the same habitat, but in the latter species, the ventral margin of the leaf lobe is revolute. 18. Diplasiolejeunea unidentata (Lehm. & Lindenb.) Schiffn. Fig. 48B Diplasiolejeunea rudolphiana Steph., syn. nov. Type: Brazil, Rio de Janeiro, Petropolis, 1890, Rudolph s.n. (G-00060872!, c. gyn.)

Fig. 48 (A) Diplasiolejeunea pluridentata, leaf. (B) Diplasiolejeunea unidentata, habit. (C and D) Drepanolejeunea andina. (C) Leaf. (D) Underleaf. (E and F) Drepanolejeunea anoplantha. (E) Underleaf. (F) Leaf. (G) Drepanolejeunea bidens, habit. (H) Drepanolejeunea crassiretis, leaf. (I and J) Drepanolejeunea cutervoensis. (I) Habit. (J) Underleaf. (K) Drepanolejeunea inchoata, habit. (L) Drepanolejeunea crucianella, habit. (M) Drepanolejeunea lichenicola, habit. (N) Drepanolejeunea fragilis, leaf. (O) Drepanolejeunea palmifolia, leaf. (P) Drepanolejeunea orthophylla, habit. (Q) Drepanolejeunea polyrhiza, habit with gynoecium. (R) Drepanolejeunea spinosa, leaf. (A redrawn from Schäfer-Verwimp 2001; C–F, H–J, O redrawn from Bischler 1964; rest from Gradstein and Ilkiu-Borges 2009). Scale bars: A  =  1000  μm; B  =  500  μm; C–F, H, J, N, O = 50 μm; G, K–M, P, Q = 250 μm; I = 200 μm; R = 100 μm

478

 

Keys and Descriptions

Sun epiphyte on twigs in the outer canopy of lowland and montane rainforest and on bark in open, disturbed forest, plantations, gardens, pastures, etc., occasionally on living leaves, from sea level to 2800 m. Colombia: Boy, Cau, Cho, Mag, Put. Ecuador: Az, EO, Ga, Or, ZC. General distribution: pantropical. Diplasiolejeunea unidentata is a very widespread and variable species characterized by ocellate leaves, underleaves with rounded to obtuse lobes, and lobules with a long, finger-like first tooth (3–15 cells long) and sometimes a small, inflexed second tooth. Diplasiolejeunea rudolphiana, which has been reported from low elevations in Colombia and Ecuador, is a synonym of D. unidentata based on the molecular study of Dong et al. (2012) who found a nested relationship of the two species. The two species have traditionally been separated by a single character (length of the lobule tooth), but the difference is gradual.

Excluded Record Diplasiolejeunea armatiloba Steph. – Colombia: Cho (Vasco et al. 2002); Ecuador: Es (Arnell 1962). Caribbean species; the records from Colombia and Ecuador are probably erroneous (e.g., Schäfer-Verwimp 2004). Drepanolejeunea (Spruce) Steph. Plants very small, to 1.5 cm long, 0.3–1(−1.3) mm wide, light green to pale brown, creeping. Stems of 7 outer cells and 3 inner cells, epidermis cells not or slightly enlarged, no hyalodermis, all cells ± thick-walled; ventral merophyte 2 cells wide. Leaf lobes suberect to rather wide-spreading, asymmetrically ovate-triangular to lanceolate, rarely elliptical, often falcate, usually apex acute-acuminate, rarely rounded, plane or recurved, margins crenulate to dentate or entire. Cells usually small, midleaf cells 10–25 μm, with trigones, cuticle smooth or coarsely papillose; oil bodies finely granular; ocelli colorless, (1–)2–4(−10) per leaf, often in a broken row (occasionally unbroken), sometimes absent, the basal ocellus not conspicuously enlarged. Lobules 1/3–1/2(−3/4) leaf length, occasionally reduced, inflated, tooth 1(−2)-celled, usually long and falcate, hyaline papilla proximal; a “preapical” tooth sometimes present at the distal end of the free margin, at the junction with the lobe. Underleaves small, remote, bifid with very slender, upright to diverging lobes, the lobes 2–4(−6) cells long and 1–2(−4) cells wide, lamina without large-celled border, insertion line straight; adhesive rhizoid discs well developed in epiphyllous plants. Gynoecia on short branches, usually with 1 pycnolejeuneoid innovation. Perianths inflated, 5-keeled above, the keels extending into toothed, laminate or horn-like projections or auricles. Sporophytes lejeuneoid. Vegetative reproduction by caducous branchlets (= cladia). A pantropical genus of about 60 species, 25 in Colombia and Ecuador. The main characters of Drepanolejeunea are (1) tiny plants, usually less than 1 mm wide, with elongate, asymmetrically ovate-lanceolate, spreading or suberect leaves with acute

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or acuminate apex (rounded in D. polyrhiza), entire or toothed margins, and often a narrow base; (2) ocelli usually present in leaf lobes, scattered or in a broken row, occasionally 2(−3) in an unbroken row at leaf base; (3) lobules with a longly falcate tooth and often with a short or long “pre-apical” tooth arising from the distal end of the free lobule margin, at the junction with the lobe; (4) underleaves distant and deeply bifid, with obliquely to widely diverging lobes; (5) gynoecia with 1 pycnolejeuneoid innovation and 5-keeled perianths with variously toothed or laciniate keels; and (6) vegetative reproduction by cladia. The underleaf lobes are usually linear, of only 1 row of cells, occasionally broader in some high-Andean taxa. Species with broad underleaf lobes may be confused with Harpalejeunea but are separated from the latter by gynoecia with pycnolejeuneoid innovation (lejeuneoid in Harpalejeunea) and ocelli in a broken row. The neotropical species of Drepanolejeunea belong in subg. Drepanolejeunea with the exception of D. polyrhiza which is a member of subg. Rhaphidolejeunea (Herzog) Grolle & R.L.Zhu. Schuster (1996b) classified the neotropical Drepanolejeunea species in eight sections based, in part, on the informal species groups recognized by Bischler (1964, 1968) in her monograph of neotropical Drepanolejeunea. Schuster’s sectional classification was preliminary and needs further work. Literature: Bischler (1964, 1968), Schuster (1996b), Gradstein and IlkiuBorges (2009) 1. Underleaf lobes 1(−2) cells wide (sometimes 2 cells wide in the lower half) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaf lobes 2–4 cells wide. Plants in páramo. . . . . . . . . . . . . . . . . . . . . 26 2. Leaf apex rounded or truncate. Underleaf lobes almost horizontally diverging. Epiphyllous plants in lowland rainforest. . . . . . . . . . . . . . . . . . . . D. polyrhiza 2. Leaf apex acute-acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Margin of all leaf lobes entire, crenulate or, occasionally, with a tooth in the lower half of the ventral margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Margin of some or all leaf lobes with more than 1 tooth . . . . . . . . . . . . . . . 13 4. Leaves of the main stem obliquely to widely spreading, at an angle of 40–90° with the stem. Ocelli (1–)3–10 per leaf. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Leaves of the main stem suberect or obliquely spreading at an of 15–45° with the stem. Ocelli 0–2 per leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5. Ocelli 1 or 2 in a broken row at the leaf base. Plants of the Chocó (epiphyllous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. evansii 5. Ocelli more than 2 per leaf, in a broken or unbroken row. . . . . . . . . . . . . . . . 6 6. Mature stem leaves 1–4–1.8× longer than wide. Plants on rock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. orthophylla subsp. saxicola 6. Mature stem leaves more elongate, ca. 2–4× longer than wide. Plants on living leaves or bark. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Leaf lobes usually with a tooth at the base of the ventral margin, near the junction with the keel . . . . . . . . . . . . . . . . . . . . . D. orthophylla subsp. orthophylla 7. Leaf lobes fully entire, without tooth . . . . . . . . . . . . . . . . . . . . . . . D. mosenii

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Keys and Descriptions

8. Stem leaves spreading at an angle of ca. 45°. Leaf cells not mammillose . . . . 9 8. Stem leaves spreading less widely (to 30°), often almost parallel to stem. Leaf cells often mammillose and with thin or thickened outer wall . . . . . . . . . . . 10 9. Leaves convex, apex often recurved. Perianth keels with a narrow wing, the wing more or less toothed. Plants common. . . . . . . . . . . . . . . . . D. biocellata 9. Leaves flat, apex plane, never recurved. Perianth keels with a very broad, hornlike wing, the wing strongly toothed. Plants very rare. . . . . . . D. ramentiflora 10. Dorsal margin of leaf lobe strongly dilated beyond the base. Leaves only 1.5–2(−2.5)× longer than wide, strongly convex . . . . . . . . . . . . D. crassiretis 10. Dorsal margin of leaf lobe not or only slightly dilated beyond the base. Leaves more elongate, 2.5–3.5× longer than wide, plane or convex . . . . . . . . . . . . . 11 11. Well-developed lobules inflated below, flattened above. Ocelli lacking or 1–2 in a broken row near leaf base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. bidens 11. Well-developed lobules fully inflated. Ocelli 2  in an unbroken row at leaf base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Underleaf lobes 4–6 cells long. Perianth keels with broad inflated wings in the upper 1/4 of the perianth. Plants of páramo of southern Colombia, above 3000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. aurita 12. Underleaf lobes 2–4 cells long. Perianth keels without broad, inflated wings. Plants of montane rainforests, below 3000 m . . . . . . . . . . . . . . D. anoplantha 13. Stem leaves obliquely to widely spreading, at an angle of 40–90° with the stem, not falcate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13. Stem leaves less spreading (to maximally 45°), often falcate . . . . . . . . . . . . 21 14. Ocelli 3–6 per leaf, occurring from leaf base to apex . . . . . . . . . . . . . . . . . . 15 14. Ocelli lacking or 1–2 in the lower half of the leaf. . . . . . . . . . . . . . . . . . . . . 17 15. Lobule tooth short and blunt, not falcate. Plants very rare (Colombia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. pterocalyx 15. Lobule tooth long, falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Mature stem leaves 1.5–2× as long as wide, widely spreading (60–90°), strongly narrowed to the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. inchoata 16. Mature stem leaves 2–3× as long as wide, obliquely spreading (45–60°), not strongly narrowed to the base . . . . . . . . . . . . . . . . . . . . . . . . . . D. orthophylla 17. Ventral margin of the leaf lobe usually with a larger tooth (1–3 cells long) near the junction with the keel. Leaf apex terminating in a row of (1–)2–3 cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Ventral margin without larger tooth near the junction with the keel. Leaf apex terminating in a single cell. Plants common. . . . . . . . . . . . . . . . . . D. inchoata 18. Plants very small, 0.3–0.45 mm wide. Basal leaf cells very small, subquadrate, ca. 20–25 μm. Plants rare (Colombia) . . . . . . . . . . . . . . . . . . . . . . . D. spinosa 18. Plants larger, 0.45–1 mm wide. Basal leaf cells larger, rectangular, ca. 30–45 μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19. Mature stem leaves lanceolate, more than 3× longer than wide, obliquely spreading, falcate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. campanulata 19. Mature stem leaves broader, rhombic, less than 3× longer than wide, widely spreading, not falcate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

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481

20. Perianth apex not contracted to a beak, perianth mouth wide open, lobed. Plants very rare (southern Ecuador, Venezuela). . . . . . . . . . . . . . . . . . . . D. urceolata 20. Perianth apex contracted to a beak, perianth mouth closed, not lobed. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. infundibulata 21. Teeth on leaf margins very large, 2–6 cells long, 1–3 cells wide at base . . . 22 21. Teeth smaller, 1–2 cells long, 1 cell wide at base . . . . . . . . . . . . . . . . . . . . . 23 22. Dorsal and ventral leaf margins toothed. Leaves plane, not falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. crucianella 22. Dorsal leaf margin toothed, ventral margin entire. Leaves convex, falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. palmifolia 23. Ventral margin of the leaf lobe often with a conspicuous tooth, (1–)2–3 cells long, near the junction with the keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. Ventral margin without conspicuous tooth near the junction with the keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24. Pre-apical tooth of lobule long and sharp, curved, often reaching the apical tooth. Stem leaves suberect, little spreading (to 30°) . . . . . . . . . . D. lichenicola 24. Pre-apical tooth short and blunt, straight, not reaching the apical tooth. Stem leaves more widely spreading . . . . . . . . . . . . . . . . . . . . . . . . . D. campanulata 25. Lobules fully inflated. Free margin of lobule not prolonged along the ventral lobe margin. Pre-apical tooth of lobule well developed, sometimes reaching the apical tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. fragilis 25. Lobules inflated below, flattened toward apex. Free margin of lobule prolonged along the ventral lobe margin. Pre-apical tooth of lobule short or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. araucariae 26. Dorsal leaf margin toothed. Leaf cells smooth or with a dorsal papilla . . . . 27 26. Dorsal leaf margin entire. Leaf cells with a dorsal papilla . . . . . . . . . . . . . . 28 27. Leaf cells with a dorsal papilla. Underleaf lobes almost upright. Ocelli lacking (?) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. subvittata 27. Leaf cells without papilla, smooth. Underleaf lobes wide-spreading. Ocelli (1–)2 in a broken row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. andina 28. Free margin of the lobule continuing along the ventral margin of the leaf lobe. Lobule without pre-apical tooth.. . . . . . . . . . . . . . . . . . . . . . . . . D. cutervoensis 28. Free margin of the lobule terminating at the junction of keel and the ventral margin of the leaf lobe, not continuing upward along the ventral margin. Lobule with a conspicuous pre-apical tooth . . . . . . . . . . . . . . . . . . . . . . D. granatensis 1. Drepanolejeunea andina Herzog Drepanolejeunea grandistipula R.M.Schust.

Fig. 48C, D

On twigs of shrubs in upper montane elfin forest and páramo, occasionally epiphyllous, 2950–4400 m. Colombia: Ris. Ecuador: Az, Ca, Na, Pi (type). General distribution: Costa Rica and tropical Andes (Venezuela, Colombia, Ecuador; new to Peru: Urubamba, Ollantaytambo, near Thastayoc, on Polylepis pepei, 4414 m, Sylvester s.n., PC). Drepanolejeunea andina (subg. Drepanolejeunea) is a characteristic northern Andean páramo species that is recognized by (1) underleaves with large,

482

 

Keys and Descriptions

wide-spreading lobes that are up to 10 or more cells long and 2–4 cells wide; (2) leaves suberect and somewhat falcate, apex acute-acuminate, margins with several 1–2 cell long teeth; (3) leaf cells smooth, ocelli 1 or 2 in a broken row; and (4) lobules large, conspicuously inflated and flattened toward apex, without pre-apical tooth. Drepanolejeunea andina approaches D. subvittata by the toothed dorsal leaf margin, the broad underleaf lobes, and the lobules without pre-apical tooth but clearly differs from the latter species in the wide-spreading underleaf lobes (suberect in D. subvittata), smooth leaf cells (papillose in D. subvittata), and presence of ocelli. Moreover, the teeth on the dorsal leaf margin are longer in D. subvittata (3–4 cells long). 2. Drepanolejeunea anoplantha (Spruce) Steph.

Fig. 48E, F

On bark and creeping over other bryophytes in montane rainforest and shrubby páramo, (200–)700–3100 m. Colombia: Ama, Caq, Cho, Put, Vau (Campos et al. 2015). Ecuador: Lo, ZC.  General distribution: West Indies, tropical South America. Drepanolejeunea anoplantha (subg. Drepanolejeunea) is a rather common neotropical species that is distinguished by (1) leaves suberect, convex, not or slightly falcate, narrowly elongate, 2.5–3× as long as wide, dorsal margin not strongly expanded above the base; (2) leaf margins and keel crenulate, leaf apex subacute to short-acuminate, terminating in a row of 1–3(−5) cells; (3) leaf base with 2 ocelli in an unbroken row; (4) leaf cells ± mammillose, the outer wall thin or thickened; (5) lobules fully inflated, apex with a 1-celled, straight or curved tooth; pre-apical tooth 1–2 cells long; (6) underleaf lobes 2–4 cells long, 1–2 cells wide at the base; (7) plants dioicous; and (8) perianth with 5 slightly winged keels, auricles lacking. Drepanolejeunea anoplantha is close to D. crassiretis and D. aurita; all three species have 2 ocelli in an unbroken row at the leaf base and are dioicous. Drepanolejeunea crassiretis is separated from D. anoplantha by the less elongate, broader leaves (2–2.5× as long as wide) with the dorsal leaf margin strongly dilated above the base and arched; D. aurita differs in longer underleaf lobes (4–6 cells long) and perianth keels with broadly inflated wings in the upper 1/4. 3. Drepanolejeunea araucariae Steph. On bark and living leaves in lowland and montane rainforests, from sea level to 3000 m. Colombia: Ama, Cas (Gradstein 12704, PC), Cho, Cun, NSa, Put, Ris, Vau. Ecuador: Na, Pi, ZC. General distribution: tropical and subtropical South America. Drepanolejeunea araucariae (subg. Drepanolejeunea) is a widespread neotropical species that is recognized by (1) leaves suberect to obliquely spreading, falcate, dorsal margin with a few short, 1(−2)-cell-long teeth, ventral margin entire or crenulate, apex terminating in a row of 2 cells, and (2) lobules ± flattened toward apex, without pre-apical tooth, apex oblique, free margin shortly continuing along the ventral margin of the leaf lobe.

Lejeuneaceae

483

Drepanolejeunea araucariae approaches D. fragilis, but in the latter species, the lobule is fully inflated, the pre-apical tooth is long and curved, and the free margin of the lobule does not continue along the ventral margin of the leaf lobe. 4. Drepanolejeunea aurita Bischl. On twigs of shrubs in páramo, 3350–3450 m. Colombia: Cau/Hui (páramo de Las Papas; type). General distribution: only known from the type locality in Colombia. Drepanolejeunea aurita (subg. Drepanolejeunea) is a rare Colombian endemic resembling D. anoplantha in leaf shape and in the presence of 2 ocelli in an unbroken row at leaf base, but differing in 4–6-cell-long underleaf lobes (2–4 cells long in D. anoplantha) and perianth keels expanded above into broad, inflated, lateral wings, forming small auricles (auricles lacking in D. anoplantha). The auricles are usually not extended beyond the perianth apex. 5. Drepanolejeunea bidens (Prantl) A.Evans

Fig. 48G

On bark, rotten wood, and living leaves in open lowland and montane rainforests, plantations, and gardens from sea level to 2650 m. Colombia: Ama, Caq, Cho, Cun, Hui, Qui, Vau. Ecuador: Az, Ga, ZC. General distribution: tropical America. Drepanolejeunea bidens (subg. Drepanolejeunea) is a common neotropical species that is distinguished by (1) leaves suberect, falcate, margins entire, apex terminating in a row of 2 cells; (2) leaf cells with small or large trigones; (3) lobules inflated below, flattened toward apex, flattened and inflated parts separated by a curved line; lobule apex truncate, not continuing along the lower half of the ventral lobe margin; and (4) pre-apical tooth long and curved, often reaching the apical tooth. Drepanolejeunea bidens is morphologically close to D. araucariae and D. fragilis, but in the latter two species, the dorsal leaf margin is toothed; moreover in D. fragilis the lobule is fully inflated, and in D. araucariae the pre-apical tooth is lacking or short and blunt. 6. Drepanolejeunea biocellata A.Evans Drepanolejeunea squarrosula Herzog On living leaves and bark in lowland and montane rainforests, often creeping between other bryophytes, from sea level to 2100 m. Colombia: Ant, Boy, Cho, Cun, Hui, NSa, Ris, San. Ecuador: MS, Pa, Tu, ZC.  General distribution: ­tropical America. Drepanolejeunea biocellata (subg. Drepanolejeunea) is a common neotropical species that is recognized by (1) leaves spreading 45–60°, convex, ovate-oblong to ovate-lanceolate and asymmetrical (dorsal margin arched, ventral margin straight to curved); (2) leaf apex subacute, recurved or plane; (3) leaf margins entire or crenulate, often with a short tooth on the ventral margin, near the junction with the keel; (4) lobules almost fully inflated (flattened only near apex), with a long falcate tooth and a short, blunt pre-apical tooth; (5) leaf cells with small or large trigones; (6) ocelli usually 2 in a broken row in the lower half of the leaf lobe, sometimes leaves

484

 

Keys and Descriptions

without ocelli or with only 1 ocellus; and (7) perianth keels with a narrow, toothed wing. 7. Drepanolejeunea campanulata (Spruce) Steph. On living leaves, bark, and rock in lowland and montane rainforests, from sea level to 3550  m. Colombia: Boy, Cas (Gradstein 12713, PC), Mag, Ris, San. Ecuador: Ch, Lo, Na, Pa, Pi, Su, Tu, ZC. General distribution: northern Andes, Central America, southeastern Brazil. Drepanolejeunea campanulata (subg. Drepanolejeunea) is a distinct neotropical species characterized by obliquely spreading, lanceolate-falcate stem leaves (more than 3× longer than wide) with shortly toothed margins and a larger tooth on the ventral margin near the junction with the keel. The species may be confused with D. lichenicola and D. infundibulata, but the leaves in D. lichenicola are less spreading (to 30°) and possess a long, curved pre-apical tooth, whereas the stem leaves in D. infundibulata are more widely spreading (to max. 90°) and broader, rhombic (less than 3× wider than long), and not falcate. 8. Drepanolejeunea crassiretis A.Evans Fig. 48H Drepanolejeunea integribracteata Bischl., syn. nov. Holotype: Brazil, Amazonas, upper Rio Negro, São Gabriel, Spruce 358 (MANCH!). Drepanolejeunea submuricata R.M.Schust., nom. inval. (herbarium not cited), syn. nov. Material: Dominica, Morne Diablotins, on bark in upper montane rainforest, Schuster 67-777a (n.v.). On bark and rotten wood in montane rainforests and mossy elfin forests, also on moist soil along river, 1500–3500 m; found at sea level in the Brazilian Amazon (Bischler 1964, type of D. integribracteata). Ecuador: Na, ZC. General distribution: West Indies, Brazil, Ecuador. Drepanolejeunea crassiretis (subg. Drepanolejeunea) is rather widespread neotropical species that is characterized by the strongly convex, suberect, asymmetrically ovate-triangular leaves (1.5–2× longer than wide) with a strongly arched dorsal leaf margin and with 2 ocelli in an unbroken row at the leaf base. The leaf cells are mammillose, and the outer cell wall is thin or thickened, with or without a low, rounded papilla; trigones and intermediate thickenings are present or absent. The leaf apex is obtuse to subacute, rarely shortly apiculate (type of D. integribracteata), the lobules are strongly inflated, and the underleaves are small, with shortlinear lobes made up of a row of 2–3 cells. The keel and ventral lobe margin usually form a straight line, and the lobule is often somewhat prolonged along the ventral margin. Occasionally, the transition between the keel and the ventral lobe margin is marked by a notch, and the lobule is not prolonged. Plants with the latter characteristics were described as D. integribractata. Because much overlap is seen in these characters, D. integribractata is treated here as a synonym of D. crassiretis. Based on the original description and illustration, D. submuricata (Schuster 1996b) from Dominica, characterized by mammillose cells with a rounded papilla, is a further synonym of D. crassiretis. The oil bodies of D. crassiretis, seen in a

Lejeuneaceae

485

recent collection from elfin forest in the páramo of Yacuambi (Benitez et al. 1308, HUTPL), are ellipsoid, finely granular, and one per cell, as described by Schuster (1996a) for D. submuricata. The material from Yacuambi had mature perianths and sporophytes, which were hitherto unknown in D. crassiretis. The perianth is oblong, ca. 0.8 × 0.4 mm, with 5 sharp and slightly winged, crenulate keels in the upper 1/4 and with a short beak, and the cells of the perianth are thin-walled or slightly evenly thick-walled, without or with few trigones and intermediate thickenings. Interestingly, numerous ocelli are present in the upper part of the perianth, between the keels. In contrast, ocelli were not seen in the female bracts. An unusual feature of old perianths with dehisced capsules was their long-stalked base, with the stalks being as long as or slightly longer than the entire perianth, up to ca. 1 cm long. The stalked perianth of D. crassiretis resembles that of Myriocoleopsis tixieri, but the perianth in the latter species lacks ocelli. 9. Drepanolejeunea crucianella (Taylor) A.Evans Drepanolejeunea capitulata sensu Spruce (non typus)

Fig. 48L

On living leaves in lowland and lower montane rainforests, occasionally on bark, 80–1600(−3000) m. Colombia: Cau, Cho, NSa, Ris, Vau. Ecuador: Lo, LR, MS, Or, Pa, Su, ZC. General distribution: widespread in tropical America. Drepanolejeunea crucianella (subg. Drepanolejeunea) is a common neotropical species that is readily recognized by the suberect, ovate-elongate leaves (1.5–2× longer than wide) with several long teeth (2–6 cells long, 1–3 cells wide at base) along the dorsal and ventral leaf margins. The teeth are usually larger toward the leaf apex. Drepanolejeunea crucianella shares long teeth with D. palmifolia, but in the latter species, the teeth are restricted to the dorsal leaf margin. Moreover, the leaves in D. palmifolia are conspicuously falcate (not falcate in D. crucianella). 10. Drepanolejeunea cutervoensis (Loitl.) Grolle  Drepanolejeunea navicularis Steph.

Fig. 48I, J

On branches and twigs of trees and shrubs in upper montane forest, scrub, and páramo, (1600–)2500–4560 m. Colombia: Ant, Boy, Cau, Hui, Qui. Ecuador: Az, Co (Cotopaxi National Park, Gradstein & Frahm 6681, GOET, U), Lo, ZC (SchäferVerwimp et al. 2013a). General distribution: tropical Andes (Venezuela to Peru). Drepanolejeunea cutervoensis (subg. Drepanolejeunea) is a characteristic northern Andean species that is distinguished by (1) leaves narrowly elongate, suberect and somewhat falcate, margins entire, apex acute-apiculate, terminating in a row of 1–2(−3) cells, ventral margin of leaf lobe plane or recurved; (2) leaf cells dorsally mamillose and with a large rounded papilla; (3) ocelli 2 in an unbroken row at leaf base or lacking; (4) lobules inflated below, flattened toward apex, without pre-apical tooth, free margin shortly or longly continuing along the ventral margin of the leaf lobe (sometimes to near the leaf apex); and (5) underleaves bifid to 1/2, with 2–4 cell broad, rounded, harpalejeuneoid lobes. The underleaves of D. cutervoensis resemble those of Harpalejeunea, but the latter genus differs from Drepanolejeunea in having lejeuneoid innovations (Grolle 1991). Drepanolejeunea cutervoensis is closely similar to D. granatensis; for

486

 

Keys and Descriptions

differences see the key. Drepanolejeunea cutervoensis also approaches D. subvittata, but in the latter species, the dorsal leaf margin is toothed (in the lower half). 11. Drepanolejeunea evansii L.Söderstr., A.Hagborg & von Konrat Drepanolejeunea infundibulata sensu A.Evans (non typus) On living leaves in wet lowland rainforest, 100 m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). General distribution: West Indies, Chocó. Drepanolejeunea evansii (subg. Drepanolejeunea) is a Caribbean species that has been recorded once from the Chocó. Its characters are (1) leaves narrowly elongate, 2.8–4.5× longer than wide, rather widely spreading (45–70°), with a short tooth in the lower half of the ventral margin of the leaf lobe, otherwise entire; (2) leaf acute or short-apiculate, terminating in 1–2 cells in a row; (3) ocelli 1 or 2 in a broken row near leaf base; (4) lobules inflated below, flattened above, inflated and flattened parts separated by a curved line; and (5) lobule tooth long, falcate. Drepanolejeunea evansii has been confused with D. infundibulata, but in the latter species, the leaf margins are toothed, by several teeth, and the leaf apex terminates in a row of 2–3 cells. 12. Drepanolejeunea fragilis L.Söderstr., A.Hagborg & von Konrat

Fig. 48N

On living leaves in Amazonian forest, 120–210 m. Colombia: Ama, Caq, Put (Campos et  al. 2014). Ecuador: Or (Mota de Oliveira and ter Steege 2013). General distribution: scattered in tropical America. Drepanolejeunea fragilis (subg. Drepanolejeunea) grows as an epiphyll in lowland rainforest and resembles D. araucariae; for differences see under the latter species. 13. Drepanolejeunea granatensis (Prantl) Bischl. Harpalejeunea granatensis (Prantl) Steph. On treelets and shrubs in upper montane rainforests, shrubby vegetation, and páramo, 1900–4000  m. Colombia: Ant (type; Benavides and Gutiérrez 2011). Ecuador: Az, Ca, Co, Im, Lo, Pi, ZC. General distribution: northern Andes, southeastern Brazil. Drepanolejeunea granatensis (subg. Drepanolejeunea) is a characteristic northern Andean-SE Brazilian disjunct that resembles D. cutervoensis in the broad, rounded, harpalejeuneoid underleaf lobes and the leaf cells with a large rounded papilla, but the free lobule margin in D. granatensis does not continue along the ventral margin of the leaf lobe and the pre-apical tooth is well developed (absent in D. cutervoensis). Drepanolejeunea granatensis is also closely related to the Caribbean D. valiae Jovet-Ast (Jamaica, Guadeloupe, Dominica). For a discussion of the differences between these two species, see Schuster (1996b). 14. Drepanolejeunea inchoata (C.F.W.Meissn.) Steph. Drepanolejeunea inchoata var. biocellata Herzog Drepanolejeunea inchoata var. roraimae (Steph.) Bischl. Lejeunea planiuscula Spruce, nom. illeg.

Fig. 48K

Lejeuneaceae

487

Common on living leaves and bark in lowland and montane rainforests, 80–3400 m. Colombia: Ant, Cau, Boy, Cas, Ces, Cho, Mag, Ris, Tol, Val. Ecuador: EO, Es, Ga, Lo, LR, MS, Na, Pa, Pi, Su, Tu, ZC. General distribution: throughout tropical America. Drepanolejeunea inchoata (subg. Drepanolejeunea) is a very common neotropical species that is recognized by (1) leaves widely spreading, margins ± toothed, apex acute, consisting of a single cell (not of a row of 2 or more cells), leaf base very narrow; (2) ocelli 0–10 per leaf, scattered through the leaf lobe or restricted to leaf base, or absent; and (3) lobules inflated below, flattened toward apex. 15. Drepanolejeunea infundibulata (Spruce) A.Evans On bark and living leaves in montane rainforests, (100–)750–2400 m. Colombia: Ant (Benavides and Gutiérrez 2011), Cho (Benavides and Sastre-de Jesús 2011). Ecuador: Ch, LR, MS, Pa, Su. General distribution: tropical America. Drepanolejeunea infundibulata (subg. Drepanolejeunea) resembles D. inchoata but differs in the leaf apex frequently terminating in a row of 2–3 cells and lobules usually fully inflated, lacking a flattened portion. Moreover, in D. infundibulata a larger tooth (1–3 cells long) is usually present at the base of the ventral margin of the leaf lobe, near the junction with the keel. In D. inchoata such a larger tooth is absent. Ocelli in D. infundibulata are 1–2 at leaf base, never scattered in the leaf lobe. 16. Drepanolejeunea lichenicola (Spruce) Steph.

Fig. 48M

On bark and rotten wood in montane rainforests, occasionally on living leaves, 450–3000 m. Colombia: Ant, Boy, Caq, Cau, Cho, Hui, Mag, Put, Ris, Val, Vau. Ecuador: Lo, LR, Na, MS, Or, Tu, ZC.  General distribution: widespread in tropical America. Drepanolejeunea lichenicola (subg. Drepanolejeunea) is a common and variable neotropical species that is distinguished by (1) leaves narrowly elongate, 2.5–4× longer than wide, suberect, ± falcate, margins with several short teeth (1–2 cells long), apex acuminate, terminating in a row of 2–4 cells; (2) ventral margin of leaf lobe usually with a longer tooth (2–3 cells long) at the base, just above the junction with the keel and opposite to the pre-apical tooth; (3) ocelli 1–2 in a broken row near the leaf base; (4) lobules fully inflated and with a long, curved pre-apical tooth, which usually reaches the apical tooth; and (5) underleaf lobes 1 cell wide. 17. Drepanolejeunea mosenii (Steph.) Bischl. Drepanolejeunea sabaliana R.M.Schust. On living leaves and bark in lowland and montane rainforests, from sea level to 2000  m. Colombia: Cun, NSA.  Ecuador: ZC.  General distribution: widespread in tropical and subtropical America. Drepanolejeunea mosenii (subg. Drepanolejeunea) is a common neotropical species close to D. orthophylla but differs in having fully entire leaf margins. In D. orthophylla the leaf margins are entire to crenulate to toothed, and a tooth (1–2 cells long) is usually present at the base of the ventral margin of the leaf lobe, near the junction with the keel. For detailed descriptions of D. mosenii, see Bischler (1968), Schuster (1980a as D. sabaliana), and Reiner-Drehwald (1995). The ocelli

488

 

Keys and Descriptions

in the leaf lobes are usually 3–6 in a broken row; occasionally, the 2 basal ocelli are in an unbroken row (Schuster 1980a; Reiner-Drehwald 1995), like in D. orthophylla subsp. saxicola. 18a. Drepanolejeunea orthophylla (Nees & Mont.) Bischl. subsp. orthophylla Fig. 48P Drepanolejeunea hamulata Steph. Drepanolejeunea hamulata var. espinulosa Herzog Leptolejeunea stenophylla (Lindenb. & Gottsche) Steph. Leptolejeunea unguiculata Steph. On living leaves in montane forest, (450–)1200–2700 m. Colombia: Boy, Cau, Qui. Ecuador: Az, Es (Burghardt 6856, 6857, 6859, QCA), Ga, Or, Pa, Pi, ZC. General distribution: tropical America. Drepanolejeunea orthophylla subsp. orthophylla (subg. Drepanolejeunea) is a widespread epiphyllous taxon in the neotropics; for detailed description and illustrations, see Bischler (1968). Characteristics are the obliquely to widely spreading, elongate leaves, (1.8–)2–3.5× longer than wide, with a crenulate to toothed margin and with 3–6 ocelli in a broken row in the leaf lobe. A tooth, 1–2 cells long, is usually seen at the base of the ventral margin of the leaf lobe, near the junction with the keel; in some populations, however, the tooth is present on only part of the leaves. Drepanolejeunea orthophylla subsp. orthophylla is close to D. mosenii, but the latter species differs by fully entire leaf margins. Moreover, D. mosenii sometimes has 2 basal ocelli in an unbroken row, like in D. orthophylla subsp. saxicola (see below). 18b. Drepanolejeunea orthophylla subsp. saxicola Gradst. & Pócs, subsp. nov. Fig. 49A–D Holotype: Colombia, Quindio, Salento, Cocora, reserva Palma de Cera, ca. 2400 m, on rock near bridge across the river, at the edge of remnant montane rainforest, S.  R. Gradstein & A.  L. López 12729 (HUQ; isotypes EGR, PC-0773220) Plants sterile, green, very small, forming small green mats on the exposed upper surface of a large rock near river in open environment, leafy stems 0.6–0.7  mm wide, densely branched, branches frequently turning into cladia with leaves becoming progressively smaller toward apex and the branch readily breaking away from the stem. Leaves obliquely to widely spreading, at an angle of ca. 45–75° with the stem, convex, lobe ovate-oblong and somewhat falcate, 1.4–1.8× longer than wide, asymmetrical with the dorsal margin arched and the ventral margin almost straight, leaf apex acute, terminating in a row of 1–2 cells, plane or recurved, margins irregularly crenulate-crenate, cells of the margin often projecting and terminating in a small papilla, ventral margin with a sharp, 1-celled tooth at the base, adjacent to the the junction with the keel. Leaf cells thin-walled to slightly thickened, trigones absent or small, intermediate thickenings scarce, ocelli 3(−6) in a broken row in the leaf lobe (ocelli vanishing in herbarium material), the 2 basalmost ocelli in an unbroken row at the base of the leaf lobe; ocelli more abundant in small leaves of cladia. Lobules large, 2/5–1/2× lobe length, strongly inflated but becoming flattened

Lejeuneaceae

489

Fig. 49  Drepanolejeunea orthophylla subsp. saxicola. (A) Leaf. (B) Cells in the lower half of the leaf lobe, showing ocelli. (C) Lobule. (D) Underleaf. (All from the type). Scale bars: A = 100 μm; B–D = 50 μm

toward the apex, made up of thin-walled cells, free margin incurved, apex with a large, 1-celled, falcate tooth, pre-apical tooth short and blunt, 1-celled; keel curved, smooth below, crenate by mammillose cells above, sinus at the junction with the ventral lobe margin notched. Underleaves small, highly distant, 2–3× stem width, deeply bifid, lobes widely spreading outward, 3–4 cells long, 2 cells wide at the base, uniseriate above. Vegetative reproduction by cladia. The new subspecies differs from Drepanolejeunea orthophylla subsp. orthophylla in having less elongate leaves, being only 1–4–1.8× longer than wide, with 3–6 ocelli in leaf lobes including 2 in an unbroken row at the base, and in its occurrence on rock. Subspecies orthophylla has leaves (1.8–)2–3.5× longer than wide and ocelli in a broken row and typically grows on living leaves. As D. orthophylla is a polymorphic species (see Bischler 1964) and because the saxicolous specimen is sterile, we prefer to describe it as a subspecies of D. orthophylla rather than as a new species. Drepanolejeunea orthophylla subsp. saxicola approaches D. mosenii in the arrangement of the 2 basalmost ocelli in an unbroken row at the leaf base but, besides its less elongate leaves, clearly differs from D. mosenii in the presence of a tooth at the base of the ventral margin of the leaf lobe, the character typical of D. orthophylla.

490

 

Keys and Descriptions

19. Drepanolejeunea palmifolia (Nees) Schiffn. Fig. 48O On bark in Amazonian rainforests, 100–350  m. Colombia: Ama, Caq, Vau (Pinzón et al. 2003; Campos et al. 2015). Ecuador: ? Pa (Bischler, 1964). General distribution: Amazonia. Drepanolejeunea palmifolia (subg. Drepanolejeunea) is a characteristic Amazonian species that is readily recognized by the strongly falcate leaves with long teeth on the dorsal margin (teeth 2–7 cells long, 1–2 cells wide at the base). The ventral leaf margin is entire or crenulate, but without teeth, and the leaf apex is apiculate to longly acuminate and strongly recurved. A further characteristic of the species is the upright lobes of the underleaves (not or little diverging). According to Bischler (1964), the underleaves lobes are 3–4 cells long and usually 1 cell wide at the base (rarely 2 cells wide). In robust plants from Vaupés (Río Apoporis, Schultes & Cabrera 11996, 12396 p.p.), however, the underleaf lobes are sometimes larger, varying from 3 cells long and 1 cell wide to 6 cells long and 2 cells wide in the lower half. 20. Drepanolejeunea polyrhiza (Nees) Grolle & R.L.Zhu Rhaphidolejeunea polyrhiza (Nees) Bischl.

Fig. 48Q

On living leaves in lowland rainforest, 100–150 m. Colombia: Caq, Cho, Vau (Schultes & Cabrera 14426, CINC, PC). General distribution: Amazonia, Guianas, Chocó. Drepanolejeunea polyrhiza (subg. Rhaphidolejeunea) is a characteristic epiphyllous species of Amazonian rainforest. As it is known from numerous collections and has never been on bark, the species may be considered an obligate epiphyll (Gradstein and Ilkiu-Borges 2009). It is the only neotropical Drepanolejeunea with rounded leaf tips. The almost horizontally spreading underleaf lobes and the ocelli in a broken line throughout the leaf lamina are further characteristics of the species. The leaf margin is irregularly crenulate, especially in the upper half, and the lobules are reduced or well developed (to 1/2× lobe length) and fully inflated. 21. Drepanolejeunea pterocalyx (Herzog) Bischl. Leptolejeunea pterocalyx Herzog On living leaves in montane rainforest, 1600–1800  m. Colombia: Val (type; Herzog 1955). General distribution: northern Andes (Colombia, Venezuela). Drepanolejeunea pterocalyx (subg. Drepanolejeunea) is a rare northern Andean species that is only known from the type locality from Colombia and a record from Venezuela. Characteristics are the rather widely spreading leaves with toothed margins, the presence of 3–4 ocelli in broken row throughout the leaf lobe, and the lobule inflated below and flattened above and with a short, blunt tooth. The species is rather similar to D. inchoata and D. infundibulata, but in the latter two species, the lobule tooth is longly falcate. Moreover, D. infundibulata has only 2 ocelli. 22. Drepanolejeunea ramentiflora Steph. On bark in montane rainforest, 1700–1800  m. Colombia: Boy (Campos and Uribe 2006). General distribution: Costa Rica (type), Colombia.

Lejeuneaceae

491

Drepanolejeunea ramentiflora (subg. Drepanolejeunea) is a very rare neotropical species that is very similar to D. biocellata but differs in the very flat leaves and the perianths with broad and strongly toothed, horn-like wings. 23. Drepanolejeunea spinosa Herzog

Fig. 48R

On living leaves in montane rainforest, 1400–3400  m. Colombia: Boy, Cau (type), Cho, Tol, Val; to be expected in Ecuador. General distribution: Colombia, French Guiana. Drepanolejeunea spinosa (subg. Drepanolejeunea) is close to D. infundibulata but differs from the latter in smaller plant size and smaller leaf cells (see key). 24. Drepanolejeunea subvittata (Herzog) Grolle Harpalejeunea subvittata Herzog On twigs in páramo, 3080–3700 m. Colombia: Cas (páramo de Chita, very wet páramo vegetation, 3080 m, Cleef 9916, det. R. Grolle, GOET, PC). Ecuador: Az (páramo de Patacocho, 3700 m, Cadena s.n., QCA, PC), Tu (type). General distribution: northern Andes (Ecuador, Colombia). Drepanolejeunea subvittata (subg. Drepanolejeunea) is a rare species from humid northern Andean páramo that is distinguished by (1) leaves suberect-falcate, acuminate, apex terminating in a row of 2–3 cells, base with a 3–4 cells long “vitta” of elongate, rectangular cells in 2 rows; (2) dorsal leaf margin with several long teeth (2–4 cells long) in the lower half; (3) leaf cells with a conspicuous conical dorsal papilla all over the surface of lobe, lobule, and keel; (4) ocelli lacking (?); (5) lobules large, 2/5–1/2× leaf length, inflated along the keel, flattened toward apex, apex with (1–)2 1–celled teeth and the hyaline papilla in between, keel arched, free margin shortly contuining along the ventral margin of the leaf lobe; (6) underleaves 1.5–2× stem width, bifid to 1/2, lobes suberect, 3–5 cells wide, apices broadly rounded; and (7) gynoecia with pycnolejeuneoid innovations (Herzog 1957; Grolle 1991). Drepanolejeunea subvittata is near to D. cutervoensis, but in the latter species, the dorsal leaf margin is entire. Herzog (1957) named D. subvittata for the short vitta of elongate rectangular cells (in 2 rows) at leaf base, but this vitta is not unique to D. subvittata and occurs also in D. cutervoensis and related taxa. 25. Drepanolejeunea urceolata R.M.Schust. On bark in montane forest, 2000–2100 m. Ecuador: ZC (Parolly et al. 2004). General distribution: northern Andes (Venezuela, southern Ecuador). Drepanolejeunea urceolata (subg. Drepanolejeunea) is a very rare northern Andean species that has been recorded only twice, from Venezuela (type) and southern Ecuador. It is habitually similar to D. infundibulata, but the perianth in D. urceolata is not contracted to a beak and has a widely open, deeply lobed mouth. Schuster (1996b) considered D. urceolata nearest to D. evansii; the latter species differs from D. urceolata and D. infundibulata, however, in the entire leaf margins (Bischler 1964).

492

 

Keys and Descriptions

Dubious Records Drepanolejeunea lancifolia (Gottsche) J.B.Jack & Steph. (Lejeunea lancifolia Gottsche)  – Colombia: Cun (type; Gottsche 1864). Ecuador: Co (Arnell 1962). Probably a synonym of D. bidens or D. araucariae (Bischler 1964); the type material is lost. Drepanolejeunea longirostris (Spruce) Steph. – Ecuador: Az, Tu, 2400–3600 m (Bischler 1964). Possibly a synonym of D. bidens or D. araucariae (Bischler 1964); the type material has not been studied. Drepanolejeunea pungens Bischl.  – Described from Colombia (“Andes Bogotense”) based on old, nineteenth-century material in the Mitten herbarium without clear indication of the locality. According to Bischler (1964), the species is very similar to the Asiatic D. ternatensis (Gottsche) Steph. Possibly, the Colombian material was mislabeled and originated from Asia. Drepanolejeunea subdissitifolia Herzog  – Colombia: Mag (Winkler 1976). Ecuador: ZC (Arnell 1962; Parolly and Kürschner 2004b). Dubious species according to León-Yánez et al. (2006); the identity of the specimens needs study. Frullanoides Raddi Plants small to large, 1–10  cm long, 1–3  mm wide, green to black, creeping or ascending, rarely pendent, often forked. Branches Frullania-type and Lejeuneatype. Stems with a hyalodermis; ventral merophyte 4–12 cells wide. Leaf lobes wide-spreading, when dry convolute, apex rounded to acute, margins entire or toothed. Cells slightly longer than wide, walls often with blackish pigmentation, trigones cordate; oil bodies homogeneous; ocelli lacking. Lobules 1/3–2/3 leaf length, never reduced, with 3–11 teeth. Underleaves undivided, margins entire, bases rounded or auriculate, insertion line straight to deeply arched. Male bracts with epistatic lobules. Gynoecia with 2 pycnolejeuneoid innovations. Perianths with 5–11 smooth keels. Sporophytes ptychanthoid. Vegetative reproduction not observed. A mainly neotropical genus of eight species (van Slageren 1985), five in Colombia and Ecuador. The main features of Frullanoides are (1) plants usually blackish-green; (2) stems with a hyalodermis; (3) leaves strongly convolute when dry; (4) lobules with numerous teeth; (5) leaf cells with cordate trigones and homogeneous oil bodies; (6) underleaves undivided and often auriculate at base; and (7) gynoecia with 2 innovations and with a pluriplicate perianth. When sterile, Frullanoides may be confused with Thysananthus (= Mastigolejeunea) which may also be blackish-green in color. However, the lobules in Thysananthus have only 1(−3) teeth; the leaf cells contain large, granular oil bodies; and a stem hyalodermis is lacking.

Lejeuneaceae

493

Literature: van Slageren (1985), Gradstein et al. (1994) 1. Leaves fully entire. Leaf apex rounded, acute or short-apiculate . . . . . . . . . . 2 1. All or some leaves toothed near apex. Leaf apex longly apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. laciniatiflora 2. Leaf apex obtuse to acute to apiculate. Underleaves with deeply arched insertion line, margins ± recurved, basal auricles large, 0.1–0.3 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (F. densifolia) 3 2. Leaf apex broadly rounded. Underleaves with straight or shallowly curved insertion line, margins plane, basal auricles lacking or small, less than 0.1 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Lobule with 6–10 teeth, the teeth 1–3 cells long. Plants dioicous, widespread in tropical America . . . . . . . . . . . . . . . . . . . . . . . . . F. densifolia subsp. densifolia 3. Lobule with 5–6 teeth, the teeth 3–4 cells long. Plants paroicous, only known from the Galápagos Islands . . . . . . . . . . . . . F. densifolia subsp. grandidentata 4. Underleaves without auricles, insertion line straight. Plants small, 1–1.5 mm wide, growing along the coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. corticalis 4. Underleaves with small auricles, insertion line curved. Plants larger, (1–)1.5–3 mm wide, growing inland, above 500 m . . . . . . . . . . . . . . . . . . . . . 5 5. Lobule with 3–4 teeth, the teeth 1(−2) cells long, free margin of the lobule wavy. Ventral margin of the leaf lobe plane. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. liebmanniana 5. Lobule with (4–)6–9 teeth, the teeth 2–3 cells long, free margin not wavy. Ventral margin of the leaf lobe upcurved. Plants paroicous . . . . . . . . F. tristis 1. Frullanoides corticalis (Lehm. & Lindenb.) van Slageren Brachiolejeunea corticalis (Lehm. & Lindenb.) Schiffn.

Fig. 50A

Sun epiphyte on bark of trees in exposed habitats in woodlands, swamps and mangroves, plantations, and on isolated trees, usually in coastal areas, from sea level to 300  m. Colombia: Cho, Nar, SAP.  General distribution: tropical America. Frullanoides corticalis is an oceanic-neotropical species that forms dense, brown green to blackish mats on bark and may grow in extensive mats on trees near the beach. Characteristic features of the species are (1) leaves with rounded apex and entire margins (strongly convolute when dry); (2) underleaves without auricles and with almost straight base; (3) lobules with 3–6 teeth, the teeth being 1–3 cells long and usually inflexed, inconspicuous; and (4) plants usually dioicous. 2a. Frullanoides densifolia Raddi subsp. densifoliaFig. 50B, C Brachiolejeunea densifolia (Raddi) A.Evans Brachiolejeunea bicolor (Nees) Schiffn. Brachiolejeunea columbica Steph. Brachiolejeunea rupestris (Gottsche) Steph. Brachiolejeunea succisa Steph.

Fig. 50 (A) Frullanoides corticalis, habit. (B and C) Frullanoides densifolia subsp. densifolia. (B) Habit with gynoecium. (C) Underleaf. (D and E) Frullanoides liebmanniana. (D) Inner side of lobule apex and free margin. (E) Leaf and lobule free margin. (F and G) Frullanoides tristis. (F) Leaf. (G) Perianth. (H–K) Fulfordianthus pterobryoides. (H) Habit. (I) Lobule. (J) Gynoecium with perianth. (K) Median cells. (L) Harpalejeunea grandis, leaf and underleaf. (M) Harpalejeunea cinchonae, habit. (N) Harpalejeunea oxyphylla, habit. (O and P) Harpalejeunea scabra. (O) Leaf apex in dorsal view. (P) Leaf. (Q and R) Harpalejeunea stricta. (Q) Underleaf. (R) Leaf. (D, E redrawn from van Slageren 1985; M redrawn from Schuster 1999; L redrawn from Grolle and Reiner-Drehwald 1999; O, P from Gradstein and Schäfer-Verwimp 2011; rest from Gradstein et al. 2001; Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009 as H. stricta). Scale bars: A, C, F, G, K, M = 250 μm; B = 1000 μm; D, I, Q, R = 50 μm; E, H, J, L = 500 μm; N, O, P = 100 μm

Lejeuneaceae

495

On bark, rock, and soil in rather open environments from the lowlands to the páramo, from sea level to 4000 m. Colombia: Ant, Boy, Cas, Cun, Hui, Mag, Nar, Ris, San, Tol. Ecuador: Az, Ca, Ch, Ga, Na, Pi, Tu, ZC. General distribution: widespread in tropical America. Frullanoides densifolia is a common neotropical species that is recognized by pointed leaves with entire margins, large lobules with numerous teeth (6–10), and imbricate underleaves with rather large auricles that are 0.1–0.3  mm long. The plants are dioicous. The plants from Colombia and the mainland of Ecuador belong to subsp. densifolia, and those from the Galápagos Islands to subsp. grandidentata; for differences between the two subspecies, see the key. 2b. Frullanoides densifolia subsp. grandidentata (L.Clark) van Slageren Brachiolejeunea grandidentata L.Clark Sun epiphyte growing pendent on bark in humid evergreen woodland and pampa in rather open situations, 450–750 m. Ecuador: Ga (Isabela, Pinzón, San Cristóbal, Santa Cruz). General distribution: only known from the Galápagos Islands. Frullanoides densifolia subsp. grandidentata is a common Galápagos endemic that is recognized by the pendent, blackish, to 4–5-cm-long plants with rather distant leaves and underleaves, an apiculate leaf apex, lobules with 5–6-cell-long teeth (usually 3–4 cells long), and paroicous sexuality. The male bracts, situated directly below the gynoecium, resemble vegetative leaves and are easily overlooked. 3. Frullanoides laciniatiflora (Loitl.) van Slageren Brachiolejeunea laciniatiflora (Loitl.) Steph. On bark in montane rainforest, 1800–2150 m. Ecuador: ZC (Nöske et al. 2003). General distribution: Peru, southern Ecuador. Frullanoides laciniatiflora is a rare Peruvian species that has been found once in southern Ecuador. The species is closely similar to F. densifolia but differs in the toothed margins of leaves and female bracts and bracteoles. In F. densifolia the margins of the female bracts may occasionally possess a few blunt teeth, but the margins of vegetative leaves are always entire. 4. Frullanoides liebmanniana (Lindenb. & Gottsche) van Slageren Brachiolejeunea liebmanniana (Lindenb. & Gottsche) Schiffn.

Fig. 50D, E

Sun epiphyte on bark in the canopy and at the margins of evergreen forests, in scrub, orchards, and on isolated trees, occasionally on rock, 150–1000 m. Colombia: Cho, Cun, Ris, Put, San, Tol; to be expected in Ecuador. General distribution: scattered in tropical America. Frullanoides liebmanniana is an uncommon neotropical species that stands out by the large lobules (1/2–2/3× lobe length) with 3–4 short teeth (1(−2) cells long), a rounded leaf apex, plane ventral margin of the leaf lobe, and underleaves with small auricles. The plants are dioicous. Although originally spelled “liebmaniana,” the epithet of this species is changed to “liebmanniana” under ICN art. 60.1 (Ex. 7) since it was named for Frederik Michael Liebmann.

496

5. Frullanoides tristis (Steph.) van Slageren Brachiolejeunea tristis Steph. Brachiolejeunea chinantlana (Gottsche) Schiffn. Brachiolejeunea wrightii Steph.

 

Keys and Descriptions

Fig. 50F, G

On bark and rock in open submontane and lower montane environments, 250–2000 m. Colombia: Cas, Cho (?), Cun, Mag, Ris. Ecuador: Ga (van Slageren 1985). General distribution: tropical America, East Africa, Himalaya. Frullanoides tristis is a widespread, almost pantropical species that resembles F. liebmanniana but differs in the lobules with more numerous teeth (5–9), the upcurved ventral margin of the leaf lobe, and paroicous sexuality. The record from the Chocó, where the species was found growing mixed with F. liebmanniana (Vasco et al. 2002), should be checked. Fulfordianthus Gradst. Plants usually large, 1–10 cm long, 1.5–4 mm wide, dull dark green with glossy light green growing points, brown with age, dendroid, erect from a rhizome-like shoot, usually densely pinnate. Branches Lejeunea-type only. Stems with a dark brown cortex of small, thick-walled cells in 4–5 layers surrounding a colorless, thinwalled medulla; ventral merophyte more than 10 cells wide. Leaf lobes widespreading, when dry not appressed or convolute, apex broad, margins toothed. Cells subisodiametric, very small, 10–15 μm long, walls uniformly thickened, trigones lacking; oil bodies absent. Lobules small, to 1/5 leaf length, never reduced, with 1 long, curved tooth. Underleaves deeply emarginate, obcuneate, margins toothed, insertion line straight. Gynoecia with 2 lejeuneoid innovations; female bract apices acute. Perianths with 3 smooth keels. Sporophytes ptychanthoid. Vegetative reproduction not observed. A small neotropical genus with two species (Gradstein 1994), one in Colombia and Ecuador. For main features of Fulfordianthus, see under the species. 1. Fulfordianthus pterobryoides (Spruce) Gradst. Lejeunea pterobryoides Spruce

Fig. 50H–K

On bark of treelets and lianas and on living leaves in the understory of virgin lowland and submontane rainforests, from sea level to 1500 m. Colombia: Ama, Cau, Cho, Put, Val. Ecuador: Es, Ma, Na, Or (J. Deleg s.n., HUTPL), Pa. General distribution: throughout the Chocó biogeographic region (Costa Rica to Peru); additionally recorded from eastern Amazonia, near the base of the Andes. Fulfordianthus pterobryoides is a very characteristic and common, robust Chocó species that is recognized by (1) erect-growing, densely (bi)pinnate plants with feather-like habit; (2) Lejeunea-type branching (no Frullania-type branches); (3) underleaves deeply emarginate; (4) leaf margins crenulate to sharply toothed; (5) leaf cells very small, 10–15  μm long in midleaf, with uniformly thickened walls without trigones; (6) oil bodies absent; and (7) gynoecia with 2 innovations. The plants resemble Bryopteris in general habit, but Bryopteris has Frullania-type

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branching, larger leaf cells with trigones and oil bodies, and a rounded or truncate underleaf apex (not emarginate) and lacks innovations. Harpalejeunea (Spruce) Schiffn. Plants small, to 1.5 cm long, usually less than 1 mm wide (rarely broader), yellowgreen to olive brown to dark brown, creeping or ascending. Stems of thick-walled cells, without distinct hyalodermis; ventral merophyte 2–4 cells wide. Leaf lobes suberect to rather wide-spreading, convex to subsquarrose, asymmetrically ovatefalcate, somewhat convolute when dry, with ocelli, apex obtuse to acute-acuminate to piliferous, plane or recurved, margins entire, crenate or toothed. Cells rather small, 15–25 μm in midleaf, with trigones, cuticle smooth; oil bodies finely granular; ocelli colorless or grayish, (1–)2–3(−4) in an unbroken row just above the leaf base (parallel to the free margin of the lobule and in ventral view often covered by the free margin), occasionally occurring scattered in the lamina of the leaf lobe (see H. tridens). Lobules well developed, 1/3–1/2 leaf length, never reduced, strongly inflated and somewhat flask-shaped, keel strongly arched, smooth or crenulate, usually forming a sharp angle with the ventral margin of the leaf lobe, sinus indented, lobule tooth 1-celled, obtuse to acute, falcate, apical free margin semicircular, often terminating in a blunt pre-apical tooth at the junction of lobule and free margin. Underleaves emarginate to bifid (rarely undivided) with broadly obtuse, diverging lobes and a wide sinus, outer margins entire, insertion line ± straight. Androecia on short-specialized branches, bracteoles limited to the base of the branch. Gynoecia on short or long branches, with 1–2 lejeuneoid innovations. Perianths inflated, sharply 5-keeled, the keels entire or denticulate or ciliate-laciniate, not extending into horns. Sporophytes lejeuneoid. Vegetative reproduction occasionally by caducous leaf lobes. A pantropical genus (ca. 15 spp.); seven species are recognized here in Colombia and Ecuador. The main features of Harpalejeunea are (1) underleaves emarginate to shallowly bifid (rarely undivided), lobes broadly rounded; (2) leaves ovate-falcate, margins entire or crenate, apex obtuse to acute-acuminate to piliferous; (3) lobules strongly inflated, always well developed, with a falcate tooth; (4) ocelli present, in a short, suprabasal, unbroken row; and (5) gynoecia with 1–2 lejeuneoid innovations and a 5-keeled perianth with smooth or ciliate keels. The genus is sister to Lejeunea and Microlejeunea (Dong et al. 2013). Harpalejeunea resembles some species of Drepanolejeunea, especially D. ­cutervoensis and D. granatensis, but the latter genus differs in having pycnolejeuneoid innovations, ocelli usually in a broken row (occasionally 2 ocelli in an unbroken basal row), and underleaf lobes mostly 1 cell wide (broader in D. cutervoensis and D. granatensis). Epiphyllous plants of Drepanolejeunea produce large, adhesive rhizoid discs; such discs have not been seen in Harpalejeunea. The genus Harpalejeunea has not been monographed, and the present treatment is a preliminary one. The genus needs more study.

498

 

Keys and Descriptions

Literature: Grolle and Reiner-Drehwald (1999), Schuster (1999), Gradstein and Ilkiu-Borges (2009) 1. Ventral merophyte 4 cells wide on main stems (2 cells wide on branches). Underleaves undivided or shallowly emarginate. Plants 1–1.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. grandis 1. Ventral merophyte 2 cells wide on main stems. Underleaves emarginate to bifid (to max. 1/2). Plants smaller, less than 1 mm wide. . . . . . . . . . . . . . . . . . . . . 2 2. Leaf cells on the dorsal surface with a large, conically elevated papilla. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. scabra 2. Leaf cells without papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Dorsal margin of leaf lobes strongly crenate and sometimes with 1 or more ciliate teeth. (Leaf apex ending in a row of 2–5 cells) . . . . . . . . . . . . . . H. tridens 3. Dorsal leaf margin entire or crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaf apex longly piliferous, rather abruptly ending in a long hair, the hair (3–)4–8 cells long. Caducous leaf lobes sometimes present . . . . . . . . . . . . . . 5 4. Leaf apex obtuse to acute to shortly acuminate, ending in a row of 1–3 cells. Caducous leaf lobes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Leaf margins entire. Perianth keels smooth or toothed by 1–3-cell-long projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. oxyphylla 5. Leaf margins crenulate. Perianth keels longly ciliate-laciniate. . . H. uncinata 6. Plants very small, 0.4–0.5 mm wide. Ventral epidermis cells less than 20 μm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. stricta 6. Plants larger. Ventral epidermis cells 25–30 μm wide. . . . . . . . . H. cinchonae 1. Harpalejeunea cinchonae (Nees) Schiffn. Fig. 50M Lejeunea cinchonae Nees. Type: Peru, on bark of Cinchona, ex hb. Nees (isotype G-00115071!) Harpalejeunea acuta S.Winkl., syn. nov. Type: Colombia, Magdalena, Cuchilla de San Lorenzo, 2550 m, 13 Jan 1967, Winkler C180 (COL? n.v.) Harpalejeunea ancistrodes (Spruce) Schiffn. (Lejeunea ancistrodes Spruce), syn. nov. Type: Ecuador, Tungurahua, Quisapincha, near Ambato, Spruce L39/2 (lectotype, designated by Grolle and Reiner-Drehwald (1999), MANCH-cc17802!, c. gyn.) Harpalejeunea cinchonae var. strigulosa Herzog Harpalejeunea grandistipula R.M.Schust., syn. nov. Type: Ecuador, Napo, “on a small peak N. of Pifo-Papallacta Rd.; in moss tundra,” 4200–4300 m, Schuster 93-220b (F n.v.); the material cannot be found (L. Briscoe, pers. comm.). Harpalejeunea tuberculata (J.B.Jack & Steph.) Steph. On trunks, branches, and twigs of trees and shrubs in montane rainforest areas up to the páramo, sometimes creeping over dense bryophyte mats, 500–4300  m. Colombia: Ant, Cas, Mag, Tol (Aguirre & Gradstein 1539, det. R. Grolle, COL, GOET). Ecuador: Az, Ca, Lo, Na, Pi, Su, Tu, ZC. General distribution: tropical America.

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The main features of Harpalejeunea cinchonae are (1) plants 0.7–1 mm wide, central merophyte 2 cells wide, ventral epidermis cells 25–30 μm wide; (2) leaves convex, margins entire to slightly crenulate, apices obtuse to acute to shortly acuminate, ending in a row of 1–2(−3) cells, plane or recurved; (3) median leaf cells 20–30 μm long, with small to large trigones and intermediate thickening, ocelli in a 2–3-cell-long suprabasal row, sometimes 1–2 additional ocelli present in the distal half of the leaves (seen in Bolivia, Gradstein 7181, PC-0706256); (4) underleaves distant to contiguous, 2–4× stem width, shallowly emarginate with a broad sinus and rounded lobes, lobes 5–10 cells wide, lamina 6–10 cells high from base to sinus; and (5) perianth elongate-obovoid, strongly 5-keeled, keels smooth or with a few cilia. Harpalejeunea cinchonae shows considerable variation in the shape of the leaf tips, which may be obtuse to subacute (type) to shortly acuminate, and underleaf size, measuring 2–4× stem width. Based on the original descriptions and illustrations, H. acuta (Winkler 1976) and H. grandistipula (Schuster 1999) are synonyms of H. cinchonae. The types of these species were not available for study. Harpalejeunea cinchonae may be confused with H. stricta, but the latter species is a smaller plant, measuring maximally 0.5 mm wide, with smaller leaf cells, narrower epidermis cells, and smaller, emarginate to short-bifid underleaves. Moreover, the leaf apex in H. stricta is always acute to shortly acuminate (terminating in a row of [1–]2–3 cells), never obtuse. 2. Harpalejeunea grandis Grolle & M.E.Reiner

Fig. 50L

On bark and humic soil in páramo, 3200–3800 m. Colombia: Boy, Cun, Ris; to be expected in Ecuador. General distribution: only known from Colombia. Harpalejeunea grandis is a Colombian páramo endemic that is recognized by (1) plants more than 1 mm wide, with rather lax foliation; (2) ventral merophyte 4 cells wide on main stems (2 cells wide on branches and poorly developed shoots); (3) leaves subimbricate, slightly falcate, apex rounded to acute to short-acuminate; (4) ocelli in 1–2 rows; (5) leaf cells with large trigones; and (6) underleaf apex undivided to shallowly retuse. In general habit H. grandis resembles a laxly growing Lejeunea rather than a Harpalejeunea. The salient features of the species are the broad ventral merophytes, the ± undivided underleaf apex, and the 1–2 rows of ocelli (Grolle and ReinerDrehwald 1999). 3. Harpalejeunea oxyphylla (Nees & Mont.) Steph. Fig. 50N Harpalejeunea buenaventurae Herzog (?) Type: Colombia, El Valle, Buenaventura, 0–10 m, Killip 11712 p.p. (JE; type not available) Harpalejeunea pilifera (Spruce) Steph. On bark and living leaves in the canopy and the understory of lowland and montane rainforests, also on rotten logs and living leaves, from sea level to 2500  m. Colombia: Cau, Cho, Qui, Ris. Ecuador: MS, Na, Or, Pa, Tu, ZC. General distribution: widespread in tropical America.

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Keys and Descriptions

Harpalejeunea oxyphylla is a common neotropical species that is readily distinguished by the apex ending rather abruptly in a long, (3–)4–8-cell-long hair. The leaf margins are entire, the leaf cells have small or medium-sized trigones and intermediate thickenings, the underleaves are shallowly to deeply bifid, and the female involucre and perianth keels are entire or toothed. Vegetative reproduction is by caducous leaf lobes. Harpalejeunea buenaventurae, described from Colombia, may be a phenotype of H. oxyphylla with shallowly bifid underleaves. The type was not available for study. 4. Harpalejeunea scabra Gradst. & Schäf.-Verw. Harpalejeunea aspera Grolle, nom. inval.

Fig. 50O, P

On rock in disturbed montane rainforest, ca. 1800 m. Colombia: Ris (Gradstein and Schäfer-Verwimp 2011). General distribution: Colombia, Costa Rica. Harpalejeunea scabra is a rare northern Andean-Costa Rican endemic that is readily recognized by the dorsal leaf surface roughened by mammillose cells crowned by a conically elevated papilla. Papillose leaf surfaces are also seen in some species of Drepanolejeunea, Lejeunea, and Prionolejeunea. Harpalejeunea scabra is readily separated from these species by the broad, diverging underleaf lobes and the short row of ocelli (2–3) near leaf base. 5. Harpalejeunea stricta (Lindenb. & Gottsche) Steph. Fig. 50Q, R Lejeunea stricta Lindenb. & Gottsche. Type: Mexico, Veracruz, Mirador, Liebmann s.n., c. gyn. (isosyntypes G-00115038!, G-0011539!) Harpalejeunea herzogii Jovet-Ast, syn. nov. Type: Guadeloupe, “Carbet, sur rocher humide,” 15 February 1936, P. & V.  Allorge s.n., c. gyn. (holotype PC-0103158!) On bark in lowland and montane rainforests, also on rotten logs, usually in wellexposed sites, from sea level to 2150 m. Colombia: Boy, Cho, Cun, Put, Ris, Vau. Ecuador: MS, Or, Pa, Pi, Tu, ZC. General distribution: widespread in tropical America. Harpalejeunea stricta is a widespread neotropical species that is morphologically close to H. cinchonae but differs in smaller plant size (0.4–0.5  mm wide), smaller cortical cells (13–18 μm wide on the ventral stem surface), and smaller leaf cells, 14–20(−23) μm long in midleaf. Moreover, the leaves in H. stricta are strongly convex; the leaf tips are acute to shortly acuminate (never obtuse), terminating in a row of (1–)2–3 cells; and the underleaves are emarginate to short-bifid (to 1/4). 6. Harpalejeunea tridens (Besch. & Spruce) Steph. Harpalejeunea heterodonta A.Evans

Fig. 51A, B

On bark, twigs, rotten logs, and living leaves in lowland and montane rainforests, from sea level to 2400  m. Colombia: Cho, Val. Ecuador: EO, Lo, Na, Or, Pa, ZC. General distribution: scattered in tropical America. Harpalejeunea tridens is a distinct neotropical species recognized by (1) leaf margins crenate and often with 1–2 ciliate teeth on the dorsal margin; (2) leaf apex

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Fig. 51 (A and B) Harpalejeunea tridens, leaves. (C) Lejeunea asperrima, habit in dorsal view. (D) Lejeunea acanthogona, habit. (E) Lejeunea adpressa, habit. (F) Lejeunea aphanes, habit with gynoecium. (G) Lejeunea bombonasensis, habit with gynoecia and sporophyte. (H and I) Lejeunea boryana. (H) Habit. (I) Underleaf. (J and K) Lejeunea cancellata. (J) Habit. (K) Underleaf. (L) Lejeunea capensis, habit with gynoecium. (M) Lejeunea catinulifera, habit with gynoecium. (N) Lejeunea cerina, habit. (D redrawn from Reiner-Drehwald and Goda 2000; J, K redrawn from Reiner-Drehwald 2000; M redrawn from Reiner-Drehwald and Schäfer-Verwimp 2008a; rest from Gradstein et al. 2001; Gradstein and Ilkiu-Borges 2009). Scales bars: A, B, K = 100 μm; C–F, H, L, N = 250 μm; G, M = 500 μm; I = 50 μm; J = 400 μm

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Keys and Descriptions

ending in a row of 2–5 cells; (3) underleaves bifid to 1/4–1/2, with diverging, rounded lobes. A specimen from El Oro (Reserva Buenaventura, on canopy branch in mat of Prionolejeunea decora, Nov 2018, Benitez et al. s.n., HUTPL) stands out by caducous leaves and by the frequent presence of scattered ocelli in the leaf lamina, additional to the ocelli (1–2) at the leaf base. In other respects the plants were quite similar to H. tridens. The laminar ocelli varied from ca. 3 to 8 per leaf and in some leaves they were lacking. Scattered laminar ocelli are rare in Harpalejeunea and have otherwise been observed in H. reflexula A.Evans, a species described from Jamaica (Schäfer-Verwimp and van Melick 2016) and quite similar to H. tridens. The relationships between these two species need study. 7. Harpalejeunea uncinata Steph. Harpalejeunea uncinata var. setulosa Herzog Colombia: Cau, Val (Herzog 1955 as var. setulosa). Ecuador: Na, Tu (SchäferVerwimp et al. 2006). General distribution: scattered in tropical America. Harpalejeunea uncinata is close to H. oxyphylla and was treated as a synonym of the latter species by Gradstein and Ilkiu-Borges (2009). However, H. uncinata can be separated from H. oxyphylla by the crenulate leaf margins and ciliate-laciniate perianth keels (Evans 1903; Schäfer-Verwimp et al. 2006).

Further Records Harpalejeunea harpaphylla (Herzog) Bischl.  – Ecuador: LR (Herzog 1952). Resembles H. tridens but underleaves more deeply bifid and leaf margins only slightly crenulate and without teeth. The type was not available for study. Lejeunea Lib. Amblyolejeunea Jovet-Ast, Amphilejeunea R.M.Schust., Crossotolejeunea (Spruce) Schiffn., Cryptogynolejeunea R.M.Schust., Dicladolejeunea R.M.Schust., Echinocolea R.M.Schust., Eulejeunea Steph., Hygrolejeunea (Spruce) Schiffn., Macrolejeunea (Spruce) Schiffn., Neopotamolejeunea M.E.Reiner, Oryzolejeunea (R.M.Schust.) R.M.Schust., Prionocolea R.M.Schust., Sphaerolejeunea Herzog, Taxilejeunea (Spruce) Steph. Plants small to rather large, stems 0.5–10 cm long, 0.4–3 mm wide, pale to bright green or whitish, often glossy, creeping or hanging. Stems thin or thick, usually with hyalodermis; ventral merophyte 2(−8) cells wide. Leaf lobes obliquely to widely spreading, apex rounded to apiculate to acute-acuminate, plane, margins entire or crenulate, rarely toothed, leaf surface smooth, rarely rough by conically projecting cells. Cells plane, usually thin-walled without or with small trigones, intermediate thickenings present or absent; oil bodies small, finely granular or homogeneous; cuticle smooth or finely papillose (“punctate”), sometimes with small wax crystals;

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ocelli lacking. Lobules often reduced, when well developed to maximally 1/2× leaf length, oval, with 1(−2) teeth; hyaline papilla proximal (distal in L. herminieri). Underleaves bifid or undivided, small or large. Androecia with bracteoles restricted to the base of the male spike or present throughout. Gynoecia on long or short shoots, with (0–)1–2 lejeuneoid innovations, the innovations sterile or fertile, bracts usually unwinged (winged in L. schultesii). Perianths with 0–5 keels, the keels smooth, crenate, toothed or ornamented with lacinia and/or cilia, apex with or without beak. Sporophytes lejeuneoid. Vegetative reproduction by caducous leaves or leaf lobes, caducous branches, shoot fragmentation, or by elongate, ribbon-like gemmae from leaf margins. A large, pantropical genus (ca. 200 spp.), at least 66 species in Colombia and Ecuador. The main generic features of Lejeunea as currently defined are (1) underleaves usually bifid, occasionally entire, lobes upright (not diverging); (2) stems fragile and with hyalodermis, occasionally rigid and without hyalodermis (especially in rheophytic species), ventral merophyte 2(−8) cells wide; (3) lobules to 1/2× lobe length, often reduced, with a proximal hyaline papilla; (4) cell walls without brownish pigmentation, oil bodies small; (5) ocelli absent; and (6) gynoecia with lejeuneoid innovations. The first lobule tooth in Lejeunea is usually 1(−2) cells long, but on reduced lobules it may be longer, 3–4(−12) cells long. Together with Plagiochila, Lejeunea is the most speciose genus in the northern Andes. Its circumscription has undergone considerable modification based on recent studies, and several genera accepted in the “Guide to the Bryophytes of Tropical America” (Gradstein et  al. 2001), such as Amblyolejeunea, Amphilejeunea, Crossotolejeunea, Echinocolea, Macrolejeunea, Oryzolejeunea, Sphaerolejeunea and Taxilejeunea, are now included in Lejeunea. On the other hand, L. subg. Otigoniolejeunea proved to be a separate genus (Wei et al. 2014). The infrageneric classification of Lejeunea is still unclear and needs more study (Heinrichs et  al. 2013a). Our understanding of the neotropical species is largely based on the important taxonomic studies of Dr. M. Elena Reiner-Drehwald (Reiner-Drehwald 2000, 2003, 2005a, b, c, 2009a, b, 2010a, b; Reiner-Drehwald and Goda 2000; ReinerDrehwald and Drehwald 2002; Reiner-Drehwald and Schäfer-Verwimp 2008a, b) and the recent treatment for Brazil (Bastos and Gradstein 2020). Nevertheless, many species remain poorly known, especially those of the former genus Taxilejeunea. The present treatment is a preliminary one. Literature: See above. Key 1. Introductory key 1. Dorsal leaf surface rough due to conically projecting cells, at least in the upper part of the leaf. Plants very small, less than 1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (sect. Echinocolea [R.M.Schust.] Gradst.) 2 1. Dorsal leaf surface smooth. Plants small or large . . . . . . . . . . . . . . . . . . . . . . 4 2. Keel strongly crenate due to conically projecting cells. Underleaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. asperrima 2. Keel smooth. Underleaf margins entire or with a tooth. . . . . . . . . . . . . . . . . . 3

504

 

Keys and Descriptions

3. Underleaf margins with a tooth. Dorsal leaf surface smooth at the base only. Plants growing above 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . L. meridensis 3. Underleaf margins entire. Dorsal leaf surface smooth up to leaf middle or more. Plants growing below 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . L. subspathulata 4. All or part of the underleaves undivided (former genera Amblyolejeunea, Amphilejeunea, Cryptogynolejeunea, Oryzolejeunea) . . . . . . . . . . . . . . . . . . 5 4. All underleaves bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 5. Insertion line of underleaves ± arched. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Insertion line of underleaves straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Lobules ca. 1/5× lobe length. with 1 small tooth. Underleaves reniform, (4–)6–8× stem width, apex usually recurved. Plants from lowland and montane forest, mostly below 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . L. reflexistipula 6. Lobules larger, 1/3–1/2× lobe length, with (1–)2 teeth. Underleaves suborbicular, 3–5× stem width, apex plane or narrowly recurved. Plants from upper montane forest and páramo, mostly above 2000 m . . . . . . . . . . . . . . L. catinulifera 7. Plants very small, 0.4–0.7 mm wide. Leaves suberect to obliquely spreading. Lobule tooth at least 4× longer than wide, acute . . . . Cheilolejeunea holostipa 7. Plants larger. Leaves wide-spreading. Lobule tooth short, obtuse . . . . . . . . . . 8 8. Cuticle smooth. Distal margin cells of lobule (between apex and keel) enlarged. Hyaline papilla proximal. Perianth terete, beak absent. Plants autoicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. fulfordiae 8. Cuticle finely papillose. Distal margin cells of lobule not enlarged. Hyaline papilla distal. Perianth keeled, beak present. Plants dioicous . . . L. herminieri 9. Underleaves very large, 5–10× stem width, frequently auriculate (former genera Macrolejeunea, Hygrolejeunea p.p., and Taxilejeunea p.p.), insertion line of underleaf usually deeply arched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Underleaves smaller, 1–5× stem width [when in doubt try both leads] . . . . . 22 10. Leaf apices broadly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10. All or some leaf apices pointed (obtuse, acute, apiculate or acuminate) . . . . 14 11. Ventral merophyte (3–)4–6 cells wide. . . . . . . . . . . . . . . . . . . . L. rotundifolia 11. Ventral merophyte 2 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Lobules large, 1/3–1/2× leaf length, with (1-)2 teeth. Underleaves shallowly bifid to maximally 1/5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. catinulifera 12. Lobules smaller, less than 1/3× leaf length, without or with 1 small tooth. Underleaves bifid to 1/5–1/3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Cuticle densely papillose. Underleaves bifid to 1/3–1/2. Leaves with a border of 5–6 rows of smaller marginal cells perpendicular to the larger central cells. Gynoecia on a short or long branch, with innovation. Perianth keeled, the keels toothed or entire. Plants autoicous, in submontane and montane forest below 2000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pulverulenta 13. Cuticle smooth. Underleaves shallowly bifid, to 1/5–1/4. Leaves without border of 5–6 rows of smaller marginal cells. Gynoecia on a very short branch, without innovation. Perianth terete. Plants dioicous, in páramo and upper montane forest above 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. elongella

Lejeuneaceae

505

Lejeunea recurva M.E.Reiner from Bolivia and Peru also keys out here; the species is characterized by reniform, shallowly bifid underleaves with a broadly recurved apex, unbordered leaves with recurved apices, papillose cuticle, and 5-keeled perianths (Reiner-Drehwald et al. 2013).

1 4. Underleaves with large auricles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14. Underleaves without large auricles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 15. Underleaf insertion line deeply arched, 0.1–0.2 mm deep. Gynoecia with fertile innovations, produced in a row. Lobules reduced or very small, to maximally 1/7× leaf length. Leaf cells without or with minute trigones, intermediate thickenings lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15. Underleaf insertion line shallowly curved, to 0.05 mm deep. Gynoecia without or with a short-sterile innovation, gynoecia not in a row. Lobules usually well developed, 1/6–1/3× leaf length. Leaf cells with trigones and intermediate thickenings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 16. Leaves strongly toothed, with up to 15 teeth. Perianth with toothed keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. chimborazensis 16. Leaves with 1–5(−7) teeth or without teeth. Perianth terete, without keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. Underleaves shallowly bifid to 1/6–1/5. Cuticle smooth. Perianth terete . . . 18 17. Underleaves more deeply bifid, to 1/4–1/3. Cuticle usually finely punctate-­ papillose (occasionally smooth). Perianth keeled . . . . . . . . . . . . . . . . . . . . . 19 18. Leaf margins crenulate, without or with 1–2 small teeth near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. serpillifolioides 18. Leaves clearly toothed, with more than 2 teeth . . . . . . . . . . . . . . L. sulphurea 19. Plants autoicous, frequently fertile. Leaves with 0–3 teeth near the apex. Perianth distinctly and irregularly toothed at apex . . . . . . . . . . . . L. perigonia 19. Plants dioicous, frequently sterile. Leaves entire or with 1 tooth, the tooth separated from the apex by a long and shallow, lunulate sinus (rarely a second, smaller tooth present). Perianth keels without teeth (rarely with 1–2 small teeth). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. debilis 20. Leaves ovate-triangular, gradually narrowed to the apex, margins entire. Lobules 1/4–1/3× leaf length. Perianth terete to weakly 3-keeled. Plants growing above 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pallescens 20. Leaves broadly ovate, not gradually narrowed to the apex, margins crenulate. Lobules smaller, ca. 1/6× leaf length. Perianth 5-keeled. Plants growing from sea level to 2500 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L.cerina 21. Insertion line of underleaves deeply arched, ca. 0.2 mm deep. Underleaves shallowly bifid to 1/5–1/4, with a very narrow incision. Perianth keels crenulate or smooth. Male bracteoles limited to the base of the male spike. . . L. caripensis 21. Insertion line of underleaves shallowly curved, to 0.05 mm deep. Underleaves more deeply bifid to 1/3–1/2, with a wide, V-shaped incision. Perianth keels dentate-laciniate. Male bracteoles present throughout the male spike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. prionoides 22. At least some leaf apices pointed (obtuse, acute, apiculate or acuminate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key 2

506

 

Keys and Descriptions

22. Leaf apices rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key 3 Key 2. Underleaves 1–5× stem width, bifid. All or some leaf apices pointed (obtuse, acute, apiculate, or acuminate) 1. Leaf margins in all or some leaves with a few teeth (or many), at least near apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 2. Lobules mostly well developed, inflated-swollen, 1/4–2/5× leaf length. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Lobules mostly reduced [when in doubt try both leads] . . . . . . . . . . . . . . . . . 5 3. Leaves sharply and irregularly toothed-ciliate along the margins, part of the teeth more than 1 cell long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Leaves with only a few small, obtuse teeth near the apex, teeth not more than 1 cell long. (Leaf apex obtuse to apiculate. Underleaves very small, 1.5–2.5× stem width. Perianth longly 5-keeled, the keels 2-winged, irregularly toothed. Plants autoicous). Plants of West Indies, Brazil, and northern Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bermudiana 4. Leaf lobes caducous. Leaf margins with scattered rhizoids and with bluntish teeth terminating in a rhizoid. Lobule tooth curved, more than 25  μm long. Plants of Colombia (Quindio), in montane forest . . . . . . . . . . . . . . L. ryszardii 4. Leaf lobes not caducous. Leaf margins without rhizoids, with acute teeth not terminating in a rhizoid. Lobule tooth straight, short, less than 25  μm long. Plants of West Indies, not known from Colombia and Ecuador. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. paucidentata (Steph.) Grolle 5. Leaves irregularly toothed in the upper half, with more than 5 teeth . . . .  L. killipii 5. Leaves with less than 5 teeth near apex (or entire) . . . . . . . . . . . . . . . . . . . . . 6 6. Leaf apex acute to shortly acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6. Leaf apex rounded to obtuse to apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Perianth terete, without keels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejeunea sp. Resembles L. sulphurea but underleaves smaller (ca. 4× stem width), distant and more deeply bifid (to ½ of underleaf length measured from the insertion to the apex) 7. Perianth with 5 short keels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. asthenica 8. Perianth keeled only near apex, in the upper 1/4–1/3. Underleaves 3–5× wider than the stem. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. flaccida 8. Perianth more longly keeled, over 1/2–2/3 of perianth length. Underleaves 2–4× wider than the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bombonasensis 9. Leaves 1.5–4× longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Leaves 1–1.5× longer than wide [when in doubt try both leads]. . . . . . . . . . 13 10. Plants very small, 0.4–0.8  mm wide. Leaf apex obtuse. Cuticle finely papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. laetevirens 10. Plants larger. Leaf apex acute-acuminate. Cuticle smooth or papillose . . . . . 11 11. Cuticle ± smooth. Gynoecia without innovation. Perianths slightly keeled at the apex, otherwise terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. osculatiana 11. Cuticle densely papillose. Gynoecia with innovation. Perianth distinctly keeled in the upper half. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Lejeuneaceae

507

12. Leaves 3–4× longer than wide. Plants autoious. Gynoecia in a row of 2–5 gynoecia on short branches. Perianth keels smooth . . . . . . . . . . L. xiphophylla 12. Leaves 1.5–2× longer than wide. Plants dioicous. Gynoecia usually single, rarely in a row of 2 gynoecia. Perianth keels toothed . . . . . . . . . . L. ramulosa 13. Gynoecia in a row of 2–6 gynoecia on short branches (sometimes also on long shoots). Lobules reduced. Plants autoicous. . . . . . . . . . . . . . . . . . . . . . . . . . . 6 13. Gynoecia not in a row or in a row of maximally 2 gynoecia or gynoecia absent. Lobules well developed or reduced. Plants autoicous or dioicous . . . . . . . . . 14 14. Cuticle finely punctate-papillose. Underleaves 3–5× stem width . . . . . . . . . 15 14. Cuticle mostly smooth. Underleaves smaller, 1–3(−4)× stem width . . . . . . . 20 15. Underleaf bases auriculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15. Underleaf bases not auriculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Leaves ovate-triangular, gradually narrowed to the apex, margins entire. Underleaves with large auricles. Lobules 1/4–1/3× leaf length. Perianth ± terete. Plants growing above 2000 m . . . . . . . . . . . . . . . . . . . . . . L. pallescens 16. Leaves broadly ovate, not gradually narrowed to the apex, margins crenulate. Underleaves with small auricles. Lobules smaller, ca. 1/6× leaf length. Perianth 5-keeled. Plants growing from sea level to 2500 m. . . . . . . . . . . . . . . L. cerina 17. Outer margins of underleaves with a tooth, bases cuneate. Leaf margins ± crenate. Cuticle conspicuously and densely papillose . . . . . . . . . . . . . L. boryana 17. Outer margins of underleaves without a tooth, underleaf bases rounded or cuneate. Leaf margins ± entire. Cuticle conspicuously to faintly punctate-papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18. Underleaf lobes finely acuminate, ending in a row of (1–)2–4 cells. Cuticle densely papillose. Perianth keeled. . . . . . . . . . . . . . . . . . . . . . . L. controversa 18. Underleaf lobes narrowly rounded to acute, ending in a row of 1–2 cells. Cuticle faintly punctate-papillose. Perianth without keels. . . . . . . . . . . . . . . . . . . . . 19 19. Perianth with a huge beak, almost as wide as the perianth . . . . L. cyathophora 19. Perianth with a small beak (as usual), much narrower than the perianth mouth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. capensis 20. Leaf apex rounded to acute to acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20. Leaf apex rounded to obtuse, never acute or acuminate [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key 3 21. Lobules when well developed to 1/3–2/5× leaf length. Perianth keels ciliatelaciniate or crenate. Leaf margins crenate. . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21. Lobules smaller, often reduced. Perianth keels entire, crenulate or crenate (rarely with lamella-like projections, L. quinqueumbonata). Leaf margins entire to crenate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22. Leaf margins and perianth keels without border of transversely oriented mammillose cells. Perianth keels ciliate-laciniate . . . . . . . . . . . . . . L. acanthogona 22. Leaf margins and perianth keels with a border of transversely oriented mammillose cells. Perianth keels crenate, occasionally with a few cilia, lacinia lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. raddiana 23. Leaf margins distinctly crenate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. Leaf margins entire or slightly crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

508

 

Keys and Descriptions

24. Leaves conspicuously bordered by transversally oriented cells. Perianth keels  strongly crenate, without lamella-like projections. Rare plants from Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. immersa 24. Leaves not conspicuously bordered by transversally oriented cells. Perianth keels not strongly crenate, with lamella-like projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. quinqueumbonata 25. Leaf apex shortly acuminate. Underleaf bases cuneate. Gynoecia without innovations. Perianth slightly keeled at the apex, otherwise terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. osculatiana 25. Leaf apex rounded to acute to shortly apiculate. Underleaf bases rounded or cuneate. Gynoecia with innovations. Perianth distinctly keeled in the upper half or over most of its length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 26. Underleaves wider than long. Branches frequently caducous. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cancellata 26. Underleaves longer than wide or orbicular. Branches not caducous. Plants autoicous or dioicous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27. Plants with numerous small, microphyllous branches, dioicous, often sterile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. flagellaris 27. Plants without microphyllous branches, autoicous, usually fertile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bombonasensis Key 3. Underleaves 1–5× stem width, bifid. Leaf apex rounded 1. Leaf lobes bordered by dead, hyaline, rectangular cells in 1–2 rows. Perianth terete, without beak. Plants epiphyllous, 1.5–2.3  mm wide, growing tightly appressed to the substrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. drehwaldii 1. Leaves not bordered. Perianth terete or with keels . . . . . . . . . . . . . . . . . . . . . 2 2. Stems robust, ventral merophyte 4–8 cells wide. Rheophytic. . . . L. topoensis 2. Stems thinner, ventral merophyte 2 cells wide . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Ventral margin of the leaf lobe strongly revolute. Plants of páramo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. inflexiloba 3. Ventral margin of the leaf lobe not strongly revolute . . . . . . . . . . . . . . . . . . . 4 4. Lobule tooth 3–12 cells long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Lobule tooth 1–2 cells long or lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Lobule tooth (4–)6–12 cells long. Leaves suborbicular, flat. Perianth obcordate, flattened, keeled over its whole length, keels smooth . . . . . . . . . . L. trinitensis 5. Lobule tooth shorter, 3–4(−6) cells long. Leaves ovate-oblong, somewhat convex. Perianth obpyriform, inflated, keeled in the upper half, keels crenate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. setiloba 6. Perianth terete (without keels). Plants autoicous, usually fertile . . . . . . . . . . . 7 6. Perianth keeled or perianth absent. Plants autoicous or dioicous . . . . . . . . . . 8 7. Underleaves very small, 1.5–2× stem width. Perianth longly exserted beyond the bracts, to more than 1/2 its length, beak absent . . . . . . . . . . . L. corynantha 7. Underleaves larger, 2.5–5× stem width. Perianth exserted less than 1/2 its length, beak present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. capensis 8. Leaves or leaf lobes caducous. Leaf margins sometimes with rhizoids (on shoots producing caducous leaf lobes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Lejeuneaceae

509

8. Leaves or leaf lobes not caducous. Leaf margins without rhizoids . . . . . . . . 15 9. Leaf cells with intermediate thickenings and trigones. Leaves widely spreading. Underleaves present throughout the male spike . . . . . . . . . . . . . . . . . . . 10 9. Leaf cells without intermediate thickenings, walls thin or ± evenly thickened (trigones present or lacking). Leaves obliquely or widely spreading. Underleaves limited to the base of the male spike (present throughout the male spike in L. rionegrensis, couplet 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10. Plants 1–1.6 mm wide. Underleaves 3–4× stem width, bases broadly rounded to rounded-quadrate. Trigones elongate, triradiate . . . . . . . . . . . . L. tapajosensis 10. Plants smaller, 0.6–1.1  mm wide. Underleaves smaller, 1.5–3× stem width, bases cuneate to narrowly rounded. Trigones not elongate, not triradiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. phyllobola 11. Plants very small, 0.4–0.8  mm wide. Underleaves scarcely wider than the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11. Plants larger, (0.8–)1–1.5 mm wide. Underleaves 2.5–5× stem width . . . . . 14 12. Leaves suberect to obliquely spreading, ovate-oblong. Cuticle smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. concinnula 12. Leaves widely spreading, ovate-orbicular. Cuticle finely punctate-papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Only leaf lobes caducous, swollen lobules remaining on stems and branches. Leaves plane, (sub)orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. galeata 13. Leaves fully caducous, lobules not remaining on stems and branches. Leaves convex, ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cochleata 14. Cells with evenly thickened walls, trigones indistinct. Leaf apex plane to recurved. Underleaves limited to the base of the male spike. Plants montane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. deplanata 14. Cell walls thin, trigones minute. Leaf apex plane. Underleaves present throughout the male spike. Plants of Amazonian lowlands (to 800 m) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. rionegrensis 15. Underleaf lobes on main stems finely acuminate, ending in a row of (1–)2–4 cells. Cuticle densely papillose . . . . . . . . . . . . . . . . . . . . . . . . . L. controversa 15. Underleaf lobes on main stems rounded to acute, ending in a row of 1–2 cells. Cuticle smooth or papillose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Underleaves 3–5(−6)× stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Underleaves 1–3× stem width [when in doubt try both leads] . . . . . . . . . . . 29 17. Underleaf bases strongly auriculate. Cuticle densely papillose. . . . . . . . . . . 18 17. Underleaf bases not or slightly auriculate. Cuticle smooth or faintly papillose (densely papillose in L. anomala, couplet 21). . . . . . . . . . . . . . . . . . . . . . . . 19 18. Underleaves suborbicular, bifid to 1/3–1/2, apex plane. Leaves with a border of 5–6 rows of smaller marginal cells, perpendicular to the larger central cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pulverulenta 18. Underleaves reniform, shortly bifid to 1/5(−1/4), apex recurved. Leaves without border of 5–6 rows of smaller marginal cells. Bolivia and Peru; not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . L. recurva M.E.Reiner 19. Leaf cells with distinct trigones. Leaf margins entire . . . . . . . . . . . . . . . . . . 20

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Keys and Descriptions

19. Leaf cells without or with minute trigones. Leaf margins entire or crenulate [when in doubt try both leads] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 20. Underleaves wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20. Underleaves not wider than long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 21. Cuticle smooth. Underleaves with a large cell at either side of the base, where the underleaf margin joins the stem. Oil bodies homogeneous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. puiggariana 21. Cuticle densely papillose. Underleaves without large cell at the base. Oil bodies finely granular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. anomala 22. Perianth terete. Leaves convex. Apical margin of lobule consisting of 3 large cells, larger than inner lobule cells. Underleaf bases cuneate to rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. capensis 22. Perianth 5-keeled. Leaves plane. Apical margin of lobule not with 3 large cells. Underleaf bases always rounded, cordate. Gynoecia on short branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. flava 23. Well-developed lobules maximally 1/4× leaf length. Plants growing from sea level to 2200 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. Well-developed lobules larger, 1/3–1/2× leaf length. Plants of high elevations, 1200–4000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 24. Gynoecia on very short branches without vegetative leaves. Perianth keels crenulate. Plants in lowland and montane rainforest, below 2000 m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. subsessilis 24. Gynoecia on elongate branches with vegetative leaves. Perianth keels smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25. Autoicous, plants superfertile. Plants ca. 2 mm wide, leaves distinctly longer than wide, oblong. Lobules reduced, tooth indistinct. Underleaves with a narrow incision, tips of the lobes often touching each other, bases subauriculate. Innovations usually fertile . . . . . . . . . L. flava (robust, hygrophytic phenotype) 25. Diocous, not superfertile. Plants smaller, 1–1.5  mm wide, leaves as wide as long, broadly ovate. Lobules 1/5–1/4× leaf length, with a long tooth (2–3 cells long). Underleaves with a wide incision, tips not touching each other, bases cuneate to rounded. Innovations usually sterile. Plants only known from the type from Ecuador, ca. 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . L. suffruticola 26. Leaf cells ± evenly thick-walled. [Plants to 1.5  mm wide, dioicous; leaves widely spreading, ovate; underleaves suborbicular] . . . . . . . . . . . L. deplanata 26. Leaf cells thin-walled [when in doubt try both leads] . . . . . . . . . . . . . . . . . . 27 27. Plants 1–2.5 mm wide, common, high-Andean. Underleaves imbricate, undivided or shallowly bifid to 1/5. . . . . . . . . . . . . . . . . . . . . . . . . . L. catinulifera 27. Plants very small, 0.5–0.8 mm wide, rare, high-Andean. Underleaves imbricate or distant, bifid to 1/3–1/2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28. Underleaves imbricate, ± as long as wide. Leaves widely spreading, suborbicular, somewhat squarrose. Lobule tooth long, made up of one large, oblong, ca. 40 μm long cell. Perianth sharply 5-keeled. Plants of páramo (Colombia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. schultesii

Lejeuneaceae

511

28. Underleaves distant, longer than wide. Leaves obliquely spreading, ovateoblong, not squarrose. Lobule tooth short. Perianth terete, immersed. Plants of mesic Polylepis forest (Ecuador) . . . . . . . . . . . . . . . . . . . . . . . . . . L. mojandae 29. Well-developed lobules up to 1/3–2/3× leaf length. . . . . . . . . . . . . . . . . . . . 30 29. Well-developed lobules smaller, maximally 1/4× leaf length . . . . . . . . . . . . 36 30. Cuticle finely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 30. Cuticle ± smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 31. Leaves obliquely spreading, ovate-oblong . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 31. Leaves widely spreading, ovate [when in doubt try both leads] . . . . . . . . . . 33 32. Underleaves as long as wide or longer than wide, frequently with a tooth on outer margins. Shoots fragile, easily fragmenting. Plants dioicous (often sterile), very common, xerotolerant, from sea level to ca. 2000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. laetevirens 32. Underleaves ± wider than long, without tooth on outer margins. Shoots not fragmenting. Plants autoicous (?), rare, in montane forest . . . . . . L. angusta 33. Perianth keels smooth. Tip cells of underleaf lobes often broken. Leaf margins entire or sligthly crenulate. Leaves not falcate . . . . . . . . . . . . . . . . L. aphanes 33. Perianth keels ciliate-laciniate. Tip cells of underleaf lobes not broken. Leaf margins crenate. Leaves falcate. . . . . . . . . . . . . . . . . . . . . . . . . . . . L. intricata 34. Lobules of stem leaves large, 1/2× leaf length. Leaves suberect, plants Microlejeunea-like. Leaf margins and keel strongly crenate. Plants from Amazonian forest of Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . L. yasuniensis 34. Lobules of stem leaves smaller, 1/3–2/5× leaf length. Leaves obliquely to widely spreading, plants not Microlejeunea-like. Leaf margins and keel entire or slightly crenulate. Plants more widespread . . . . . . . . . . . . . . . . . . . . . . . . 35 35. Tip cells of underleaf lobes often broken or collapsed. Perianth keels smooth or slightly crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. aphanes 35. Tip cells of underleaf lobes never broken. Perianth keels ciliate-laciniate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. acanthogona 36. Underleaves bifid to 2/3 of underleaf length, lobes triangular to lanceolate. Leaf cells with intermediate thickenings and trigones . . . . . . . . . . . . . . . L. adpressa 36. Underleaves bifid to maximally 1/2, lobes triangular. Leaf cells with or without trigones and intermediate thickenings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 37. Underleaf margins conspicuously toothed by obtuse elongate cells, tooth apex elevated away from the underleaf plane. Leaf margins entire. Plants of Panama (Barro Colorado Island, on rock), not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . L. tamasii M.E.Reiner, N.Salazar & C.Chung 37. Underleaf margins not toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38. Perianth keels ciliate-laciniate. [Leaf margins entire]. . . . . . . L. acanthogona 38. Perianth keels smooth, crenate or with lamella-like projections, not ciliatelaciniate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39. Gynoecia on elongate branches with bracts and vegetative leaves. Leaf margins entire. Perianth keels smooth or slightly toothed. [Leaf apex rounded to obtuse, flat, or recurved.] Plants in or near rivers . . . . . . . . . . . . . . . . . . . . . . . L. laeta

512

 

Keys and Descriptions

39. Gynoecia mostly on very short branches with bracts only (no vegetative leaves). Leaf margins and perianth keels crenulate or crenate. Plants not in rivers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 40. Leaf and underleaf margins strongly crenate. Leaf apex rounded to narrowly obtuse. Perianth keels at apex with lamella-like projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. quinqueumbonata 40. Leaf and underleaf margins crenulate only, not strongly crenate. Leaf apex always broadly rounded. Perianth keels without lamella-like projections . . . 41 41. Underleaves as long as wide or longer than wide, 1.5–2× stem width. Perianth keels conspicuously crenate. Plants on dead wood . . . . . . . . . . L. glaucescens 41. Underleaves wider than long, more than 2× stem width. Perianth keels crenulate. Plants on living leaves and bark. . . . . . . . . . . . . . . . . . . . . . L. subsessilis 1. Lejeunea acanthogona Spruce Figs. 51D, 52K Harpalejeunea acanthogona (Spruce) Steph. Type: Ecuador, Mt. Tunguragua, 3000 m, “supra truncos putrescentes,” Spruce L528 (MANCH-000001!) Crossotolejeunea bogotensis Steph. Crossotolejeunea cristatella Steph. Crossotolejeunea cristulata Steph. (fide Bastos and Gradstein 2020) Crossotolejeunea cristuliflora Steph. (fide Bastos and Gradstein 2020) Crossotolejeunea grossiretis Steph. (fide Bastos and Gradstein 2020) Lejeunea blepharogona Spruce, syn. nov. Type: Ecuador, Mt. Azuay, 3000 m, “in arborum ramulis,” Spruce s.n. (MANCH-000003!) On rotten wood, twigs, and living leaves in montane rainforest, 2000–3500 m; found at lower elevation in southeastern Brazil and Panama (Reiner-Drehwald and Goda 2000; Schäfer-Verwimp 2014). Colombia: Cas (Gradstein 12705, 12706 p.p., PC), Cun, Ris (Gradstein 3576, U). Ecuador: Az, Ca, Ch (Herzog 1952 as L. blepharogona), Lo (Schäfer-Verwimp et  al. 2013a as L. cristuliflora), Tu (type), ZC (Parolly et al. 2004 as L. grossitexta Steph.). General distribution: southeastern Brazil and tropical Andes northward to Panama and Costa Rica. Lejeunea acanthogona is a rather widespread and variable neotropical taxon that is characterized by (1) plants autoicous, small, 0.5–1.5  mm wide; (2) leaf lobes widely spreading, ovate, plane, apex rounded to obtuse to subacute to apiculate, margins entire or crenulate, bordered by small quadrate cells; (3) leaf cells without or with small trigones, intermediate thickenings lacking or few, cuticle smooth; (4) lobules frequently reduced, when well developed 1/4–1/2× leaf length, usually fully inflated and with involute free margin, tooth ± elongate, obtuse, keel arched, angle between keel and ventral margin of leaf lobe usually sharp; (5) underleaves small, distant, bifid 1/3–2/5 with a V-shaped incision, outer margins arched, entire, lobes broadly triangular, obtuse, bases narrow, cuneate, insertion line straight; and (6) gynoecia on a long shoot with a sterile innovation, perianth keels 2-winged, ciliatelaciniate or with a few cilia only, beak 2–6 cells long. The small plants with rounded to obtuse to apiculate leaf tips, inflated lobules (but lobules often reduced), small underleaves, smooth cuticle, and ciliate-laciniate perianths with a 2–6-cell-long beak are the main features of L. acanthogona.

Lejeuneaceae

513

Considerable variation is seen in the shape of the leaf tip (rounded to apiculate), size and shape of lobules, perianth ornamentation, and length of perianth beak, and several species were described based on these features by Stephani (1896, 1898–1924; see also Reiner-Drehwald and Goda 2000). Most of these species are now considered synonyms of L. acanthogona. Lejeunea acanthogona is closely related to L. intricata, but the latter species has a finely papillose cuticle and crenate leaf margins. Lejeunea acanthogona is also related to the widespread paleotropical L. pulchriflora (Pearson) G.E.  Lee et  al. (2016), but the latter species produces elongate, ribbon-like gemmae on leaf margins. Such gemmae have not yet been observed in neotropical Lejeunea species. 2. Lejeunea adpressa Nees Fig. 51E Lejeunea anisophylla Mont., syn. nov. Type: Hawaii, “ad cortices inter muscos in insulis Sandwich,” Gaudichaud s.n. (PC-Mont.-MB1679!) Lejeunea caespitosa auct. (non typus; e.g., Schuster 1980a; Wigginton 2004) Lejeunea carolensis Spruce Lejeunea cauapunensis Spruce Lejeunea drymophila Spruce Lejeunea longifissa Spruce Lejeunea magnoliae Lindenb. & Gottsche Lejeunea orbicularis Spruce On bark and living leaves in dry and moist lowland and lower montane forests, often in open and disturbed vegetation, in plantations, gardens, on isolated trees, etc., from sea level to 2100 m. Colombia: Cho, Hui, Mag, Put. Ecuador: Es, MS, Or, Pa, Pi, Su, Tu, ZC. General distribution: pantropical; widespread in tropical and subtropical America. Lejeunea adpressa is a common and widespread pantropical species that has been described under many different names (Reiner-Drehwald 2009a). The species is recognized by (1) plants autoicous, small, 0.7–1.1 mm wide, light green to yellowish-green; (2) leaf lobes widely spreading, ovate, apex rounded, margins entire; (3) leaf cells with conspicuous trigones and intermediate thickenings, cuticle smooth, oil bodies finely segmented, ca. 3–12 per cell; (4) lobules small, ovoid, ca. 1/4× leaf length, strongly inflated, free margin incurved, tooth short, obtuse; (5) underleaves distant, small, deeply bifid (to 2/3) with a broad incision, lobes narrowly lanceolate, acute, margins sometimes with a short, blunt tooth; and (6) gynoecia on short or long branches with innovation, perianth keels smooth, lateral keels somewhat expanded above. Lejeunea adpressa sometimes grows together with L. laeta, and these two species may be confused; for differences see under L. laeta. Lejeunea minutiloba A.Evans, a widespread Caribbean species not known from Colombia and Ecuador, has sometimes been confused with L. adpressa but differs in fully thin-walled leaf cells, without any trigones and intermediate thickenings, homogenous oil bodies, and less deeply bifid underleaves (maximally to 1/2). Heinrichs et al. (2013a) found that a specimen named L. minutiloba was nested in L. adpressa in a molecular analysis. The identification of the specimen should be checked.

514

 

Keys and Descriptions

In tropical Asia and Africa, L. adpressa has been called L. anisophylla. Several authors have noted the close similarity between these two species (Zhu and So 2001; Reiner-Drehwald 2009a; Pócs 2010), and the only difference found to separate them has been the number of oil bodies per cell (6–20 in L. anisophylla, 3–7 in L. adpressa). The oil body data of L. adpressa were based on one specimen only, however (Schuster 1980a; as L. caespitosa). Recently, I have examined oil bodies in fresh material of L. adpressa collected near Santa Marta in several localities (Gradstein 12482, 12509, PC) and found 3–12 oil bodies per leaf cell. This indicates that the oil body characters of the two species overlap and that the two are morphologically identical. The conspecificity of L. anisophylla and L. adpressa has been confirmed by molecular analysis (R.L. Zhu, pers. comm.). 3. Lejeunea angusta (Lehm. & Lindenb.) Mont. Jungermannia angusta Lehm. & Lindenb. Type: Mexico, Xalapa, without collector (isotype PC-Mont.-2118!, ster.) On bark of trees in montane rainforest, 1400–2900  m. Colombia: Boy, Cun. Ecuador: Pi, Tu, ZC (Schäfer-Verwimp et  al. 2013a). General distribution: scattered in tropical America. Lejeunea angusta is little-known neotropical species that is very similar to L. laetevirens. Both have obliquely spreading, ovate-oblong leaves; well-developed, swollen lobules; a densely punctate-papillose cuticle, and small bifid underleaves. However, the stems of L. angusta are not fragmenting like in L. laetevirens, and the underleaves are ± wider than long and never produce a tooth on the outer margins. Moreover, the species is presumably autoicous (Ilkiu-Borges and Oliveira-da-Silva 2018). Gradstein et  al. (2018c) reported the occurrence of caducous leaves in L. angusta, but the the latter observation was erroneous; the plants with caducous leaves belonged to L. cochleata (which is not a synonym of L. angusta as proposed by the latter authors). The published records of L. angusta from the Sierra Nevada de Santa Marta and Mt. Chimborazo belong to L. cochleata. 4. Lejeunea anomala Lindenb. & Gottsche Ceratolejeunea anomala (Lindenb. & Gottsche) Steph. Lejeunea matteola Spruce On bark and wet rock in lowland and submontane rainforest, up to 1000  m. Ecuador: Pa (Dauphin 2003). General distribution: Suriname, Ecuador, Brazil. Lejeunea anomala is a rare and little-known neotropical species that was associated with Ceratolejeunea in the past because of its brownish color. Reiner-Drehwald (2003) and Dauphin (2003) confirmed that it is a genuine member of Lejeunea. Based on the description by the latter author and our observations (Bastos and Gradstein 2020), the species is briefly characterized as follows (1) plants dioicous, brownish-green, 1–1.5  mm wide; (2) leaf lobes widely spreading, symmetrically ovate-oblong, ca. 1.5× longer than wide, apex broadly rounded, plane, margins entire; (3) leaf cells with small but distinct trigones and intermediate thickenings, cuticle sligthly and finely papillose; (4) lobules mostly reduced, less than 1/5× leaf length, free margin plane, tooth short, obtuse; (5) underleaves wider than long, ca.

Lejeuneaceae

515

3–5× stem width, not imbricate, bifid to about 1/4, lobes acute, margins entire, bases rounded (not auriculate), insertion line curved; and (6) gynoecia variable, without or with 1–2 innovations, sometimes in a short row of two gynoecia, bracts and bracteole with obtuse to acute to acuminate tips, perianth 5-keeled, keels entire, crenulate or weakly toothed. Lejeunea anomala may be confused with phenotypes of L. cerina with a poorly developed apiculus and small underleaves (e.g., Gradstein and Ilkiu-Borges 2009, Fig. 59), but the latter species differs in having somewhat pointed leaves (apex not fully rounded), crenulate leaf margins, and gynoecia without innovation or with a short, sterile innovation. Lejeunea anomala approaches L. pulverulenta in the darkish plant color, leaves with a broadly rounded apex, papillose cuticle, rather large and wider than long underleaves with a cordate base, and gynoecia sometimes two in a row. The latter species differs from L. anomala, however, in autoicous sexuality, crenulate leaf margins, and underleaves with large auricles. Moreover, the leaf cells in L. pulverulenta become conspicuously smaller toward to the leaf margin, forming a border of 5–6 rows of smaller cells. Lejeunea anomala seems strikingly similar to L. sordida (Nees) Nees, a species common in tropical Asia, but the tips of the female bracts in the latter species are obtuse. The relationship of L. anomala and L. sordida deserves further study. 5. Lejeunea aphanes Spruce Lejeunea autoica R.M.Schust. Lejeunea filipes Spruce

Fig. 51F

On smooth bark of trees and on living leaves in primary and secondary lowland and montane rainforests up to the forest line, from sea level to about 3800  m. Colombia (Gradstein et al. 2019a): Met, Qui. Ecuador: Ch, Co, EO, Ga (?), MS, Or, Pi, Tu, ZC. General distribution: tropical America, Africa. Lejeunea aphanes a widespread Afro-American species that is recognized by (1) plants autoicous, small, 0.4–1.0 mm wide, pale green to yellowish-green; (2) leaf lobes obliquely to widely spreading, ovate, convex, apex rounded to obtuse, margins entire; (3) leaf cells with small trigones and few intermediate thickenings, cuticle smooth or slightly punctate-papillose, oil bodies finely granular, few per cell (2–4); (4) lobules often reduced, large when well developed and strongly inflated, 1/3–2/5× leaf length, ovoid, 1–1.5× longer than wide, free margin incurved or flat, tooth obtuse, slightly curved outwards; (5) underleaves very small, 1–2× stem width, usually appressed to the stem, as long as wide or slightly longer than wide, bifid to 2/5–1/2, lobes subacute, tip cells of underleaf lobes often broken or collapsed, outer margins curved, entire, without tooth, bases cuneate, insertion line weakly curved; (6) male bracteoles restricted to the base of the male shoot; (7) gynoecia on short branches with 1 innovation, perianths with 3–5 ± smooth keels; and (8) vegetative reproduction lacking (Schuster 1980a as L. autoica; Reiner-Drehwald 2000, 2009b as L. filipes). By the small plants with somewhat convex, obliquely to widely spreading leaves with rounded apex and entire to crenulate margins; large, swollen, ovoid lobules (1–1.5× longer than wide, but sometimes lobules reduced); and very small

516

 

Keys and Descriptions

underleaves (1–2× stem width) with often broken or collapsed tip cells, L. aphanes is unmistakable. The underleaves in plants from high elevation (above 2000 m) are slightly larger than described by Schuster and Reiner-Drehwald and are not closely appressed to the stem. Small underleaves with broken or collapsed tip cells are also seen in L. yasuniensis, but the latter species differs in having suberect leaves with larger lobules, ca. 1/2× lobe length, and strongly crenate margins. 6. Lejeunea asperrima Spruce Echinocolea asperrima (Spruce) R.M.Schust.

Fig. 51C

On bark, living leaves, and rotten wood in undisturbed lowland and submontane rainforests, in very humid and shaded environments, from sea level to 1500  m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). Ecuador: MS, Na, Or, ZC. General distribution: tropical America. Lejeunea asperrima (sect. Echinocolea) is a minute neotropical rainforest species that is readily distinguished by the keel, margins, and dorsal surface of leaf lobes strongly crenate due to conically projecting cells crowned by a papilla. The plants are dioicous, 0.2–0.5  mm wide; the leaf lobes are suberect to obliquely spreading with a very narrow base and rounded apex; the leaf cells are quite thinwalled; the lobules are ca. 1/2× leaf length and fully inflated, with a falcate tooth and semicircular mouth; and the underleaves are very small, 1–2× stem width, deeply bifid to 1/2–2/3, with acute-acuminate lobes and entire or crenulate margins. The perianth has 5 crenate keels, and vegetative reproduction is by cladia, leaf fragmentation, and linear gemmae (Ilkiu-Borges 2005). By the dorsal leaf surface and margins roughened by conically projecting cells, L. asperrima is similar to L. meridensis and L. subspathulata; for differences see the key. 7. Lejeunea asthenica Spruce Taxilejeunea asthenica (Spruce) Steph. On bark, soil, and living leaves in rather open habitats, to ca. 1800 m; on the Galápagos Islands (Santa Cruz) occurring on fern fronds in moist woodland and scrub near the forest line, 500–600  m. Ecuador: Ga, MS, Or, Su, ZC (SchäferVerwimp et al. 2013a; Mota de Oliveira and ter Steege 2013). General distribution: scattered in tropical America. Lejeunea asthenica is a little-known neotropical species that is characterized by (1) plants autoicous and usually fertile, loosely creeping to pendent; (2) leaf lobes broadly ovate-triangular, apex sharply acute to apiculate to shortly acuminate, margins crenulate, entire or with 1(−3) small teeth near apex; (3) leaf cells with distinct trigones and intermediate thickenings, cuticle densely punctate-papillose; (4) lobules small, ovate, to 1/7 of leaf length, often reduced; (5) underleaves distant, orbicular or slightly wider than long, 2–3× stem width, bifid to 1/3–1/2, bases cuneate or rounded, basal margin cell large; and (6) gynoecia 1–2 on long branches, bracts toothed, perianth 5-keeled in the upper 1/3, keels crenulate.

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Lejeunea asthenica is morphologically close to L. flaccida, but in the latter species, the leaf tips are rounded to obtuse to apiculate, the underleaves are larger (3–5× stem width), the trigones are smaller, and the cuticle is ± smooth. 8. Lejeunea bermudiana (A.Evans) R.M.Schust. In Colombia on living leaves in secondary humid montane forest, 2040 m; elsewhere on tree bases, rotten wood, and rock from sea level to 2000 m. Colombia: Mag (San Lorenzo, Gradstein et al. 12568, “cf.”, CBUMAG). General distribution: southeastern USA, West Indies, Panama, Brazil, Colombia. Lejeunea bermudiana is a Caribbean species characterized by (1) plants autoicous ca. 1.3  mm wide; (2) leaf lobes obliquely to widely spreading, slightly convex, ovate, apex obtuse to apiculate, terminating in 1–2 cells in a row, margins crenulate, ventral leaf margin usually with 1–3 small teeth near apex; (3) leaf cells transparent, very thin-walled, trigones very small, intermediate thickenings absent, cuticle smooth; (4) lobules well developed, ovate-trapezoid, 1/4(−1/3)× leaf length, inflated-swollen, free margin plane, with 1(−2) teeth, keel smooth; (5) underleaves distant, 2–3.5× stem width, bifid to 1/2, apices subacute, margins crenulate, bases rounded-cordate; and (6) gynoecia on short or long branches, with 1(−2) innovations, perianth ellipsoid, longly 5-keeled, keels irregularly 2-winged, toothed, beak short. Lejeunea bermudiana stands out by the obtuse to apiculate leaf apex, leaf margins with 1–3 small teeth near apex, well-developed lobules, small underleaves, and perianth keels with a few short teeth. The Colombian specimen of L. bermudiana is a relatively robust, epiphyllous plant with unusually large lobules (ca. 1/3 of leaf length) and large underleaves, 3.5× stem width. The material is scanty and the identification is tentative. 9. Lejeunea bombonasensis Spruce Fig. 51G Taxilejeunea bombonasensis (Spruce) Steph. Type: Ecuador, Río Bombanasa, Spruce L213, autoicous (lectotype, designated here, MANCH-cc18215!; isolectotypes MANCH-cc18214!, MANCH-cc18216!) Taxilejeunea crebriflora (Spruce) Steph. Taxilejeunea implexa (Spruce) Steph. Taxilejeunea lusoria auct. (non typus) Taxilejeunea tenax (Spruce) Steph. On bark, rock, and living leaves at rather low elevations in disturbed rainforest and plantations, from sea level to ca. 2000 m. Colombia: Ant, Cau, Cho, Hui, Mag, San, Tol. Ecuador: Es, MS, Na, Or, Pa, Pi, Su, Tu. General distribution: widespread in tropical and subtropical America. Lejeunea bombonasensis is the most common neotropical member of the former genus Taxilejeunea with small underleaves. The species has generally been called Taxilejeunea lusoria, but the type of the latter species belongs to L. flaccida. Characteristic features of L. bombonasensis are (1) plants autoicous and frequently fertile, loosely creeping to pendent; (2) leaf lobes asymmetrically ovate-triangular, apex variable, rounded to acute to short-apiculate, margins entire or with a few teeth

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near apex; (3) leaf cells with minute trigones and without intermediate thickenings, cuticle smooth, oil bodies finely segmented, 4–10 per cell (Gradstein et al. 1977); (4) lobules usually reduced; (5) underleaves distant, 2–4× stem width, orbicular, bifid to 1/3–1/2, bases cuneate to rounded, insertion line slightly curved; (6) gynoecia terminal on the stem or up to 6 in a row on innovating branches, bracts entire or slightly toothed; and (7) perianths sharply and longly 5-keeled over (1/3–)1/2–2/3 of perianth length, keels smooth or crenulate, often widened in the upper third (Gradstein and Costa 2003). Lejeunea bombonasensis is morphologically close to L. flaccida and L. asthenica, but L. asthenica has sharply acute-acuminate leaf tips and L. flaccida has larger underleaves (3–5× stem width) and very short perianth keels. Lejeunea bombonasensis also resembles L. flagellaris, but the latter species is dioicous and usually sterile and has numerous microphyllous branches. 10. Lejeunea boryana Mont. Fig. 51H, I Lejeunea viridis L.Söderstr. & A.Hagborg (Prionocolea viridissima R.M.Schust., nom. inval.), syn. nov. Type: Colombia, Cauca, Isla Gorgona, Mirador, Schuster 88-1621b, with Cystolejeunea lineata (holotype F-CO172148F!) On trunk bases and rotten wood in lowland and submontane rainforests, 100–650 m. Colombia: Ama, Caq, Cau, Cho, Put, Vau (Pinzón et al. 2003; Campos et  al. 2015). Ecuador: MS, Or (Mota de Oliveira and ter Steege 2013; SchäferVerwimp et  al. 2013a). General distribution: rather widespread in tropical America. Lejeunea boryana is a characteristic neotropical lowland species that is distinguished by (1) plants autoicous, to 1 mm wide, green; (2) leaf lobes ovate-falcate, slightly convex, apex obtuse to apiculate, margins usually crenate; (3) leaf cells with trigones, with or without intermediate thickenings, cuticle finely and densely papillose, oil bodies finely segmented, 2–8 per cell, globose to narrowly elongate (Schuster 1992c as Prionocolea viridissima); (4) lobules when well developed ca. 1/3× leaf length, ovate, fully inflated, tooth straight to falcate; (5) underleaves distant to contiguous, 2–3× stem width, deeply bifid to 2/3, lobes acute-acuminate, terminating in a row of (1–)2 cells, outer margins with a tooth (1–3 long), bases cuneate; and (6) gynoecia on short or long shoots, with innovations, perianth 4–5-keeled, the keels crenate to dentate to ciliate-laciniate and slightly expanded above, beak short (Reiner-Drehwald and Goda 2000; Reiner-Drehwald 2009b). By the densely papillose cuticle, apiculate leaves with crenate margins, and small, deeply bifid underleaves with a tooth on the outer margins, L. boryana is readily separated from other Lejeunea species. The perianth is 4–5-keeled, with or without dorsal keel. A phenotype of L. boryana with reduced dorsal perianth keel has been described as Prionocolea viridissima R.M.Schust. 11. Lejeunea cancellata Nees & Mont. Lejeunea cladiophora (R.M.Schust.) R.M.Schust.

Fig. 51J, K

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On bark in primary and secondary lowland and montane rainforests, plantations, gardens, etc. from sea level to 2800  m. Colombia: Boy (Uribe 3661, 3722, det. M.  E. Reiner-Drehwald, COL), Cho, Ris (Gradstein 8631, Wolf 197, GOET). Ecuador: Az, EO, MS, Pi, ZC. General distribution: scattered throughout tropical America. Lejeunea cancellata is an uncommon neotropical species that is recognized by (1) plants dioicous, glossy pale green, often densely branched, branches caducous; (2) leaves obliquely spreading, somewhat convex, apex rounded to apiculate, margins crenulate; (3) leaf cells fully thin-walled, cuticle smooth, oilbodies homogenous, very small and numerous, Massula-type; (4) lobules ca. 1/4× leaf length, inflated, keel crenulate; (5) underleaves small, distant, 3–4× stem width, slightly wider than long, bifid to 1/3–1/2, with obtuse lobes and rounded bases (Schuster 1980a as L. cladiophora; Reiner-Drehwald 2000). Caducous branches are very characteristic of L. cancellata. They are also seen in L. laetevirens and L. osculatiana, but L. osculatiana is readily separated from L. cancellata by the slender and usually pendent stems with lanceolate-acuminate leaves, while L. laetevirens is a smaller plant with ovate-oblong leaves; larger lobules; smaller, deeply bifid underleaves; finely papillose cuticle; and segmented oil bodies. Moreover, in L. laetevirens all shoots are fragile, also the main stems. In a molecular analysis, L. cancellata was sister to L. cerina (Heinrichs et al. 2013a), but the latter species lacks caducous branches and has wide-spreading, apiculate leaves, larger underleaves, a densely papillose cuticle, granular oil bodies, and gynoecia frequently without innovation. 12. Lejeunea capensis Gottsche Fig. 51L Lejeunea caespitosa Lindenb. Taxilejeunea gomphocalyx Herzog, syn. nov. Type: Colombia, Cundinamarca, Boqueron near Bogotá, on treelets, 3200 m, C. Troll 2031 (JE n.v.). Taxilejeunea lindbergiana Steph. (fide Bastos and Gradstein 2020) On bark of trees in rather open, montane forests, scrub, and plantations up to the páramo, 1800–3900 m. Colombia: Boy (Gradstein 2019a), Mag (?; Winkler 1976 as Taxilejeunea lindbergiana). Ecuador: Co, Lo, Pi (laguna Mojanda, Polylepis forest, 3800 m, León-Yánez s.n., QCA), ZC. General distribution: scattered in tropical America, common in Africa. Lejeunea capensis is a widespread Afro-American species characterized by (1) plants autoicous and usually fertile, small, 0.6–1  mm wide, yellowish-green; (2) leaf lobes widely spreading, convex, ovate to ovate-oblong, apex rounded to narrowly obtuse; (3) leaf cells with small trigones and few intermediate thickenings, cuticle finely punctate-papillose; (4) lobules when well developed 1/3–2/5× lobe length, inflated, tooth very short, obtuse, apical margin of 3 large cells (larger than other lobule cells); (5) underleaves distant to imbricate, rather large, 2.5–5× stem width, suborbicular, bifid to 1/3–1/2, lobes obtuse to acute, margins entire, bases usually cuneate, becoming rounded in large underleaves; and (6) gynoecia on long shoots with 1–2 innovations, perianths obovoid, terete, without keels, exserted

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Keys and Descriptions

beyond the bracts to less than 1/2 its length, beak narrow (Reiner-Drehwald and Schäfer-Verwimp 2008a). Lejeunea capensis resembles L. flava but differs from the latter by eplicate perianths on long shoots (perianths 5-keeled and on short branches in L. flava), convex leaves, lobules with 3 large cells along the apical margin, and underleaves with narrow, cuneate to slightly rounded bases (underleaves with distinctly rounded or auriculate bases in L. flava). In its eplicate perianth, L. capensis approaches L. corynantha, but the latter species is a more delicate plant; the perianth in L. corynantha is much more longly exserted and lacks a beak, the lobules are larger (ca. 1/2 × leaf length), and the underleaves are smaller, ca. 2× stem width. Based on the original description and illustration, Taxilejeunea gomphocalyx is a synonym of L. capensis. Plants from high elevation in the Sierra Nevada del Cocuy (3900 m) stand out by large, rounded to elongate underleaves, ca. 5× stem width, with rounded bases. 13. Lejeunea caripensis Lindenb. & Gottsche Taxilejeunea caripensis [Lindenb. & Gottsche] Steph.). Type: Venezuela (“Colombia”), Prov. Monagas, Caripe, Moritz s.n. (lectotype (Bastos and Gradstein 2020), W-Lindenb. 6385!, ster.) On bark in moist montane forests and scrub, 700–2600 m. Ecuador: Ca (HolmNielsen et al. 6233, AAU, PC), Na (Holm-Nielsen 26514, AAU, PC), MS, Pa (HolmNielsen et al. 464, AAU, PC), Tu (Holm-Nielsen 16304, AAU, PC); to be expected from Colombia. General distribution: recorded from Venezuela (type), French Guiana (Gradstein and Ilkiu-Borges 2009, as “Taxilejeunea sp.”), Ecuador, and E Brazil (Bastos & Gradstein 2020); the species is probably more widespread in tropical America. Lejeunea caripensis is a little-known species that is recognized by (1) plants dioicous and often sterile, 1–1.3 mm wide, stems long and slender, little branched or with several small-leaved branches; (2) leaf lobes obliquely spreading, asymmetrically ovate-triangular, tapering to an obtuse to apiculate apex ending in a row of 1–3 cells, margins crenulate, sometimes with a small teeth near apex; (3) leaf cells with conspicuous trigones and intermediate thickenings, cuticle finely and densely papillose; (4) lobules ca. 1/6× leaf length, inflated, short bottle-shaped, keel crenate, at an oblique angle with the axis and forming a sharp angle with the ventral lobe margin, apex with a short, blunt tooth; (5) underleaves very large, 5–10× stem width, contiguous to imbricate, wider than long, broadly ovate to reniform, shallowly bifid to 1/4 with a very narrow incision, margin crenulate, insertion line deeply arched, ca. 0.2 mm high, bases not distinctly auriculate; and (6) gynoecia in rows of 2–3 on branches and main stems, perianths 5-keeled over most of their length, keels crenulate, beak short. Lejeunea caripensis has been confused with L. cerina, (Bastos and Gradstein 2020) but the leaves in the latter species are more symmetrical with a broad apex ending suddenly in a short apiculus (not tapering); the underleaves in L. cerina are smaller and much more deeply bifid (to 1/3–1/2) with a broad, V-shaped incision; the insertion line of the underleaves is shallowly curved (to 40 μm high; not deeply

Lejeuneaceae

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arched) and the bases more or less auriculate; and the gynoecia are not produced in a row. 14. Lejeunea catinulifera Spruce Amphilejeunea catinulifera (Spruce) R.M.Schust. Amphilejeunea patellifera (Spruce) R.M.Schust. Amphilejeunea viridissima R.M.Schust. Taxilejeunea obtusifolia Steph. (fide Bastos and Gradstein 2020)

Fig. 51M

Sun epiphyte on tree trunks and branches in upper montane forests, scrub, and páramo, also on rock, 1800–4350 m. Colombia: Ant, Ces, Cun, Mag, Qui, Ris, San, Tol. Ecuador: Az, Lo, Na, Pi, Tu, ZC. General distribution: Mexico to Bolivia (new to Peru: Prov. Urubamba, Huayllabamba, laguna Quellacocha, 4327  m, Sylvester s.n., PC), southeastern Brazil (Serro do Itatiaia), Dominican Republic. Lejeunea catinulifera is a common high-Andean species that is readily distinguished by the large, orbicular to reniform, undivided to shortly bifid (to max. 1/5) underleaves, which are ca. 3–6× wider than the stem and with cuneate to rounded bases, and the usually well-developed lobules (1/3–1/2× leaf length) with an inflexed free margin and (1-)2 teeth separated by a distinct sinus. The plants are dioicous, glossy green when fresh (pale brownish when dry), 1–2.5 mm wide; the leaves are convex, ovate, with a rounded and somewhat recurved apex and entire to crenulate margins; the leaf cells are thin-walled with a smooth to slightly papillose cuticle and numerous small, homogeneous oil bodies (20–40 per cell); the gynoecia are on long shoots with 1–2 innovations; and the perianth is 5-keeled, with smooth keels and a frequently elongate, stalk-like base (Reiner-Drehwald 2005a). Lejeunea catinulifera may be confused with L. reflexistipula; for differences see under the latter species. 15. Lejeunea cerina (Lehm. & Lindenb.) Lehm. & Lindenb. Fig. 51N Macrolejeunea cerina (Lehm. & Lindenb.) Gradst. Lejeunea longiuscula Taylor Lejeunea subsimplex Mont. & Nees (Macrolejeunea subsimplex [Mont. & Nees] Schiffn.) On bark, rock, living leaves, and soil in primary and secondary montane forests and in roadside vegetation, from sea level to 2500 m. Colombia (erroneously listed as doubtful in Gradstein and Uribe 2016a): Cho, Cun, Hui, Mag, Ris, Tol. Ecuador: EO, Lo, MS, Pi, Tu, ZC. General distribution: common in the West Indies, scattered elsewhere in the neotropics; part of the published records of this species may be erroneous. Lejeunea cerina is a variable neotropical species that is recognized by (1) plants dioicous and often sterile, 1–2 mm wide, pale green to brownish, stems long and slender, little branched; (2) leaf lobes widely spreading, concave or flat, broadly ovate and with a broad apex that becomes suddenly obtuse to shortly apiculate, margins usually crenulate, sometimes with one or a few teeth near apex; (3) leaf cells with distinct trigones and intermediate thickenings, cuticle finely and densely papillose, oil bodies small, finely granular; (4) lobules small, ca. 1/6× leaf length (or

522

 

Keys and Descriptions

reduced), tooth short, obtuse; (5) underleaves rather large, 4–6× stem width (but smaller on branches), contiguous to subimbricate, wider than long, broadly ovate, bifid to 1/3–1/2, lobes acute, margins entire, insertion line shallowly curved, to ca. 40 μm high, bases rounded to conspicuously auriculate; and (6) gynoecia on a short branch, without innovation or with a short sterile innovation, perianth 5-keeled over most of its length, keels smooth or crenulate. In the slender habit, wide-spreading ovate leaves with a broadly rounded apex that is rather suddenly narrowed to a short or long apiculus, the densely papillose cuticle, the rather large underleaves with a shallowly curved insertion line and rounded to conspicuosly auriculate bases, and the gynoecia on short branches without or with a short sterile innovation, L. cerina is unmistakable. Branches of L. cerina, however, may have much smaller underleaves than main stems, and branch material of the species cannot be identified with certainty when main stems are lacking. Lejeunea cerina has been confused with L. caripensis (Bastos & Gradstein 2020), but the latter species is readily distinguished from L. cerina by the obliquely spreading, ovate-triangular leaves tapering to a narrow apex; the very large, orbicular underleaves with a shallow incision (to 1/4); a deeply arched insertion line (0.1–0.2 mm high) and lack of auricles; and gynoecia in rows, with fertile innovations. Sterile material of Lejeunea cerina has been confused with L. debilis, but the latter species is readily separated by the much more deeply arched underleaf insertion (0.1–0.2 mm deep). 16. Lejeunea chimborazensis Spruce Taxilejeunea chimborazensis (Spruce) Steph. Taxilejeunea speciosa Herzog (?). Type: Colombia, Cauca, El Valle, Córdoba, Dagua Valley, Killip 11790 (JE n.v.) On fern leaves and rock in lowland and lower montane rainforests, 80–1500 m. Colombia (?): Val (Herzog 1955 as T. speciosa); Cho (Rangel and Gradstein 2004 as T. speciosa). Ecuador: Ch (type). General distribution: Ecuador, western Colombia (?), Bolivia. Lejeunea chimborazensis is a rare, robust Andean species that is morphologically close to L. pterigonia but differs in leaf margins more strongly toothed, with ca. 3–15 teeth in the upper half of the leaf, and more shallowly bifid underleaves, to only about 1/8 of underleaf length (measured from the insertion to the apex). The plants are autoicous, to 6 cm long and 2 mm wide; the leaf tips are acute to apiculate; the leaf cells are fully thin-walled; the lobules are reduced; the underleaves are very large, ca. 5–7× stem width, orbicular or slightly wider than long, with plane to recurved margins and cordate-auriculate bases; the bracts and bracteoles are toothed; and the perianths are in long rows on branches and keeled in the upper part, with toothed keels and a distinct, narrow beak. Taxilejeunea speciosa from Pacific coastal forests of Colombia is very similar to L. chimborazensis and may be a synonym but was described as dioicous (Herzog 1955).

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17. Lejeunea cochleata Spruce Type: Ecuador, Mt. Tungurahua, Spruce L480 (lectotype, designated by Gradstein et al. 2018c, MANCH-000119!; isolectotype MANCH-0000118!) On bark of trees in montane rainforest, ca. 1800 m. Colombia: Mag (Gradstein et al. 2018c as L. angusta). Ecuador: Ch, Tu (Spruce 1884). General distribution: northern Andes (Colombia, Ecuador). Lejeunea cochleata is a little-known northern Andean species that is characterized by (1) plants autoicous, very small, 0.4–0.9 mm wide, with denuded, flagelliform branches bearing underleaves only, due to caducous leaves; (2) leaf lobes wide-spreading, distinctly convex, ovate, apex rounded, margins entire; (3) leaf cells thin-walled or with ± uniformly thickened walls, cuticle finely punctate-papillose; (4) lobules well developed, ca. 1/3× leaf length, ovoid, strongly inflated, with a curved, obtuse tooth; (5) underleaves distant, small, 1.5–2× stem width, bifid to 1/2, lobes triangular, 3–4 cells wide at base, apices obtuse, ± squarrose on flagelliform branches; (6) male bracteoles restricted to the base of the male shoot; (7) gynoecia on short branches with 1 innovation, perianths with smooth keels; and (8) vegetative reproduction by caducous leaves (lobe + lobule caducous). Caducous whole leaves (including lobule), strongly convex leaf lobes, large swollen lobules, finely punctate-papillose cuticle, very small underleaves with obtuse apices, and denuded branches without any trace of the lateral leaves (only with underleaves) are the main characters of L. cochleata. The species resembles L. aphanes, but in the latter species, the apical cells of underleaves are very thinwalled and often broken, and the leaves are not caducous. By its caducous leaves and ± uniformly thickened cell walls, L. cochleata is similar to L. deplanata, but the latter species is dioicous and a larger plant (to 1.5 mm wide); the underleaves in L. deplanata are broader, 2.5–5× stem width; the cuticle is smooth; and only the leaf lobes are caducous (the lobules in L. deplanata are persistent). Gradstein et  al. (2018c) erroneously treated L. cochleata as a synonym of L. angusta. 18. Lejeunea concinnula Spruce & Steph. Microlejeunea concinnula (Spruce & Steph.) Steph. On bark of trees in the understory of humid montane forest, ca. 1900–2500 m. Colombia: Boy, Mag, Qui (Gradstein 12722, 12725, HUQ), Val; to be expected from Ecuador. General distribution: northern Andes (Venezuela, Colombia) and Guayana Highland. Lejeunea concinnula is an uncommon northern Andean species that is characterized by (1) plants dioicous, small, 0.4–0.8 mm wide, little branched; (2) leaf lobes suberect to obliquely spreading, caducous, ovate-oblong, apex rounded or obtuse, margins entire; (3) leaf cells ± uniformly thick-walled, cuticle very finely punctatepapillose (almost smooth), oil bodies 4–8 per cell, rounded, finely segmented; (4) lobules often reduced, when well developed ovoid and strongly swollen, ca. 1/3× lobe length, with a short tooth; (5) underleaves very small, scarcely wider than the stem, deeply bifid, lobes very narrow; and (6) vegetative reproduction by caducous leaf lobes (Gradstein et al. 2018c).

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Keys and Descriptions

Lejeunea concinnula resembles L. laetevirens in the suberect to obliquely spreading ovate-oblong leaf lobes, rather large lobules, and finely punctate-papillose cuticle, but L. laetevirens lacks caducous leaf lobes and grows in dryer environments. 19. Lejeunea controversa Gottsche Crossotolejeunea angulata S.Winkl. Crossotolejeunea lindeniana Steph. Lejeunea paucispina Spruce Lejeunea paucispina var. rufescens Spruce

Fig. 52A

Shade epiphyte on trunk bases, rock, and living leaves in primary and secondary lowland and montane forests, from sea level to 2200 m. Colombia: Mag (Winkler 1976 as Crossotolejeunea angulata). Ecuador: EO, Es, LR, Na, Or, Pa, Pi, ZC. General distribution: tropical America. Lejeunea controversa is a common neotropical species that is distinguished by (1) plants autoicous, ca. 1 mm wide, green; (2) leaf apex rounded to apiculate, margins ± entire; (3) leaf cells with conspicuous trigones and intermediate thickenings, cuticle finely and densely papillose; (4) lobules small, to ca. 1/5× leaf length, with an elongate, straight, or falcate tooth; (5) underleaves distant to subimbricate, (2–)3–4× stem width, bifid to 1/2–2/3, lobes acuminate, ending in a row of (1-)2–4 cells, outer margins entire, bases rounded to subauriculate, insertion line curved; and (6) gynoecia on short or long branches, sometimes in a short row, perianths (4–)5-keeled (dorsal keel smaller), keels toothed to ciliate-laciniate, rarely subentire, often somewhat expanded to auriculate at apex, beak short (Reiner-Drehwald and Goda 2000; Reiner-Drehwald 2009b). The strongly papillose cuticle, underleaves (2–)3–4× stem width and with acuminate tips, and toothed perianth keels are the main characters of Lejeunea controversa. Some variation is seen in these characters, however, and sometimes the uniseriate underleaf tips are only 1–2 cells long and the perianth keels subentire. Lejeunea controversa resembles L. boryana in the papillose cuticle and deeply bifid underleaves, but the latter species differs in the crenate leaf margins and cuneate underleaves with a tooth on the outer margins. 20. Lejeunea corynantha Spruce 

Fig. 52B

On twigs in open environments in upper montane cloud forest areas and páramo, 3000–3700 m. Colombia: Cas (Gradstein et al. 2018c). Ecuador: Az (Gradstein and Benitez 2017). General distribution: West Indies, Panama, northern Andes (Colombia, Ecuador). Lejeunea corynantha is a rare Caribbean-northern Andean species that resembles L. capensis in the terete perianth but differs from the latter in the larger lobules (1/2× lobe length), smaller underleaves (ca. 2× stem width), strongly convex leaf lobes, ± smooth cuticle, and a longly exserted perianth without beak (ReinerDrehwald and Schäfer-Verwimp 2008a). 21. Lejeunea cyathophora Mitt. On bark of trees and treelets in montane rainforests, also found on Citrus leaves in orchard (Schäfer-Verwimp et  al. 2013a), 1800–3000  m. Colombia: Ant, Cho,

Fig. 52 (A) Lejeunea controversa, habit with gynoecium. (B) Lejeunea corynantha, habit with gynoecium. (C) Lejeunea debilis, habit. (D and E) Lejeunea deplanata. (D) Median cells. (E) Habit with gynoecium. (F) Lejeunea flaccida, habit with gynoecia. (H and I) Lejeunea herminieri. (H) Leaf. (I) Lobule apex. (J) Lejeunea flava, habit with gynoecia. (K) Lejeunea acanthogona, habit with gynoecium. (L) Lejeunea inflexiloba, habit with gynoecium. (M) Lejeunea laeta, habit. (N) Lejeunea laetevirens, habit. (B redrawn from Reiner-Drehwald and Schäfer-Verwimp 2008a; L redrawn from Reiner-Drehwald and Goda 2000, M redrawn from Reiner-Drehwald 2000; rest from Gradstein et al. 2001; Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009). Scale bars: A, G, I, J, M  =  250  μm; B, C, E  =  500  μm; D  =  25  μm; F, H  =  100  μm; K  =  1000  μm; L = 400 μm

526

 

Keys and Descriptions

Mag. Ecuador: Pi, Tu, ZC.  General distribution: tropical Andes, Central America (Panama, Costa Rica). Lejeunea cyathophora resembles L. capensis but differs in the peculiar perianth with a huge cup-shaped beak, almost as wide as the perianth (Reiner-Drehwald and Schäfer-Verwimp 2008a). 22. Lejeunea debilis (Lehm. & Lindenb.) Nees & Mont. Fig. 52C Taxilejeunea debilis (Lehm. & Lindenb.) Steph. Lejeunea heterocheila Taylor, syn. nov. Type: Ecuador, Pichincha, Jameson s.n. (isosyntype G-00115683!, ster.) On shrubs in open vegetation, on road banks, and on rock, ca. 400–3500  m. Colombia: Cun, Qui (Gradstein 12726a, HUQ), Ris, San (Gil 2088, UPTC). Ecuador: EO, Ga, Lo, Pi (Thiers 4337, QCA), ZC.  General distribution: throughout tropical America. Lejeunea debilis is a common dioicous neotropical member of the former genus Taxilejeunea that is recognized by (1) plants frequently sterile, 1–1.5 mm wide, pale whitish-green; (2) leaf lobes obliquely spreading, asymmetrically ovate-triangular, apex acute-apiculate to short-acuminate, margins crenulate, usually with a conspicuous tooth near apex that is separated from the apex by a lunulate sinus, sometimes with a smaller, second tooth, dorsal leaf base auriculate; (3) leaf cells very thinwalled, trigones minute, intermediate thickenings absent, cuticle densely punctatepapillose, oil bodies segmented, 3–10 per cell; (4) lobules very small, ca. 1/7× leaf length, ovate-triangular, keel crenulate; (5) underleaves large, ca. (4–)5–7× stem width, imbricate or contiguous, orbicular, bifid to 1/4 with a sharp to rounded incision, underleaf lobes not overlapping, margin crenulate, base with very large auricles, insertion line deeply arched (0.1–0.2 mm deep); and (6) gynoecia in a row on innovating branches, perianths with five keels in the upper 1/3, keels smooth or crenulate (Evans 1921; Gradstein and Ilkiu-Borges 2009). Lejeunea debilis is morphologically close to L. pterigonia; for differences see under the latter species and Evans (1921). Both species have a punctate-papillose cuticle, but in L. debilis the cuticle is more densely papillose. Sterile material of Lejeunea debilis has been confused with L. cerina; both species may produce a conspicuous tooth near apex that is separated from the apex by a lunulate sinus. The two species are readily separated, when sterile, by the shape of the underleaf insertion line which is deeply arched (0.1–0.2  mm deep) in L. debilis and shallowly curved (to 0.05 mm deep) in L. cerina. 23. Lejeunea deplanata Nees Fig. 52D, E Hygrolejeunea paramicola Herzog Lejeunea hebetata Spruce Lejeunea megalostipa Spruce Lejeunea montana Gottsche Lejeunea radicans Lindenb. & Gottsche, syn. nov. Type: Mexico, Veracruz, Mirador, Mar 1842, Liebmann s.n. (lectotype, designated here, W-Lindenb. 6686!; isolectotype C-M9213!) Rectolejeunea longiloba Steph. Rectolejeunea maxonii A.Evans (Lejeunea maxonii [A.Evans] X.L.He)

Lejeuneaceae

527

Common on branches and twigs of trees and shrubs in rather open situations, occasionally on rotten wood and living leaves, (200–)1200–3800 m; one of commonest liverworts in the Galápagos Islands in evergreen woodlands and scrub at 300–1200  m (Gradstein 2009). Colombia: Ant, Boy, Cun, Ris, San, Tol, Val. Ecuador: Az, Ga, Im, MS, Or, Pa, Pi, Su, Tu, ZC. General distribution: widespread in tropical and subtropical America. Lejeunea deplanata is a common neotropical species that is distinguished by (1) plants dioicous, 0.8–1.5 mm wide, glistening pale green, branches usually Lejeuneatype, with very large leafy collars at the base; (2) leaf lobes wide-spreading, ovatesuborbicular, convex or plane, apex rounded, recurved or plane, margins entire to crenulate; (3) leaf cells ± uniformly thick-walled, trigones small or obscure, cuticle smooth, oil bodies segmented, few per cell; (4) lobules when well developed ovoid to oblong, variable in size, 1/5–1/2× lobe length, fully inflated, free margin incurved, tooth 1–2 cells long; (5) underleaves distant to subimbricate, 2.5–5× stem width, orbicular to slightly wider than long, bifid to 1/2, margins entire, bases cuneate to rounded, insertion line curved; (6) male bracteoles restricted to the base of the male spike; (7) gynoecia terminal on the stem or on branches, with 1–2 innovations, perianths obovoid, keels smooth, beak short; and (8) vegetative reproduction present or absent, when present by caducous leaf lobes and caducous microphyllous branches (Reiner-Drehwald 2010b). The small, uniformly thick-walled cells and the branches with large leafy collars at the base are particularly characteristic of L. deplanata. The species is morphologically close to L. phyllobola and L. tapajosensis; all are dioicous and may produce caducous leaf lobes. The latter two species differ from L. deplanata by leaf cells with distinct trigones and intermediate thickenings, male bracteoles present throughout the male spike, flat leaves with a plane apex, and absence of caducous microphyllous branches. Moreover, L. deplanata occurs at higher elevations. 24. Lejeunea drehwaldii Heinrichs & Schäf.-Verw. Sphaerolejeunea umbilicata Herzog On living leaves in humid submontane and lower montane rainforest, 1000–1800  m. Colombia: Cau (type). Ecuador: ZC (Schäfer-Verwimp et  al. 2013a) General distribution: northern Andes (Colombia to Peru). Lejeunea drehwaldii is a rare northern Andean epiphyll that stands out by the presence of a leaf border of hyaline rectangular cells in 1–2 rows and by the small, terete perianth without beak and with the mouth displaced to the dorsal, adaxial side of the perianth (Reiner-Drehwald and Drehwald 2002 as Sphaerolejeunea umbilicata). Further characteristics of this unusual species are (1) plants autoicous and usually highly fertile, mostly densely pinnately branched, rather large, 1.5–2.3 mm wide, very flat and growing strongly appressed to the substrate, stems conspicuously zig-zag; (2) leaves suborbicular, apex broadly rounded, margins entire; (3) lobules often reduced, when well developed shortly rectangular, ca. 1/5× leaf length, with 2 teeth, first tooth large, 3–4 cells long and 2–4 cells wide, second tooth 1-celled; (4) underleaves distant, 2–3× stem width, slightly longer than wide, lobes obtuse, margins entire but somewhat irregular, bases narrowly rounded; and (5)

528

 

Keys and Descriptions

androecia and gynoecia numerous, on very short branches hidden behind the leaves, gynoecia with a short, sterile innovation. By its very flat habit, rather large size, pinnate branching, bordered leaves, small 2-toothed lobules, small underleaves and terete, and beakless perianths, this species cannot be confused with other epiphyllous Lejeunea species. Heinrichs et  al. (2013a) found L. drehwaldii in a clade together with members of the former genera Macrolejeunea and Taxilejeunea. 25. Lejeunea elongella Gottsche Hygrolejeunea elongella (Gottsche) Steph. Lejeunea fulva Spruce (Hygrolejeunea fulva [Spruce] Steph.), syn. nov. Type: Ecuador, Baños, Mt. Tungurahua, Spruce s.n. (isotype G-00282426!) On bark in montane forest and on Espeletia trunks in páramo, 2100–3450  m. Colombia: Boy (Gil s.n., UPTC), Cun (type), Qui (páramo El Campanario, Téllez s.n., UDBC). Ecuador: Tu. General distribution: northern Andes (Colombia, Ecuador). Lejeunea elongella is a rare, little-known northern Andean species that is recognized by (1) plants dioicous, long and slender, little branched, whitish-green, ventral merophyte 2 cells wide; (2) leaves ovate-orbicular, with a broadly rounded and frequently somewhat recurved apex; (3) leaf cells with small trigones and intermediate thickenings, cuticle smooth; (4) lobules very small or reduced, maximally 1/6× leaf length; (5) underleaves large, 5–7× stem width, orbicular, shallowly bifid, bases auriculate; and (6) gynoecia on short branches without innovation, perianth terete. Lejeunea elongella resembles L. rotundifolia but differs from the latter in the ventral merophyte being 2 cells wide (4–6 cells wide in L. rotundifolia) and cell walls with intermediate thickenings (intermediate thickenings lacking in L. rotundifolia) (Reiner-Drehwald 2005b). Moreover, the gynoecia of L. elongella are developed on very short branches without innovation, and the perianth is terete (J. Gil, pers. comm.). In L. rotundifolia the gynoecia are on short or long branches with innovations, and the perianth is 5-keeled. By the large, auriculate underleaves, smooth cuticle, gynoecia on very short branches without innovation, and eplicate perianth, L. elongella is similar to L. pallescens, but the latter species has pointed leaf apices and larger lobules. 26. Lejeunea flaccida Lindenb. & Gottsche Fig. 52F Type: Mexico, Veracruz, Río Nautla, finca Cabrestras, Apr 1841, Liebmann 476, c. gyn. (lectotype (Bastos and Gradstein 2020), W-Lindenb. 6389!; isolectotype C-M9200!) Lejeunea florida Spruce (Taxilejeunea florida [Spruce] Steph.), syn. nov. Type: Ecuador, Amabato, 2500 m, Spruce L29, autoicous (MANCH-cc18250!) Lejeunea lusoria (Lindenb. & Gottsche) Steph. (Taxilejeunea lusoria [Lindenb. & Gottsche] Steph.) (fide Bastos and Gradstein 2020) Lejeunea obtusangula Spruce (Taxilejeunea obtusangula [Spruce] A.Evans) (fide Bastos and Gradstein 2020) Taxilejeunea foliicola Steph. (fide Bastos and Gradstein 2020)

Lejeuneaceae

529

Taxilejeunea macroloba Steph. Taxilejeunea multiflora Steph. (fide Bastos and Gradstein 2020) Taxilejeunea terricola (Spruce) Steph. (fide Bastos and Gradstein 2020) Sun epiphyte on bark of trees and shrubs in rather open areas, also on living leaves, from sea level to ca. 2800 m. Colombia: Boy (Uribe 3736, COL), Hui, Mag, Qui, Ris, Vau. Ecuador: Ch, Ga (Gradstein H16, H362, GOET, U), MS, Pa, Pi (Thiers 4335, 4342, 4345, QCA), Or (Mota de Oliveira and ter Steege 2013), Su, Tu. General distribution: widespread in tropical and subtropical America from Florida to northern Argentina (Buenos Aires). Lejeunea flaccida is a common and widespread neotropical species that was usually called Taxilejeunea obtusangula, which is a synonym. Taxilejeunea lusoria is a further synonym; the latter name has been misapplied and was used for L. bombonasensis (see there). The main character of L. flaccida is the perianth with 5 short keels in the upper (1/5–)1/4–1/3 of the perianth. Other important morphological features are (1) plants autoicous, 1–1.5 mm wide, pale whitish-green, loosely creeping or pendent; (2) leaf lobes ovate-triangular, somewhat asymmetrical, apex narrowly rounded to obtuse to apiculate, margins entire or with a few small teeth near apex; (3) leaf cells with very small trigones, intermediate thickenings scarce, cuticle ± smooth; (4) lobules usually reduced, when well developed ca. 1/4× leaf length, with a short tooth; (5) underleaves distant to imbricate, rounded or wider than long, 3–5× stem width, bifid to 1/2 with a wide incision, margins entire, bases cordate, rounded or with low auricles; (6) gynoecia on the main stem or in a row on short innovating branches (row up to 5 gynoecia), bracts and bracteoles slightly and remotely toothed to entire; and (7) mature perianth oblong, keeled near apex, over maximally 1/4(−1/3) its length, keels smooth or crenulate, occasionally with a few small teeth, somewhat inflated above (Schuster 1980a as T. obtusangula; ReinerDrehwald 2000 as T. obtusangula). 27. Lejeunea flagellaris Spruce Type: Ecuador, Mt. Chimborazo, Spruce L525 (MANCH-000213!) Lejeunea leptalea Spruce, syn. nov. Type: Ecuador, Mt. Altar, 3350 m, Spruce L30 (lectotype, designated here, MANCH-000270!; isolectotypes MANCH-000271!, MANCH-000272!) Taxilejeunea galapagensis Onr., syn. nov. Type: Galápagos Is., Floreana, on trunk of avocado tree, 350 m, 27 Apr 1976, Gradstein H186 (holotype U!) On living leaves, twigs, and branches in moist evergreen woodlands, scrub, and orchards, rarely on rock, ca. 2000–3350 m on the mainland; at 350–800 m in the Galápagos Islands. Colombia (?): Cho (Winkler 1970). Ecuador: Ch (type), Ga, MS. General distribution: only known with certainty from Ecuador. Lejeunea flagellaris is a little-known Ecuadorian endemic that is common on the Galápagos Islands. The species is recognized by the slender stems with numerous small, microphyllous branches; in addition, upper parts of the stems may also become etiolated and microphyllous (e.g., in the type). The lobules on microphyllous branches are reduced or sometimes large and swollen (in the type of L. leptalea). Further characteristics of L. flagellaris are (1) plants dioicous, 1–1.5 mm wide,

530

 

Keys and Descriptions

glistening green, creeping or pendent; (2) leaf lobes ± distant, flat, elongate ovatetriangular, (sub)acute to apiculate, in lower part of stem sometimes obtuse to rounded, margins entire or crenulate; (3) leaf cells with small trigones and intermediate thickenings, cuticle smooth; (4) lobules to 1/4× leaf length, often reduced, tooth obscure, keel entire or crenulate; (5) underleaves distant, 2–3× stem width, ovate and somewhat longer than wide to suborbicular, bifid to 1/2 with a wide incision, lobes acute or narrowly obtuse, margins entire, bases cuneate; (6) gynoecia on short or long shoots, with 1–2 innovations that may again become fertile, perianth sharply keeled over most of its length, keels smooth, beak short (Onraedt 1982 as T. galapagensis); and (7) vegetative reproduction not observed. Lejeunea flagellaris resembles L. bombonasensis in general habit but differs from the latter in the numerous microphyllous branches and dioicy. In the presence of microphyllous branches, L. flagellaris is similar to L. cancellata, but the latter species has caducous branches and broader underleaves (3–4× stem width) which are slightly wider than long. Lejeunea leptalea is a phenotype of L. flagellaris with subacute leaves and microphyllous branches with large, swollen lobules (to 2/3× leaf length). 28. Lejeunea flava (Sw.) Nees Fig. 52J Lejeunea albida (Spruce) Steph. Lejeunea gaudichaudii Lindenb. Lejeunea granatensis Steph., nom. illeg. (≡ Lejeunea silvatica Gottsche), syn. nov. Type: Colombia, Lindig s.n. [= lectotype of L. silvatica Gottsche, designated here] (G-00280484!) Lejeunea leucophaea Spruce Lejeunea pallida Lindenb. & Gottsche (Hygrolejeunea pallida [Lindenb. & Gottsche] Steph.), syn. nov. Type: Mexico, Veracruz, St. Maria Tlapacoyo, Liebmann 577 (isotype C-M9878!) Lejeunea symphoreta Spruce Taxilejeunea tenuiplica Steph., syn. nov. Type: Ecuador, Morona Santiago, Gualaquiza, Allioni s.n., Bryotheca Levier 6647 (holotype G-00047597!) Common on bark, rock, living leaves, and dead wood in rather open habitats, in primary and secondary forests, scrub, plantations, gardens, and on isolated trees, in primary forest mainly in the canopy, from sea level to about 3000 m. Colombia: Ama, Ant, Boy, Caq, Cas, Cau, Cho, Cun, Hui, Mag, Met, Put, Qui, Ris, San, Tol, Vau. Ecuador: EO, Ga, MS, Na, Or, Pa, Tu, ZC. General distribution: pantropical, western Europe, southeastern USA. Lejeunea flava is one of the most common and widely distributed species in the genus and is particularly characteristic of open, disturbed habitats. The species is distinguished by (1) plants autoicous and usually copiously fertile, 0.8–1.5(−2) mm wide, light green to yellowish-green; (2) leaf lobes widely spreading, ovate-oblong to oblong, plane, apex rounded, margins entire; (3) leaf cells usually with conspicuous trigones (sometimes absent) and few intermediate thickenings, cuticle smooth or faintly papillose due to the presence of wax crystals, oil bodies segmented, ca. 2–10 per cell; (4) lobules ca. 1/4× leaf length, sometimes reduced, inflated, ovoid,

Lejeuneaceae

531

free margin incurved, tooth short, obtuse; (5) underleaves rather large, contiguous to imbricate, slightly longer than wide, 3–4× stem width, bifid to 1/2, lobes acute, margins entire, bases rounded to subauriculate, cordate; and (6) gynoecia on short branches, single or two in a row, perianth 5-keeled, keels smooth (Schuster 1980a; Reiner-Drehwald 2000, 2009b). The ovate-oblong leaf lobes with rounded tips, leaf cells with conspicuous trigones, rather large, frequently longer than wide underleaves with a cordate base, gynoecia single or two in a row on short branches, and perianths with five smooth keels are the main characters of L. flava. In plants from very humid habitats, trigones may be lacking and leaf cells fully thin-walled. Lejeunea silvatica was described based on two specimens collected by Lindig in the surroundings of Bogotá. The two syntypes are lost, but a duplicate of one of the syntypes (which one is unclear), annotated by Stephani as “Kolumbien, leg. Lindig,” is kept in G and is the type of L. granatensis Steph. Although the earlier name L. silvatica is not mentioned in the protologue, it is referred to on the label of the type of L. granatensis (“olim L. silvatica”). The syntype in G is chosen here as the lectotype of L. silvatica; the name L. granatensis is considered illegitimate. 29. Lejeunea fulfordiae (Jovet-Ast) R.L.Zhu Amblyolejeunea fulfordiae Jovet-Ast Epiphytic on Baccharis in roadside scrub, 2220  m. Ecuador: Na (SchäferVerwimp et al. 2006; Zhu and Cheng 2008). General distribution: Guadeloupe, Ecuador. Lejeunea fulfordiae is a rare neotropical species that is recognized by (1) plants autoicous, 1–1.5 mm wide; (2) leaf lobes widely spreading, convex, apex rounded, usually recurved, margins slightly crenulate, ventral margin of the leaf lobe frequently recurved; (3) leaf cells thin-walled, trigones very small, intermediate thickenings ± absent, cuticle smooth; (4) lobules rather large, 2/5–1/2× lobe length, truncate, apex short, obtuse, hyaline papilla proximal; (5) underleaves small, undivided, orbicular, insertion line straight; and (6) gynoecia on short branches with innovation, perianth terete, without beak. Lejeunea fulfordiae resembles L. herminieri in the undivided underleaves, ± crenulate leaf margins, and rather large, truncate lobules. However, L. herminieri differs in dioicy, distal hyaline papilla, finely papillose (“punctate”) cuticle, and 5-keeled, beaked perianths. The oil bodies in L. herminieri are homogeneous; those in L. fulfordiae are unknown. 30. Lejeunea galeata Spruce Harpalejeunea galeata (Spruce) Steph. On bark, rock, and rotten wood in lowland and montane rainforest, 200–1820 m. Colombia: Cho (Gradstein et  al. 2018c). Ecuador: Ch, ZC (Schäfer-Verwimp et al. 2013a). General distribution: northern Andes (Colombia to northern Peru). Lejeunea galeata is a little-known northern Andean species that is readily recognized by the very small plants (0.4–0.8 mm wide) with strongly caducous, orbicular leaf lobes. Further characters are (1) plants dioicous und usually sterile, irregularly

532

 

Keys and Descriptions

branched, stems and branches partly denuded due to leaf lobes dropping off; (2) leaf lobes wide-spreading, (ovate-)orbicular, plane, apex rounded, margins entire, leaf base very narrow; (3) leaf cells fully thin-walled, cuticle finely punctate-papillose; (4) lobules ovoid, swollen, ca. 1/4× lobe length, with a conspicuous tooth; (5) underleaves distant, very small, scarcely wider than the stem, suborbicular, bifid to 1/2, lobes obtuse to acute, bases cuneate; (6) male bracteoles restricted to the base of the male shoot; and (7) gynoecia are on short branches with an innovation, perianths sharply keeled in the upper half, keels smooth, beak short. 31. Lejeunea glaucescens Gottsche Lejeunea caulicalyx (Steph.) M.E.Reiner & Goda (Crossotolejeunea caulicalyx Steph.) (fide Bastos and Gradstein 2020) Lejeunea tarapotensis Spruce, syn. nov. Type: Peru, near Tarapoto, “ad rivuli Cumbaea ostia, in cortice vetusta,” 300 m, Spruce L18 (lectotype, designated here, MANCH-000435!). Paratype: ibid., Spruce L3 (MANCH-000436!, MANCH-0004348!) Lejeunea tarapotensis var. microstipa Spruce, nom. inval. (in sched.). Material: Peru, near Tarapoto, Mt. Campana, Spruce s.n. (MANCH-000439!, MANCH-000440!, G!) Otigoniolejeunea reducta S.Winkl. (fide Uribe and Gradstein 1998) On rotten or dead wood in lowland and submontane primary rainforest, from sea level to 650 m. Colombia: Mag (Winkler 1976 as Otigoniolejeunea reducta), Met (Río Guayabero, Bischler 1607, U). Ecuador: MS, Na (Schäfer-Verwimp et  al. 2013a), Or. General distribution: widespread in tropical America. Lejeunea glaucescens is a characteristic neotropical lowland species from dead wood that stands out by crenulate to crenate leaf margins, small underleaves (usually 1.5–2× stem width), gynoecia on very short branches, and perianths with crenate keels. The species may be characterized as follows: (1) plants autoicous, 0.8–1.5 mm wide; (2) leaf lobes wide-spreading, ovate, flat, apex broadly rounded, margins crenulate; (3) leaf cells fully thin-walled or with small trigones and minute intermediate thickenings, cuticle smooth or slightly punctate-papillose; (4) lobules 1/5(−1/3)× leaf length, swollen, with incurved margin and a short, obtuse tooth; (5) underleaves small, distant, 1.5–2(−2.5)× stem width, as wide as long or slightly longer than wide, bifid to 1/2, lobes acute, margins crenulate, sometimes with a short tooth, bases cuneate, insertion line curved; and (6) gynoecia on very short branches without vegetative leaves and with 1 sterile innovation, perianths obovoid to obpyriform, slightly flattened, with 4–5 short keels in the upper half (dorsal keel low, sometimes lacking), the keels sharp and somewhat expanded above, crenate by mammillose cells, beak very short (Schuster 1980a; Reiner-Drehwald and Goda 2000 as L. caulicalyx). Lejeunea glaucescens is closely similar to L. subsessilis and L. quinqueumbonata; all of them produce gynoecia on very short branches with bracts only (no vegetative leaves) and have perianths keeled only in the upper half. Lejeunea subsessilis differs from L. glaucescens in the lesser crenation of perianth keels and slightly broader underleaves, 2–3(−4)× stem width. Moreover, the two species grow

Lejeuneaceae

533

in different habitats, L. glaucescens on dead wood and L. subsessilis on living leaves and bark. Lejeunea quinqueumbonata differs from L. glaucescens in the rounded to subacute leaf apex, strongly crenate leaf and underleaf margins, broader underleaves with an occasional blunt tooth on the outer margins, and perianth keels terminating above in a lamellate projection. 32. Lejeunea herminieri (Steph.) R.L.Zhu Oryzolejeunea antillana (R.M.Schust.) R.M.Schust. Oryzolejeunea saccatiloba (Steph.) Gradst.

Fig. 52H, I

On bark in montane rainforests, especially in the forest canopy, 1500–2650 m. Colombia: San. Ecuador: Lo, Na, Tu, ZC.  General distribution: tropical America. The unique character of this rather common neotropical species is the distal hyaline papilla; other characters include (1) plants dioicous, 1–1.3 mm wide; (2) leaf lobes widely spreading, convex, apex rounded, margins crenulate; (3) leaf cells thin-walled, trigones very small, cuticle finely papillose (“punctate”), oil bodies homogeneous, numerous per cell, Massula-type; (4) lobules large, 2/5–1/2× lobe length, truncate, apex short, obtuse, hyaline papilla distal; (5) underleaves small, distant, undivided, orbicular, insertion line straight; and (6) gynoecia on short branches with innovation, perianth with 5 smooth keels and a short beak. Lejeunea herminieri may be confused with L fulfordiae; for differences see under L. fulfordiae. By the distal papilla, L. herminieri is similar to members of Cheilolejeunea, especially C. holostipa; both species share undivided underleaves. However, C. holostipa is a much smaller plant (ca. 0.5 mm wide) with suberect leaf lobes, lobules with a long and sharp tooth, and leaf cells with a smooth cuticle and finely granular oil bodies. 33. Lejeunea immersa Spruce Trachylejeunea immersa (Spruce) Steph. Type:  – PERU: Yurimaguas, fl. Huallaga, Spruce L489 (lectotype, MANCH-000007! isotype MANCH000008!) On bark and rotten wood in lowland rainforest, 100–250 m. Colombia (new): Vau (Río Apoporis, ca. 250  m, Schultes & Cabrera 14419, det. M.E.  ReinerDrehwald, GOET). General distribution: Amazonia (Brazil, Peru, Colombia) and E Brazil (Bahia). Lejeunea immersa is an uncommon Amazonian lowland species that is characterized by (1) plants autoicous, 1.0–1.4 mm wide, usually fertile; (2) leaves obliquely to widely spreading, ± flat, ovate, not falcate, apex rounded to acute to apiculate, margins strongly crenate by transversally oriented quadrate cells; (3) leaf cells without or with minute trigones, intermediate thickenings absent, cuticle smooth; (4) lobules often reduced, when well developed small, to maximally 1/5 of leaf length, tooth short, keel crenate; (5) underleaves distant, orbicular, 2–3× stem width, bifid to 1/2, apices subacute, margins crenate, outer margins without tooth; and (6) gynoecia on short branches with vegetative leaves and with 1 sterile innovation, perianths oblong to obovoid, 5-keeled, keels strongly crenate, without lamella-like projections or cilia, beak short (Bastos and Gradstein 2020).

534

 

Keys and Descriptions

Lejeunea immersa is recognized by the strongly crenate leaf margins and perianth keels. The species resembles L. quinqueumbonata and L. raddiana but differs from L. quinqueumbonata in the strongly crenate perianth keels without lamellalike projections and from L. raddiana in the non-falcate leaf lobes, smaller lobules, and larger underleaves with acute tips. 34. Lejeunea inflexiloba Prantl  Crossotolejeunea inflexiloba J.B.Jack & Steph., nom. inval. Crossotolejeunea cavifolia Steph.

Fig. 52L

On bark in shrubby páramo, in the type locality growing over Plagiochila cucullifolia, ca. 3000  m. Colombia: Ant (type). Ecuador: ZC (Parolly et  al. 2004). General distribution: northern Andes, southeastern Brazil. Lejeunea inflexiloba is a rare páramo species that is only known from the type from Colombia and a record from southern Ecuador. The species is distinguished by (1) plants dioicous (?), ca. 1.5 mm wide; (2) leaf lobes wide-spreading, ovate, apex rounded, ventral margin of the leaf lobe strongly recurved; (3) leaf cells with distinct trigones and intermediate thickenings, cuticle smooth or slightly and finely papillose; (4) lobules rather large, ca. 2.5× leaf length, subrectangular, fully inflated or flattened to the free margin, apex truncate, tooth obtuse, keel forming an almost straight line with the revolute margin of the leaf lobe; (5) underleaves distant, ca. 2.5–3× stem width, suborbicular, bifid to 1/5–1/3, lobes subacute, margins slightly crenate, bases cuneate, insertion line slightly curved; and (6) gynoecia on long shoots with 1–2 innovations, perianths pyriform, keels 2-winged, entire or sparsely ciliate, beak short, 2–3 cells long (Reiner-Drehwald and Goda 2000). By the strongly revolute ventral margin of the leaf lobe, this species cannot be confused with other neotropical Lejeunea species. 35. Lejeunea intricata Prantl Crossotolejeunea intricata J.B.Jack & Steph., nom. inval. Crossotolejeunea pustulosoalata Herzog Crossotolejeunea rotundata S.Winkl. On bark of tree in montane rainforests, 650–3000 m. Colombia: Ant, Cau, Ris. Ecuador: MS (Schäfer-Verwimp et  al. 2013a). General distribution: Central America, northern Andes, Dominican Republic. Lejeunea intricata is an uncommon neotropical species that is recognized by (1) plants autoicous, very small, 0.5–0.8  mm wide; (2) leaf lobes widely spreading, ovate-falcate, plane, apex rounded, margins crenate; (3) leaf cells with small trigones and few intermediate thickenings, cuticle finely papillose; (4) lobules when well developed 2/5–1/2× leaf length, ca. 1.5× longer than wide, fully inflated or flattened above, free margin incurved or plane, tooth slightly elongate and obtuse, keel arched, sharp angle between keel and ventral margin of the leaf lobe; (5) underleaves small, distant, 1.5–2.5× stem width, bifid 1/3–1/2 with a wide V-shaped incision, outer margins arched, entire, lobes broadly triangular, obtuse, bases narrow, cuneate, insertion line straight; and (6) gynoecia on a long shoot with a sterile innovation, perianths keels 2-winged, ciliate-laciniate, beak short, 1–2 cells long (Reiner-Drehwald and Goda 2000).

Lejeuneaceae

535

Lejeunea intricata resembles L. acanthogona, but the latter species has a smooth cuticle, leaf margins slightly crenulate only, and a 2–6-cell-long perianth beak. 36. Lejeunea killipii (Herzog) Gradst., comb. nov. Taxilejeunea killipii Herzog, Feddes Repert. Spec. Nov Regni Veg. 57: 190. 1955 On living leaves in wet lowland rainforest along the Pacific coast, 300–500 m. Colombia: Val (Herzog 1955). General distribution: only known from the type specimen from Colombia. Lejeunea killipii is a rare Colombian endemic that is close to L. bombonasensis but differs in the leaf margins irregularly toothed in the upper half, especially on branch leaves, and the strongly toothed perianth keels. The leaf apex in L. killipii is broadly rounded to apiculate, the lobules are reduced, and the underleaves are small and distant, ca. 2–3× stem width. 37. Lejeunea laeta (Lehm. & Lindenb.) Lehm. & Lindenb. Lejeunea geophila Spruce

Fig. 52M

On moist rock and moist soil in or near rivers and waterfalls, and along trails, often in running water, from sea level to 2000  m. Colombia: Ant, Boy, Cho (Gradstein 8667, det. M. E. Reiner-Drehwald, GOET), Mag (Stephani 1898–1924 as Taxilejeunea papuliflora Steph. [not the lectotype], G-00113382!, G-00113385!; Winkler 1976 as “L. filipes”), Met (La Macarena, Schultes 11130, 11168 (U), Bischler 1652, 1659 (U); Villavicencio, C.  Romero s.n., LLANOS), Ris (ReinerDrehwald et al. 2018). Ecuador: EO (Reserva Buenaventura, Benitez et al. 1314, HUTPL), Es, Tu. General distribution: widely distributed in tropical America (Reiner-Drehwald et al. 2018). Lejeunea laeta is a widespread hygrophytic species that is characterized by (1) plants autoicous and usually fertile, ca. 1  mm wide, yellowish-green; (2) stems rather fragile, of very thin-walled cells; (3) leaf lobes widely spreading, ovate, plane or slightly convex, apex broadly rounded to obtuse, often recurved, margins entire; (4) cells very thin-walled, trigones minute, intermediate thickenings absent, cuticle smooth; (5) lobules usually reduced, when well developed ca. 1/4× leaf length, ovate, with a short, obtuse tooth; (6) underleaves small, slightly wider than long, 1.5–2× stem width, bifid to 1/2, lobes obtuse, margins rounded, entire; (7) male bracteoles 2–3 in the lower half of the male spike or present throughout (variable!); and (8) gynoecia on elongate shoots, with innovation (innovation usually sterile), perianth oblong, 5-keeled, keels irregularly winged and variously ornamented with teeth and short cilia or almost entire, beak short (Reiner-Drehwald 2000). Although only few collections are available from Colombia and Ecuador, L. laeta is probably widespread in the region and undercollected. The species grows sometimes together with L. adpressa, and the two may be confused but the leaf apex in L. adpressa is always rounded and flat, the leaf cells have conspicuous trigones and intermediate thickenings, and the underleaves are more deeply bifid (to 2/3), with narrower, lanceolate lobes and acute tips. Moreover, the male bracteoles in L. adpressa are always restricted to the base of the male spike. Lejeunea laeta also approaches L. glaucescens, but the latter species differs in having constantly rounded, flat leaf apices (not obtuse or recurved), gynoecia on very short branches,

536

 

Keys and Descriptions

perianth keels strongly crenate by mammillose cells, and underleaves usually slightly longer than wider (not wider than long). 38. Lejeunea laetevirens Nees & Mont. Fig. 52N Microlejeunea laetevirens (Nees & Mont.) A.Evans Lejeunea cladobola Spruce Lejeunea glaucophylla Gottsche Lejeunea lepida Lindenb. & Gottsche Lejeunea lucens Taylor Lejeunea subhyalina Lindenb. & Gottsche Lejeunea subrufula Spruce, syn. nov. Type: Ecuador, Mt. Chimborazo, on living leaves, Spruce L547 (MANCH-000011!) Very common on bark of trees and shrubs in rainforests and dry woodlands, scrub, plantations, gardens, parks, and on isolated trees, also on living leaves and rock, 0–2000(−2650) m; abundant on the Galápagos Islands. Colombia: Ama, Cas, Ces, Cun, Mag, Put, Ris, San, Vau. Ecuador: Az, Ch, EO, Es, Ga, Lo, LR, MS, Na, Or, Pa, Pi, Su, Tu, ZC. General distribution: common and widespread in tropical America; also on Ascension Island (Pressel et al. 2017). Lejeunea laetevirens is one of the most common neotropical Lejeunea species and one of the most xerotolerant epiphytic liverworts in the region. The species is distinguished by (1) plants dioicous and often sterile, small, 0.4–0.8 mm wide, dull pale green to bluish-green, stems fragile, often broken; (2) leaf lobes obliquely spreading, straight or slightly falcate, flat to slightly convex, ovate-oblong, 1.5–2× longer than wide, somewhat gradually narrowed to a rounded to obtuse to subacute apex, margins entire or crenulate; (3) cells very thin-walled, without or with small trigones, intermediate thickenings scarce, cuticle densely punctate-papillose, oil bodies segmented, few per cell; (4) lobules when well developed 1/3–2/5× leaf length, ovoid and fully inflated, free margin incurved, tooth short, obtuse; (5) underleaves distant, 1.5–3× stem width, as long as wide or slightly longer than wide, bifid to 1/2, lobes acute, outer margins frequently with a blunt tooth; (6) male bracteoles present throughout the male spike; (7) perianth weakly flattened, with five smooth keels and a short beak; and (8) vegetative reproduction by shoot fragmentation (Schuster 1980a; Reiner-Drehwald 2000, 2009b). The small, dioicous plants with obliquely spreading, ovate-oblong leaves; welldeveloped, swollen lobules; a densely punctate-papillose cuticle; and small bifid underleaves with or without tooth on the outer margins are unmistakable. The stems are fragile and vegetative reproduction is by shoot fragmentation. Lejeunea laetevirens is very similar to L. angusta, and the two species may be confused; for differences see under L. angusta. Lejeunea laetevirens also resembles L. concinnula, but the latter species has caducous leaf lobes and uniformly thick-walled cells and occurs in humid montane forests. 39. Lejeunea meridensis Ilk.-Borg. On bark and rotten wood in upper montane rainforest, 2550–2850 m. Ecuador: Lo, ZC (Schäfer-Verwimp et al. 2013a); not known from Colombia but expected there. General distribution: northern Andes (Venezuela, southern Ecuador).

Lejeuneaceae

537

Lejeunea meridensis (sect. Echinocolea) is a rare northern Andean species that is recognized by the dorsal leaf surface roughened by mammillose-papillose cells and the underleaf margins with a conspicuous tooth (Ilkiu-Borges 2005). The species resembles L. asperrima and L. subspathulata, but the latter two species occur at lower elevation, below 1500 m, and have entire underleaf margins. 40. Lejeunea mojandae Gradst., sp. nov.  Fig. 54A Type: Ecuador, Pichincha, Mojanda, crater lake, on flaky bark of Polylepis in mixed, mesic Polylepis forest at east side of lake, ca. 3800  m, 12 Oct 2018, S. R. Gradstein, S. León-Yánez & A. Moscoso 12749, c. per. (holotype QCA; isotype PC). Paratype: ibid., 2017, S. León-Yánez et al. s.n., ster. (QCA) Lejeunea mojandae is quite common on the flaky bark of Polylepis in mixed Polylepis forests on the slopes of the crater lake of Mojanda at 3800 m. The species is distinguished by (1) plants dioicous (androecia not seen), very small, 0.5–0.8 mm wide, light green, very transparent, prostrate; (2) stems fragile, of thin-walled cells, in cross section (probably) of 7 epidermis cells surrounding 3 smaller and more elongate medullary cells, epidermis cells rectangular on the ventral surface; (3) leaf lobes obliquely spreading, flat to convex, ovate-oblong to oblong, 0.75–1.25 mm long, 0.5–0.75 mm wide, 1.5–2× longer than wide, apex narrowly rounded, margins entire; (4) leaf cells slightly mammillose, 20–25 × 18–20 μm in midleaf, slightly smaller to the margin and becoming larger and more elongate to the base, walls thin to slightly thickened, with minute trigones, intermediate thickenings absent or obscure, cuticle slightly and finely punctate-papillose; oil bodies small, finely segmented, ca. 5–10(−15) per cell in midleaf, more numerous to the leaf base; (5) lobules occasionally reduced, when well developed ca. 2/5× leaf length, ellipsoid, fully inflated, ca. 1.5–1.8× longer than wide, free margin incurved, tooth short, erect, obtuse, hyaline papilla narrowly ellipsoid, marginal-proximal; keel curved, crenulate by mammillose cells, forming a very wide angle or almost straight line with the ventral margin of the leaf lobe; (6) underleaves distant, relatively large, ovate-oblong, longer than wide, 0.2–0.3  mm long, 0.15–0.2  mm wide, 1.3–1.5× longer than wide, 2.5–4× stem width, bifid to 1/2, lobes narrowly triangular, 4–7 cells wide at the base, apices narrowly rounded to obtuse, formed by 2 cells side by side, outer margins entire, bases cuneate, insertion line slightly curved; (7) gynoecia on long shoots with 1–2 lejeuneoid innovations, bracts elliptical, lobe 2× longer than the lobes or vegetative leaves, apex rounded to subacute, margins remotely bluntly angled (rudimentarily toothed), lobule ca. 3/4 × lobe length, slightly elongate beyond the keel, apex rounded, margin entire; keel ca. 2/3× lobe length, crenulate; bracteole slightly shorter than lobe (ca. 4/5× lobe length), elliptical, bifid to 1/3, margins entire; perianth immersed, slightly shorter than the bracts, obovoid, terete, apex broadly truncate, beak short; and (8) vegetative reproduction by shoot fragmentation. Lejeunea mojandae is closely similar to L. globosiflora (Steph.) Steph., a species endemic to dry, Mediterranean forests of central Chile (for description and illustration, see Gradstein and Cuvertino 2015). The latter species differs, however, in having suborbicular underleaves and a longly exserted perianth. A molecular comparison

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Keys and Descriptions

of the two species would be worthwhile; possibly L. mojandae might be a subspecies of L. globosiflora. The combined range of these two species resembles that of Frullania pluricarinata, a species common in central Chile where it grows together with L. globosiflora and, in addition, occurring at high elevations in the northern Andes (and extending northwards to Mexico). Lejeunea mojandae habitually resembles L. laetevirens, but the latter species differs by smaller, as long as wide underleaves (1.5–3× stem width) with acute lobes and outer margins often with a blunt tooth and a 5-keeled perianth. Moreover, L. laetevirens occurs at much lower elevation. 41. Lejeunea osculatiana De Not.  Fig. 53A Hygrolejeunea osculatiana (De Not.) Steph. Lejeunea flavicans Gottsche Taxilejeunea lancifolia Steph. (Macrolejeunea lancifolia [Steph.] Herzog) Pendent on bark of tree trunks and twigs in montane forest, also on rock and living leaves, 1250–2700  m. Colombia: Ris. Ecuador: Lo, Na, Pa, Su, Tu, ZC. General distribution: Central America, tropical South America. Lejeunea osculatiana is a characteristic neotropical-montane species that stands out by distant, lanceolate leaves and pendent growth. Further characters are (1) plants dioicous and often sterile, 1–2.5 mm wide, pale green to whitish-green, very slender, often growing pendent; (2) leaf lobes obliquely to widely spreading, 1.5–3× longer than wide, acuminate, margins entire; (3) lobules very small, less than 1/4× lobe length, tooth obscure; (4) leaf cells with small trigones and 0–1 intermediate thickenings, cuticle smooth or slightly punctate-papillose; (5) underleaves 2–3.5× stem width, bifid to 1/2–2/3, lobes narrowly lanceolate, acute, bases cuneate, insertion line almost straight; (6) gynoecia on short branches without innovation, bracts toothed, perianths mostly eplicate, slightly 5-keeled only at apex; and (7) vegetative reproduction by caducous branches (Reiner-Drehwald and Schäfer-Verwimp 2008a). Lejeunea osculatiana is close to L. pallescens (Heinrichs et al. 2013a), but the latter species has broader, ovate-triangular leaf lobes, 1.1–1.5× longer than wide, with an auriculate dorsal base, and much larger underleaves, 3.5–7× stem width, with large auricles. Moreover, L. pallescens lacks caducous branches. Lejeunea osculatiana is habitually similar to L. xiphophylla, but in the latter species, the leaf cells are more densely papillose, the gynoecia have an innovation and are usually arranged in a short row, and the perianths are longly 5-keeled. 42. Lejeunea pallescens Mitt.  Fig. 53B Macrolejeunea pallescens (Mitt.) Schiffn. Lejeunea prominula Gottsche (Taxilejeunea prominula [Gottsche] Steph.), syn. nov. Type: Colombia, Fusagasuga, Lindig s.n. (n.v.); the type could not be located Taxilejeunea laevis (Gottsche) Steph. (Omphalanthus laevis Gottsche) (?) see Reiner-Drehwald & Schäfer-Verwimp (2008a, p. 413) Taxilejeunea leioscypha (Gottsche) Schiffn. (Omphalanthus leioscyphus Gottsche). Lectotype: Colombia, Monte de las Escaleras, Lindig 1706

Fig. 53 (A) Lejeunea osculatiana, habit. (B) Lejeunea pallescens, habit with gynoecia. (C and D) Lejeunea phyllobola. (C) Habit with gynoecium. (D) Median cells. (E) Lejeunea pterigonia, habit with gynoecia. (F) Lejeunea pulverulenta, habit. (G) Lejeunea ramulosa, habit. (H) Lejeunea reflexistipula, habit. (I) Lejeunea rotundifolia, habit. (J) Lejeunea serpillifolioides, habit with gynoecia. (K and L) Lejeunea subspathulata. (K) Habit. (L) Leaf, dorsal view. (M) Lejeunea subsessilis, habit with gynoecia and androecia. (N and O) Lejeunea trinitensis. (N) Lobule. (O) Habit. (P) Lejeunea sulphurea, habit with gynoecia. (A from Reiner-Drehwald and Schäfer-Verwimp 2008a; C, D, M redrawn from Reiner-Drehwald 2000; F redrawn from Reiner-Drehwald 2005c; G redrawn from Reiner-Drehwald 2010a; I redrawn from Reiner-Drehwald 2005b; K, L from IlkiuBorges 2005; rest from Gradstein et  al. 2001; Gradstein and Costa 2003; Gradstein and IlkiuBorges 2009). Scale bars: A–C, F, H–K = 500 μm; D = 25 μm; E, P = 1000 μm; G, L = 250 μm; N = 100 μm; M, O = 400 μm

540

 

Keys and Descriptions

(PC-0102708!). Reiner-Drehwald and Schäfer-Verwimp (2008a) designated a duplicate in MANCH (“ex herb. Bescherelle, Paris”) as the lectotype, considering it was the only specimen of the species available. However, an excellently preserved specimen from the Lindig herbarium, with Gottsche’s handwriting, is in PC and should be the lectotype; the specimen in MANCH may be considered an isolectotype. Taxilejeunea peruviana Steph., syn. nov. Type: Ecuador, Mt. Tungurahua, Spruce s.n. (lectotype, designated here, G-00047598!, c. gyn.) Pendent on tree trunks and twigs in upper montane cloud forests and páramo, in moist, shaded environments, (1000–)2000–3800  m. Colombia: Boy, Ces, Cun, Mag, Qui, Ris, San. Ecuador: Az, Ca, Ch, Pi, Tu. General distribution: tropical Andes, Costa Rica. Lejeunea pallescens is common pendent epiphyte of tropical montane cloud forests above 2000 m in the Andes; records from lower elevation need verification. The species is distinguished by (1) plants dioicous, 1–2 mm wide, pale whitish-green; (2) leaf lobes obliquely spreading, asymmetrically ovate-triangular, 1.1–1.5× wider than long, apiculate to shortly acuminate, sometimes rounded on branches, margins entire or crenulate, dorsal base shortly auriculate; (3) leaf cells with conspicuous trigones and 0–2 intermediate thickenings, cuticle finely punctate-papillose, oil bodies finely segmented, 4–6 per cell; (4) lobules usually well developed, 1/4–1/3× lobe length, tooth obtuse, disk cell below the tooth larger than adjacent cells; (5) underleaves large, 4–7× stem width, bifid to 1/3, lobes acute, bases with large auricles; and (6) gynoecia on very short branches without innovations, bracts toothed, perianths terete or 3-plicate with 3 weak, broadly rounded keels, with a short beak (Reiner-Drehwald and Schäfer-Verwimp 2008a). In its pendent habit and underleaves with large auricles, L. pallescens resembles former members of the genus Taxilejeunea such as L. pterigonia and L. serpillifolioides. The latter species differ, however, in having reduced lobules, very thin-walled cells with minute trigones, and gynoecia in a row on innovating branches. Lejeunea pallescens is very close to L. osculatiana (Heinrichs et al. 2013a), and the two have been confused, but the latter species has narrower, ovate-lanceolate leaves, smaller lobules, and smaller, more deeply bifid underleaves without auricles. Some Ecuadorian records of L. pallescens belong to L. osculatiana (Schäfer-Verwimp et al. 2013a). Although the type of L. prominula has not been located, the species is considered a synonym because its original description fits L. pallescens very well and material in G identified as L. prominula (Colombia, Irmscher 23) belongs to L. pallescens. 43. Lejeunea phyllobola Nees & Mont. Rectolejeunea phyllobola (Nees & Mont.) A.Evans Lejeunea cephalandra Spruce Lejeunea polycephala Spruce Rectolejeunea brittoniae A.Evans Rectolejeunea colombiana R.M.Schust., nom. inval.

Fig. 53C, D

Lejeuneaceae

541

On bark of trees and shrubs, occasionally on rock and rotten wood, in lowland and montane rainforests, scrub, and plantations, often in rather open environments, from sea level to 2150 m. Colombia: Ama, Caq, Hui, Put, Suc, Val, Vau. Ecuador: LR (Gradstein 6951, 6970, GOET, U), MS, Pi (Thiers 4339, QCA), Su, ZC (SchäferVerwimp et al. 2013a). General distribution: tropical and subtropical America, Africa, India. Lejeunea phyllobola is one of the most common neotropical Lejeunea species with caducous leaf lobes. The main characters of the species are (1) plants dioicous but often freely fertile, 0.6–1 mm wide, growing appressed in dense mats; (2) leaves wide-spreading, ovate, quite flat, apex rounded, margins entire; (3) leaf cells with small, triangular trigones, cuticle smooth, oil bodies segmented, few per cell; (4) lobules when well developed 1/4–1/3× leaf length, inflated, free margin incurved, apical tooth obtuse; (5) underleaves distant, 1.5–3(−4)× stem width, suborbicular, bifid to 1/2, margins entire, bases cuneate to narrowly rounded, insertion line curved; (6) male bracteoles present throughout the male spike; (7) gynoecia terminal on the stem or on branches, with 1–2 innovations, perianth obovoid, ± flattened, longly 4–5-keeled (dorsal keel weak or reduced, lateral keels broadly expanded), keels smooth or crenulate, beak short; and (8) vegetative reproduction by caducous leaf lobes (Schuster 1980a as R. brittoniae; Reiner-Drehwald 2000). Schuster (1980a) described L. phyllobola as autoicous (following Evans), but Reiner-Drehwald (2000) showed that the species is dioicous; the identity of the autoicous plants is unclear. Lejeunea phyllobola resembles L. deplanata and L. tapajosensis, but L. deplanata differs in the frequently recurved leaf apex, ± uniformly thick-walled cells, and restriction of male bracteoles to the base of the male spike, whereas L. tapajosensis is a larger plant (1–1.5 mm wide) with larger underleaves (3–4× stem width) and conspicuously elongate, triradiate trigones. Lejeunea oligoclada Spruce, a caducous-leaved species endemic to the Atlantic coast of Brazil, is also close to L. phyllobola but differs in having obliquely spreading leaves with rounded to acute leaf apices, ± uniformly thick-walled cells with a finely papillose cuticle, and larger lobules (Reiner-Drehwald 2000). 44. Lejeunea prionoides Spruce On bark of trees in primary and seconday lower montane rainforests, 1000–1825  m. Ecuador: Pi, Tu, ZC.  General distribution: northern Andes (Ecuador, northern Peru). Lejeunea prionoides is a rare northern Andean species that is distinguished by (1) plants dioicous and often sterile, small, 1–1.8 mm wide, whitish-green, not fragile; (2) leaf lobes widely spreading, narrowly ovate-triangular, ca. 1.5× longer than wide, apex acute, plane or recurved, margins entire; (3) leaf cells very thin-walled, without or with minute trigones, intermediate thickenings usually lacking, cuticle finely papillose, oil bodies segmented, 5–12 per cell; (4) lobules ca. 1/4× leaf length, not reduced, ovoid and fully inflated, free margin incurved, tooth short, obtuse; (5) underleaves large, imbricate, 5–8× stem width, suborbicular, bifid to 1/3–1/2, lobes obtuse, margins entire, bases rounded, insertion curved; (6) male bracteoles present throughout the male spike; and (7) gynoecia on short branches with a usually sterile

542

 

Keys and Descriptions

innovation, perianth keels 2-winged, dentate-laciniate and undulate, beak short (Reiner-Drehwald 2010a). Lejeunea prionoides is morphologically close to L. ramulosa; both species share a finely papillose cuticle, male bracteoles present throughout the male spike, dentate-laciniate keels, and dioicy (Reiner-Drehwald 2010a). Lejeunea ramulosa differs from L. prionoides, however, in having more elongate, acuminate leaf tips (ending in a row of 2–3[−6] cells), smaller underleaves (2.5–4× stem width) with acute-acuminate tips, and perianth keels not undulate. Moreover, the shoots of L. ramulosa are fragile and easily fragmented. 45. Lejeunea pterigonia (Lehm. & Lindenb.) Mont. Fig. 53E Taxilejeunea pterigonia (Lehm. & Lindenb.) Schiffn. “pterogonia” Taxilejeunea densiflora A.Evans, syn. nov. Type: Jamaica, vicinity of Moody’s Gap, “near spring and waterfall,” 1908, E. G. Britton 938 [913? cf. isotype in NY] (YU n.v.) On bark, rock, rotten wood, and litter in montane rainforest, (300–)600–3400 m. Colombia: Boy, Cal, Cas, Cun, Hui, Mag, Nar, Qui, Ris, San (Gil 2626, UPTC), Tol, Val. Ecuador: Es, Ga, MS, Na, Pa, Pi, Tu, ZC. General distribution: widespread in tropical America. Lejeunea pterigonia is the most common member of the former genus Taxilejeunea and is recognized by (1) plants autoicous, pale whitish-green, often robust, stems long and slender, frequently pendent; (2) leaf lobes rather widely spreading, ovate-triangular, apex acute to apiculate, margins crenulate and frequently with 1–3 small, irregular teeth near the apex; (3) leaf cells thin-walled, trigones minute or well developed, intermediate thickenings absent or present, cuticle finely papillose (“punctate”) to smooth, oil bodies segmented, variable in number, 4–6 or about 10–16 per cell; (4) lobules very small, often reduced; (5) underleaves large, 5–7× stem width, imbricate, shallowly bifid to (1/5–)1/4–1/3, underleaf lobes frequently overlapping, tips acute-apiculate, margins crenulate, base with large auricles; and (6) gynoecia arranged in a row on short branches, perianths 5-keeled in the upper half, the keels with teeth-like projections near apex, beak well developed (Evans 1921 as Taxilejeunea pterogonia; Reiner-Drehwald 2000 as T. pterigonia). Lejeunea pterigonia resembles L. debilis, but the latter species is dioicous and mostly sterile; the leaf lobes in L. debilis have 1 prominent tooth near the apex and separated from the apex by a long and shallow, lunulate sinus; the cuticle is more densely papillose; and the perianth keels are smooth or crenulate, not toothed. Judging from the original description and illustration, Taxilejeunea densiflora, described from Jamaica and additionally recorded from Colombia (Evans 1921), is a synonym of L. pterigonia. 46. Lejeunea puiggariana Steph. On bark of trees in submontane rainforest, 600  m. Ecuador: MS (SchäferVerwimp et  al. 2013a). General distribution: scattered in tropical America (Central America, Ecuador, southeastern Brazil, northern Argentina).

Lejeuneaceae

543

Lejeunea puiggariana is a rare neotropical species of low elevation that is characterized by (1) plants autoicous and usually copiously fertile, 0.8–1.5 mm wide, yellowish-green; (2) leaf lobes widely spreading, ovate to ovate-oblong, plane, apex rounded, margins entire; (3) leaf cells with small trigones and intermediate thickenings, cuticle smooth, oil bodies homogeneous, small, numerous per cell; (4) lobules usually reduced, when well developed ca. 1/4× leaf length, inflated, oblong, free margin incurved, tooth obscure; (5) underleaves rather large, contiguous, 3–4× stem width, ± wider than long, bifid to 1/2, lobes acute, margins entire, bases rounded and somewhat cordate, insertion curved, with a large cell at the angles; and (6) gynoecia on short branches with vegetative leaves and with 1 frequently fertile innovation, perianth 5-keeled in the upper half, keels entire (Reiner-Drehwald 2000). Lejeunea puiggariana resembles L. flava in the ovate to ovate-oblong leaves, contiguous underleaves (3–4× stem width), and 1–2 gynoecia in a row but differs from L. flava in underleaves ± wider than long (longer than wide in L. flava), lobules mostly reduced, presence of a large cell at either side of the underleaf base, and homogeneous oil bodies. In a molecular analysis, these two species were unrelated (Heinrichs et al. 2013a); instead, L. puiggariana was sister to L. subsessilis. The latter species differs in crenulate leaf margins, granular oil bodies, and gynoecia single on very short branches, without vegetative leaves and with a sterile innovation. 47. Lejeunea pulverulenta (Steph.) M.E.Reiner Fig. 53F Taxilejeunea pulverulenta Steph. Type: Guadeloupe, l’Herminier s.n. (lectotype, designated by Reiner-Drehwald (2005c), G-00052572!) Taxilejeunea linguifolia Steph. (“linguaefolia”), syn. nov. Type: Guadeloupe, without collector (holotype G n.v.). Based on the description and illustration of the holotype by Jovet-Ast (1949), T. linguifolia and T. pulverulenta are fully identical. Taxilejeunea steyermarkii H.Rob. (Gradstein et al. 2018c) On bark, rotten wood, rock, and living leaves in submontane and lower montane rainforest and cloud forest, 700–1900 m. Colombia: Ris (Gradstein et al. 2018c). Ecuador: Pa (Reiner-Drehwald 2005c), ZC (Reserva Biologica San Francisco, Burghardt 6987, QCA, small phenotype). General distribution: Guadeloupe (type), northern Andes and Guyana; in addition recorded from Mexico by Fulford and Sharp (1990) as T. linguifolia; distribution insufficiently known. Lejeunea pulverulenta is a little-known neotropical species that is distinguished by (1) plants autoicous, 1–2 mm wide; (2) leaf lobes obliquely spreading, ovate, apex rounded, plane, margins entire or crenulate; (3) leaf cells elongate, with conspicuous trigones and intermediate thickenings, cuticle densely papillose, cells becoming rather suddenly smaller toward the leaf margin, forming a border of 5–6 rows of smaller marginal cells perpendicular to the larger central cells; (4) lobules less than 1/5× leaf length, with an elongate, straight or falcate tooth, keel curved, forming a sharp angle with the ventral margin of the leaf lobe; (5) underleaves imbricate, ca. 5–6× stem width, plane, suborbicular, bifid to 1/3–1/2, lobes shortly acuminate, apices of 2–3 cells in a row, bases strongly auriculate, insertion line deeply arched; and (6) gynoecia single or in a short row, perianth pyriform, keels ±2-winged, toothed or subentire (Reiner-Drehwald 2005c).

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Keys and Descriptions

By the rather large, strongly auriculate underleaves, the rounded leaf tips, the densely papillose cuticle, and the leaf cells becoming conspicuously smaller toward to the leaf margin, forming a border of 5–6 rows of smaller cells, Lejeunea pulverulenta is unmistakable. The species is genetically close to L. controversa (Heinrichs et  al. 2013a), but the latter species has much smaller underleaves, without large auricles, and a rounded to apiculate leaf apex. 48. Lejeunea quinqueumbonata Spruce Otigoniolejeunea quinqueumbonata (Spruce) Steph. Lejeunea cyrtotis Spruce (fide Bastos and Gradstein 2020) On tree trunks, ca. 600 m. Ecuador: MS (Herzog 1952). General distribution: scattered in tropical America. Lejeunea quinqueumbonata is an uncommon neotropical species that is recognized by (1) plants autoicous, ca. 1.2–2 mm wide, usually fertile; (2) leaves obliquely to widely spreading, ovate-triangular to ovate-oblong, apex rounded to subacute, margins strongly crenate; (3) leaf cells convex, bulging outward, rather large, without or with small trigones, intermediate thickenings absent, cuticle smooth; (4) lobules usually reduced, when well developed to maximally 1/4 of leaf length, tooth 1(−2) cells long, keel crenate; (5) underleaves distant, 2–4× stem width, bifid to 1/3–1/2, apices subacute, margins crenate, outer margins sometimes with a blunt tooth, bases cuneate to rounded, basal margin attached to the stem by a large cell; and (6) gynoecia single or in a short row on short branches without or with a few series of vegetative leaves, perianths with 5 short keels in the upper half, the keels terminating above in narrow lamella-like projections or cilia, perianth base sometimes stalk-like elongated (Reiner-Drehwald 2000). Lejeunea quinqueumbonata was originally described in Otigoniolejeunea but is placed in Lejeunea because of the absence of ocelli and pycnolejeuneoid innovations. Characteristics are the strongly crenate leaf and underleaf margins, the rounded to subacute leaf apices, and the perianth keels with a lamella-like projection. Lejeunea cyrtotis Spruce is identical to L. quinqueumbonata and a synonym. Lejeunea quinqueumbonata is closely similar to L. immersa, but the latter species has strongy crenate perianth keels without lamella-like projections or cilia. Lejeunea quinqueumbonata resembles L. raddiana in the strongly crenate leaf margins, but the latter species has falcate leaves with an acute and usually reflexed apex, larger lobules, small underleaves with rounded lobes, and perianth keels without lamellalike projection. In the gynoecia on very short branches, L. quinqueumbonata is similar to L. glaucescens and L. subsessilis, but the leaf and underleaf margins in the latter species are less crenate, the leaf apices are always rounded, the underleaves are smaller, and the perianth keels lack lamella-like projections. 49. Lejeunea raddiana Lindenb. Harpalejeunea pellucida Herzog Lejeunea gomphocalyx Spruce (fide Bastos and Gradstein 2020) On twigs and rotten wood in lowland and lower montane rainforests, up to 2000 m. Colombia: Boy, Hui (Campos and Uribe 2006). Ecuador: Es (Dauphin

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545

2718, GOET), ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical Andes, southeastern Brazil, Panama. Lejeunea raddiana is characterized by (1) plants autoicous and usually fertile, 0.7–1 mm wide, whitish-green when fresh, becoming brownish in herbarium; (2) leaf lobes wide-spreading, convex, ovate-falcate, apex plane broadly recurved, narrowly obtuse to acute to apiculate, margins ± crenate by mammillose cells that are usually transversely oriented to the margin, forming a border; (3) leaf cells thinwalled with minute trigones, intermediate thickenings absent, cuticle smooth; (4) lobules large, ca. 1/3–2/5× leaf length, inflated, free margin incurved, tooth slightly elongate, obtuse, keel arched, forming a sharp angle with the ventral margin of the leaf lobe; (5) underleaves very small, distant, 1–2× stem width, bifid to 2/5, tips rounded to obtuse; and (6) perianths subcylindrical, longly exserted, 5-keeled, keels 2-winged, wings entire or with a few teeth or cilia, usually bordered by a row of transversely oriented, subrectangular mammillose cells (cells higher than wide) (Reiner-Drehwald and Goda 2000; Reiner-Drehwald 2009b). The ovate-falcate leaves with acute tips, the tiny underleaves with rounded lobes and the crenate borders of perianth keels, and leaves made up of transversely oriented mammillose cells are the most characteristic features of L. raddiana. The species is close to L. acanthogona, L. immersa, and L. quinqueumbonata; for differences see under the latter species. Lejeunea gomphocalyx Spruce is identical to L. raddiana and a synonym. 50. Lejeunea ramulosa Spruce

Fig. 53G

On bark of trees, treelets, and shrubs, often pendulous, also on rotten wood, in primary and secondary montane rainforests, usually in rather open environments, (600–)1000–3300 m. Colombia: Ant, Boy, Cas, Ris, Tol. Ecuador: Ch, Lo, LR, MS, Na, Pi, Su, Tu, ZC. General distribution: scattered in tropical America. Lejeunea ramulosa is a rather widespread neotropical species that is characterized by (1) plants dioicous, 1–1.3 mm wide, whitish-green, stems fragile; (2) leaf lobes widely spreading, ovate-triangular to lanceolate, 1.5–2× longer than wide, apex acute-acuminate, terminating in a row of (1–)2–3(−6) cells, plane or recurved, margins entire; (3) leaf cells very thin-walled, without or with small trigones, intermediate thickenings few, cuticle finely punctate-papillose, oil bodies finely granular, 5–10 per cell; (4) lobules ca. 1/4× leaf length, ovoid and fully inflated, free margin incurved, tooth short, obtuse; (5) underleaves distant to subimbricate, 2.5–4× stem width, suborbicular, bifid to 1/3–1/2, lobes acute-acuminate, margins entire or with 1–2 teeth, bases rounded; (6) male bracteoles present throughout the male spike; and (7) gynoecia on short branches with innovation, perianth keels 2-winged, dentate-laciniate but not undulate-crisped, beak short (Reiner-Drehwald 2010a). The delicate, prostrate, very translucent whitish-green plants with wide-spreading ovate-lanceolate, acute-acuminate leaves are unmistakable. Lejeunea ramulosa is most similar to L. prionoides; for differences see under the latter species. 51. Lejeunea reflexistipula (Lehm. & Lindenb.) Lehm. & Lindenb.  Fig. 53H Amphilejeunea reflexistipula (Lehm. & Lindenb.) Gradst. Cryptogynolejeunea reflexistipula (Lehm. & Lindenb.) R.M.Schust. Dicladolejeunea reflexistipula (Lehm. & Lindenb.) R.M.Schust.

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Keys and Descriptions

On bark of trees in primary and secondary rainforests in rather open situations, mainly in the forest canopy and at forest margins, 150–2000(−3000) m. Colombia: Boy, Cas, Cau, Huila, Put, Ris, San, Tol, Val. Ecuador: Ca, Lo, MS, Na, Or, Pa, Pi, Tu, ZC. General distribution: tropical South America, Central America (Panama, Costa Rica). Lejeunea reflexistipula is a common tropical South American species that has been placed in at least six different genera. It is readily distinguished by the broad, reniform, undivided underleaves with usually recurved apex and deeply arched insertion, the crenate leaf margins, small lobules, finely papillose cuticle, and homogeneous oil bodies (12–30 per cell). Further characteristics are (1) plants dioicous, 1–2  mm wide, deep green, ventral merophyte 2 cells wide (4–6 cells wide in L. reflexistipula var. costaricensis [R.M.Schust.] M.E.Reiner); (2) leaf lobes widely spreading, ovate, apex broadly rounded; (3) cells with small trigones and intermediate thickenings; (4) lobules small, to 1/5× leaf length (or less), inflated, free margin incurved, tooth rounded to elongate, keel arched, smooth, forming a sharp angle with the ventral margin of the lobe; (5) underleaves (4–)6–8× stem width, undivided, apex usually recurved (sometimes plane), bases rounded to auriculate; and (6) perianth keels smooth (Reiner-Drehwald 2005a, 2009b). The material from Colombia and Ecuador belongs to L. reflexistipula var. reflexistipula, characterized by a 2-cell-wide ventral merophyte. Lejeunea reflexistipula may be confused with L. catinulifera; both species share undivided underleaves, homogeneous oil bodies, and dioicy. Lejeunea catinulifera, however, has larger lobules (1/3–1/2× lobe length) with 2 teeth, smaller underleaves (3–5× stem width) with a more shallowly curved insertion, and a smooth cuticle. 52. Lejeunea rionegrensis Spruce On bark in Amazonian rainforest and on isolated trees, preferably in rather open habitats, 200–800 m. Ecuador (new): MS (Bomboiza, growing in large, pure, pale green mats on trunk of isolated tree along Río Bomboiza, 1 Oct 2019, Pérez et al. 11570, QCA, PC), Or (Yasuni National Park, near Yasuni station, on tree trunk in valley, ca. 200 m, 2019, M. Berdugo et al. 1VV, QCA); to be expected in Colombia. General distribution: inner Amazonia (Venezuela, Brazil, Ecuador, Peru, Bolivia), SE Brazil. Lejeunea rionegrensis is a rare Amazonian species with caducous leaves that is recorded as new to Ecuador. The species resembles L. tapajosensis but differs from the latter in (1) leaf cells very thin-walled, with minute trigones and without intermediate thickenings; (2) underleaves 3–5× stem width, bases cordate-subauriculate; and (3) caducous leaves produced on flagelliform shoots (like in Rectolejeunea), smaller than ordinary leaves, suborbicular, with reduced lobules, falling off entirely (lobules not remaining on shoot), margins with short rhizoids and frequently with a regenerant shoot, underleaves on flagelliform shoots persistent or caducous. For a description and illustration of L. rionegrensis, see Reiner-Drehwald and SchäferVerwimp (2008b). The plants from Bomboiza were purely male, and bracteoles were present throughout the male spikes although 1–2 bracteoles were usually lacking in the middle of the spike. Oil bodies, which were not yet observed in the

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species, are 5–10 per leaf cell, small, rounded to elongate, coarsely granular (Pérez et al. 11570). 53. Lejeunea rotundifolia Mitt. Fig. 53I Dicladolejeunea rotundifolia (Mitt.) León-Yánez et al., nom. inval. Dicladolejeunea anomala R.M.Schust. Dicladolejeunea saccatiloba (Steph.) R.M.Schust. Taxilejeunea auriculata Steph., syn. nov.  – Ecuador, Morona Santiago, Gualaquiza, “in silva Churiacu, ad arborum corticum,” Sep 1909, Allioni s.n., Bryotheca Levier 6534 (lectotype, designated here, G-00064193!; Icon. Ined. 9997) On bark of trees and on humose soil in montane rainforests, 600–2900  m. Colombia: Bol, Cun, San, Val. Ecuador: Lo, MS, Na, Pa, Pi, Tu, ZC. General distribution: tropical Andes, Central America (Panama, Costa Rica). Lejeunea rotundifolia is a very distinct, robust Taxilejeunea-like species that is recognized by (1) plants dioicous, 1.5–3  mm wide, pale yellowish-green, stems robust, ventral merophyte (3–)4–6 cells wide; (2) leaf lobes widely spreading, convex, suborbicular, apex widely rounded, margins crenulate; (3) leaf cells thinwalled, trigones small, intermediate thickenings lacking, cuticle ± smooth; (4) lobules 1/4–1/3× lobe length, hidden behind the large underleaves, ovate, inflated along the keel, flattened toward the free margin, with 2 small teeth, keel crenulate, angle between keel and ventral margin of the leaf lobe sharp; (5) underleaves imbricate, 5–9× stem width, wider than long, shallowly bifid to 1/5(−1/4), lobes subacute, margins entire, bases with small or large auricles, the auricles overlapping or not; and (6) gynoecia on short or long branches with innovation, bracts entire, perianths cylindrical, longly 5-keeled, keels inflated and crenulate, apex depressed, beak very short (Reiner-Drehwald 2005b). Lejeunea rotundifolia may be confused with L. elongella but in the latter species the ventral merophyte is only 2 cells wide, the cell walls have intermediate thickenings, the gynoecia lack innovations and the perianth is terete. 54. Lejeunea ryszardii Gradst. & Ilk.-Borg. 

Fig. 65A_C

On base of large tree trunk in old secondary montane rainforest on steep slope, growing rather exposed in a dense Lejeuneaceae mat, 1900–2000 m. Colombia: Qui (Gradstein and Ilkiu-Borges 2019). General distribution: only known from the type from Colombia. Lejeunea ryszardii is a distinct species that is recognized by the strongly caducous, asymmetrically ovate-falcate, acuminate leaf lobes with a very narrow base and with irregularly crenate-toothed margins with scattered rhizoids and with a few irregular, blunt teeth terminating in a rhizoid. The leaf margin cells are mammillose and occasionally crowned by a small papilla and the lobules are small but never reduced, strongly swollen, flask-shaped, and with a long and curved, 30–40 μm long tooth. The stem epidermis is brownish and somewhat thick-walled. The plants are male and are probably dioicous like in most other Lejeunea species with caducous leaves or leaf lobes (Gradstein and Ilkiu-Borges 2019).

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Keys and Descriptions

Lejeunea ryszardii approaches the Caribbean L. paucidentata (Steph.) Grolle but the leaf lobes in the latter species are not caducous, the leaf margins in L. paucidentata have sharp teeth not terminating in a rhizoid and they lack marginal rhizoids and papillate mammillae, the lobule tooth is short and straight (15–25 μm long), and the stem epidermis is colorless, thin-walled (Reiner-Drehwald and Goda 2000). Moreover, L. paucidentata is autoicous and is usually copiously fertile. 55. Lejeunea schultesii Gradst., sp. nov.Fig. 54B Holotype: Colombia, Cundinamarca, páramo de Chisacá, southeast of Bogotá, ca. 3500 m [“11.000–12.000 feet”], on twigs among Metzgeria dorsipara, Jul 1953, R. E. Schultes 20212a, c. gyn. (PC-0774214) Plants dioicous, very small, 0.5–0.6  mm wide, very pale-colored, transparent. Stems very thin, ca. 50 μm in diameter, of 7 rows of epidermal cells and 3 rows of medullary cells. Leaves wide-spreading, subimbricate, convex and standing upward, subsquarrose, ovate-suborbicular, ca. 0.3 × 0.25–0.3 mm, apex rounded to subtruncate, ventral and dorsal margin arched, plane, crenulate, ventral lobe margin and upper part of the leaf tending to become subparallel to the stem. Leaf cells thinwalled, with minute trigones, intermediate thickenings lacking, cuticle smooth. Lobules ovate-oblong, ca. 1/2× leaf length, inflated, flattened to the apex, free margin plane, apex with a large, blunt, oblong tooth, the tooth ca. 40 × 25 μm, large disc cell present, elongate, extending from the tooth to the end of the free margin; keel arched, crenulate, forming a continuous line with the ventral margin of the leaf lobe. Underleaves imbricate, large, ca. 4× stem width, ovate, slightly longer than wide, ca. 0.25–0.3 × 0.2–0.23 mm, smaller on branches, bifid to 1/3–2/5, lobes triangular, apices narrowly rounded, margins entire, plane, bases cuneate, with a large cell in the basal angles at the junction with the stem, insertion slightly curved. Gynoecia on the main stem or on elongate branches, with 2 innovations, bracts and bracteole larger than vegetative leaves and underleaves, as long as or shorter than the perianth, with irregularly crenate margins, bract lobe apex rounded, lobule slightly shorter than lobe, apex obtuse, keel ca. 2/3× bract length, winged over much of its length, wing to 6 cells wide, wing margin irregularly crenate; female bracteole short-bifid. Perianths oblong to cylindrical, 0.7–1.0 × 0.25–0.35 mm, sometimes stalked at the base, sharply 5-keeled in the upper half, keels rudimentarily winged, beak 40–80 μm long, made up of 1–2 rows of narrowly elongate cells, beak cells 2–3× longer than wide. Vegetative reproduction not observed. Lejeunea schultesii is a small twig epiphyte that is known only from the páramo de Chisacá near Bogotá, where it was collected by Richard E. Schultes in the 1950s. The species is readily recognized by the very small plants with suborbicular, subsquarrose leaf lobes; very thin-walled leaf cells; large lobules with a prominent, oblong tooth; large, imbricate, bifid underleaves (ca. 4× stem width) with cuneate bases and rounded lobe tips; gynoecia with 2 innovations and winged bracts; and sharply 5-keeled perianths (keeled in the upper half) with a beak made up of 1–2 rows of narrowly rectangular cells. By its small size, large lobules, suborbicular leaf lobes which tend to become subparallel to the stem in the upper part, and the winged female bracts, L. schultesii is reminiscent of the genus Microlejeunea, but the

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Fig. 54 (A) Lejeunea mojandae, leaf and underleaf. (B) Lejeunea schultesii, leaf and underleaf. (C–E) Lejeunea yasuniensis. (C) Leaf and underleaf. (D) Underleaf, showing collapsed tip cell. (E) Habit, dorsal view. (F) Habit, ventral view. (A from the isotype, B–F from the holotypes). Scale bars: A, B = 250 μm; C, D = 100 μm; E, F = 500 μm

absence of ocelli and the very large underleaves separate it from Microlejeunea. The new species somewhat resembles L. catinulifera, but the latter differs in the much larger plant size (1–2.5  mm wide), lobules with 2 teeth made up of small cells, underleaves undivided or shallowly bifid (to maximally 1/4), female bract keels without wing, and a perianth mouth made up of small, quadrate cells.

550

 

Keys and Descriptions

56. Lejeunea serpillifolioides (Raddi) Gradst. Fig. 53J Taxilejeunea serpillifolioides (Raddi) D.P.Costa Taxilejeunea isocalycina (Nees) Steph. (Lejeunea isocalycina [Nees] Spruce) On bark of trees and shrubs and on rock, often in open environments, also along rivers and withstanding periodical inundation, (100–)900–2000 m. Colombia: San (Gradstein et  al. 2018c); probably more common in the country. Ecuador: EO (250–400  m), Pa, Tu, ZC (Nöske et  al. 2003). General distribution: tropical America. Lejeunea serpillifolioides is a common neotropical member of the former genus Taxilejeunea that is recognized by (1) plants autoicous, whitish-green, stems long and slender; (2) leaf lobes obliquely spreading, ± asymmetrically ovate, apex rounded to obtuse to obtusely apiculate, margins crenulate, without teeth (rarely with 1–2 small teeth); (3) leaf cells thin-walled, trigones minute or well developed, intermediate thickenings absent, cuticle smooth; (4) lobules very small, usually reduced; (5) underleaves large, imbricate, 5–7× stem width, bifid to 1/6–1/4 (measured from the middle of the underleaf insertion) with a narrow U-shaped incision, underleaf lobes frequently overlapping, tips acute, bases with very large auricles; and (6) gynoecia arranged in a row on innovating branches, bracts smaller than leaves, apices acute, margins crenulate, entire or with 1–2 small teeth near apex, perianths terete, mouth recessed, beak minute. In its terete perianths arranged in rows and the large, shallowly bifid underleaves L. serpillifolioides is similar to L. sulphurea. In the latter species, however, the leaves are always toothed and, more conspicuously so, with (2–)3–7 teeth near the apex. Since the difference between the two species is slight, L. sulphurea may be a variety of L. serpillifolioides; this needs study. In Ecuador Lejeunea serpillifolioides is very common along the Pastaza river and its tributaries, growing on small subshrubs along the rivers between 900 and 1600 m (Gradstein 12130, 12133, 12138, 12142, 12144, GOET). 57. Lejeunea setiloba Spruce On tree trunks, living leaves, rock, and soil in rather dry evergreen and semideciduous forests, from sea level to 1500 m. Colombia (new): Bol (det. M.E. ReinerDrehwald, GOET). Ecuador: Ga (Cerro Azul), ZC.  General distribution: tropical America. Lejeunea setiloba is an uncommon neotropical species that is distinguished by (1) plants autoicous, ca. 1–1.5  mm wide, pale yellowish-green; (2) leaves widespreading, somewhat convex, ovate to ovate-oblong, apex broadly rounded, margins entire; (3) leaf cells thin-walled with minute trigones and a smooth cuticle, oil bodies segmented; (4) lobules very small, with a long-linear tooth, 3–6 cells long, 1–2 cells wide, tipped by a slime papilla; (5) underleaves distant, 2–3× stem width, bifid to 1/2, incision wide, V- or U-shaped, lobes obtuse, bases cuneate, insertion line straight; and (6) gynoecia on short or long branches 1–3 in a row, perianths obpyriform, inflated, with short, crenate keels in the upper half and a short beak (ReinerDrehwald 2000). The species grows in rather dry environments. Lejeunea setiloba resembles L. trinitensis; for differences see under the latter.

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58. Lejeunea subsessilis Spruce Fig. 53M Type: Ecuador, Pastaza, Río Bombonasa, Spruce L557 (MANCH-000410!, autoicous) Lejeunea monimiae (Steph.) Steph. (Eulejeunea monimiae Steph.) (fide Bastos and Gradstein 2020) On living leaves or bark in lowland and montane rainforest, 250–1880  m. Colombia: Boy, Mag (?), Vau (Schultes & Cabrera 14429, 15275, CINC, PC). Ecuador: Ga (new: Santa Cruz, Fortuna, Howell 266, det. M.E. Reiner-Drehwald [GOET]), Pa (type), ZC (Parolly et al. 2004). General distribution: subtropical South America (southern Brazil, northern Argentina), northern Andes. Lejeunea subsessilis is manly distributed in southern Brazil and northern Argentina (Reiner-Drehwald 2000). The species has been recorded a few times from Colombia and Ecuador, but the record from Magdalena (Winkler 1976) is doubtful and needs verification. Lejeunea subsessilis is recognized by (1) plants autoicous, 0.7–1.1  mm wide, very delicate and translucent, growing tightly appressed to the substrate, often highly fertile with numerous androecia and gynoecia; (2) leaves obliquely to widely spreading, ovate with a rather narrow base, apex rounded to obtuse, margins crenulate; (3) leaf cells thin-walled with minute trigones, intermediate thickenings absent, cuticle smooth; oil bodies granular, 2–5 per cell (Reiner-Drehwald 2000); (4) lobules ovate, ca. 1/4× leaf length, with a short blunt tooth, reduced lobules usually with a larger, 1–4 cells long, erect tooth; (5) underleaves distant, 2–3(−4)× wider than the stem, wider than long, bifid to 1/2(−2/3), lobes obtuse, sinus between the lobes very wide, V- or U-shaped, margins crenulate and sometimes with a blunt short tooth, bases cuneate to slightly rounded; and (6) gynoecia on very short branches without vegetative leaves, with a short sterile innovation, perianth pyriform, 5-keeled but dorsal keel usually smaller, keels crenulate, somewhat expanded above (Reiner-Drehwald 2000 as L. monimiae). Lejeunea subsessilis is closely similar to L. glaucescens; for differences see under L. glaucescens. Lejeunea tonduzana (Steph.) M.E.Reiner (basionym, Hygrolejeunea tonduzana Steph.) from Costa Rica (Río Naranja, on fern leaf, Tonduz 3077a, G) resembles L. subsessilis but differs in leaf cells with distinct intermediate thickenings, smaller suborbicular underleaves (ca. 2× stem width) with acute lobes, and perianths keels conspicuously inflated and expanded in the upper half. Moreover, the dorsal perianth keel in L. tonduzana is not smaller than the other keels. Interestingly, L. tonduzana was also described by Stephani as Taxilejeunea tonduzana Steph. (syn. nov.) based on two different Tonduz collections (15612, 15614, G) from the same locality (fide Icones Ined. 10126). Lejeunea seriata Lindenb. & Gottsche from Mexico (Type: Mexico, Veracruz, Mirador, Liebmann 236 p.p., isotype C-M9875!) approaches L. tonduzana in the presence of intermediate thickenings, the large dorsal perianth keel, etc., but the perianth keels in L. seriata are not expanded above and the innovations are usually fertile, with the gynoecia being arranged in pairs on short branches.

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59. Lejeunea subspathulata Spruce Echinocolea subspathulata (Spruce) Grolle

 

Keys and Descriptions

Fig. 53K, L

On bark, rotten wood, soil, and living leaves in primary lowland and submontane rainforest, 400–1000(−3000?) m. Colombia: Cho, Val. Ecuador: Na (SchäferVerwimp et al. 2013a), Pa (type). General distribution: widespread in tropical America. Lejeunea subspathulata (sect. Echinocolea) is a widespread neotropical species from lowland and submontane rainforests, up to ca. 1000 m; records from above 2000 m (Colombia) might refer to L. meridensis and should be verified. Lejeunea subspathulata is related to L. asperrima but differs from the latter in larger plant size (0.5–1.0 mm wide), being about twice the size of L. asperrima, the smooth keel and lower dorsal surface of the leaves, and the perianth with cilia and lacinia near apex. In L. asperrima the whole dorsal leaf surface and keel are strongly crenatedenticulate, and the perianth lacks cilia or lacinia (Ilkiu-Borges 2005). 60. Lejeunea suffruticola Spruce Type: Ecuador, hacienda Guataxi, “Andium Quitensium, in suffruticibus,” Spruce L26 (lectotype, designated here, MANCH-000411!; isolectotype MANCH-000412!) On small subshrubs (Malvaceae, Solanum, etc.), 2200 m. Ecuador: Ch (type). General distribution: only known from the type from Ecuador. Lejeunea suffruticola is a very little-known Ecuadorian species that is recognized by rather large, bifid underleaves (4–5× stem width), rounded leaf tips, and lobule tooth 2–3 cells long. Other characters are (1) plants dioicous, pale green, 1–1.5  mm wide, delicate, irregularly branched, most branches rather short and small-leaved; (2) leaves distant to subimbricate, widely spreading, broadly ovate, margins ± crenulate (?); (3) leaf cells very thin-walled, trigones minute, intermediate thickenings absent, cuticle smooth; (4) lobules small, 1/5–1/4× leaf length, ovate-subquadrate, slightly inflated, tooth conspicuous, 2–3 cells long, 2–3 cells wide at the base; (5) underleaves distant, 4–5× stem width, ovate, about as long as wide, bifid to 1/2 with a wide incision, apices acute or obtuse, bases cuneate to rounded, insertion line shallowly curved; and (6) gynoecia on elongate branches with 1(−2) innovations that remain sterile or, occasionally, become again fertile, bracts and bracteoles entire, perianth (Spruce 1884) obovate, 5-keeled, keels high and smooth, apex rounded to retuse, beak short. 61. Lejeunea sulphurea (Lehm. & Lindenb.) Spruce Fig. 53P Taxilejeunea sulphurea (Lehm. & Lindenb.) Schiffn. Taxilejeunea renistipula (Lindenb.) Steph. (Omphalanthus renistipulus Lindenb.) (?) On bark, rotten woodrock, and soil in humid montane forests and scrub, 400–3200  m. Colombia: Ant, Boy, Cho, Hui, Ris, San. Ecuador: MS, Pi, Tu (Spruce 1884 as L. renistipula and L. sulphurea). General distribution: tropical America.

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Lejeunea sulphurea is a common neotropical member of the former genus Taxilejeunea that is recognized by (1) plants lax, often pendent, whitish-green; (2) leaf lobes broadly ovate-triangular, margins crenate and with 2–7 teeth near apex (teeth 1–3 cells long), apex obtuse to apiculate to shortly acuminate; (3) underleaves very large, imbricate, rounded, very shallowly bifid, strongly auriculate; and (4) gynoecia arranged in a row on innovating branches, perianths terete, usually small and somewhat broadened at the apex, with or without beak, apex irregularly crenate by mammillose cells. Most of the collections seen had perianths without beak, but in some specimens the perianth beak was well developed. The taxonomic significance of this variation needs further study. Lejeunea sulphurea closely resembles L. serpillifolioides, but in the latter species, the leaf margins are usually entire (occasionally with 1–2 small teeth near apex). Since the differences between the two species are slight, L. sulphurea may be a variety of L. serpillifolioides. Taxilejeunea renistipula, described from the West Indies and recorded from Ecuador by Spruce (1884), is probably a synonym of L. sulphurea (type not seen). 62. Lejeunea tapajosensis Spruce On bark of trees in lowland and montane rainforests, preferably in rather open habitats, from sea level to 2150  m. Colombia: Ris (Gradstein et  al. 2018c). Ecuador: Es, Or, ZC.  General distribution: tropical South America, West Indies, Panama. Lejeunea tapajosensis is a rather common neotropical species with caducous leaf lobes that is distinguished by (1) plants 1–1.5 mm wide, green, dioicous; (2) leaf lobes ovate, plane, apex widely rounded, margins entire; (3) leaf cells with conspicuously elongate, triradiate trigones and intermediate thickenings, 1(−2) per wall, cuticle smooth; (4) lobules very small, ca. 1/5× leaf length, inflated, free margin incurved, apical tooth obtuse; (5) underleaves contiguous to imbricate, 3–4× stem width, ± wider than long, bifid to 1/2, lobes acute, margins entire, bases broadly rounded-quadrate, insertion line curved; (6) androecia with underleaves present throughout the male spike; (7) perianth obovate, with 4–5 long, smooth keels; and (8) vegetative reproduction via caducous leaf lobes common (Reiner-Drehwald 2000, 2009b). Lejeunea tapajosensis resembles L. phyllobola, but the latter species is smaller in size (ca. 0.6–1 mm wide), the underleaves of L. phyllobola are smaller, and the trigones are simple-triangular, not radiate. Lejeunea rionegrensis Spruce, recorded from the Amazon region of Brazil, Venezuela, Peru, and Bolivia and not yet recorded from Colombia and Ecuador, differs from L. tapajosensis in the thin-walled leaf cells with small trigones and without intermediate thickenings and in producing caducous whole leaves, with or without lobules (Reiner-Drehwald and SchäferVerwimp 2008b). Bastos & Gradstein (2020) showed that the correct name for L. tapajosensis is L. parviloba Ångstr.

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Keys and Descriptions

63. Lejeunea topoensis Gradst. & M.E.Reiner On twigs and branches in rivers, usually growing on the woody stems of the small subshrub Cuphaea bombonasae and periodically submerged, 1000–1600 m. Ecuador: Tu (Río Topo), ZC (Río Nangaritza; Gradstein and Reiner-Drehwald 2007); to be expected in Colombia. General distribution: Ecuador, southern Brazil (Santa Catarina); in Ecuador only known from the Topo and Nangaritza rivers. Lejeunea topoensis is a rare neotropical rheophyte with robust, rigid stems with a 4–8-cell-wide ventral merophyte and pinnate branching, copiously fertile, with numerous male and female branches. These features are interpreted as adaptations to the rheophytic habitat and are the distinctive characters of Lejeunea subg. Neopotamolejeunea (M.E.Reiner) Gradst. & M.E.Reiner, a small neotropical group of rheophilous species (Gradstein and Reiner-Drehwald 2007). Molecular evidence, however, does not support recognition of this subgenus (Heinrichs et  al. 2013a). Further characters of L. topoensis include (1) plants autoicous, 1.1–2.2 mm wide, ascending from the substrate; (2) leaves obliquely to widely spreading, ovate-triangular, apex narrowly rounded to obtuse, margins entire; (3) leaf cells thin-walled with minute trigones and without intermediate thickenings, cuticle smooth; (4) lobules usually reduced (short-rectangular and to 1/4× leaf length when well developed); (5) underleaves distant, rather small (and often damaged), 1–2.5× stem width, narrowly bifid to 2/5, with obtuse lobes and cuneate bases; and (6) gynoecia in long rows on short innovating branches, perianth with 5 smooth keels and a short beak. Lejeunea topoensis is closely similar to L. juruana Gradst. & M.E.Reiner and L. polyantha Mont. from the Brazilian Amazon, two further members of subg. Neopotamolejeunea, but the latter two species have longer leaf lobe insertions, almost as long as the leaf width. Heinrichs et al. (2013a) found that L. topoensis is sister to L. debilis, but morphologically these two species are very different. Lejeunea topoensis is known in Ecuador from only two localities and should be more widespread; its distribution resembles that of Colura irrorata. At the Río Topo, L. topoensis and C. irrorata may grow mixed. 64. Lejeunea trinitensis Lindenb. Fig. 53N, O Lejeunea pililoba Spruce (Stylolejeunea pililoba [Spruce] A.Evans) Lejeunea prorepens Spruce On living leaves, bark, rotten logs, or shaded rock in rather dry evergreen woodlands, degraded rainforests, and plantations, from sea level to 800  m. Colombia (new): Cun (along road Bogota – Honda, 1300 m, 15 Jul 1980, Gradstein 3501, det M.E. Reiner-Drehwald, U). Ecuador: Es (Burghardt 2019), Ga (Gradstein 2009). General distribution: peri-Afroamerican (tropical America, tropical Africa: Comores); the species is particularly common in the West Indies, southeastern Brazil, and northern Argentina, less frequent elsewhere. Lejeunea trinitensis is a widespread species that is readily distinguished by the long-linear lobule tooth, about 6–11 cells long and 1–2 cells wide and tipped by a hyaline papilla. Further characters of the species are (1) plants autoicous, ca. 1–1.5  mm wide; (2) leaves wide-spreading, very flat, suborbicular, apex broadly

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rounded, margins entire; (3) leaf cells very thin-walled, trigones minute, intermediate thickenings lacking, cuticle smooth; (4) lobules very small; (5) underleaves distant, orbicular, 2–3× stem width, bifid to 1/2–3/5, with a wide V-shaped incision, lobes narrowly triangular, tips of 1–3 cells in a row, margins entire, bases cuneate; and (6) gynoecia on long shoots with 1–2 innovations, single or in a short row, perianths somewhat flattened, obcordate, longly 5-keeled, lateral and ventral keels very wide, dorsal keel small, keels smooth, beak short (Reiner-Drehwald 2000). Lejeunea trinitensis shares a long lobule tooth with L. setiloba, but in the latter species, the leaves are more elongate and convex, the tooth is only 3–6 cells long, and the perianths are inflated, obpyriform, and narrowly keeled in the upper half, with crenate keels. Both species grow in rather dry environments and have been very little collected in the northern Andean region and are not known from Colombia. 65. Lejeunea xiphophylla (Herzog) M.E.Reiner Taxilejeunea xiphophylla Herzog On bark in submontane rainforest, 750 m. Ecuador: LR (Hacienda Clementina, Samama Forest Reserve, type; Herzog 1952), Su. General distribution: only known from the foothills of the Andes in western Ecuador. Lejeunea xiphophylla is a rare Ecuadorian endemic that is characterized by (1) plants autoicous, 1.5–2 mm wide, whitish-green; (2) leaf lobes obliquely to widely spreading, ovate-lanceolate, 3–4× longer than wide, apex acute-acuminate, plane, margins entire; (3) leaf cells with small trigones and intermediate thickenings, cuticle densely and finely papillose; (4) lobules very small, 1/8–1/6× leaf length, tooth short, obtuse; (5) underleaves distant, ca. 2× stem width, longer than wide, deeply bifid to 2/3–3/4, lobes lanceolate, acute-acuminate, margins entire, bases cuneate; (6) androecia very small, male bracteoles restricted to the base of the male spike; and (7) gynoecia in a short row on short branches, perianth obovoid, with 5 smooth keels in the upper half, beak well developed (Herzog 1952). Lejeunea xiphophylla resembles L. osculatiana in habit and leaf shape but differs in the narrower underleaves, which are ca. 2× wider than the stem, the densely papillose cuticle, the gynoecia usually arranged in a row, and the keeled perianths. 66. Lejeunea yasuniensis Gradst. & C.Bastos, sp. nov.Fig. 54C_F Type: Ecuador, Orellana, Yasuni National Park, 0°40′S 76°24–28′W, lowland rainforest in valley, 200–230 m a.s.l., on tree trunks at ca. 2–12(−20) m height, creeping scattered over Ceratolejeunea, Plagiochila or Symbiezidium, March– April 2019, M. Berdugo, J. Deleg, L. Guérot & K. Suarez 10VV-22VV, c. gyn. (holotype PC-0763782; isotype QCA) Plants dioicous (?) (connection between male and female plants not seen), very small, 0.5–0.7  mm wide, whitish-green, transparent, little branched. Stems 40–50 μm in diameter, of 7 rows of epidermal cells and 3 rows of medullary cells. Leaves suberect, distant to contiguous, lobes asymmetrically obovate, ca. 0.3 × 0.2 mm on stem leaves, smaller on branches, sometimes reduced to an obtrapezioid to narrowly triangular lamina less than 0.1 mm long; upper half of the lamina of well-developed lobes flat, lower half (opposite the lobule) convex, margins

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plane and conspicuously crenate by rounded projecting cells (sometimes conical on reduced leaves), ventral margin short and curved, dorsal margin strongly arched and somewhat angulate. Leaf cells mostly isodiametrical-hexagonal, mammillose, slightly large in the upper lamina than in the lower lamina, ca. 20–35 μm in largest diameter in midleaf and at the base, marginal cells slightly smaller, 15–20 μm, all cells thin-walled, without or with minute trigones, intermediate thickenings lacking, cuticle smooth; oil bodies not observed. Lobules sometimes reduced, when well developed ovoid and almost fully inflated, ca. 0.12–0.15 × 0.1 mm, 1/2× leaf length, on branch leaves sometimes almost as long as the whole leaf (due to reduction of lobe length), free margin incurved and appressed to the lobe, apex of lobule incurved or flat, with a 1-celled tooth, the tooth oblong, ca. 25 μm long; keel arched, crenate, forming a rather sharp angle with the ventral margin of the leaf lobe, sinus indented. Underleaves small, 1–2.5× stem width, distant, ovate-obovate, as long as wide or longer than wide, shallowly bifid to 1/3–2/5, undivided lamina 3–4 cells high, lobes triangular, terminating in a fragile, rounded tip cell, the tip cell sometimes broken or collapsed, underleaf margins irregularly crenulate to crenate, plane, bases cuneate, with a larger cell in the basal angles at the junction with the stem, insertion line curved. Rhizoids absent. Androecia on short specialized branches, 2–7 pairs of bracts, the bracts strongly swollen and imbricate, hypostatic, bract keel crenate and with a conspicuous, 1-cellwide wing of large hyaline, outward bulging cells, bracteoles limited to the base of the male spike. Gynoecia on a short or elongate branch, with 1 sterile or fertile innovation, 1–4 gynoecia in a row; female bracts and bracteole much larger than vegetative leaves and underleaves, bract oblong, 0.4–0.5 × 0.2 mm, apex obtuse, margin irregularly crenate, lobule ca. 4/5× lobe length, keel about 3/4 of lobe length, unwinged, margin irregularly crenate; bracteole narrowly oblong-rectangular, about as long as the bract, bifid to 1/4, margin crenulate. Perianth (only one seen) obpyriform, longly exserted beyond the bracts, ca. 1.0 mm long, gradually narrowed to the base, sharply 5-keeled in the upper half, keels irregularly crenulate, beak short, 2 cells long, beak cells subquadrate. Vegetative reproduction not observed. Lejeunea yasuniensis is thus far only known from Yasuni National Park, Ecuador, where it grows on trees in the understory and canopy of lowland rainforest, usually creeping over other bryophytes. The species is recognized by its very small size (0.5–0.7 mm wide), the suberect leaves with large lobules 1/2× leaf length on stem leaves (lobules subequal to lobes on branch leaves), and the strongly crenate leaf margins and keel. Sometimes the leaves are reduced to a small fold. A very unusual feature of the species is the winged male bracts, which have not yet been reported in neotropical Lejeunea and are characteristic of the genus Prionolejeunea (IlkiuBorges 2016). By its branch leaves with subequal lobe and lobule, L. yasuniensis may be confused with Microlejeunea, but the leaves of Microlejeunea have ocelli and lobules more than 1/2× leaf length on main stems. Other features of the new species that may cause confusion are the reduced leaves with conically projecting cells

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resembling those of Prionolejeunea muricatoserrulata (but the latter species has very different mature leaves and is less pale-colored) and the underleaves with collapsed tip cells reminding of L. aphanes. The leaves of L. aphanes, however, are more widely spreading, the lobules smaller, and the leaf margins entire. Overall, L. yasuniensis is perhaps closest to the West Indian L. sporadica Besch. & Steph., but the leaves in the latter species are ovate-falcate and more widely spreading, the lobules smaller, the keels ± smooth, the male bracts unwinged, and the perianth keels narrowly 2-winged and ciliate. Further Records Lejeunea apiculata (Gottsche) Steph., nom. illeg. (Omphalanthus apiculatus Gottsche, Taxilejeunea apiculata [Gottsche] Steph.) – Colombia: Cun (Gottsche 1864, type); Ecuador: Su (Arnell 1962). The type is lost; specimens in the Stephani herbarium (G) identified as “T. apiculata” are L. pallescens or L. pterigonia. Lejeunea biformis Gottsche – Colombia: Tol (Gottsche 1864). The type has not been located. Lejeunea diaphana Spruce (Harpalejeunea diaphana [Spruce] Steph.)  – Ecuador: Tu (Spruce 1884). The type is lost (Grolle and Reiner-Drehwald 1999). Lejeunea dictyocalyx Spruce (Taxilejeunea dictyocalyx [Spruce] Steph.) – Ecuador: Ga (Gradstein 2009, “cf.”), Tu (Spruce 1884, type). The type has not been located. Lejeunea fusagasugana Gottsche – Colombia: Cun (Gottsche 1864, type). The type has not been located. Lejeunea hyalina (Steph.) L.Söderstr. & A.Hagborg (Pycnolejeunea hyalina Steph.)  – Colombia: Cundinamarca, Tequendama, 2500  m, Lindig 1716 p.p. (G-00128273!). The type material is too scanty to permit identification; the plant resembles L. capensis, but the cuticle is fully smooth, the leaf apex is obtuse, the leaf cells are very small, and the apical margin of the lobule has a large, rounded disc cell. Lejeunea hygrophila Gottsche – Colombia (Gottsche 1864, type). The type has not been located. Lejeunea macrorhyncha Spruce (Taxilejeunea macrorhyncha [Spruce] Steph.) – Colombia: Val (Herzog 1955 as T. macrorhyncha). The type (Peru, Tavalosos, 800 m, Spruce s.n.) has not been examined. Lejeunea semiscabrida Gottsche – Colombia: Mag (Gottsche 1864). The possible isotype in NY has not been examined. Lejeunea sporadica Besch. & Steph. (= L. elliottii Spruce = Echinocolea prionocalyx [Schiffn.] R.M.Schust.) – Colombia: Mag (Winkler 1976). Caribbean species (Schäfer-Verwimp and Reiner-Drehwald 2009); the record from Colombia is dubious and should be checked. Taxilejeunea cuervi (Gottsche) Steph. (Omphalanthus cuervi Gottsche)  – Colombia (Gottsche 1864, type). The type has not been located.

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Keys and Descriptions

Excluded Records Hygrolejeunea grossereticulata Steph.  – Colombia: Mag (Winkler 1976). Described from Cuba, considered a doubtful name by Uribe and Gradstein (1998) Lejeunea ambigua Lindenb. & Gottsche – Colombia: Ris (Wolf 1993). Described from Mexico, considered a doubtful name by Gradstein and Uribe (2016a) Lejeunea cladogyna A.Evans – Ecuador: Ga (Clark 1953). North American species; the records from the Galápagos Islands have been rejected (Gradstein 2009). One of the specimens cited by Clark (1953), Howell 266 from Santa Cruz (Fortuna), belongs to Lejeunea subsessilis Spruce (det. M.E. Reiner-Drehwald). Lejeunea caracensis Lindenb. (Taxilejeunea caracensis [Lindenb.] Schiffn.)  – Colombia: Mag (Winkler 1976 as T. “caracasensis”). Considered a doubtful record by Gradstein and Uribe (2016a). The type has not been found (not in W or G). Lejeunea diversicuspis Spruce  – Colombia: Ris (Wolf 1993 as L. megalantha Spruce). Rare northern Amazonian species, close to L. sporadica (Reiner-Drehwald and Goda 2000 as L. erostrata; Reiner-Drehwald et  al. 2018); the material from Risaralda, from montane forest, is presumably misidentified. Lejeunea grossitexta Steph.  – Ecuador: ZC (Parolly et  al. 2004). The plants belong to L. acanthogona (fide M. E. Reiner-Drehwald). Lejeunea hahnii Solari (= Microlejeunea grandistipula Steph.) – Colombia: Mag (Winkler 1976). A Patagonian species, not known from the tropics (Bischler et al. 1963). The Colombian material is presumably misidentified. Lejeunea remotifolia Steph. – Colombia: Cho (Winkler 1970). Described from the Guianas, considered a doubtful name by Gradstein and Uribe (2016a) Lejeunea smaragdina Besch. & Spruce  – Ecuador: ZC (Parolly et  al. 2004). Doubtful Caribbean species (Schäfer-Verwimp and Reiner-Drehwald 2009); the record from Ecuador needs verification. Taxilejeunea dissitifolia Steph. – Colombia: Boy, Mag (Winkler 1976). Dubious record; the type (Guadeloupe, Duss s.n.) and the Colombian specimen have not been seen. Taxilejeunea herzogiana Steph.  – Colombia: Mag (Winkler 1976). Dubiuous species from Bolivia, not accepted in the World Checklist (Söderström et al. 2016); the Colombian specimen is probably misidentified. Lepidolejeunea R.M.Schust. Plants small- to medium-sized, 1–4 cm long, 0.6–2 mm wide, yellowish-green to pale brown, loosely creeping to pendent. Stems of thin-walled cells, with hyalodermis; ventral merophyte 2 cells wide (rarely 4–10 cells wide). Leaf lobes widespreading, with scattered ocelli, apex rounded to acute-apiculate, plane or recurved, margins entire to crenulate to ± toothed. Cells plane, hexagonal, small, ca. 15–25(−30) μm long in lobe middle, walls thin, uniformly thickened or with small trigones, cuticle smooth or finely papillose; oil bodies lacking in green cells; ocelli numerous, colorless (sometimes pale brown in herbarium material), scattered in leaves, underleaves, bracts, bracteoles and perianths. Lobules ca. 1/5–1/3 leaf length, ovate, free margin usually incurved, tooth 1-celled, short-obtuse to

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somewhat elongate-acute, hyaline papilla proximal. Underleaves bifid, occasionally undivided, 3–8× stem width, bases usually cordate, insertion line arched or almost straight. Androecial bracteoles restricted to the base or present throughout the spike. Gynoecia on elongate shoots, with 1–2 pycnolejeuneoid innovations. Perianths inflated, 5-keeled, the keels sometimes conspicuously swollen above. Sporophytes lejeuneoid. Vegetative reproduction occasionally, by caducous leaf lobes (in L. involuta). A pantropical genus of 13 species, 7 in Colombia and Ecuador. The main features of Lepidolejeunea are (1) stems usually fragile, of thin-walled cells; (2) leaf apex acute to apiculate (occasionally obtuse); (3) leaf margins usually crenulate; (4) ocelli numerous in leaves and underleaves, scattered; (5) leaf cells small, hexagonal, often without trigones, without oil bodies; (6) underleaves bifid or ± undivided; and (7) perianth keels often swollen above, with small auricles. Literature: Piippo (1986), Heinrichs et al. (2015) 1. Underleaf apex truncate to shallowly emarginate . . . . . . . . . . . . . . . . . . . . . . 2 1. Underleaf apex clearly bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Ventral merophyte 4–6(−10) cells wide. . . . . . . . . . . . . . . . . . . . . . . . . L. eluta 2. Ventral merophyte 2 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Underleaf bases with large auricles. Ocelli larger than adjacent cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. auriculata 3. Underleaf bases without auricles. Ocelli smaller than adjacent cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sullivantii 4. Underleaves imbricate, 5–6× stem width, bases auriculate. . . . . . . . . . . . . . . 5 4. Underleaves distant to contiguous, 3–4× stem width, bases cuneate to rounded (not auriculate). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5. Leaf apex rounded to obtuse. Leaf cells with trigones. Plants of Chocó . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. grossepapulosa 5. Leaf apex acute. Leaf cells without trigones . . . . . . . . . . . . . . . . . L. cordifissa 6. Ocelli smaller than adjacent cells . . . . . . . . . . . . . . . . . . . . . . . . . L. cuspidata 6. Ocelli about as large as adjacent cells . . . . . . . . . . . . . . . . . . . . . . . L. involuta 1. Lepidolejeunea auriculata Schäf.-Verw. & Heinrichs On bark in upper montane forests, 2770–2870 m. Colombia: Ris. Ecuador: Lo (type). General distribution: northern Andes (Colombia, Ecuador). Lepidolejeunea auriculata is a little-known northern Andean species that is only known from two collections (Heinrichs et al. 2015). The species resembles L. sullivantii in the large underleaves with ± undivided apex and 2-cell-wide ventral merophyte but differs in ocelli larger than adjacent cells (smaller than adjacent cells in L. sullivantii) and underleaf bases with large auricles (without auricles in L. sullivantii). 2. Lepidolejeunea cordifissa (Taylor) M.E.Reiner Fig. 55A–D Kingiolejeunea ornata H.Rob. (Lepidolejeunea ornata [H.Rob.] R.M.Schust.) Lejeunea pentotantha Spruce

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Keys and Descriptions

Fig. 55 (A–D) Lepidolejeunea cordifissa. (A) Habit (ocelli shown in part of the plant). (B) Leaf. (C) Underleaf. (D) Median leaf cells. (E and F) Lepidolejeunea cuspidata. (E) Median leaf cells. (F) Habit. (G and H) Lepidolejeunea grossepapulosa. (G) Habit (ocelli not shown). (H) Underleaves (ocelli not shown in one of the underleaves). (I and J) Lepidolejeunea eluta. (I) Habit (ocelli not shown). (J) Median leaf cells. (K–N) Lepidolejeunea involuta. (K) Leaf apex. (L) Habit (ocelli not shown). (M) Lobule. (N) Leaf. (E, F redrawn from Heinrichs et al. 2015; G–J redrawn from Piippo 1986; rest from Gradstein et  al. 2001; Gradstein and Ilkiu-Borges 2009). Scale bars: A, H, L = 500 μm; B, C = 250 μm; D, K, M = 25 μm; E = 50 μm; F = 750 μm; G, I = 1000 μm; J, N = 100 μm

Lejeuneaceae

561

On bark and living leaves in lowland and lower montane rainforests, from sea level to 1600 m. Colombia: Cho, Put, Ris. Ecuador: Ecuador: Bo, Ch, Es, LR, MS, Na, Or, ZC. General distribution: northern South America, Costa Rica. Lepidolejeunea cordifissa is a distinct neotropical species that is recognized by (1) leaves with acute, recurved apices and crenulate margins; (2) leaf cells thinwalled without trigones; (3) ocelli similar in size to adjacent cells; and (4) underleaves large, imbricate, 5–6× stem width, bases cordate-auriculate. The species may be confused with L. involuta and L. cuspidata, but the latter two species have smaller, distant to contiguous underleaves (3–4× stem width) with cuneate to rounded bases (not auriculate) and apiculate leaf tips. 3. Lepidolejeunea cuspidata (Gottsche) Heinrichs & Schäf.-Verw. Fig. 55E, F Lejeunea cuspidata Gottsche Hygrolejeunea punctata Herzog (Lepidolejeunea punctata [Herzog] R.M.Schust.) On bark in lowland and montane rainforest, 200–2000 m. Ecuador: Or (Parque Nac. Yasuni, near Yasuni Biological station, Gradstein, Deleg & Bader s.n., HUTPL), Tu, ZC (Heinrichs et al. 2015). General distribution: tropical America. Lepidolejeunea cuspidata resembles L. involuta but differs in ocelli distinctly smaller than adjacent cells. Although differing in only one morphological character, L. cuspidata and L. involuta are separate lineages and should therefore be considered different species, as shown by Heinrichs et  al. (2015). The distribution of L. cuspidata is insufficiently known (see under L. involuta); the species is not known from Colombia but is expected there. The plants from Yasuni National Park stand out by apiculate or rounded leaf apices. 4. Lepidolejeunea eluta (Nees) R.M.Schust.  Fig. 55I, J Hygrolejeunea devoluta (Spruce) Steph. Hygrolejeunea pastasensis (Spruce) Steph. Hygrolejeunea planiloba Steph. Pycnolejeunea granatensis Steph. (Perilejeunea granatensis [Steph.] H.Rob.) On bark in lowland and lower montane rainforests, from sea level to 1800 m. Colombia: Cho, San, Tol. Ecuador: MS.  General distribution: tropical America. Lepidolejeunea eluta is a distinct neotropical species that is readily separated from the other members of the genus by the rigid stems with a broad, 4–10-cellwide ventral merophyte. The leaf apices are acute and frequently reflexed, the leaf margins are entire, ocelli are numerous in leaves and underleaves, and the underleaf apices are very shallowly bifid (to 1/8 of underleaf length). A unique feature of L. eluta is that herbarium material, when soaked in water, turns rose-reddish in color, especially near the stem apices; this has not been reported in any other member of Lejeuneaceae. The chemical basis of the color reaction is unknown.

562

5. Lepidolejeunea grossepapulosa (Steph.) Piippo Hygrolejeunea obtusiserrulata Herzog

 

Keys and Descriptions

Fig. 55G, H

On bark in wet lowland and lower montane rainforests along the Pacific coast, from sea level to 1500 m. Colombia: Cho, Ris, Val. General distribution: western Colombia, Guadeloupe. Lepidolejeunea grossepapulosa resembles L. cordifissa in the large, cordateauriculate underleaves but differs in the rounded to obtuse leaf apices (acute in L. cordifissa) and leaf cells with trigones (trigones absent in L. cordifissa). The species is common in the Chocó. 6. Lepidolejeunea involuta (Gottsche) Grolle  Lejeunea torticuspis Spruce

Fig. 55K–N

On bark, leaves, rock, and soil in lowland and montane rainforests, plantations, and isolated trees, from sea level to 3000 m. Colombia: Caq, Cho, Hui, Qui, Put, Ris, Vau. Ecuador: Ga, MS, Or, Pa, Tu, ZC.  General distribution: tropical America. Lepidolejeunea involuta is a common neotropical species that is distinguished by (1) plants delicate, ca. 1 mm wide, pale green; (2) leaves ovate-falcate, apex apiculate, margins crenulate-denticulate; (3) leaf cells small, subquadrate, without trigones; (4) ocelli numerous in leaves and underleaves, about as large as adjacent cells; and (5) underleaves 3–4× stem width, bases cuneate to rounded, insertion line straight to curved. Heinrichs et al. (2015) showed that L. involuta is genetically heterogeneous and consists of two well-defined species, L. involuta and L. cuspidata. Differences are shown in the key. Both species seem to be widespread in the neotropics, but their precise distributions remain unknown because the herbarium collections of L. involuta (Piippo 1986 with updates) have not been revised. The above distribution of L. involuta in Colombia and Ecuador refers to the species in the broad sense, and some records may belong to L. cuspidata. 7. Lepidolejeunea sullivantii (Gottsche) M.E.Reiner Hygrolejeunea sullivantii (Gottsche) Steph. Lepidolejeunea spongia (Spruce) B.Thiers

Fig. 56A, B

On bark in upper montane forests, 2450–3500  m. Colombia: Cau, Cun, Qui, Ris. Ecuador: Tu, ZC. General distribution: northern Andes, Cuba. Lepidolejeunea sullivantii is a characteristic northern Andean species that is recognized by (1) leaf apex rounded or obtuse; (2) ocelli smaller than adjacent leaf cells; and (3) underleaves imbricate, very large, 7–9× stem width, apex ± undivided, truncate to shallowly emarginate, bases cuneate or rounded (not auriculate). Leptolejeunea (Spruce) Steph. Plants very small, to 1(−1.5) cm long, 0.3–1(−1.5) mm wide, green to pale brown to blackish in herbarium, creeping, with a strong odor when fresh. Stems of 7 outer cells and 3 inner cells, epidermis cells weakly enlarged; ventral merophyte 2 cells wide. Leaf lobes obliquely spreading, often standing up when dry, elliptical or

Lejeuneaceae

563

Fig. 56 (A and B) Lepidolejeunea sullivantii. (A) Habit. (B) Underleaf. (C–E) Leptolejeunea elliptica. (C) Habit. (D) Leaf. (E) Underleaf. (F and G) Leptolejeunea exocellata. (F) Leaf. (G) Habit. (H) Leptolejeunea jamaicensis, leaf. (I) Leptolejeunea radicosa, habit. (J) Leptolejeunea tridentata, habit. (K) Lindigianthus cipaconeus, habit. (L) Lopholejeunea nigricans, habit. (M and N) Lopholejeunea subfusca. (M) Habit. (N) Median cells. (O and P) Luteolejeunea herzogii. (O) Leaf. (P) Habit. (Q) Lopholejeunea eulopha, habit. (A, B redrawn from Piippo 1986; H–J redrawn from Bischler 1969; rest from Gradstein and Ilkiu-Borges 2009; Gradstein et al. 2001). Scale bars: A, B, K = 1000 μm; C, G, O = 250 μm; D–F, H = 50 μm; I, J = 200 μm; L, M, P, Q = 500 μm; N = 25 μm

564

 

Keys and Descriptions

obovate from a narrow base, with ocelli, apex rounded or acute-apiculate, plane, margins entire or toothed above. Cells with small trigones, cuticle smooth; oil bodies small, few per cell, simple or composed of a few droplets; ocelli colorless (rarely yellowish to reddish), 1 very large ocellus present at leaf base (ca. 2.5–4× larger than adjacent cells), smaller additional ocelli usually present, scattered and/or in a broken row in the lamina, becoming progressively smaller to the apex. Lobules 1/3–1/2 leaf length, not reduced, inflated, tooth 1-celled, short and blunt, hyaline papilla proximal. Underleaves small, remote, often with a large, adhesive rhizoid disc, bifid with very slender, upright or diverging lobes, the lobes 2–4 cells long and 1(−3) cells wide, lamina usually with a border of 6 or more large outer cells surrounding smaller inner cells, insertion line almost straight. Gynoecia on short branches, without innovation. Perianths inflated, 5-keeled above, the keels extending into short, horn-like projections. Sporophytes lejeuneoid. Vegetative reproduction by caducous branchlets. A pantropical genus of about 30 species, 7 in Colombia and Ecuador. The main features of Leptolejeunea are (1) plants usually epiphyllous; (2) leaves obliquely spreading, often standing away upright from the substrate, obovate to elliptical with a rounded to acute apex; (3) ocelli present in leaf lobes, colorless or yellowish to reddish, in a broken row, scattered or only one at leaf base, the basal ocellus very large, 2.5–4× larger than adjacent green cells and much larger than other ocelli; (4) lobules with 1 short, blunt tooth, hyaline papilla proximal; (5) underleaves with 2 subulate, upright or diverging lobes and a lamina bordered by larger cells; and (6) gynoecia without innovations. Literature: Bischler (1969) 1. Leaf margins crenate to toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaf margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Teeth large, more than 1 cell long . . . . . . . . . . . . . . . . . . . . . . . . . L. tridentata 2. Teeth or crenations smaller, not more than 1 cell long . . . . . . . . . . . . . . . . . . 3 3. Leaves with only 1 ocellus (at the base). Plants very small, ca. 0.4–0.5  mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. serratifolia 3. Leaves with more than 1 ocellus. Plants larger . . . . . . . . . . . . . . L. jamaicensis 4. Leaves with only 1 large ocellus (at the base). . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Leaves with several ocelli, scattered or in a row . . . . . . . . . . . . . . . . . . . . . . . 6 5. Leaf apex of at least some leaves acute or obtuse . . . . . . . . . . . . . L. exocellata 5. Leaf apex always broadly rounded. . . . . . . . . . . . . . . . . . . . . . . . . L. obfuscata 6. Leaves elliptical, ca. 2× as long as wide. Ocelli in a row . . . . . . . . . L. elliptica 6. Leaves obovate, less than 2× as long than wide. Ocelli scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. radicosa 1. Leptolejeunea elliptica (Lehm. & Lindenb.) Besch.

Fig. 56C–E

Very common on living leaves, occasionally on bark, from sea level to 2750 m. Colombia: Ama, Caq, Ces, Cho, Cun, NSa, Put, Ris, San, Vau. Ecuador: Az, Es, Ga, Gu, MS, Na, Or, Pi, Su. General distribution: common and widespread in tropical and subtropical America.

Lejeuneaceae

565

Leptolejeunea elliptica is one of the most common epiphyllous bryophyte species in tropical America. It is readily distinguished by the elliptical leaves (ca. 2× longer than wide) with entire margins, a rounded apex, and 3–4 ocelli in a broken row throughout the leaf. Fresh plants have a strong smell of anise. The characteristic fragrance of L. elliptica is due to the aromatic ether p-ethylanisol, which is unique to this species (Asakawa et al. 2013). 2. Leptolejeunea exocellata (Spruce) A.Evans

Fig. 56F, G

On bark and living leaves in lowland rainforest, 185–500  m. Colombia: Put (Campos et al. 2014). Ecuador: Es (Schäfer-Verwimp et al. 2006). General distribution: tropical America. Leptolejeunea exocellata is characterized by the obtuse to acute leaves with entire margins and only 1 large ocellus (at the leaf base). 3. Leptolejeunea jamaicensis R.M.Schust. Fig. 56H Leptolejeunea maculata auct. (non typus) Leptolejeunea convexistipa Bischl. (?) (fide L. Shu & R.L. Zhu, pers. comm.) On living leaves in lowland and montane rainforest remnants and scrub, from sea level to ca. 2000 m. Colombia: Cho (?) (Benavides and Sastre-de Jesús 2011 as L. maculata). Ecuador: Pi (Río Guajalito, Gradstein 12622, det. L. Shu & R. L. Zhu, HSNU, PC, QCA), ZC (Schäfer-Verwimp et al. 2013a as L. convexistipa). General distribution: scattered in tropical America, distribution insufficiently known. Leptolejeunea jamaicensis is an insufficiently known neotropical species that is characterized by leaf lobes with crenate to toothed margins and several ocelli and underleaves with a broad lamina and tiny, 2–3-cell-long lobes (Schuster 1967). The species was recorded from Ecuador as L. convexistipa (Schäfer-Verwimp et  al. 2013a), but according to Dr. R.L. Zhu (pers. comm.), these plants belong to L. jamaicensis and the two species may be conspecific. Grolle (1976) aequated L. convexistipa with L. maculata (Mitt.) Schiffn., a species described from Asia, but the latter species is polyphyletic and may not occur in the neotropics (Shu 2016). The taxonomy and distribution of L. jamaicensis are being investigated by L. Shu and R.L. Zhu in the framework of an integrative study of neotropical Leptolejeunea. 4. Leptolejeunea obfuscata (Spruce) Steph. On living leaves, very rarely on bark, 100–200 m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). Ecuador: Or (Mota de Oliveira and ter Steege 2013). General distribution: Amazonia, Chocó, Guianas. Leptolejeunea obfuscata is a rare neotropical species resembling L. elliptica in leaf shape but differing in leaves with only 1 ocellus. 5. Leptolejeunea radicosa (Mont.) Grolle Leptolejeunea obovata Bischl.

Fig. 56I

On living leaves in lowland and montane rainforests, 200–1700 m. Colombia: Ant, Cho, San; to be expected in Ecuador. General distribution: scattered in tropical America. Leptolejeunea radicosa is a rather widespread neotropical species that is readily distinguished by the relatively short, obovate leaves (1.5–1.8× longer than wide)

566

 

Keys and Descriptions

with a broadly rounded apex, entire margins, and several small ocelli scattered in the leaf lobe (additional to the large basal ocellus). 6. Leptolejeunea serratifolia Schiffn. On living leaves in wet lowland rainforests along the Pacific coast, 100  m. Colombia: Cho (Benavides and Sastre-de Jesús 2011). General distribution: scattered in tropical America. Leptolejeunea serratifolia is a rare neotropical species that is recognized by the obovate leaves with toothed margins (teeth usually 1-celled), acute-acuminate apex, and only 1 ocellus (at the leaf base). 7. Leptolejeunea tridentata Bischl.

Fig. 56J

On living leaves in wet lowland rainforests along the Pacific coast, 80  m. Colombia: Cho (Bischler 1969). General distribution: Chocó (type), Brazilian Amazon. Leptolejeunea tridendata is a rare Amazonian-Chocó species that stands out by the acuminate leaf with 2–3-cell-long teeth in the upper half (teeth more than 1 cell long). Further characters include the crenulate keel and the presence of 2–3 ocelli in the leaf: 1 large ocellus at the base and 1–2 small ocelli higher up in the lamina. Further Record Leptolejeunea trigonostipa (Spruce) Steph. – Ecuador: Chimborazo (Spruce 1884 as Lejeunea trigonostipa Spruce). The type is lost (Bischler 1969). Excluded Records Leptolejeunea convexistipa Bischl. – Ecuador: ZC (Heinrichs et al. 2014b). The material belongs to L. jamaicensis R.M.Schust. (R.L. Zhu, pers. comm.). Leptolejeunea maculata (Mitt.) Schiffn.  – Colombia: Cho (Benavides and Sastre-de Jesús 2011). Paleotropical species, probably absent in the neotropics (Shu 2016; R.L. Zhu, pers. comm.); the identity of the material from the Chocó needs verification. Lindigianthus Kruijt & Gradst. Plants to 10  cm long, 1.5–3  mm wide, glossy brown, creeping, often forked. Branches Lejeunea-type or Frullania-type. Stems with a hyalodermis; ventral merophyte 4 cells wide. Leaf lobes wide-spreading with broadly recurved apex, when dry ± convolute, apex acute, margins entire or toothed. Cells subisodiametric, trigones radiate; oil bodies homogeneous. Lobules 1/3 leaf length, never reduced, with 2 teeth, second tooth smaller. Underleaves undivided, large, margins entire, recurved, insertion line deeply arched. Dioicous. Male bracts on a long branch, with epistatic lobules. Gynoecia with 2 pycnolejeuneoid innovations. Perianths flat, smooth or toothed above. Sporophytes lejeuneoid. Vegetative reproduction not observed.

Lejeuneaceae

567

A neotropical genus with a single species; for main features of Lindigianthus, see under L. cipaconeus. Literature: Gradstein et al. (1994) 1. Lindigianthus cipaconeus (Gottsche) Kruijt & Gradst. Dicranolejeunea cipaconea (Gottsche) Steph. Dicranolejeunea circinnata (Spruce) Steph.

Fig. 56K

On bark, humus, leaf litter, and soil in upper montane forests and páramo, sometimes growing between the rosette leaves of the tree fern Blechnum loxense, 2000–3500  m. Colombia: Ant, Cun, Hui, Ris, Tol. Ecuador: Lo, Pi, Tu, ZC. General distribution: northern Andes, Costa Rica. Lindigianthus cipaconeus is a very distinct northern Andean-Costa Rican species that is recognized by (1) plants glossy brown and rather transparent, large, to 3 mm wide; (2) leaf apex acute and broadly recurved; (3) lobules ovate-truncate, with 2 teeth; (4) underleaves very large, imbricate, reniform, with reflexed margins and deeply arched insertion line; and (5) perianth flat, without dorsal and ventral keels, entire or with a few small teeth. Lopholejeunea (Spruce) Steph. Plants rather small, 1–2 mm wide, glossy green to black (or brown), usually creeping. Branches Lejeunea-type, occasionally Frullania-type. Stems with a hyalodermis; ventral merophyte 4 cells wide. Leaf lobes wide-spreading, when dry not appressed or convolute, apex rounded to apiculate, margins entire. Cells (sub)isodiametric, walls often with blackish pigmentation, trigones simple-triangular or radiate; oil bodies homogeneous; ocelli lacking. Lobules 1/4–1/2 leaf length, sometimes reduced, strongly inflated in the lower half, with 0–1 teeth. Underleaves undivided, margins entire, insertion line curved. Gynoecia without true innovations. Perianths with 4–5 dentate-laciniate keels (rarely entire). Sporophytes ptychanthoid. Vegetative reproduction not observed. A pantropical genus of about 35 species, most of them in the paleotropics, 3 species in the neotropics including Colombia and Ecuador. The main features of Lopholejeunea are (1) plants dark green to brown to blackish, forming flat mats on bark, rock, or living leaves; (2) stems with a hyalodermis; (3) leaves wide-spreading when dry; (4) leaf cells isodiametric, often with darkish-pigmented walls, trigones small, simple-triangular to radiate, oil bodies homogeneous; and (5) gynoecia without innovations, perianth 4–5-keeled, the keels usually dentate-laciniate (rarely subentire). Although true Radula-type innovations (originating between inner bract and perianth, without collar at the base) are lacking in Lopholejeunea, the occurrence of short or long “pseudo-innovations” is frequently observed. These pseudo-innovations resemble true innovations but originate below the inner female bract, away from the perianth, and are of the Lejeunea-type, with a collar at the base. Literature: Gradstein et al. (1994), Gradstein and Ilkiu-Borges (2009) 1. Underleaves of main stems 1.5–2× wider than long. Female bracteole strongly toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. eulopha

568

 

Keys and Descriptions

1. Underleaves 1–1.5× wider than long. Female bracteole entire . . . . . . . . . . . . 2 2. Perianth ± immersed. Distal end of the free margin of the lobule truncate, attached to the lobe surface across 2–3 cells. Well-developed lobules ± fully inflated. Leaf apex rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. subfusca 2. Perianth distinctly emergent, to half of its length. Distal end of the free margin acute, attached to lobe surface by only 1 cell. Well-developed lobules partially flattened. Leaf apex apiculate or rounded . . . . . . . . . . . . . . . . . . . L. nigricans 1. Lopholejeunea eulopha (Taylor) Schiffn. Lopholejeunea cocosensis Clark Lopholejeunea quelchii Steph.

Fig. 56Q

On trees in lowland rainforest, from sea level to 200 m. Colombia: Ama, Cho, Put, Vau. Ecuador: Or (Mota de Oliveira and ter Steege 2013). General distribution: pantropical. Lopholejeunea eulopha is a widespread pantropical species that is very common in Africa and Asia but rather rare in the neotropics where it is limited to the West Indies, Amazonia (incl. Guianas) and the Chocó region (Gradstein 1994). The species is readily recognized by the broad underleaves (1.5–2× wider than long) and strongly toothed female bracteole. 2. Lopholejeunea nigricans (Lindenb.) Schiffn.  Lopholejeunea andersonii Steph. Lopholejeunea muelleriana (Gottsche) Schiffn.

Fig. 56L

On bark and rock in lowland and montane rainforests, from sea level to 1800 m. Colombia: Boy, Cas, Cho, Hui, Mag, Met, Put, Ris, San, Tol, Vau. Ecuador: Ga, MS, Na, Or, Pa, Pi, Tu. General distribution: pantropical. Lopholejeunea nigricans is the only neotropical species of the genus Lopholejeunea with acute leaves. However, rounded leaf tips are also seen in this species and may lead to confusion with L. subfusca. For differences with the latter species, see the key. 3. Lopholejeunea subfusca (Nees) Schiffn. Lopholejeunea sagreana (Mont.) A.Evans

Fig. 56M, N

Common sun epiphyte on bark in the canopy and at the margins of lowland and lower montane rainforests, in plantations, gardens, and on isolated trees, from sea level to 1500 m. Colombia: Ama, Cas, Cho, Put, Qui, Ris, Val, Vau. Ecuador: Az, Es, MS, Na, Or, ZC. General distribution: pantropical. Lopholejeunea subfusca is a very common and widespread, xerotolerant species that grows in large black patches on smooth bark of trees in open locations. Fertile plants are readily recognized by the immersed perianths, which are almost fully covered by the female bracteole, with only the coarse lacinia of the perianth keels protruding. When sterile, the species can be recognized by the truncate distal end of the free margin of the lobules (at the junction with the keel), which is connate with the surface of the lobe along 2–3 cells. This characteristic is also seen in L. eulopha. In contrast, in L. nigricans, a common species sometimes confused with L. subfusca, the distal end of the lobule free margin is acute and attached to the lobe surface by only 1 cell.

Lejeuneaceae

569

Luteolejeunea Piippo Plants rather robust, to 4 cm long, 1.5–2 mm wide, pale yellowish-green to yellowbrown, loosely creeping. Stems rigid, of thick-walled cells, with enlarged epidermis; ventral merophyte 2 cells wide. Leaf lobes wide-spreading, with scattered ocelli as well as 1–2 at the base, apex rounded, usually recurved, margins crenulate to slightly toothed above. Cells 25–40  μm long in lobe middle, with swollen trigones; oil bodies present in all cells, 2–4 per cell, coarsely granular. Ocelli conspicuously larger than the green cells, occurring scattered in leaves, underleaves, bracts, bracteoles, and perianths. Lobules very small, ca. 1/10–1/7 leaf length, bottle-shaped, curved downward, free margin incurved, tooth inconspicuous, hyaline papilla proximal. Underleaves undivided, large, reniform, 6–10× stem width, insertion line deeply arched. Androecial bracteoles present throughout the spike. Gynoecia on short branches, with 1 pycnolejeuneoid innovation. Perianths inflated, 5-keeled, the keels horn-like swollen above. Sporophytes lejeuneoid. Vegetative reproduction not observed. A neotropical genus with a single species. For main features of Luteolejeunea, see under L. herzogii. Literature: Piippo (1986) 1. Luteolejeunea herzogii (Buchloh) Piippo Stictolejeunea herzogii Buchloh

Fig. 56O, P

On bark or trees in wet lowland rainforest along the Pacific coast, from sea level to 400  m. Colombia: Cau, Cho, Val; to be expected in Ecuador (Esmeraldas). General distribution: Chocó biogeographic region. Luteolejeunea herzogii is a characteristic Chocó endemic that is distinguished by (1) stems rigid, made up of thick-walled cells, ventral merophyte 2 cells wide; (2) leaves and underleaves with numerous large, scattered ocelli; (3) leaf cells with large trigones; (4) underleaves large, reniform (6–10× stem width), undivided; (5) lobules very small, downward-curved, hidden behind the underleaves; and (6) perianth 5-keeled, keels horn-like and swollen above. Marchesinia Gray Homalolejeunea (Spruce) Steph. by S. Robbert Gradstein & Jaime Uribe-M. Plants often large, 2–10  cm long, 2–4  mm wide, deep green to black or brown, ascending or pendent, often forked. Branches Lejeunea-type, occasionally Frullania-type. Stems with weakly enlarged epidermis, the epidermis usually brownish; ventral merophyte 6–11 cells wide. Leaf lobes wide-spreading, when dry plane or reflexed (not convolute), ovate-oblong, apex apiculate to cuspidate (rounded in African species), margins entire or toothed. Cells (sub)isodiametric, trigones usually triradiate; oil bodies homogeneous (in tropical American spp.); ocelli lacking. Lobules 1/5–1/3 leaf length, sometimes reduced, usually flat, with (1–)2–4 teeth, free margin usually plane. Underleaves undivided, margins entire or toothed, often

570

 

Keys and Descriptions

recurved, bases decurrent, insertion line deeply arched. Gynoecia with 2 pycnolejeuneoid innovations. Perianths flat, margins entire. Sporophytes ptychanthoid. Vegetative reproduction by caducous or fragmenting leaves. A small genus of six species in tropical America, Africa, western Europe, and Hawaii, three in Colombia and Ecuador, in subg. Marchesiniopsis R.M.Schust. (Söderström et al. 2016). Marchesinia is readily recognized in the neotropics by (1) plants dark green when fresh; (2) leaves elongate, ovate-oblong, apiculate to cuspidate, with ± isodiametric leaf cells and radiate trigones; (3) lobules small, flat, with 2–4 teeth; (4) underleaves with deeply arched insertion line (depth of underleaf insertion equals underleaf length as measured from rhizoid disc to apex); and (5) perianths flat, with 2 innovations. Marchesinia may be confused with Symbiezidium when sterile; both have large underleaves with deeply arched insertions and ovate-oblong leaves with a broadly rounded apex. However, Symbiezidium is usually pale brown in color, never blackish-green, the leaf apex is not apiculate, and the lobules in Symbiezidium are strongly inflated, ball-shaped, with involute free margin and without discrete teeth. When fertile, the two genera are immediately separated by the completely different gynoecium. Gradstein and Geissler (1994) recognized two Marchesinia species in tropical America, the widespread and highly variable M. brachiata (underleaves entire) and M. robusta found in the Andes and Costa Rica and characterized by toothed underleaves. A molecular-phylogenetic analysis of the two species revealed the existence of four taxa, in two clades. One clade contained M. brachiata and M. robusta; the other clade contained M. languida and M. bongardiana, which were previously included in M. brachiata (Heinrichs et al. 2009). Our revision of the Marchesinia collections in COL and GOET showed that all four taxa occur in Colombia and Ecuador. Since M. robusta differs morphologically in only one character from M. brachiata, we treat M. robusta as a variety of M. brachiata. Literature: Gradstein and Geissler (1994), Heinrichs et  al. (2009), Gradstein (2012a) 1. Underleaf margins plane. Lobules with 1–2(−4) teeth. Leaves ± fragile . . . . . 2 1. Underleaf margins recurved. Lobules with 3–4 teeth. Leaves not fragile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (M. brachiata) 3 2. Leaf apex longly apiculate to cuspidate and with 1–7 distinct teeth, the teeth 1–6(−10) cells long (but teeth often broken). Underleaf base cuneate to auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. languida 2. Leaf apex shortly apiculate, with 1–4 small teeth, the teeth only 1–2 cells long. Underleaf base cuneate to rounded, not auriculate . . . . . . . . . M. bongardiana 3. Underleaf apex toothed . . . . . . . . . . . . . . . . . . . . . . . M. brachiata var. robusta 3. Underleaf apex entire on sterile or male shoots (but sometimes toothed on female shoots) . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. brachiata var. brachiata

Lejeuneaceae

571

1. Marchesinia bongardiana (Lehm. & Lindenb.) Trevis.  Fig. 57C Lejeunea bongardiana Lehm. & Lindenb. Type: Brazil, Rio de Janeiro, Bongard s.n. (lectotype, designated here, S-B53738!; isolectotypes see Gradstein and Geissler 1994) Marchesinia aquatica Herzog Marchesinia galapagona (Ångstr.) Steph. Marchesinia longirostris (Sw.) Schiffn. Marchesinia nigrescens (Ångstr.) Steph. On bark and rock in semi-deciduous and evergreen woodlands and forests, plantations, gardens, and on isolated trees, (200–)900–1500 m; on the Galápagos Islands common in the mesic, semi-deciduous belt at 200–300 m. Colombia: Cas, Cun, Mag, Met, NSa (Herzog 1955 as M. aquatica), Qui, Ris, Val (Gradstein and Uribe 2016a). Ecuador: Az, EO (250–400 m), Ga (Gradstein 2009), LR (Spruce 1884), MS. General distribution: tropical America. Marchesinia bongardiana (subg. Marchesiniopsis) is a widespread neotropical species that is genetically close to M. languida (Heinrichs et  al. 2009). It differs from the latter species in the shortly apiculate leaves with 1–4 small teeth (maximally 2 cells long) and the underleaves with cuneate to rounded bases (not auriculate). Marchesinia bongardiana was considered a synonym of M. brachiata (Evans 1908; Gradstein and Geissler 1994) but differs from M. brachiata in the flat underleaf apex (not recurved), the somewhat fragile leaves, and the lobule usually with only 1–2 teeth, rarely more. 2a. Marchesinia brachiata (Sw.) Schiffn. var. brachiataFig. 57A, B Jungermannia brachiata Sw. Type: Jamaica, Swartz s.n. (lectotype, designated here, S-B54152!; isolectotype BM!) Lopholejeunea saxatilis (Ångstr.) Steph. Marchesinia guilleminiana (Nees & Mont.) Trevis. Marchesinia longistipula Steph. For further synonyms, see Gradstein and Geissler 1994; some of them may belong to M. languida or M. bongardiana. On bark and rock in montane rainforests, scrub, and plantations, (250–) 500–3300 m; on the Galápagos Islands common in the humid belt between 300 and 750 m. Colombia: Boy, Cal, Cas, Cho, Hui, Mag, Met, Qui, Ris, San, Tol. Ecuador: Ch, Ga, Gu, Lo, MS, Or, Pa, Pi, Tu, ZC. General distribution: tropical America, St. Helena, East Africa, Hawaii. Marchesinia brachiata var. brachiata (subg. Marchesiniopsis) is a widespread Afro-American taxon that is recognized by underleaves with entire, recurved margins. The margins may be recurved at the apex or all around. The characters of the underleaves should be examined on sterile shoots or male shoots; underleaves on female shoots may be toothed like in the next variety. The leaves are imbricate and not fragile, the leaf apex is rounded or apiculate and finely toothed, and the lobules have 3–4 teeth. The plants may be autoicous or dioicous.

Fig. 57 (A and B) Marchesinia brachiata var. brachiata (A) Habit. (B) Leaf. (C) Marchesinia bongardiana, habit. (D and E) Metalejeunea cucullata. (D) Habit. (E) Portion of stem with leaf and underleaf. (F) Microlejeunea acutifolia, habit. (G and H) Microlejeunea bullata. (G) Leaf. (H) Habit. (I) Microlejeunea epiphylla, habit. (J and K) Microlejeunea globosa. (J) Habit, dorsal view. (K) Habit, ventral view. (L) Microlejeunea diversiloba, habit. (M–O) Myriocoleopsis gymnocolea. (M) Habit. (N) Cladograph. (O) Leaf, dorsal view. (P and Q) Myriocoleopsis minutissima subsp. myriocarpa. (P) Gemma. (Q) Habit. (R) Neurolejeunea breutelii, habit. (C redrawn from Herzog 1955; J, K redrawn from Reiner-Drehwald 1994a; L redrawn from Bischler et al. 1963; M–O from Reiner-Drehwald and Gradstein 1995; rest from Gradstein and Ilkiu-Borges 2009; Gradstein et al. 2001). Scale bars: A = 1000 μm; B, D, J, K, Q, R = 250 μm; C = 1100 μm; E, F, H, I = 100 μm; G, P = 50 μm; L, O = 200 μm; M = 400 μm

Lejeuneaceae

573

We have revised the material of M. brachiata in COL and have found that several Colombian records belong to M. bongardiana. The Ecuadorian records of M. brachiata have not yet been revised. 2b. Marchesinia brachiata var. robusta (Mitt.) Gradst. & Uribe, comb. nov. Lejeunea robusta Mitt., Hooker’s J.  Bot. Kew Gard. Misc. 3: 359. 1851. (Marchesinia robusta [Mitt.] Schiffn.). Type: Ecuador, vicinity of Quito, Jameson s.n., herb. Mitten 150a (lectotype, designated here, NY-900775!; isolectotypes see Gradstein and Geissler 1994) On bark in montane rainforests, scrub, and plantations, also on rock, 650–2500 m. Colombia: Ant (?), Boy, Cas, Hui, Ris, San. Ecuador: Es, Pi, Tu, ZC. General distribution: tropical Andes, Costa Rica. Marchesinia brachiata var. robusta differs from var. brachiata by toothed underleaf margins. Genetically the two varieties are very near; their close genetic relationship is illustrated by the fact that one of the collections of var. brachiata (Risaralda, Gradstein 8372, GOET) is clustered with var. robusta in a molecular analysis (Heinrichs et al. 2009). 3. Marchesinia languida (Nees & Mont.) Steph. Lejeunea languida Nees & Mont. Type: Peru, “Yungas,” d’Orbigny s.n., herb. Mus. Paris (lectotype, designated here, PC-0103222!; isolectotypes see Gradstein and Geissler 1994) Lejeunea palaeflora Spruce On tree trunks in montane rainforest, 1200–2100  m. Colombia: Cun, Mag (Gradstein et al. 2018c). Ecuador: Ch, Na, Pi, Tu. General distribution: tropical Andes, Central America. Marchesinia languida (subg. Marchesiniopsis) is an uncommon neotropicalmontane species that is characterized by the longly apiculate to cuspidate leaf tips with 1 to several teeth (up to 7) on the apical margin. The teeth are 1–6(−10) cells long and are often broken in dried material. The species is dioicous, the leaves are rather fragile, the lobules are small and possess 1–2(−4) teeth, and the underleaves are distant or subimbricate, ca. 4–6× stem wide, with a flat apex, and the underleaf bases vary from cuneate to auriculate. The variation seen in the underleaves correlates with the size of the underleaves: large, imbricate underleaves have auriculate bases, whereas small, distant underleaves have cuneate to narrowly rounded bases. Both types of underleaves may occur on single stems. The presence of large underleaves with auriculate bases and the longly apiculate to cuspidate leaf apex with several long teeth (1–6[−10] cells long) separates M. languida from the closely related M. bongardiana (see also under M. bongardiana). Metalejeunea Grolle Plants very small, 0.2–0.3  mm wide, whitish-green to dull yellowish, creeping. Stems zig-zag, rather rigid, of thick-walled cells, without hyalodermis, in cross section of 7 outer rows of cells and 3 smaller inner rows; ventral merophyte 2 cells wide. Leaf lobes distant, suberect, elongate, apex rounded, margins entire. Cells

574

 

Keys and Descriptions

small, 12–17 μm long in midleaf, trigones usually small; oil bodies unknown; ocelli lacking. Lobules 1/2–2/3× leaf length, occasionally reduced, strongly swollen, keel crenate, tooth long, 1-celled, straight or curved, hyaline papilla proximal. Underleaves bifid, distant, small, 1.5–2.5× stem width. Autoicous. Gynoecia on long or short shoots, with 1 pycnolejeuneoid innovation, the innovation sterile or with androecia; bracts ± winged. Perianths inflated, with (3–)5 smooth keels. Sporophytes lejeuneoid. Vegetative reproduction not observed. A small pantropical genus (3 spp.), one species in Colombia. Metalejeunea resembles Microlejeunea and the two genera may be confused; for differences see under Metalejeunea cucullata. Literature: Gradstein et al. (2001) 1. Metalejeunea cucullata (Reinw., Blume & Nees) Grolle Microlejeunea monoica Bischl.

Fig. 57D, E

On twigs and branches in the forest canopy and on tree trunks in lowland rainforest, 100–350 m. Colombia: Ama, Caq, Cho, Put, Vau (Pinzón et al. 2003; Campos et al. 2015); to be expected in Ecuador. General distribution: pantropical. Metalejeunea cucullata resembles Microlejeunea but differs in autoicous sexuality, leaves without ocelli, and, especially, gynoecia with pycnolejeuneoid innovations (lejeuneoid in Microlejeunea). The leaf lobes in M. cucullata are 1.5–1/8× longer than wide, with a rounded apex, entire margins, and a crenate keel, and the leaf cells are small, ca. 15 μm long in midleaf, with small trigones. The plants are frequently fertile. Microlejeunea (Spruce) Steph. Plants very small, 0.2–0.6 mm wide (rarely larger), pale green to pure green, usually somewhat dull-colored, creeping. Stems zig-zag, very thin, of thick-walled cells, without hyalodermis, of 7 outer rows of cells and 3 smaller inner rows; ventral merophyte 2 cells wide. Leaf lobes usually distant, suberect or obliquely spreading, with large lobules (usually more than 1/2 the size of the lobe), apex rounded to acute, margins entire or crenulate. Cells small, 15–25 μm long in middle of lobe, thin-walled or thick-walled; oil bodies small, finely granular; ocelli usually present, 1–3 at lobe base, grayish, sometimes more numerous and brown, scattered in the leaf (M. globosa). Lobules 0.5–0.8 leaf length, sometimes reduced, strongly swollen, with 1 rather long and curved tooth, hyaline papilla proximal. Underleaves bifid, small. Dioicous (sometimes autoicous in paleotropical species). Gynoecia on long or short shoots, with 1–2 lejeuneoid innovations; bracts ± winged. Perianths inflated-pyriform, with 0–5 smooth keels. Sporophytes lejeuneoid. Vegetative reproduction very rare, by caducous branchlets. A pantropical genus of about 25 species, 6 in Colombia and Ecuador. The genus is readily recognized in the region by the very small plants (0.2–0.6 mm wide) with stems zig-zag, leaves upright ± parallel to the stem, and very large lobules, more than half the size of the leaf. Usually there are 1–3 ocelli in the leaves (near the base), rarely more (M. globosa).

Lejeuneaceae

575

Literature: Bischler et  al. (1963), Schuster (1980a as Lejeunea subg. Microlejeunea), Reiner-Drehwald (1994a) 1. Plants 0.2–0.3  mm wide. Leaves suberect, almost parallel to the stem. Welldeveloped lobules 0.6–0.8× lobe length. Ocelli grayish, 1–3 at leaf base, rarely absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants larger, 0.3–0.6 mm wide. Leaves obliquely spreading. Well-developed lobules smaller, 0.5–0.6× lobe length. Ocelli brown, scattered in the leaf lobe and sometimes forming a row. Plants in dry montane forest . . . . . . M. globosa 2. Mature stem leaves less than 1.5× longer than wide. Ventral margin of leaf lobe sometimes recurved in the lower half (due to prolongation of the lobule along the margin). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Mature stem leaves more than 1.5× longer than wide. Ventral margin of leaf lobe plane. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 3. Lobules fully inflated. Well-developed underleaves made up of 12–25 cells ............................................................. 4 3. Lobules partially inflated, distal portion flat (along free margin and apex). Welldeveloped underleaves larger, made up of (20–)25–50 cells. . . . . M. capillaris 4. Keel crenate by mammillose cells. Rudimentary lobules frequently present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. epiphylla 4. Keel smooth. Rudimentary lobules usually absent, very scarce . . . . M. bullata 5. Leaf apex broadly rounded. Keel smooth . . . . . . . . . . . . . . . . . M. diversiloba 5. Leaf apex obtuse or acute. Keel crenate . . . . . . . . . . . . . . . . . . . . M. acutifolia 1. Microlejeunea acutifolia Steph. Microlejeunea triangulata Herzog

Fig. 57F

On bark, living leaves, and dead wood in lowland and montane rainforests, at forest margins, and on roadside trees, in Colombia mainly along the Pacific and Atlantic coasts, from sea level to 2800 m. Colombia: Cho, Mag, Val. Ecuador: Az, Es, Lo, Na, Or, Pi (Schäfer-Verwimp et al. 2013a). General distribution: tropical America. Microlejeunea acutifolia is characterized by elongate leaves with an obtuse or acute tip, entire leaf margins, flat ventral margin of the leaf lobe, and crenate keel. 2. Microlejeunea bullata (Taylor) Steph. Fig. 57G, H Lejeunea ulicina (Taylor) Gottsche, Lindenb. & Nees subsp. bullata (Taylor) R.M.Schust. On bark in montane rainforests, in scrub, and on isolated trees, less common in the lowlands, up to 3000 m. Colombia: Ama, Boy, Caq, Cas, Ces, Cho, Cun, Hui, NSa, Put, Qui, Ris, San, Vau. Ecuador: Az, Ch, Es, Ga, Lo, MS, Na, Or, Pi, Tu, ZC. General distribution: common and widespread in tropical America. Microlejeunea bullata is a common species of neotropical montane rainforests that is recognized by the short-ovate, hardly longer than wide leaves with a rounded to slightly emarginate apex, fully inflated lobules ca. 3/4× leaf length, a smooth keel, and the usual absence of rudimentary leaf lobules.

576

 

Keys and Descriptions

3. Microlejeunea capillaris (Gottsche) Steph. Lejeunea capillaris Gottsche. The type specimen is lost (fide Bischler et al. 1963). Microlejeunea colombiana Bischl. Type: Colombia, Cundinamarca, Salto de Tequendama, 2500 m, Bischler 2297b (isotype PC-0103188!) On bark, living leaves, rotten wood, and humose soil in montane forests and scrub up to the páramo, (700–)2000–4780 m. Colombia: Boy, Cun, Mag. Ecuador: Ca, Co, EO, Im, Lo (Schäfer-Verwimp et al. 2013a), Na, Pi, ZC. General distribution: scattered in tropical America (new to Peru, Urubamba, Mantanay, above laguna Manalloqsa, 4780 m, Sylvester s.n., PC). Microlejeunea capillaris is a rather widespread neotropical-montane species. Characteristic features of the species are (1) leaves somewhat longer than wide, with a broadly rounded leaf apex and a shallow depression at the proximal side of the apex; (2) lobules 0.6–0.7× leaf length, with a flattened apical portion and a large, curved tooth, keel slightly crenate, rudimentary lobules scarce; (3) underleaves rather large, 1.5–2.5× stem width, made up of (20–)25–50 cells. Our knowledge of Microlejeunea capillaris, a species described from southern Brazil, is essentially based on the treatment by Bischler et al. (1963); the type specimen of the species is lost. Microlejeunea colombiana is a robust phenotype of M. capillaris from high elevation. The conspecificity of these two species was already suggested by Bischler et al. (1963), and Gradstein and Uribe (2016a) treated them as synonyms. 4. Microlejeunea diversiloba (Spruce) Müll.Frib. Microlejeunea stricta Steph. (fide Söderström et al. 2015a)

Fig. 57L

On bark of trees in submontane and lower montane rainforests, 700–3100  m. Colombia: Mag, Qui, Ris. Ecuador: Lo, Na, ZC (Schäfer-Verwimp et al. 2013a). General distribution: tropical America. Microlejeunea diversiloba is an uncommon neotropical species characterized by elongate leaves, more than 1.5× longer than wide, with a plane ventral margin, broadly rounded apex, and smooth keel. 5. Microlejeunea epiphylla Bischl. Lejeunea dimorphophylla R.M.Schust.

Fig. 57I

On bark and living leaves in evergreen and semi-deciduous forests and scrub, from sea leavel to 2500 m. Colombia: Ama, Cho, Qui (Gradstein 12726a, HUQ), Put, Ris. Ecuador: MS, Or, ZC. General distribution: throughout tropical and subtropical America. Microlejeunea epiphylla is a widespread neotropical species that is rather similar to M. bullata but differs in the crenate keel and the frequent presence of reduced lobules. Microlejeunea epiphylla may be confused with Lejeunea yasuniensis, which has branch leaves strikingly similar to those of M. epiphylla. However, L. yasuniensis lacks ocelli, stem leaves have lobules only half the length of the lobe, and the leaf margins of L. yasuniensis are strongly crenate (not crenate in M. epiphylla).

Lejeuneaceae

6. Microlejeunea globosa (Spruce) Steph. Microlejeunea cardotii Steph.

577

Fig. 57J, K

On bark of trees in dry montane woodlands and in urban areas, 2300 m. Ecuador: Or (?), Pi (León-Yánez et  al. 2006 as L. cardotii; Werner and Gradstein 2008). General distribution: subtropical regions of tropical America (Mexico, Florida, southern Brazil, Paraguay, northern Argentina) and above 2000 m in the Ecuadorian Andes; also in tropical Africa (Kenya). Microlejeunea globosa is a characteristic species of dry subtropical woodlands that has been found at high elevation in Ecuador. It differs from other Microlejeunea species in the Andes in its larger size (to 0.6  mm wide), more widely spreading leaves, and smaller lobules (to 0.6× lobe length). By these characters the species is more similar to Lejeunea; the presence of ocelli clearly separates the species from Lejeunea, however. The Ecuadorian material stands out by rather large underleaves, with lobes 5–6 cells wide at base, and numerous brown ocelli in leaves, bracts, and perianth, which are sometimes arranged in a row (Schäfer-Verwimp et al. 2013a). The record of M. globosa from Amazonian lowland rainforest (Mota de Oliveira and ter Steege 2013) needs verification. Further Record Microlejeunea perpusilla (Spruce) Steph.  – Colombia: Cho (Winkler 1970); Ecuador: Ca, Ch (Spruce 1884). The identity of the species is unclear (Bischler et al. 1963). Myriocoleopsis Schiffn. Plants small- to medium-sized, 0.5–3 cm long, 0.8–1.2 mm wide, greenish to blackish, sometimes rheophytic (sect. Myriocoleopsis); leafy shoots irregularly branched, ascending from a creeping, stolon-like shoot (sect. Myriocoleopsis) or creeping without stoloniform shoot (sect. Protocolea). Branches Lejeunea-type, rather long and usually fertile. Stems fragile or rigid, of thin-walled or thick-walled cells, in cross section of 5–30 cell rows. Leaf attached to stem by only few cells, leaf insertion very short, transverse. Leaf lobes obliquely spreading, obovate, apex rounded, plane, margins entire or crenate, without hyaline border. Cells thin-walled with small trigones; oil bodies finely granular; ocelli lacking. Lobules usually reduced to a long, flat fold along the ventral margin of the lobe (to 2/3 of lobe length), sometimes large and inflated, with 2 teeth. Underleaves absent; rhizoids in small bundles from the ventral side of the stem, near the base of each leaf. Autoicous. Androecia in long spikes of (4–)10–20 pairs of bracts. Gynoecia single, with 1 sterile innovation, or in clusters with repeatedly fertile inovations. Perianths inflated, with 5 smooth keels, base often stalk-like elongate. Sporophytes lejeuneoid. Vegetative reproduction by multicellular, disciform gemmae from leaf surfaces. A small pantropical genus (5 spp.), three species in Colombia and Ecuador. Myriocoleopsis lacks underleaves and is close to Cololejeunea but is separated from the latter by frequent occurrence of upright growth, with leafy stems arising from a

578

 

Keys and Descriptions

creeping stoloniform shoot, by the lobules often reduced to a long flat fold, and by the copiously produced perianths. The two genera constitute different molecular lineages (Yu et  al. 2014). Myriocoleopsis is subdivided into two sections: (1) Myriocoleopsis sect. Myriocoleopsis characterized by upright growth, rigid stems without hyalodermis, presence of a creeping stoloniform shoot, very long androecial spikes, and clustered gynoecia with fertile innovations and (2) Myriocoleopsis sect. Protocolea (R.M.Schust.) Gradst. characterized by creeping growth, fragile stems with a hyalodermis, absence of a creeping stoloniform shoot, short androecial spikes, and gynoecia single, with sterile innovations (Gradstein 2018). The members of subg. Myriocoleopsis grow as rheophytes in rivers, while those of subg. Protocolea occur on bark or rock in open and often rather dry environments. Literature: Reiner-Drehwald and Gradstein (1995, 1997), Yu et al. (2014) 1. Lobules strongly inflated, with 2 well-devoped teeth . . M. minutissima subsp. minutissima 1. Lobules flat or reduced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Leafy shoots arising from a creeping, stoloniform stem. Stems rigid, made up of rather thick-walled cells. Plants on rock in rivers, rheophilous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. gymnocolea 2. Leafy shoots not arising from a creeping, stoloniform stem, stoloniform stem absent. Leaves more or less quarrose. Stems fragile, made up of thin-walled cells. Plants on bark or rock, not rheophilous . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Leaves ovate-lanceolate. Leaf apex obtuse to acute. Perianth on a long stalk (to 0.5 mm long). Plants in páramo, above 3000 m . . . . . . . . . . . . . . . . M. tixieri 3. Leaves ovate. Leaf apex broadly rounded. Perianth on a shorter stalk or not stalked. Plants below 3000 m, in dry environments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. minutissima subsp. myriocarpa 1. Myriocoleopsis gymnocolea (Spruce) M.E.Reiner & Gradst. Myriocoleopsis riparia M.E.Reiner & Gradst.

Fig. 57M–O

On periodically submerged rocks, twigs, and living leaves in blackwater rivers, 500–1300 m. Ecuador: MS (Río Quimi, Pérez et al. 11576, QCA, PC), Tu (tributaries of upper Río Pastaza), ZC (Río Nangaritza; Gradstein et  al. 2011); to be expected in Colombia. General distribution: Ecuador (type), Bolivia (Stephani 1916), southeastern and southern Brazil, northern Argentina (Misiones). Myriocoleopsis gymnocolea (sect. Myriocoleopsis) is a characteric rheophilous tropical South American species of blackwater rivers with strongly fluctuating water levels. The main characters of the species are (1) leafy shoots arising from a creeping, stolon-like stem; (2) stems rigid, made up of rather thick-walled cells, cortex of 5 rows of cells; (3) male spikes long, with 10–20 pairs of bracts; and (4) gynoecia in clusters with up to 6 perianths in a row. 2a. Myriocoleopsis minutissima (Sm.) R.L.Zhu, Y.Yu & Pócs subsp. minutissima Cololejeunea minutissima (Sm.) Schiffn. Cololejeunea turbinifera (Spruce) León-Yánez, Gradst. & Wegner (fide Pócs et al. 2014)

Lejeuneaceae

579

On bark in dry woodlands, parks, gardens, and on isolated trees, 1500–2800 m. Ecuador: Gu, Lo, Pi, Tu; to be expected in Colombia. General distribution: pantropical, extending into warm-temperate regions of the Northern and Southern Hemisphere. Myriocoleopsis minutissima (sect. Protocolea) subsp. minutissima is a very widespread pantropical taxon recognized by the fragile, creeping stems without a stoloniform base, ovate leaves with a broadly rounded apex, and a very large, strongly inflated lobule with 2 distinct teeth. 2b. Myriocoleopsis minutissima subsp. myriocarpa (Nees & Mont.) R.L.Zhu, Y.Yu & Pócs Fig. 57P, Q Cololejeunea myriocarpa (Nees & Mont.) A.Evans (Cololejeunea minutissima subsp. myriocarpa [Nees & Mont.] R.M.Schust.) On bark, rock, and living leaves in dry evergreen forest, semi-deciduous and deciduous woodlands, scrub, plantations, and on isolated trees, always in dry environments, 100–3000  m. Colombia: Tol (Gradstein and Uribe 2016a). Ecuador: Az, EO (Benitez 539, HUTPL), Ga, Im, Pi, ZC.  General distribution: pantropical. Myriocoleopsis minutissima (sect. Protocolea) subsp. myriocarpa is readily recognized by the strongly squarrose leaf lobes and the more or less reduced lobules, which are developed as a long, flat fold along the ventral margin (Schuster 1980a as Cololejeunea minutissima subsp. myriocarpa). In Colombia and Ecuador, subsp. myriocarpa has a wider elevational range than subsp. minutissima and is found from almost sea level to over 3000 m. The general distribution of subsp. myriocarpa is more restricted than that of subsp. minutissima and is largely confined to the tropics and subtropics. 3. Myriocoleopsis tixieri Pócs & Gradst., nom. nov. Cololejeunea tixieri M.I.Morales & C.Dauphin, nom. illeg. [“tixierii”], Tropical Bryology 14: 133. 1998 Cololejeunea dauphinii R.L.Zhu On bark in dwarf forest in páramo, 3900  m. Colombia (new): Boy (Sierra Nevada del Cocuy, Corralitos, 10 Jun 1959, Bischler 2932, det. T.  Pócs, PC). General distribution: Costa Rica, Panama, Colombia, E Brazil (Bahia). Myriocoleopsis tixieri (sect. Protocolea) is a rare neotropical species that is newly reported here from Colombia. The species occurs in páramo, above 3000 m, in Central America and Colombia but at much lower elevation, in submontane forest at 500–600  m, in eastern Brazil (Bôas-Bastos and Bastos 2016). Myriocoleopsis tixieri is recognized by (1) plants autoicous and copiously fertile, creeping, without stoloniform base, stems fragile; (2) leaf lobes elongate, ovate-lanceolate, ca. 2× longer than wide, leaf apex obtuse, margin crenulate; (3) lobules developed as a long flat fold parallel to the leaf lobe and about half the length of the lobe; (4) leaf cells rather large, ca. 30–50 μm long, thin-walled; and (5) perianth sharply 5-keeled, with a long stalk at the base, the stalk almost as long as the perianth. Note that the correct spelling of the epithet is “tixieri” (not “tixierii”) (see Turland 2013, pp. 91–93).

580

 

Keys and Descriptions

Neurolejeunea (Spruce) Schiffn. Plants small, 1–2 cm long, 0.7–1.5 mm wide, glossy olive green to brown or black, creeping, often (bi)pinnate or forked. Branches Lejeunea-type. Stems simple, without enlarged epidermis; ventral merophyte 4–6(−10) cells wide. Leaf lobes widespreading, when dry spreading or slightly convolute, sometimes with a row of ocelli, apex rounded, sometimes white-bordered, margins entire. Cells (sub)isodiametric, 10–20 μm long, walls with trigones or uniformly thickened; oil bodies smooth to faintly granular; row of ocelli in leaf lobes present or absent. Lobules often darker in color than leaf lobes, strongly inflated and usually flask-shaped, 1/5–1/2 of leaf length, never reduced, first tooth usually reduced, second tooth long, straight or falcate, hyaline papilla distal-ental. Underleaves undivided, suborbicular to reniform, apex plane or recurved, margins entire, insertion line straight or curved. Androecia on short, specialized branches, bracteoles limited to base of spike, bracts with hypostatic lobules. Gynoecia with 1–2 pycnolejeuneoid innovations. Perianths with 3–4 keels, lateral keels often auriculate. Sporophytes lejeuneoid. Vegetative reproduction not observed. A small neotropical genus of four species (Gradstein 1994), two species in Colombia and Ecuador, both in sect. Aneurolejeunea Gradst. characterized by the leaf lobes without a row of ocelli. The record of Neurolejeunea from Africa (Pócs et al. 2015) belongs to Cheilolejeunea sect. Cyrtolejeunea (S. R. Gradstein, pers. obs.). The main features of Neurolejeunea are (1) plants greenish-brown to blackish; (2) epidermis cells not larger than medullary cells; (3) lobules darker in color than leaf lobes, flask-shaped, conspicuously narrowed to the apex (with a distinct “neck”) and with a long, curved tooth; and (4) perianth 3–4-keeled, lateral keels often expanded above into auricles. The hyaline papilla of the lobule is distal to the tooth, which technically represents the second tooth. The first tooth is reduced or produced as a large papilla (N. sastreana). A vitta composed of a row of ocelli is present in sect. Neurolejeunea (not known from Colombia and Ecuador). Literature: Gradstein et al. (1994) 1. Leaf lobes with a row of ocelli. Plants canopy epiphytes in Amazonian rainforest; not yet known from Colombia and Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. seminervis (Spruce) Schiffn. 1. Leaf lobes without ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Lobule apex with a conspicuous papilla-like cell (25–30 μm long), which projects backward to the stem and has a very thick, colorless apical wall; underleaves plane. Plants rare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. sastreana 2. Papilla-like cell at lobule apex lacking or rudimentary, less than 10 μm long; underleaves concave or plane. Plants common . . . . . . . . . . . . . . . . N. breutelii 1. Neurolejeunea breutelii (Gottsche) A.Evans 

Fig. 57R

On bark and rock in lowland and montane rainforest, cloud forest, and scrub, in rather open environments or in the canopy of dense forest, from sea level to 3000 m. Colombia: Boy, Cas, Cau, Cho, San. Ecuador: Ga, LR, MS, Or, ZC. General distribution: tropical America, southeastern USA.

Lejeuneaceae

581

Neurolejeunea breutelii (sect. Aneurolejeunea) is a widespread neotropical species that is distinguished by (1) plants greenish-brown to blackish in color, ventral merophyte 4–6(−10) cells wide; (2) leaves ovate, apex rounded, margins entire; (3) leaf cells 15–20 μm, isodiametric, walls almost uniformly thickened, trigones confluent; (4) lobules darker-colored than leaf lobes, rather large and conspicuously elongate, 1/4–1/2× leaf length, never reduced, narrowed to the mouth and flaskshaped, apex with a long, curved tooth made up of a row of narrowly elongate cells, papilla-like cell at lobule apex lacking or rudimentary, less than 10 μm long; (5) underleaves distant, orbicular to reniform, undivided, 2–4× stem width, concave or plane, apex plane or recurved, bases cuneate, insertion line curved; and (6) gynoecia with 1–2 pycnolejeuneoid innovations, perianths with 3–4 keels which are broadly expanded above into horn-like auricles. Plants from low elevations are usually small and do not exceed 1 mm in width, whereas mountain plants may be more robust and up to 1.5 mm wide. Unusually robust plants, with 10-cell-row wide ventral merophytes and reniform underleaves up to 0.7 mm wide, occur in cloud forests at about 3000 m in the Eastern Cordillera of Colombia (Boyacá, Casanare; Gradstein 1994). The high-Andean populations seem to be invariably sterile. 2. Neurolejeunea sastreana Gradst.

Fig. 58A

On bark and rock in rainforest and scrub, 1500 m. Colombia: Ris (Uribe and Gradstein 1998). General distribution: West Indies, Central America, northern South America. Neurolejeunea sastreana (sect. Aneurolejeunea) is close to N. breutelii but is readily separated from the latter by the peculiar, large, papilla-like cell (= first tooth) at the lobule apex, with a very thick outer wall. Odontolejeunea (Spruce) Schiffn. Plants small- to medium-sized, 1–4  cm long, 0.8–3.5  mm wide, glossy green to brown, creeping. Branches Lejeunea-type, occasionally Frullania-type (robust vegetative branches). Stems with a hyalodermis; ventral merophyte 2 cells wide. Leaf lobes widely spreading, when dry irregularly crisped, ± convolute or revolute, apex rounded, margins sharply toothed. Cells (sub)isodiametric, trigones simple-triangular or cordate; oil bodies homogeneous, small and numerous per cell (Massulatype); ocelli lacking. Lobules 1/5–2/5 leaf length, occasionally reduced, with 1–6 teeth. Underleaves undivided, margins toothed or entire, bases decurrent and often folded, insertion line deeply arched. Androecia on short or long branches, bracteoles present throughout, bracts with epistatic lobules. Gynoecia without or with 1–2 pycnolejeuneoid innovations. Perianths 3-keeled with a low ventral keel, conspicuously winged and ciliate-laciniate above. Sporophytes lejeuneoid. Vegetative reproduction by caducous branchlets. A small Afro-American genus of three species (Teeuwen 1989), all three in Colombia and Ecuador. Odontolejeunea is a common epiphyllous genus that is recognized by undivided underleaves, toothed leaves, and dentate-ciliate perianths. The teeth on the ventral leaf margin are usually larger and more widely spaced than those on the dorsal margin. Odontolejeunea resembles Cyclolejeunea, especially

582

 

Keys and Descriptions

Fig. 58 (A) Neurolejeunea sastreana, leaf. (B) Odontolejeunea decemdentata, habit. (C and D) Odontolejeunea lunulata. (C) Leaf. (D) Habit. (E) Odontolejeunea rhomalea, leaf. (F) Otigoniolejeunea huctumalcensis, habit. (G and H) Pictolejeunea picta. (G) Leaf. (H) Habit. (I–K) Prionolejeunea aemula. (I) Leaf margin cells. (J) Habit. (K) Perianth. (L and M) Prionolejeunea arguta. (L) Habit. (M) Leaf margin cells. (N–P) Prionolejeunea denticulata. (N) Habit. (O) Leaf, dorsal view. (P) Leaf margin cells. (Q) Prionolejeunea decora, leaf. (From Gradstein and IlkiuBorges 2009; Ilkiu-Borges 2016). Scale bars: A, G, O = 100 μm; B–E, L, N = 500 μm; F, H, J, K, Q = 250 μm; I, P = 25 μm; M = 50 μm

Lejeuneaceae

583

C. convexistipa, and the two groups often grow together on living leaves. Odontolejeunea is separated from Cyclolejeunea by the undivided underleaves with rounded apex (bifid in Cyclolejeunea, except C. convexistipa), frequently brownish plant color (purely green in Cyclolejeunea and O. decemdentata), leaves often somewhat crisped-undulate, absence of gemmae, and seta with 16 rows of outer cells (12 rows of outer cells in Cyclolejeunea). Literature: Gradstein et al. (1994), Gradstein and Ilkiu-Borges (2009) 1. Underleaf margins sharply toothed. Lobules very high, much broader than long, with (4–)5–6 teeth. Cells in midleaf arranged in transverse rows, trigones small or large. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. rhomalea 1. Underleaf margins entire or with a few small teeth. Lobules not much broader than long, with 0–4 teeth. Cells in midleaf arranged in longitudinal rows (as usual in liverworts), trigones small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants 0.8–1.5  mm wide, purely green. Lobules with 0–2 teeth. Underleaves distant, 1.5–2.5× stem width, margins entire, bases without auricles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. decemdentata 2. Plants larger, (1–)1.5–3 mm wide, brownish-green. Lobules with 2–4(−5) teeth. Underleaves distant to imbricate, (2–)3–6× stem width, margins usually toothed, bases auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. lunulata 1. Odontolejeunea decemdentata (Spruce) Steph. Lejeunea hapalochroa Spruce Odontolejeunea longispica A.Evans

Fig. 58B

On living leaves and twigs in lowland and lower montane rainforests, from sea level to 2000 m. Colombia: Cau, Cho, Cun, Qui, San, Val. Ecuador: Az, Ch, EO, LR, MS, Na, Pa, Pi, Tu, ZC. General distribution: scattered in tropical America. Odontolejeunea decemdentata differs from other Odontolejeunea species in the rather small plant size (usually less than 1.5 mm wide), purely green color, lobules with only 0–2 teeth, and small, distant underleaves with entire margins and a cuneate base (without auricles). According to Gradstein et al. (1994), O. decemdentata also stands out by having granular oil bodies, but fresh material of O. decemdentata from Ecuador (Río Guajalito, Gradstein 12625, QCA) had homogeneous, Massulatype oil bodies like the other two Odontolejeunea species. Since Massula-type oil bodies may quickly degenerate upon drying of the material and become granular before disintegrating (Gradstein et  al. 1977), the record of granular oil bodies in O. decemdentata might have been based on degenerated ones. 2. Odontolejeunea lunulata (F.Weber) Schiffn.  Dicranolejeunea grossiloba Steph. Odontolejeunea calcarata (Spruce) Steph. Odontolejeunea chaerophylla (Spruce) Schiffn. Odontolejeunea ecuadorensis Steph. Odontolejeunea nigrescens Steph. Odontolejeunea sieberiana (Gottsche) Schiffn. Odontolejeunea spiniloba Steph.

Fig. 58C, D

584

 

Keys and Descriptions

Very common on living leaves and bark in low and and montane rainforests and scrub, from sea level to 3200 m. Colombia: Ama, Boy, Cal, Cho, Cun, Hui, Mag, Put, Qui, Ris, San, Tol, Val, Vau. Ecuador: Ch, EO, Es, LR, MS, Na, Or, Pa, Pi, Su, Tu, ZC. General distribution: tropical America, East Africa. Odontolejeunea lunulata is one of the most common epiphyllous bryophyte species in the neotropics and has been described under many different names. The species is recognized by (1) plants 1.5–3  mm wide, brownish-green; (2) leaves irregularly crisped-undulate when dry; (3) leaf margins with many sharp teeth, the teeth along the ventral margin of the leaf lobe larger and more widely spaced; (4) lobule with 2–4 teeth; and (5) underleaf margins usually with a small teeth, rarely entire, underleaf bases auriculate, the auricles folded. For differences with other members of Odontolejeunea, see the key. Records cited from “Azuay” (Gualaquiza and Bomboiza, leg. Allioni) are from Morona Santiago. 3. Odontolejeunea rhomalea (Spruce) Steph. Odontolejeunea grandiloba Steph. Odontolejeunea obversilobula Herzog

Fig. 58E

On living leaves and twigs in the understory and canopy of ± undisturbed lowland and lower montane rainforests, from sea level to 1500(−3000?) m. Colombia: Cho, San, Val. Ecuador: Es, LR, MS, Na, Or, Pa, ZC.  General distribution: tropical America. Odontolejeunea rhomalea is a characteristic neotropical epiphyllous liverwort that is distinguished by (1) plants rather robust, 2–3.5 mm, usually reddish-brown to blackish-brown when dry; (2) lobules very high, much broader than long, with 4–6 teeth; (3) leaf cells arranged in transverse rows, usually with large trigones; and (4) underleaf margins (and leaf margins) sharply toothed. The species may be confused with O. lunulata but is separated from the latter by the higher lobule with more teeth (usually 5–6), very sharply toothed underleaves, and frequently reddish-brown plant color. Records cited from “Azuay” (Gualaquiza and Bomboiza, leg. Allioni) are from Morona Santiago. Otigoniolejeunea (Spruce) Schiffn. Plants small, 1–2  cm long, 0.8–1.5  mm wide, pale green to dark green, sparsely branched, creeping. Stems of 7–9 outer cell rows surrounding 7–15 smaller inner rows, cells thin-walled; ventral merophyte 2 cells wide. Leaf lobes ovate, apex rounded to subacute, plane or recurved, margins entire to crenulate. Leaf cells convex, smooth, thin-walled with small trigones and intermediate thickenings, cells becoming larger toward the leaf base; ocelli 1–4(−10) per leaf, adjacent or spaced, suprabasal, 2–3 cells up from leaf base, ocelli as large as or slightly larger than green cells. Lobules ca. 1/4 of leaf lobe, sometimes reduced, inflated, short bottleshaped, apex with 1 slightly elongate tooth, hyaline papilla proximal to the first tooth, keel smooth. Underleaves distant, 1.5–3× stem width, bifid to 1/2,

Lejeuneaceae

585

suborbicular, insertion line almost transverse. Androecia usually on short branches, bracteoles restricted to the base of the spike. Gynoecia on main stem or on elongate branches, with 1–2 lejeuneoid and pycnolejeuneoid innovations. Perianth 5-keeled near apex, keels produced as flat, entire or branched laciniae. Sporophytes lejeuneoid. Vegetative reproduction by elongate, ligulate gemmae and by caducous leaf lobes. A small neotropical genus of two species, one in Colombia and Ecuador. For main characters of the genus, see under the species. Literature: Reiner-Drehwald and Ilkiu-Borges (2008 as Lejeunea huctumalcensis), Wei et al. (2014) 1. Otigoniolejeunea huctumalcensis (Lindenb. & Gottsche) Y.M.Wei, R.L.Zhu & Gradst.Fig. 58F Lejeunea huctumalcensis Lindenb. & Gottsche Ceratolejeunea dentatocornuta Steph. Ceratolejeunea dussiana (Steph.) Dauphin Lejeunea acanthotis Spruce Lejeunea schizotis Spruce Lejeunea xiphotis Spruce Rectolejeunea intermedia R.M.Schust., nom. inval. (?) (fide Reiner-Drehwald and Grolle 2012) On bark of tree trunks, tree bases, lianas, twigs, and rotten wood in undisturbed and disturbed rainforest, occasionally on living leaves, from sea level to about 2000 m. Colombia: Ama, Cho, Mag, Met, Qui (Filandia, Gradstein 12721, 12724, HUQ). Ecuador: Ga (new: Santa Cruz, 500  m, Howell 258, det. M.E.  ReinerDrehwald, CAS, GOET; ibid., Miconia belt, van der Werff 1757, GOET), MS, Tu (Gradstein 12124a, GOET), ZC (Río Nangaritza, Gradstein 12101, GOET). General distribution: tropical America. Otigolejeunea huctumalcensis has been a misunderstood neotropical species that was described in four different genera. Reiner-Drehwald and Ilkiu-Borges (2008 as Lejeunea huctumalcensis) provided a detailed treatment of the species; characteristics are (1) plants autoicous or dioicous, delicate, growing in low, flat mats, stem cells thin-walled; (2) leaves ovate to ovate-triangular, sometimes caducous, apex rounded to subacute, plane or reflexed, margins entire or crenulate, ocelli present in the lower half of the leaf above the base, 1–4 per leaf; (3) leaf cells mammillose, with small trigones and intermediate thickenings; (4) lobules small, bottle-shaped, sometimes reduced, apex with 1 tooth; (5) underleaves distant, bifid, suborbicular, 1.5–3× stem width; (6) gynoecia with 1–2 lejeuneoid and/or pycnolejeuneoid innovations; (7) perianth 5-keeled near apex, keels produced as flat, entire or branched (antler-like) laciniae; and (8) vegetative reproduction by ligulate gemmae (without rhizoids) and by caducous leaf lobes. Otigoniolejeunea huctumalcensis is somewhat similar to Xylolejeunea, but the stem cells in Xylolejeunea are thick-walled; the perianth keels lack lacinia; the lobules are never reduced and with 2 teeth; 2–5 ocelli are present above the leaf base in a transverse, suprabasal row; and the innovations are always pycnolejeuneoid (never lejeuneoid).

586

 

Keys and Descriptions

Pictolejeunea Grolle Plants small, creeping, 1–2 cm long, 0.5–1 mm wide, pale green to brown, often conspicuously dark-reddish spotted. Stems with hyalodermis; ventral merophyte 2 cells wide. Leaf lobes wide-spreading, with ocelli, apex rounded, often recurved, margins crenulate-papillose or entire. Cells with uniformly thickened walls, trigones lacking, cuticle with or without large, lens-shaped papilla; oil bodies unknown; ocelli scattered in leaves and underleaves, bracts and perianths, protruding from the leaf surface, in dried plants dark reddish to pale brown, in fresh plants at least in part grayish, usually larger than adjacent cells. Lobules ca. 1/3 of leaf length, ovateinflated, with a blunt, 1-celled tooth, hyaline papilla on the inner side of the tooth. Underleaves small, 1–2× stem width, bifid or undivided, with ocelli, margins entire or toothed, insertion line curved. Autoicous. Androecial bracteoles limited to the base of the spike. Gynoecia on a very short branch without innovations. Perianths ± flattened with 2 broad lateral keels expanded above into short auricles and sometimes with 2 ventral keels. Sporophytes lejeuneoid. Vegetative reproduction (present in P. picta) by means of 2-cell-wide linear gemmae from leaf margins. A predominantly neotropical genus (6 spp.), two species in Colombia and Ecuador. The main characters of Pictolejeunea are the numerous, reddish or brown ocelli in leaves and underleaves of dried plants, the gynoecia on short branches without innovations, and the 2(−4)-keeled perianths with lateral keels expanded above into short auricles. The ocelli are usually larger than adjacent cells. Pictolejeunea resembles Prionolejeunea but differs from the latter in the presence of ocelli. Literature: Grolle (1977), Pócs (2007) 1. Leaf margins finely toothed. Leaf cells with a broad papilla. Ocelli at least in part dark reddish. Ventral surface of perianth without keels . . . . . . . . . P. picta 1. Leaf margins entire. Leaf cells without papilla. Ocelli brown. Ventral surface of perianth with 2 keels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sprucei 1. Pictolejeunea picta (Steph.) Grolle Prionolejeunea cuatrecasii H.Rob.

Fig. 58G, H

On bark and rotten wood in lowland rainforest, from sea level to 200  m. Colombia: Ama, Cho, Val. Ecuador: Or (Mota de Oliveira and ter Steege 2013). General distribution: tropical America. Pictolejeunea picta is a widespread neotropical species that is readily recognized by the numerous large, reddish ocelli in leaves and underleaves (some ocelli may be pale-colored), finely toothed leaf margins, and papillose leaf cells. 2. Pictolejeunea sprucei Grolle On bark and rotten wood in Amazonian rainforest, 100–350 m. Colombia: Ama (Pinzón et al. 2003); to be expected in Ecuador. General distribution: Amazonia. Pictolejeunea sprucei is a rare Amazonian species that is readily separated from P. picta by the characters given in the key.

Lejeuneaceae

587

Prionolejeunea (Spruce) Schiffn. Plants very small- to medium-sized, 0.2–1.5(−2) mm wide, pale green to yellowishbrown, creeping, rarely pendent. Branches Lejeunea-type. Stems with hyalodermis; ventral merophyte 2 cells wide. Leaf lobes suberect to widely spreading, often with a narrow base and abruptly dilated to distal portions, apex acute-apiculate to rounded, margins crenulate-denticulate by conically projecting cells, the tips of these cells often thickened by a papilla, marginal cells projecting outward to upward, lobe surface smooth or roughened by papillose or thick-walled cells. Cells with trigones and intermediate thickenings, cuticle smooth to verruculose; oil bodies small, finely granular, often only one per cell; ocelli lacking. Lobules ovate, ca. 1/3 of leaf length, sometimes reduced, inflated, with a 1-celled tooth, hyaline papilla proximal, keel usually strongly arched. Underleaves 1–3× stem width, deeply bifid, margins entire to denticulate, occasionally 1 angular tooth on outer margins, insertion line ± curved. Androecia on short branches, bracteoles limited to the base. Gynoecia usually on short branches, without innovations or with 1 short, lejeuneoid innovation, the innovation often with androecia. Perianths obcordate to obpyriform, with 2 flattened lateral keels which are denticulate to ciliate-laciniate and sometimes expanded into short auricles, ventral and dorsal keels lacking. Sporophytes lejeuneoid. Vegetative reproduction by means of caducous leaf lobes, caducous branchlets, or elongate, ribbon-like gemmae without rhizoids. An Afro-American genus of 24 species, most of them (22) in tropical America, 12 species in Colombia and Ecuador. This treatment follows the monograph by Ilkiu-Borges (2016). Prionolejeunea is readily recognized by the leaf margins crenulate-denticulate by conically projecting cells; leaves without ocelli; gynoecia on short branches without or with a short, lejeuneoid innovation; and flattened perianths with 2 broad, denticulate to ciliate-laciniate keels. The genus is particularly speciose in the Chocó. Literature: Ilkiu-Borges (2016) 1. Leaves suberect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Leaves obliquely to widely spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Plants tiny, 0.2–0.6 mm wide. Leaf apex rounded to acute. Dorsal leaf margin witthout prominent tooth near the apex. . . . . . . . . . . . . . P. muricatoserrulata 2. Plants larger, more than 0.6 mm wide. Leaf apex axcuminate. Dorsal leaf margin with a prominent, 1–4-cell-long tooth near the apex . . . . . . P. schlimiana 3. Plants with caducous leaf lobes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Plants without caducous leaf lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Leaf apex acute to acuminate. Leaves asymmetrically ovate, dorsal margin sometimes with a 1–4-cell-long tooth near the apex . . . . . . . . . . . . . . P. decora 4. Leaf apex rounded to subacute. Leaves orbicular to oblong, dorsal margin never with a cell tooth near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. scaberula 5. Cells on dorsal leaf surface and keel usually with a papilla . . . . . . . . . . . . . . 6 5. Cells on dorsal leaf surface and keel smooth . . . . . . . . . . . . . . . . . . . . . . . . . . 8 6. Leaf apex rounded to subacute. . . . . . . . . . . . . . . . . . . . . . . . . . . . P. scaberula

588

 

Keys and Descriptions

6. Leaf apex acute to acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Lobule strongly papillose throughout. Underleaves 1.5–2× stem width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. trachyodes 7. Lobule smooth or slightly papillose near the keel only. Underleaves larger, 2–4× stem width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mucronata 8. Underleaves of main stem large, 3–5× stem width, bases auriculate. Plants robust, pendent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Underleaves of main stem smaller, 1.5–3× stem width, bases not auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Teeth on leaf margins pointing outward, perpendicular to the margin. Lobules often reduced. Underleaf lobes acute to acuminate, margins toothed, rarely entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. arguta 9. Teeth on leaf margins pointing to the leaf apex, obliquely to the margin. Lobules never reduced. Underleaf lobes rounded to acute, margins entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. magnistipula 10. Dorsal leaf margin sometimes with a 1–4-cell-long tooth near the apex . . . . 11 10. Dorsal leaf margin without tooth near the apex . . . . . . . . . . . . . . . . . . . . . . . 12 11. Plants ascending from the substrate or pendent. Leaves obliquely spreading. Underleaves longer than wide. Perianth keels expanded above into short auricles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. schlimiana 11. Plants creeping. Leaves widely spreading. Underleaves wider than long (or as long as wide). Perianth keels not expanded into auricles . . . . P. decora 12. Leaf margins and teeth ± reflexed . . . . . . . . . . . . . . . . . . . . . . . . . P. recurvula 12. Leaf margins and teeth plane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Teeth on leaf margins 1–2 cells long, variable in length. Median leaf cells (20–)25–40 μm long. Plants rare, of lowland rainforest . . . . . . . . P. ampliretis 13. Teeth on leaf margins up to 1 cell long, not variable in length. Median leaf cells smaller, 15–25 μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Vegetative reproduction by means of caducous leaf lobes. Leaves falcate, apex rounded. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. scaberula 14. Caducous leaf lobes lacking. Leaves not falcate, leaf apex rounded to acute to apiculate. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15. Leaves from a narrow base abruptly dilated to the lamina. Dorsal leaf base recurved. Gynoecia without innovations or, rarely, with 1 sterile innovation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aemula 15. Leaves not abruptly widened. Dorsal leaf base plane. Gynoecia with 1–2 innovations, the innovations often fertile and with androecia . . . . . . P. denticulata 1. Prionolejeunea aemula (Gottsche) A.Evans Prionolejeunea gemmata (Gottsche) Steph. Prionolejeunea prionodes Steph.

Fig. 58I–K

On bark and rock in lowland and montane rainforests and woodlands, from sea level to 1800  m. Colombia: Cau, Put, Ris. Ecuador: Ch, EO, Ga, MS, Or, Pi, ZC. General distribution: widespread in tropical America.

Lejeuneaceae

589

Prionolejeunea aemula is a common neotropical species that is recognized by ovate-subfalcate leaves with a narrow base that is abruptly widened to the lamina, a rounded to acute-apiculate apex, finely denticulate margins, a narrowly recurved dorsal leaf base, and the ventral margin forming a sharp angle with the keel. The underleaves are small, suborbicular, 1–2.5× stem width, with obtuse to acute lobes. The gynoecium is on short branch usually without innovation. 2. Prionolejeunea ampliretis Herzog On bark in lowland rainforests, 200–300 m. Colombia: Val (type). Ecuador: Or (Schäfer-Verwimp et  al. 2013a). General distribution: West Indies, Panama, northern South America. Prionolejeunea ampliretis is a rare neotropical species distinguished by ovatefalcate leaves with irregularly toothed margins, the teeth being variable in length (1–2 cells long). The leaf cells in the species are rather large, (20–)25–40 μm long in midleaf, and the perianth apex is broadly truncate, not auriculate. 3. Prionolejeunea arguta (Nees) Steph.

Fig. 58L, M

Pendent or creeping on tree trunks and twigs in lowland and montane rainforest, occasionally on rotten logs, living leaves, or rock, from sea level to 1800  m. Colombia: Cho, Ris, Val. Ecuador: MS, Na, Pa, ZC.  General distribution: West Indies, northern South America. Prionolejeunea arguta is a robust, ± pendent neotropical species with large, auriculate underleaves. The underleaf margins are usually toothed (occasionally entire), and the underleaf lobes are acute to acuminate. The lobules are frequently reduced. 4. Prionolejeunea decora (Taylor) Steph.

Fig. 58Q

On bark of trees, twigs, rotten wood, and living leaves in lowland and montane rainforests, 200–1800 m. Colombia: Cho, Met, Ris, San, Val. Ecuador: EO, MS, Or, Pi. General distribution: tropical America. Prionolejeunea decora is a widespread neotropical species recognized by leaf lobes frequently caducous and with a 1–4-cell-long teeth on the dorsal margin near the leaf apex. The latter character is otherwise only seen in P. schlimiana; for differences between the two species, see under P. schlimiana. Fertile plants of P. decora are recognized by the longly ciliate-laciniate wings of the lateral perianth keels. 5. Prionolejeunea denticulata (F.Weber) Schiffn. Prionolejeunea helleri A.Evans Prionolejeunea macrocardia (Spruce) Steph. Prionolejeunea microdonta (Gottsche) Steph.

Fig. 58N–P

On bark of trees, twigs, rotten wood, and living leaves in lowland and montane rainforests, from sea level to 2300 m. Colombia: Ama, Ant, Boy, Caq, Cau, Cho, Hui, Put, Vau. Ecuador: Ch, Es, Ga (?), MS, Na, Or, Pa, Tu, ZC. General distribution: widespread and common in tropical America. Prionolejeunea denticulata is one of the most common species of Prionolejeunea in tropical America. Characteristics are the wide-spreading, ovate leaves with a

590

 

Keys and Descriptions

narrow base that is gradually (not abruptly) widened to the lamina, a rounded to acute-apiculate leaf apex, finely crenulate-denticulate margins, a plane dorsal leaf base, and the ventral margin forming a rather wide angle with the keel. The underleaves are like in P. aemula. The gynoecium is situated on a short branch and usually has 1–2 innovations with androecia. 6. Prionolejeunea magnistipula Herzog 

Fig. 59B, C

Pendent on twigs in wet lowland and submontane rainforests along the Pacific coast, 200–700 m. Colombia: Cho, Val (Ilkiu-Borges 2016). General distribution: only known from the Chocó region. Prionolejeunea magnistipula is a rare Chocó endemic characterized by large auriculate underleaves and a pendent, Taxilejeunea-like habit. The species resembles P. arguta and has been confused with the latter; for differences see the key. 7. Prionolejeunea mucronata (Sande Lac.) Steph. Prionolejeunea circinulata Herzog On tree trunks, twigs, living leaves, rotten logs, and rock in lowland and montane rainforests, from sea level to 1500  m. Colombia: Cho, Put, Ris. Ecuador: Pa (Ilkiu-Borges 2016). General distribution: tropical America. Prionolejeunea mucronata is recognized by the dorsal surface of the leaf lobe covered with papillae (one per cell), a character shared with P. trachyodes. However, the papillae in P. mucronata are smaller and are mostly lacking on the lobule, and the underleaves are larger (2–4× stem width) and usually wider than long, with rounded bases (as long as wide and with cuneate bases in P. trachyodes) (IlkiuBorges 2016). 8. Prionolejeunea muricatoserrulata (Spruce) Steph.

Fig. 59A

On bark and rotten wood in lowland and montane rainforests in the Western Cordillera of the Andes and along the Pacific coast, from sea level to 1800  m. Colombia: Cho, Ris. Ecuador: Es, Or. General distribution: tropical South America. Prionolejeunea muricatoserrulata is the smallest species of Prionolejeunea (to 0.6  mm wide). It is distinguished by the suberect, ovate-oblong leaves with a rounded to acute apex and denticulate to serrate margins. The lobules are large, ca. 1/2× lobe length, but are frequently reduced and sometimes fully absent. 9. Prionolejeunea recurvula (Spruce) Steph. On twigs in lowland rainforest along the Pacific coast, 200 m. Colombia: Cho (Ilkiu-Borges 2016). General distribution: northern Amazonia (type), Chocó. Prionolejeunea recurvula is a rare species from the upper Rio Negro region and the Chocó. The species is mainly recognized by the narrowly recurved leaf margins with reflexed teeth. The plants are very small, less than 1 mm wide, the leaf apices are acuminate and usually broadly reflexed, and the gynoecia lack innovations.

Lejeuneaceae

591

Fig. 59 (A) Prionolejeunea muricatoserrulata, habit. (B and C) Prionolejeunea magnistipula. (B) Underleaf. (C) Habit. (D and E) Prionolejeunea scaberula. (D) Habit. (E) Leaf margins cells. (F) Prionolejeunea schlimiana, habit. (G and H) Pycnolejeunea contigua. (G) Habit. (H) Leaf. (I–K) Prionolejeunea trachyodes. (I) Cross section of part of a leaf. (J) Habit. (K) Leaf apex, dorsal view. (L and M) Pycnolejeunea decurviloba. (L) Leaf. (M) Habit. (N) Pycnolejeunea macroloba, leaf. (from Ilkiu-Borges 2016; Gradstein and Ilkiu-Borges 2009). Scale bars: A, D, F, G, J, M, N = 250 μm; B, H = 100 μm; C = 500 μm; E, I, K, L = 25 μm

592

10. Prionolejeunea scaberula (Spruce) Steph.

 

Keys and Descriptions

Fig. 59D, E

On bark of trees and shrubs in lowland and (sub)montane rainforests, occasionally on dead logs or rock, from sea level to 1500  m. Colombia: Cho, Put, Val. Ecuador: EO, Na, Or, ZC. General distribution: scattered in tropical America. Prionolejeunea scaberula is an uncommon neotropical species recognized by falcate leaves with rounded tips and by caducous leaf lobes. The ventral margin of the leaf lobe usually forms a sharp angle with the keel, and the leaf cell walls are thickened and sometimes with a low papilla on the dorsal leaf surface. An unusual characteristic of the species is the frequently reddish- or rose-colored branch tips (Ilkiu-Borges 2016). 11. Prionolejeunea schlimiana (Gottsche) Steph. Prionolejeunea fabroniaefolia (Spruce) Schiffn.

Fig. 59F

Ascending or pendent on tree trunks, twigs, lianas, and living leaves in montane rainforests, occasionally on rotten wood and rock, 500–1500 m. Colombia: Cho, Gui, Mag (type), Val. Ecuador: Bo, Ch, Es, LR, MS, Pi. General distribution: Central and South America. Prionolejeunea schlimiana is a rather common neotropical species distinguished by obliquely spreading leaves with an acuminate apex and a prominent tooth, 1–4 cells long, on the dorsal leaf margin, near the leaf apex. The latter character is also seen in P. decora, but in the latter species, the leaf lobes are widely spreading often caducous, the underleaves are usually wider than long (longer than wide in P. schlimiana), and the perianth lacks auricles. 12. Prionolejeunea trachyodes (Spruce) Steph.

Fig. 59I–K

Creeping on bark and palm leaves in lowland and montane rainforests in the Colombian Western Cordillera and along the Pacific coast, 200–1500 m. Colombia: Cho, Ris (Ilkiu-Borges 2016). General distribution: Dominica, Panama, northern South America (Bastos and Ilkiu-Borges 2011). Prionolejeunea trachyodes is a very small neotropical species (0.3–0.7  mm wide) with leaf lobes, lobules, and keel covered by large papillae; each papilla extends over almost the entire surface of a cell. Papillae are also seen in P. mucronata and P. scaberula, but in the latter species, the papillae are much less prominent and are absent on the lobule surface and most of the keel. Moreover, P. mucronata is larger than P. trachyodes. Pycnolejeunea (Spruce) Schiffn. Plants small to rather robust, 1–4 cm long, 0.6–2.5 mm wide, dull and pale whitishgreen to brownish, creeping. Stems of thick-walled cells, hyalodermis lacking; ventral merophyte 2–8 cells wide. Leaf lobes wide-spreading, with or without ocelli, apex rounded to acute to apiculate, often recurved, margins entire, rarely crenulate. Cells convex, trigones small to large, cuticle smooth or papillose; oil bodies large,

Lejeuneaceae

593

1–3 per cell, coarsely granular; ocelli present or absent, when present large, scattered in the leaf lobe or, more often, restricted to the base of the lobe, usually larger than surrounding green cells, lacking in underleaves. Lobules small or large, 0.1–0.5× leaf length, free margin straight or curved, tooth 1-celled, short-obtuse to somewhat elongate-acute, hyaline papilla proximal. Underleaves orbicular to reniform, bifid, imbricate (rarely distant), insertion line arched or almost straight. Androecial bracteoles usually restricted to the base of the spike. Gynoecia on elongate shoots, with 1–2 pycnolejeuneoid innovations. Perianths inflated, with 5 smooth or crenulate keels. Sporophytes lejeuneoid. Vegetative reproduction by caducous or fragmented leaf lobes. A pantropical genus of about 14 species, five in Colombia and Ecuador. The main features of Pycnolejeunea are (1) stems rigid, made up of thick-walled cells, ventral merophyte 2(−8) cells wide; (2) leaf lobes with rounded apex, entire margins and with or without ocelli; (3) cells with large trigones and large, coarsely granular oil bodies; (4) lobules with a well-developed first tooth and proximal hyaline papilla; and (5) gynoecia with pycnolejeuneoid innovations. The genus was monographed by He (1999) and is closely similar to Cheilolejeunea but differs from the latter genus in the proximal hyaline papilla (distal in Cheilolejeunea) and the frequent presence of ocelli (ocelli lacking in P. chocoensis, P. decurviloba, and P. remotistipula C.J.Bastos & C.E.Zartman). Many Pycnolejeunea species are sun epiphytes and grow in open habitats. The plants may be dull pale-colored to brownish. Literature: He (1999), Reiner-Drehwald and Gradstein (2018) 1. Lobules decurved, lobule apex ± touching the ventral leaf margin. Ocelli lacking������������������������������������������������������������������������������������������������������������� 2 1. Lobules not decurved, lobule apex not touching the ventral leaf margin. Ocelli present. Mostly in Amazonian rainforest, occasionally in montane forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Underleaves reniform, 6–7× stem width, shallowly bifid to ca. 1/4. Ventral merophyte 4–6 cells wide. Plants in lowland and montane forests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. decurviloba 2. Underleaves ovate, 3× stem width, bifid to 1/2. Ventral merophyte 2 cells wide. Plants in lowland rainforest of the Chocó . . . . . . . . . . . . . . . . . . P. chocoensis 3. Lobule large, 1/2–2/3× leaf length. . . . . . . . . . . . . . . . . . . . . . . . P. macroloba 3. Lobule smaller, 1/5–1/3× leaf length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Ventral merophyte 4–8 cells wide. Plants 2–3 mm wide. . . . . . P. densistipula 4. Ventral merophyte 2 cells wide. Plants 1–2 mm wide. . . . . . . . . . . P. contigua 1. Pycnolejeunea chocoensis M.E.Reiner & Gradst. On bark in everwet lowland rainforest along the Pacific coast, 30 m. Colombia: Cho (Reiner-Drehwald and Gradstein 2018). General distribution: only known from the type from western Colombia (Chocó).

594

 

Keys and Descriptions

Pycnolejeunea chocoensis is a rare Chocó endemic recognized by decurved lobules with the apex touching the ventral leaf margin; small distant underleaves, ca. 3× wider than the stem; and absence of ocelli. Decurved lobules and absence of ocelli are shared with P. decurviloba, but the latter species is a much more robust plant with broadly reniform underleaves (6–7× stem width) and a 4–6-cell-wide ventral merophyte (2 cells wide in P. chocoensis). 2. Pycnolejeunea contigua (Nees) Grolle Pycnolejeunea spruceana Schiffn.

Fig. 59G, H

On canopy branches in lowland and submontane rainforest or lower down on tree trunks in forest gaps, at forest margins, and in open environments, from sea level to 650(−2000) m; Schäfer-Verwimp et al. (2013a) recorded the species from unusually high elevation (1830–2000 m) in the Reserva Biológica San Francisco. Colombia: Ama, Caq, Cho, Put, Vau. Ecuador: Or, ZC.  General distribution: pantropical. Pycnolejeunea contigua is a common canopy epiphyte of lowland rainforests throughout the tropics. The species is recognized by (1) plants pale-colored to glossy brownish, autoicous but sometimes seemingly dioicous; (2) ventral merophyte 2 cells wide; (3) leaves imbricate, strongly convex, ovate, margins entire, apex rounded; (4) leaf cells rounded, smooth, with conspicuous trigones; (5) ocelli 1–8 in the lower half of the leaf, larger than ordinary cells, sometimes indistinct in herbarium material; (6) lobules small, hidden by the underleaves, with a short, blunt tooth and a smooth to crenate keel; (7) underleaves (sub)imbricate, broadly ovate to reniform, 3–5× stem width, bases cuneate to rounded, insertion line arched; and (8) asexual reproduction by caducous leaf lobes. 3. Pycnolejeunea decurviloba Steph. Fig. 59L, M Cheilolejeunea decurviloba (Steph.) X.L.He (Trachylejeunea decurviloba [Steph.] X.L.He & Grolle) On bark, from sea level to 1700 m. Colombia: Boy, Cho, Nar, Val. Ecuador: unknown locality (syntype). General distribution: Central America, tropical South America. Pycnolejeunea decurviloba is an uncommon neotropical species that is readily distinguished by the very broad, reniform, strongly imbricate underleaves (6–7× stem width) with a broadly truncate and very shallowly bifid apex and the decurved lobules with the lobule apex strongly curved downward and ± touching the ventral leaf margin. The plants are relatively robust, 2–4 mm wide, and the ventral merophyte is 4–6 cells wide. 4. Pycnolejeunea densistipula (Lehm. & Lindenb.) Steph. On bark and logs in open habitats in lowland and montane rainforests, 300–2280 m. Ecuador: MS (He 1999), ZC (Parolly et al. 2004); to be expected in Colombia. General distribution: scattered in tropical America, most common in SE Brazil. Pycnolejeunea densistipula resembles P. contigua but differs in larger plant size (2–3 mm wide), broader ventral merophytes (4–8 cells wide), and more numerous ocelli, 7–21 per leaf lobe (He 1999).

Lejeuneaceae

5. Pycnolejeunea macroloba (Nees & Mont.) Schiffn. Pycnolejeunea callosa (Lindenb.) Steph.

595

Fig. 59N

Xerotolerant epiphyte, on canopy branches in lowland rainforests and on isolated trees, 100–350 m. Colombia: Ama, Caq, Cho, Put, Vau. Ecuador: Or (Mota de Oliveira and ter Steege 2013). General distribution: tropical America. Pycnolejeunea macroloba is a common canopy species of Amazonian rainforests (Campos et al. 2015). The species is readily separated from other members of the genus by the large, rectangular lobules, which are half or more of the leaf length. The plants are usually very pale green to almost whitish in color but may become brownish when dry. Sometimes the leaves in the species are fragmented. Rectolejeunea A.Evans Plants small, 0.5–1 mm wide, dull pale green, creeping, often with tiny, upright flagelliform shoots. Stems fragile, with hyalodermis; ventral merophyte 2 cells wide. Leaf lobes wide-spreading, plane, apex rounded or obtuse, occasionally acute to apiculate, margins entire, sometimes producing short rhizoids (on stems terminating in a flagelliform shoot), ventral margin forming a straight line with the keel. Cells plane, small, ca. 14–20 μm long in the middle of the lobe, walls thin or uniformly thickened, without trigones; oil bodies lacking in green cells or minute; ocelli present in a group at the leaf base (usually forming a short row) and scattered through the lobe. Lobules 1/4–1/3× leaf length, strongly inflated, ovoid to ellipsoid, mouth truncate, tooth 1-celled, long-falcate, hyaline papilla proximal. Underleaves bifid, small, with or without ocelli, orbicular to obovate, margins angulate, insertion line almost straight. Gynoecia on long shoots, with 1–2 pycnolejeuneoid innovations. Perianths compressed, 3-keeled, with a broad 2-angled ventral keel. Sporophytes lejeuneoid. Vegetative reproduction by caducous leaves produced on upright flagelliform shoots, caducous leaves usually small, without lobules or with a small, filiform lobule (R. versifolia), margins of caducous leaves with rhizoids or cilia; underleaves on flagelliform shoots flat or subsquarrose, margins entire or toothed. A small neotropical genus of four species, all of them in Colombia and Ecuador (Gradstein and Ilkiu-Borges 2019). Many further species were assigned to Rectolejeunea in the past, but all of them have been transferred to other genera, especially to Lejeunea, or treated as synonyms (Reiner-Drehwald and Grolle 2012; Zhu et al. 2018). The main features of the genus Rectolejeunea as currently defined are (1) plants very small, less than 1 mm wide; (2) lobules small, ca. 1/4 leaf length, strongly inflated, with a truncate apex and a long, falcate tooth; (3) leaf cells very small, less than 20  μm long in midleaf, with thin or uniformly thickened walls, without trigones, cuticle smooth, oil bodies usually absent; (4) ocelli present at leaf base (scattered or in a short row) and scattered through the leaf lobe, sometimes also present in underleaves (R. emarginuliflora, R. halinae); (5) gynoecia with pycnolejeuneoid innovations; and (6) vegetative reproduction by small caducous leaves produced on upright flagelliform shoots (species with caducous leaves produced on normal shoots are now excluded from Rectolejeunea).

596

 

Keys and Descriptions

Literature: Reiner-Drehwald and Grolle (2012), Zhu et al. (2018), Gradstein and Ilkiu-Borges (2019) 1. Leaf apex obtuse to acute-apiculate. Underleaves with ocelli. Plants of western Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. halinae 1. Leaf apex rounded (sometimes acute-apiculate on leaves just below the flagella). Underleaves with or without ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Margins of caducous leaves with long cilia, the cilia composed of a row of 2–7 cells. Underleaves on flagella subsquarrose, toothed. Plants autoicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. flagelliformis 2. Margins of caducous leaves without cilia (but with unicellular rhizoids). Underleaves on flagella flat (appressed to stem), entire. Plants dioicous . . . . 3 3. Underleaves with ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. emarginuliflora 3. Underleaves usually without ocelli . . . . . . . . . . . . . . . . . . . . . . . . R. versifolia 1. Rectolejeunea emarginuliflora (Schiffn.) A.Evans On bark and living leaves in lowland and submontane rainforests, 80–1000 m. Colombia: Ama, Caq, Put, Ris (Reiner-Drehwald and Grolle 2012), Val, Vau (Campos et  al. 2015). Ecuador: Es, Or, ZC (Schäfer-Verwimp et  al. 2013a). General distributon: West Indies, tropical South America. Rectolejeunea emarginuliflora is a misunderstood species. Traditionally this species was distinguished by caducous leaf lobes produced on normal leafy shoots, but Reiner-Drehwald and Grolle (2012) showed that the caducous leaves of R. emarginuliflora are produced on flagelliform branches, like in all other genuine members of the genus. According to the latter authors, R. emarginuliflora is closely similar to R. versifolia but differs from the latter in the presence of ocelli in underleaves. The published records of R. emarginuliflora not cited in Reiner-Drehwald and Grolle (2012) must be verified. 2. Rectolejeunea flagelliformis A.Evans

Fig. 60K, L

Sun epiphyte on twigs in the high canopy of primary rainforests and lower down in more open environments, from sea level to about 600 m. Colombia: Cho (ReinerDrehwald and Grolle 2012). Ecuador: MS (Schäfer-Verwimp et al. 2013a). General distributon: scattered in tropical America. Rectolejeunea flagelliformis is a widespread neotropical species that seems to be rare in Colombia and Ecuador, being known from two locations. The species is characterized by (1) plants autoicous, ventral merophyte 2 cells wide; (2) leaf apex mostly rounded, becoming acute to apiculate on leaves below the flagelliform branches; (3) ocelli present in leaves but generally absent in underleaves (exceptionally a few underleaves with one or a few ocelli); (4) lobule tooth long and sharp, falcate, rarely short and obtuse; (5) caducous leaves produced on specialized, flagelliform branches, without lobule, margins of caducous leaves with several long cilia, the cilia 2–7 cells long and often terminating in a short rhizoid; and (6) underleaves on flagelliform branches subsquarrose, margins with a sharp tooth.

Lejeuneaceae

597

Fig. 60 (A–C) Reinerantha foliicola. (A) Habit. (B) Lobule. (C) Underleaf. (D–F) Schiffneriolejeunea polycarpa. (D) Leaf. (E) habit. (F) Median cells. (G–J) Rectolejeunea versifolia. (G) Leaf. (H) Habit. (I) Caducous leaf. (J) Habit with flagelliform shoot producing caducous leaves. (K and L) Rectolejeunea flagelliformis. (K) Habit with flagelliform shoot producing caducous leaves. (L) Caducous leaf. (M–O) Spruceanthus theobromae. (M) Median leaf cells. (N) Leaf. (O) Habit. (A–C from the type; rest from Gradstein et al. 2001; Gradstein and Ilkiu-Borges 2009). Scale bars: A = 300 μm; B, C = 50 μm; D, H, J, L = 250 μm; E, O = 1000 μm; F, M = 25 μm; G, I, K = 100 μm; N = 500 μm

598

 

Keys and Descriptions

Rectolejeunea flagelliformis stands out by autoicy; flagella with subsquarrose, toothed underleaves; caducous leaves with cilia; and underleaves without ocelli (Reiner-Drehwald and Grolle 2012). Schuster (1992c, 2000b) described R. flagelliformis subsp. hamata R.M.Schust. from the island of Gorgona characterized by cilia on caducous leaves with hooked, uncinate tips. A deviant specimen with ocelli in underleaves (Reiner-Drehwald and Grolle 2012, p.  458) is described here as R. halinae. 3. Rectolejeunea halinae Gradst. & Ilkiu-Borges

Fig. 65F–K

On bark in the canopy of submontane rainforest on the western side of the Cordillera Occidental, 1200 m. Colombia: Ris (Gradstein and Ilkiu-Borges 2019). General distribution: only known from the type from the western Colombia. Rectolejeunea halinae is a rare Colombian endemic recognized by the pointed leaf apex (acute to apiculate to shortly acuminate). In other Rectolejeunea species, the leaf apex is rounded; exceptionally, leaves just below the flagelliform shoot in R. flagelliformis have pointed tips. The latter species shares ciliate caducous leaves with R. halinae and shows greatest similarity to R. halinae. Indeed, R. halinae was tentatively identified as R. flagelliformis by Reiner-Drehwald and Grolle (2012, p. 458). Rectolejeunea halinae clearly differs from R. flagelliformis, however, by the presence of ocelli in underleaves, flagelliform shoots with underleaves appressed and with entire margins, and pointed leaf tips. In R. flagelliformis ocelli are lacking in underleaves, the underleaves on flagelliform shoots are squarrose and have toothed margins, and the leaf tips are mostly rounded. The two species possibly differ also in sexuality as R. flagelliformis is autoicous and is usually copiously fertile, whereas R. halinae lacks gametoecia and may be dioicous. By the flat, entire-margined underleaves of flagelliform shoots, R. halinae is similar to R. emarginuliflora and R. versifolia, but the latter two species have rounded leaf tips and caducous leaves without cilia. Moreover, R. versifolia lacks ocelli in underleaves. 4. Rectolejeunea versifolia (Schiffn.) L.Söderstr. & A.Hagborg Fig. 60G–J Pycnolejeunea flagellifera S.W.Arnell Rectolejeunea berteroana (Steph.) A.Evans (fide Söderström et al. 2015a) On bark, rotten wood, or living leaves in lowland and montane forests, in scrub, and on isolated trees, usually in open locations, from sea level to 2700 m. Colombia: Boy, Cho, Ris, Vau. Ecuador: Es, MS, Na, Or, ZC. General distribution: widespread in tropical and subtropical America. Rectolejeunea versifolia is a common neotropical species that is distinguished by (1) plants dioicous, ventral merophyte 2 cells wide; (2) leaf apex rounded; (3) ocelli present in leaves but generally absent in underleaves (exceptionally a few underleaves with 1 or a few ocelli); (4) lobule tooth long and sharp, falcate, rarely short and obtuse; (5) caducous leaves produced on specialized, flagelliform branches, caducous leaves with a small, filiform lobule and with unicellular rhizoids on the margins, cilia absent; and (6) underleaves on flagelliform branches appressed, flat, margins without teeth (Reiner-Drehwald and Grolle 2012).

Lejeuneaceae

599

Reinerantha Gradst. & R.L.Zhu Plants 1.5–2.5 mm wide, epiphyllous, prostrate. Stems with 7 epidermis cells and 3 medullary cells, the cells thin-walled, hyalodermis present. Leaf segmentation spiral. Leaf insertion very short, transverse. Leaf cells mammillose, thin-walled, without trigones, ocelli lacking, oil bodies very small, faintly granular. Lobules with deeply involute free margin, 1 tooth, and an ental hyaline papilla. Stylus present, unicellular. Underleaves one per leaf pair, distant, bifid, with slightly diverging, narrowly lanceolate lobes and a widely lunulate sinus. Autoicous. Androecia on elongate, leafy shoots, male bracts epistatic, bracteoles present throughout the spike. Gynoecia with a pycnolejeuneoid innovation and a 4–5-keeled perianth with rounded, smooth keels. Sporophytes lejeuneoid. Asexual reproduction not observed. A neotropical genus with a single species. For main features of Reinerantha, see under R. foliicola. Literature: Gradstein et al. (2018b) 1. Reinerantha foliicola Gradst. & R.L.Zhu

Fig. 60A–C

On leaves of Piper and Euphorbiaceae in bushes along river, in open environment of meadows and remnant montane rainforest, 1800  m. Ecuador: Pi (Río Guajalito; Gradstein et al. 2018b); not known from Colombia but expected there. General distribution: northern Andes (Ecuador, Venezuela; Pócs 2018). Reinerantha foliicola is a rare northern Andean epiphyllous liverwort that is recognized by (1) plants pale green, 1.5–2.5 mm wide; (2) leaves widely spreading, ovate-orbicular, insertion very short, transverse, margins crenulate; (3) leaf cells very thin-walled, mammillose; (4) lobules 2/5× leaf length, strongly inflated, margin involute, tooth conspicuous, erect, (1–)2 cells long; (5) underleaves 2–3× stem width, distant, broadly ovate (wider than long), Leptolejeunea-like, with a wide, lunulate sinus and narrowly lanceolate-acute lobes; and (6) male bracts epistatic, similar to normal leaves, bracteoles present throughout the male spike. In the very short leaf insertion, R. foliicola resembles Cololejeunea and Diplasiolejeunea, but the leaf segmentation in the latter two genera is pendular (one underleaf per leaf, or underleaves lacking), whereas R. foliicola has spiral leaf segmentation (one underleaf per leaf pair). Reinerantha foliicola is morphologically most similar to the Asiatic genus Tuyamaella, but the underleaves in Tuyamaella are much larger, imbricate, and with broad lobes, and the male bracts are hypostatic. Molecularly, Reinerantha is sister to Diplasiolejeunea and not closely related to Tuyamaella (Gradstein et  al. 2018b). Tuyamaella has been recorded once from South America (Tixier 1973), but the record was based on a mislabelled Asiatic specimen (Gradstein et al. 2018b). Schiffneriolejeunea Verd. Plants 1–2 mm wide, green to rather glossy brown, creeping or ascending. Branches Lejeunea-type, occasionally Frullania-type. Stems with enlarged dorsal epidermis (ventral epidermis not enlarged), epidermis brown; ventral merophyte 4–8 cells

600

 

Keys and Descriptions

wide. Leaf lobes wide-spreading, when dry convolute and wrapped around the stem, apex rounded, margins entire. Cells slightly longer than wide, trigones cordate; oil bodies coarsely granular; ocelli lacking. Lobules 1/3–1/2 leaf length, sometimes reduced, with 1–2 teeth, free margin not inflexed. Underleaves undivided, with entire margins and straight or weakly curved insertion line. Gynoecia without innovations. Perianths with 4(−6) keels, the keels inflated-swollen or sharp. Sporophytes ptychanthoid. Vegetative reproduction by regeneration from leaf cells. A pantropical genus of 14 species, 2 in Colombia and Ecuador. The principal characters of neotropical Schiffneriolejeunea are (1) plants glossy green to brown; (2) stems rigid, with enlarged dorsal epidermis; (3) lobules with 1–2 teeth; (4) cordate trigones and granular oil bodies; (5) obdeltoid underleaves with cuneate bases; and (6) gynoecia without innovations, acute-acuminate bracts and a perianth with 4–5 smooth keels. Literature: Gradstein (1994, 2015b) 1. Lobules ovate, 1/3–2/5× lobe length, with 1 tooth (occasionally with a very small second tooth). Leaves convolute when dry. Ventral merophyte 5–8 cell rows wide. Perianth with rounded keels and a short beak (2–3 cells long). Plants widespread. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. polycarpa 1. Lobules narrow rectangular, 1/2–3/5× lobe length, with 2 distinct teeth. Leaves not convolute when dry. Ventral merophyte 4 cell rows wide. Perianth with 4 sharp keels and a long beak (7–10 cells long). Plants of Amazonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amazonica 1. Schiffneriolejeunea amazonica Gradst. Sun epiphyte on canopy branches in Amazonian forest and in scrub, 100–200 m. Colombia: Put, Vau (Campos et  al. 2014, 2015). General distribution: Amazonia, Guianas. Schiffneriolejeunea amazonica is a rare Amazonian canopy epiphyte that is recognized by the pale brown plants with longly rectangular lobules (ca. 2/3 of leaf length) with 2 distinct teeth, gynoecia on short branches without innovations, boatshaped female bracts with lobe and lobule of equal length, and a sharply 4-keeled perianth with a long (7–10 cells long) beak, without innovations. Dry leaves are rather flat and spreading and are not convolute like in S. polycarpa. 2. Schiffneriolejeunea polycarpa (Nees) Gradst. Ptychocoleus polycarpus (Nees) Trevis.

Fig. 60D–F

Sun epiphyte on bark and dead wood in disturbed lowland and lower montane rainforests, in rather open situations, from sea level to 1700 m. Colombia: Ant, Cas, Cho, Hui, Met, Qui, Ris. Ecuador (new): Es (Burghardt 7285, QCA). General distribution: nearly pantropical. Schiffneriolejeunea polycarpa is a widespread tropical species that is known from several localities in Colombia but has been found in Ecuador only once. The species is distinguished by its glossy brownish color, leaves convolute when dry, ovate lobules with usually only 1 tooth, gynoecia without innovations, acute female bracts, and perianths with 4–5 short, rounded keels near apex.

Lejeuneaceae

601

Schusterolejeunea Grolle Plants small, 1–1.3  mm wide, pale green, creeping to loosely pendent. Stems of thick-walled cells, epidermis hardly enlarged, hyalodermis lacking; ventral merophyte 2 cells wide. Leaf lobes obliquely spreading, oval to narrowly elliptical, apex rounded, plane, margins entire, base very narrow, insertion line 2–3 cells long. Cells elongate-hexagonal, very translucent, thin-walled, without trigones; oil bodies unknown; ocelli lacking. Lobules flat, with a very short insertion, triangular to ligulate, variable in size and shape, standing obliquely upward, apex rounded, tooth absent, hyaline papilla terminal at the rounded apex. Underleaves narrower than the stem, longer than wide, narrowly bifid with erect, lanceolate lobes. Autoicous. Gametoecia numerous, on very short branches (shorter than leaves) at both sides of the stem. Gynoecia 2–4 in a short row, each gynoecium with 1 pycnolejeuneoid or lejeuneoid innovation which is again fertile (except the outermost one). Perianths inflated, with 5 smooth keels. Vegetative reproduction not observed. A neotropical genus with a single species. For main features of Schusterolejeunea, see under the species. Literature: Gradstein et al. (2001), Gradstein and Costa (2003) 1. Schusterolejeunea inundata (Spruce) Grolle Lejeunea inundata Spruce Potamolejeunea sprucei Steph. Semi-aquatic on rock and logs along rivers, rarely on living leaves, 250  m. Colombia (new): Vau (Río Apaporis, Cachivera de Jirijirimo, Soratama, 26–27 Mar 1951, Schultes & Cabrera 11954, 11964, 11970, CINC, PC); to be expected in Ecuador. General distribution: peri-Amazonian, occurring along the upper reaches of tributaries of the Amazon River in Brazil, Venezuela, Colombia, and Peru; also in the Guianas. Schusterolejeunea inundata is a characteristic peri-Amazonian rheophyte that is reported here new to Colombia. The species is recognized by (1) plants growing near running water, leaves obliquely spreading, oval to elliptical, 1.5–3.5× as long as wide, with a very short insertion; (2) lobules flat, triangular to ligulate, obliquely spreading, keel ca. 1/6–1/3× leaf length; (3) leaf cells thin-walled; (4) underleaves longer than wide, small, not wider than the stem, deeply bifid; and (5) gynoecia in clusters on very short branches. The species should be more common in the Amazon region of Colombia and Ecuador and has probably been overlooked. Spruceanthus Verd. Plants robust, 3–4 mm wide, green to brownish, ascending or pendent, forked when fertile. Branches Lejeunea-type only. Stems with a brown cortex of small, thickwalled cells in 2–3 layers surrounding a colorless, thin-walled medulla; ventral merophyte ca. 16 cells wide. Leaf lobes wide-spreading, when dry not appressed or convolute, apex rounded, margins entire. Cells 1–2× longer than wide, trigones radiate, intermediate thickenings 0–3 per wall; oil bodies homogeneous. Lobules very small, reduced to small fold, with 1 tooth. Underleaves undivided, with entire margins and straight insertion line. Gynoecia with 1–2 lejeuneoid innovations;

602

 

Keys and Descriptions

female bract apices acute. Perianths with 5–8 keels. Sporophytes ptychanthoid. Vegetative reproduction not observed. A mainly Asian-Australasian genus of 16 species, 1 species in the neotropics (Ecuador). For main features of Spruceanthus, see under the species. The genus has not yet been monographed. Literature: Gradstein et al. (1994), Kautz and Gradstein (2001), Shi et al. (2015), Sukkharak and Gradstein (2017) 1. Spruceanthus theobromae (Spruce) Gradst. Ptychanthus theobromae (Spruce) Schiffn.

Fig. 60M–O

On bark of old cacao trees in cacao plantations, 300–600  m. Ecuador: Gu (Spruce 1884), LR (Kautz and Gradstein 2001). General distribution: only known from western Ecuador. Spruceanthus theobromae is a rare Ecuadorian endemic that is readily recognized by ± dichotomous branching, robust stems with ca. 16-cell-wide ventral merophytes and 5–8-plicate perianths, which are often arranged in a series. The species resembles members of the genus Dibrachiella, especially D. auberiana, but differs in more robust stems with a differentiated epidermis and subepidermis, presence of small flagelliform branches at stem bases, elongate leaf cells with homogeneous oil bodies (granular in Dibrachiella), and 5–8-keeled perianths (4–5-keeled in Dibrachiella). Stictolejeunea (Spruce) Schiffn. Plants small to large, 1–15 cm long, 1–3 mm wide, pale to dark green, brown or black with age, creeping, ascending or pendent, often (bi)pinnate. Branches Frullania-type or Lejeunea-type. Stems with or without a brown cortex of small, thick-walled cells in 1–3 layers surrounding a colorless medulla of thin-walled cells; ventral merophyte 4–18 cells wide. Leaf lobes wide-spreading, little altered when dry, with scattered ocelli, apex rounded, margins entire, often white-bordered. Cells (sub)isodiametric, walls uniformly thickened; oil bodies absent in green cells; ocelli numerous, scattered in leaves, underleaves, bracts, bracteoles and perianths. Lobules small, to 1/4 leaf length, sometimes reduced, straight or curved backward, with 0–4 teeth. Underleaves undivided, margins entire, sometimes white-bordered, insertion line straight to deeply arched. Gametoecia on very short branches. Male bracteoles limited to base of spike. Gynoecia without innovations or with 1 short, pycnolejeuneoid innovation. Perianths with 2 large auricles above, margins of the auricles sometimes white-bordered. Sporophytes lejeuneoid. Vegetative reproduction not observed. A small pantropical genus of three species, two in Colombia and Ecuador, in two subgenera, subg. Leptostictolejeunea R.M.Schust. and subg. Stictolejeunea (Gradstein 1994). The main features of Stictolejeunea are the presence of scattered ocelli in leaves and underleaves, undivided underleaves, small leaf cells with uniformly thickened walls, frequent presence of a white border on leaves (consisting of dead cells), and perianths with 2 large auricles. Robust plants have stems with a 2-layered brown subepidermis, but delicate plants lack a subepidermis.

Lejeuneaceae

603

Literature: Gradstein (1985, 1994) 1. Branching mainly Frullania-type, usually (bi)pinnate. Underleaves (sub)imbricate, 4–10× stem width, insertion line arched. Plants 1.3–2.5 mm wide, common, growing on bark, rotten wood, and living leaves . . . . . . . . . S. squamata 1. Branching mainly Lejeunea-type, irregular. Underleaves distant, 2–3× stem width, insertion line straight. Plants 1–1.3 mm wide, rare, growing on tree bases and rock along creeks in moist lowland forest . . . . . . . . . . . . . . . . . S. balfourii 1. Stictolejeunea balfourii (Mitt.) E.W.Jones

Fig. 61A

On tree bases and moist rock along creeks in the shaded understory of undisturbed lowland and submontane rainforests, from sea level to 800 m. Colombia: Ama, Cau, Cho, Met; to be expected in Ecuador. General distribution: pantropical. Stictolejeunea balfourii (subg. Leptostictolejeunea) is a widespread but littlecollected pantropical species that is separated from the common S. squamata by the characters given in the key. The species grows in shaded locations along creeks in lowland and submontane rainforests and has probably been overlooked. 2. Stictolejeunea squamata (F.Weber) Schiffn. Stictolejeunea kunzeana (Gottsche) Schiffn.

Fig. 61C, D

On tree trunks, lianas, twigs, logs, and living leaves in lowland and montane rainforests, plantations and gardens, occasionally on moist rock, from sea level to 2400 m. Colombia: Ama, Cau, Cho, Mag, Nar, Qui, Ris, San, Val. Ecuador: Az, Chi, EO, LR, Na, Or, Pa, Pi, ZC. General distribution: widespread in tropical America. Stictolejeunea squamata (subg. Stictolejeunea) is a common neotropical species that is readily recognized by (1) scattered ocelli in leaves and underleaves; (2) (bi) pinnate habit and predominantly Frullania-type branching; (3) small, uniformly thickened leaf cells without trigones and without oil bodies; and (4) leaves and perianths often with whitish margins. Symbiezidium Trevis. Platylejeunea (Spruce) Schiffn. Plants usually large, to 15 cm long, 2–5 mm wide, green to brown, creeping to pendent. Branches Lejeunea-type, sometimes Frullania-type. Stems rigid, epidermis cells thick-walled, larger than medullary cells; ventral merophyte 4–8 cells wide. Leaf lobes wide-spreading, flat or convex, little altered when dry, apex rounded to apiculate, margins entire, rarely toothed near apex. Cells isodiametric, trigones small, triradiate, intermediate thickening frequently present, small; oil bodies large, homogeneous; ocelli lacking. Lobules strongly inflated, 1/10–1/4(−1/3)× leaf length, globose to elongate, fully inflated or with a flat apex, free margin strongly involute, with 0–1 tooth. Underleaves undivided, large, (3–)4–10× stem width, apex flat or recurved, rounded to truncate to emarginate, margins entire,

604

 

Keys and Descriptions

Fig. 61 (A) Stictolejeunea balfourii, habit. (B) Symbiezidium barbiflorum, habit. (C and D) Stictolejeunea squamata. (C) Habit. (D) Leaf. (E) Symbiezidium transversale, habit. (F–H) Thysananthus amazonicus. (F) Underleaf. (G) Habit. (H) Leaf. (I and J) Thysananthus auriculatus. (I) Underleaf. (J) Habit. (From Gradstein and Ilkiu-Borges 2009; Gradstein et al. 2001). Scale bars: A, D, F, H, I = 250 μm; B, C, G = 500 μm; E, J = 1000 μm

Lejeuneaceae

605

insertion line deeply arched. Androecia on short-specialized or elongate branches, male bracteoles limited to base of spike. Gynoecia on very short branches, without innovations or with 1 short, pycnolejeuneoid innovation. Perianths ± flat, apex ciliate-laciniate, ventral perianth surface smooth or with cilia and/or lacinia. Sporophytes lejeuneoid. Vegetative reproduction not observed. An Afro-American genus of four species, three in Colombia and Ecuador, in subg. Symbiezidium (Gradstein and van Beek 1985). The neotropical species of Symbiezidium are readily recognized by the rather robust plants with oblong leaves; isodiametric cells with triradiate trigones and intermediate thickenings; strongly inflated, globose to elongate lobules; and large undivided underleaves (3–)4–10× stem width) with deeply arched insertion. The gynoecia are always on very short branches, without or with a short innovation, and the perianths are ciliate-laciniate. Identification of the species may require material with mature perianths. Sterile plants frequently cannot be identified with certainty. Literature: Gradstein and van Beek (1985), Gradstein et al. (1994) 1. Leaf apex finely toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dentatum 1. Leaf apex entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 3. Plants autoicous (fertile plants bisexual). Androecia on short-specialized male branches, not on long branches. Ventral surface of the perianth with lacinia or short cilia distributed randomly over the surface (and sometimes, additionally, in 1–2 rows). Plants 1.5–2.5(−3) mm wide. Ventral merophyte 4 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. barbiflorum 3. Plants heteroicous (fertile plants unisexual or bisexual). Androecia on shortspecialized male branches or on long branches with vegetative leaves. Ventral surface of perianth smooth or with lacinia arranged in 1–2 rows (never arranged randowly over the surface). Plants larger, 2.5–5 mm wide. Ventral merophyte 4–6 cells wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (S. transversale) 4 4. Ventral surface of perianth smooth . . . . . . . . S. transversale var. transversale 4. Ventral surface of perianth with cilia or lacinia in 1–2 rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. transversale var. hookerianum 1. Symbiezidium barbiflorum (Lindenb. & Gottsche) A.Evans Symbiezidium pogonopterum (Spruce) Steph. Symbiezidium setosum (Steph.) Steph.

Fig. 61B

On bark and decaying wood in lowland and montane rainforests, plantations, and gardens, occasionally on living leaves, from sea level to 2800 m. Colombia: Ama, Cau, Ces, Cho, Hui, Mag, Nar, Put, Ris, San. Ecuador: Az, EO, Es, LR, MS, Na, Or. General distribution: widespread in tropical America; also in Madagascar (peri-Afroamerican). Symbiezidium barbiflorum (subg. Symbiezidium) is a very common and variable species that closely resembles S. transversale but differs from the latter species in being constantly autoicous (S. transversale is heteroicous and may produce unisexual plants). The androecia are always on short-secialized branches, not on elongate branches, and the ventral surface of the perianth is roughened by randomly

606

 

Keys and Descriptions

distributed cilia or lacinia. In S. transversale, androecia are produced on short-specialized or elongate branches, and the ventral perianth surface is smooth or with cilia or lacinia in 1–2 rows, never randomly distributed. Perianth ornamentation in S. barbiflorum varies, however, and perianths with distinctly ciliate-laciniate and with almost smooth ventral surface may be found on the same plant. For identification examination of several perianths may therefore be necessary. Symbiezidium barbiflorum is also separated from S. transversale by smaller plant size and narrower ventral merophyte width (see key), but these characters overlap with S. transversale and are less reliable. Variation is also observed in the size and shape of the lobules, which measure 1/10–1/4 of leaf length and may be rounded to somewhat elongate in shape, and in the width of the underleaves, which varies from 3 to 10× stem width. The observed variation seems to be correlated with the environment, as plants of exposed, rather dry habitats were found to have larger lobules and broader underleaves than plants of shaded, moist habitats (Cornelissen and ter Steege 1989; Gradstein 1994). 2. Symbiezidium dentatum Herzog Symbiezidium transversale (Sw.) Trevis. subsp. dentatum (Herzog) Gradst. & van Beek On bark of trees in lowland and submontane rainforests, from sea level to 900 m. Colombia: Ama, Cho, Put, Val, Vau. Ecuador: Es, Pi. General distribution: Pacific coastal region from Costa Rica to northern Ecuador; Colombian Amazon. Symbiezidium dentatum (subg. Symbiezidium) is readily recognized by the robust plants (3.5–5 mm wide) with a finely toothed apex. The species is common in the Chocó region and, in addition, has been found in the Colombian Amazon (Campos et al. 2015). 3. Symbiezidium transversale (Sw.) Trevis. Symbiezidium subrotundum (Kunth) Trevis.

Fig. 61E

Common on bark and decaying wood in lowland rainforests, plantations, and gardens, less common in montane forests, from sea level to 2600 m. Colombia: Ama, Caq, Cau, Cho, Cun, Mag, Nar, Put, Qui, Ris, San, Tol. Val, Vau. Ecuador: Az, Bo, Es, Ch, Gu, Na, Or, Pa, ZC. General distribution: tropical America. Symbidium transversale (subg. Symbiezidium) is a common, robust epiphyte. The plants are heteroicous (autoicous or dioicous), to 10 cm long and 2.5–5 mm wide, the leaves are oblong with entire margins and a broadly rounded apex, the lobules are globose to elongate, and the underleaves are usually large, (4–)5–10× wider than the stem, with a deeply arched insertion line. Like in S. barbiflorum, the size of the underleaves and of the lobule may vary in relation to the environment (see under S. barbiflorum). Two varieties are recognized in S. transversale based on the ornamentation of the perianth (see below). The distribution of the varieties in Colombia and Ecuador has not been studied, however. Sterile material cannot be identified to varietal level.

Lejeuneaceae

607

Symbiezium transversale is closely related to S. barbiflorum; for a discussion of the differences between the two, see under S. barbiflorum. Material without mature perianths cannot always be identified with certainty. An integrated molecular and morphological investigation into the separation of the species and varieties of Symbiezidium is recommended. 3a. Symbiezidium transversale (Sw.) Trevis. var. transversale Symbiezidium hobsonianum (Lindenb.) Trevis. Habitat as the species. Ecuador: Na (Schäfer-Verwimp et al. 2013a). General distribution: tropical America. Recognized by the smooth perianth surface; cilia and lacinia are in this variety restricted to the perianth apex. Symbiezidium transversale var. transversale is probably widespread in Colombia and Ecuador, but the variety can only be distinguished when perianths are available. 3b. Symbiezidium transversale var. hookerianum (Gottsche) Gradst. & van Beek Lejeunea transversalis (Sw.) Nees var. hookeriana Gottsche Symbiezidium taeniopsis (Lindenb.) Trevis. Symbiezidium vincentinum (Gottsche) Trevis. Habitat as the species. Ecuador: Gu (León-Yánez et al. 2006), Or (leg. J. Deleg). General distribution: tropical America. Recognized by 1–2 rows of cilia or lacinia on the ventral surface of the perianth. Although scarcely recorded, S. transversale var. hookerianum is probably widespread in Colombia and Ecuador. The perianth ornamentation is somewhat variable in this variety. Plants with a single, short row of cilia near the perianth apex approach var. transversale, whereas those with 1–2 random cilia on the ventral surface, additional to the 2 rows, approach S. barbiflorum. The latter species may usually be separated by its smaller size (plants maximally 3 mm wide) and the randomly distributed spines, cilia, and/or lacinia on the ventral surface of the perianth, not in rows. In case of doubt, several perianths per plant should be examined (Gradstein and van Beek 1985; Gradstein 1994). Thysananthus Lindenb. Dendrolejeunea (Spruce) Lacout., Thysanolejeunea (Spruce) Steph.

Mastigolejeunea

(Spruce)

Steph.,

Plants 1.5–6 cm long, 2–4 mm wide, glossy dark green to olive green to black or dark brown, creeping, ascending or pendent; stem bases frequently with flagelliform branches. Branches Lejeunea-type. Stems without enlarged epidermis or with enlarged dorsal epidermis cells (ventral epidermis cells never enlarged), the epidermis strongly thick-walled and brown; ventral merophyte 5–10 cells wide. Leaf lobes wide-spreading, when dry convolute, apex acute, margins toothed or entire. Cells longer than wide, trigones cordate; oil bodies coarsely granular; ocelli lacking. Lobules 1/5–1/2× leaf length, not reduced, with 1–3 teeth, free margin plane or

608

 

Keys and Descriptions

inflexed. Underleaves undivided, obovate to spathulate, plane or squarrose, apex rounded to emarginate, margins entire or toothed, insertion line ± straight. Gynoecia with 1(−2) lejeuneoid innovations. Perianths with 3(−10) keels, entire or toothed. Sporophytes ptychanthoid. Vegetative reproduction by regeneration from leaf cells. A pantropical genus of 30 species (Sukkharak and Gradstein 2017), 4 species in Colombia and Ecuador, in two subgenera, subg. Mastigolejeunea (Spruce) Sukkharak & Gradst. (3 spp.) and subg. Thysananthus (1 sp.) (Sukkharak and Gradstein 2017). The main features of Thysananthus are (1) dark green to blackish or dark brown plants with purely Lejeunea-type branching and rigid stems; (2) leaves when dry suberect and appressed to the stem, the apical portion plane or convolute; (3) leaf cells elongate-prosenchymatic, with cordate trigones; (4) lobules not reduced, with 1–3 teeth; and (5) gynoecia with 1–2 lejeuneoid innovations and a 3(−10)-keeled perianth. Sterile specimens of Thysananthus may be difficult to separate from Schiffneriolejeunea, but the species of the latter genus are typically glossy brown in color and may produce Frullania-type branches. Fertile specimens are readily separated from Schiffneriolejeunea by the presence of innovations (absent in Schiffneriolejeunea) and the mostly sharply 3-keeled perianths (4–5-keeled in Schiffneriolejeunea, with broadly rounded keels). Literature: Sukkharak (2015), Sukkharak and Gradstein (2015, 2017) 1. Leaf apex in most leaves acute-apiculate. Underleaves ± toothed (sometimes entire). Female involucre toothed. Dorsal epidermis cells not larger than medullary cells and ventral epidermal cells. . . . . . . . . . . . . . . . . . . . . . T. amazonicus 1. Leaf apex rounded, never acute-apiculate. Underleaves entire. Female involucre entire. Dorsal epidermis cells larger than medullary cells and ventral epidermal cells. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Lobule of stem leaves with 2–3 teeth . . . . . . . . . . . . . . . . . . . . . . . . T. innovans 2. Lobule of stem leaves with 1 tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Ventral surface of perianth with 1 keel. Leaves 1.2–1.5× longer than wide, ventral margin of the leaf lobe upcurved. Underleaves concave, apex ± recurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. auriculatus 3. Ventral surface of perianth with 2–5 keels. Leaves more elongate, 1.5–2× longer than wide, ventral margin of the leaf lobe plane. Underleaves flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. plicatiflorus 1. Thysananthus amazonicus (Spruce) Schiffn. 

Fig. 61F–H

Sun epiphyte on bark in lowland and submontane rainforests and scrub, usually in the forest canopy, 100–350 m. (Colombia: Ama, Put, Vau (Campos et al. 2015). Ecuador (new): Or (Yasuni National Park, near Yasuni station, in rainforest canopy, ca. 225  m, 2019, M.  Berdugo et  al. 4VH, QCA). General distribution: northern South America, Panama, Costa Rica, Cuba. Thysananthus amazonicus (subg. Thysananthus) is a characteristic, robust Amazonian species that is readily recognized by (1) plants 2–4  mm wide, dark green to blackish-brown, branches only Lejeunea-type, sometimes flagelliform; (2) stems rigid, epidermis cells not larger than medullary cells; (3) leaves convex, oblong, apex in most leaves acute-apiculate (a few leaves may have rounded

Lejeuneaceae

609

apices), margins entire or with a few teeth in the upper half; (4) leaf cells elongateprosenchymatic, with large, cordate trigones that are often coalesced; (5) lobules rectangular, with 0–2 teeth, free margin usually continuing along the ventral margin of the leaf lobe; (6) underleaves spathulate, with emarginate apex and entire to toothed margins; and (7) gynoecia with 1 innovation, female bracts and perianths toothed, perianth sharply 3-keeled. 2. Thysananthus auriculatus (Wilson & Hook.) Sukkharak & Gradst. Fig. 61I, J Mastigolejeunea auriculata (Wilson & Hook.) Schiffn. Mastigolejeunea pittieri Steph. Sun epiphyte on bark and rock in disturbed evergreen lowland and montane forests, scrub, plantations, gardens, and on isolated trees, in rather open situations, from sea level to ca. 1500 m. Colombia: Ant, Ces, Cho, Hui, Met, Put, Qui, Ris, Tol. Ecuador: EO, LR, MS, Or, Pa. General distribution: widespread in tropical and subtropical America but rare in Amazonian rainforest where it is replaced by T. plicatiflorus; also in Africa. Thysananthus auriculatus (subg. Mastigolejeunea) is a common Afro-American epiphyte that is recognized by (1) dark green to blackish color; (2) leaves rather strongly convex to subsquarrose, ovate-oblong, 1.2–1.5× longer than wide, with upcurved to recurved ventral margin; (3) apical margin of the lobule oblique and frequently continuing into the ventral margin of the lobe; (4) underleaves concave to subsquarrose, apex recurved; and (5) perianth 3-keeled. Thysananthus auriculatus may be confused with T. plicatiflorus but is separated from the latter by the less elongate leaves, the upcurved ventral margin of the leaf lobe (rather flat in T. plicatiflorus), blackish-green plant color (usually brown in T. plicatiflorus), and 3-keeled perianths (pluriplicate in T. plicatiflorus). Moreover, T. plicatiflorus grows in more sheltered habitats than T. auriculatus and is largely restricted to the lowland rainforests of northern South America. The record of T. auriculatus from the Galápagos Islands (Clark 1953 as M. auriculata) refers to Dicranolejeunea axillaris (Gradstein 1994). 3. Thysananthus innovans (Spruce) Sukkharak & Gradst. Mastigolejeunea innovans (Spruce) Steph. Sun epiphyte on canopy branches in Amazonian rainforest, from sea level to 700 m. Ecuador (?): MS (Herzog 1952). General distribution: northern South America. Thysananthus innovans (subg. Mastigolejeunea) is a very rare Amazonian canopy epiphyte that has been recorded once from Ecuador; the collection is lost (J. Hentschel, pers. comm.). The species is readily separated from other neotropical Thysananthus species by the large lobules with 2–3 teeth. The plants are autoicous, 2–2.5 mm wide and often brownish in color; the leaves are convex but not squarrose; the lobules are rectangular, to 2/5× lobe length; the underleaves are rather flat, obdeltoid; and the androecia are usually produced on short-specialized branches (rarely on long branches).

610

 

Keys and Descriptions

Fig. 62 (A and B) Thysananthus plicatiflorus. (A) Leaf. (B) Habit. (C and D) Verdoornianthus griffinii. (C) Habit. (D) Median cells. (E) Verdoornianthus marsupifolius, leaf. (F and G) Xylolejeunea crenata. (F) Leaf. (G) Habit. (From Gradstein et al. 1977, 1994; Gradstein and IlkiuBorges 2009). Scale bars: A, E, F = 250 μm; B = 1000 μm; C, G = 500 μm; D = 20 μm

4. Thysananthus plicatiflorus (Spruce) Sukkharak & Gradst. Fig. 62A, B Mastigolejeunea plicatiflora (Spruce) Steph. (Lejeunea plicatiflora Spruce) Lejeunea plicata Spruce On bark in the canopy and understory of lowland and submontane rainforests, from sea level to 800 m. Colombia: Cho, Guj, Mag, Ris; to be expected in Ecuador. General distribution: northern South America. Thysananthus plicatiflorus (subg. Mastigolejeunea) is a characteristic tropical South American lowland rainforest species that is distinguished by (1) leaves rather flat and elongate, ca. 1.5–2× longer than wide, with plane margins; (2) underleaves rather flat and appressed to the stem; (3) lobules small (ca. 1/4× leaf length); and (4) perianth pluriplicate, with a broad, swollen ventral keel split into 2–5 folds. The species has been confused with T. auriculatus; for differences see under the latter species.

Lejeuneaceae

611

Verdoornianthus Gradst. Plants 1–2 mm wide, dull olive green to dark brown when dry, loosely creeping. Branches Lejeunea-type only. Stems with enlarged dorsal epidermis (ventral epidermis not enlarged); ventral merophyte 4–8 cells wide. Leaf lobes obliquely spreading to falcate, when dry weakly convolute, apex rounded, margins entire. Cells subisodiametric, trigones simple-triangular to radiate; oil bodies coarsely granular; ocelli lacking. Lobules 1/2 leaf length, never reduced, with 1 tooth, free margin often inflexed. Underleaves undivided, with entire margins and curved insertion line. Gynoecia without innovations. Perianths with 4–5 sharp keels, the keels slightly toothed above. Sporophytes ptychanthoid. Vegetative reproduction not observed. A small Amazonian genus of two species; both occur in Colombia, and one is known from Ecuador. Verdoornianthus is a characteristic sun epiphyte of the rainforest canopy and has been little collected. The genus is closely related to Archilejeunea but differs in the dull brownish color, enlarged dorsal epidermis cells, obliquely spreading leaves which are convolute when dry, large swollen lobules with ± inflexed margins, and, especially, lack of innovations. The leaf cells are isodiametric and the trigones are radiate. Literature: Gradstein et al. (1994) 1. Lobule ovate, less than 1.5× longer than wide, apex folded, tooth ± invisible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. marsupiifolius 1. Lobule rectangular, more than 1.5× longer than wide, apex plane, tooth clearly visible. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. griffinii 1. Verdoornianthus griffinii Gradst.

Fig. 62C, D

On bark on canopy branches in Amazonian rainforest and scrub, 100–220  m. Colombia: Ama, Caq, Put, Vau (Campos et al. 2014, 2015). Ecuador: Or (Mota de Oliveira and ter Steege 2013). General distribution: Amazonia, Guianas. Verdoornianthus griffinii is a characteristic Amazonian canopy epiphyte that is distinguished by its rectangular lobules with a flat apex and a clearly visible tooth, 2–3 cells long, which is not inflexed. 2. Verdoornianthus marsupiifolius (Spruce) Gradst. Ptychocoleus marsupiifolius (Spruce) Steph.

Fig. 62E

On bark on canopy branches in Amazonian rainforest and scrub, 100–220  m. Colombia: Ama, Caq, Put, Vau (Campos et al. 2014, 2015). General distribution: Amazonia. Verdoornianthus marsupiifolius is a characteristic Amazonian canopy epiphyte that is readily recognized by the ovate lobule with a folded apex (not plane). The lobule tooth is inflexed due to the folded apex and is invisible in optical view; it can only be observed by dissection of the lobule.

612

 

Keys and Descriptions

Vitalianthus R.M.Schust. & Giancotti Plants very small, 0.3–0.7 mm wide, pale green to brown, creeping. Stems of 7 outer cell rows and 3 smaller inner rows, cells thick-walled, hyalodermis lacking; ventral merophyte 2 cells wide. Leaf lobes obliquely rather widely spreading, with ocelli in a row, apex acute to obtuse, plane or recurved, margins entire or crenulate. Cells small, 15–20  μm in leaf middle, with trigones, cuticle smooth; oil bodies finely granular; ocelli brownish or gray, 3–6 in an unbroken row near the leaf base, forming a short vitta, also in female bracts and perianths. Lobules strongly swollen, 2/5–1/2× leaf length, sometimes reduced, tooth long-falcate, a pre-apical tooth sometimes present at the junction of the lobule and ventral margin of the leaf lobe, leaving a circular opening to the interior of the lobule, hyaline papilla proximal. Underleaves very small, narrowly bifid to 1/2, lobes erect or slightly spreading, outer margins angular, insertion line straight. Gynoecia on short branches, with 1–2 pycnolejeuneoid innovations. Perianths pyriform, with 0–5 smooth or crenulate keels. Sporophytes lejeuneoid. Vegetative reproduction by fragmentation. A small pantropical genus of four to five species, one in Colombia. The main characters of Vitalianthus are (1) plants very small (less than 1  mm wide), with obliquely spreading leaves; (2) leaf lobes with a short vitta of 3 gray to brownish ocelli in a row; (3) large inflated lobules with a falcate tooth; and (4) small, bifid underleaves with erect to slightly diverging, lanceolate lobes. Literature: Bischler et al. (1963), Bechteler et al. (2016) 1. Vitalianthus aphanellus (Spruce) Bechteler et al. Microlejeunea aphanella (Spruce) Steph. On bark in lowland and submontane rainforest, 100–800  m. Colombia: Vau (Campos et al. 2014). General distribution: Amazonia, French Guiana, southeastern Brazil (São Paulo). Vitalianthus aphanellus is a delicate Amazonian species that is distinguished by obliquely spreading, oblong leaves with an obtuse apex, large lobules, a vitta of 3 ocelli in the lower half of the leaf, and small, bifid underleaves (1–2 × stem width) with lanceolate-acute, slightly spreading lobes. The records from high elevation in Ecuador (Arnell 1961, 1962) are probably erroneous (Bischler et al. 1963). Xylolejeunea X.L.He & Grolle Plants 1–1.5 mm wide, dull whitish to yellow green to brown, creeping. Stems cells thick-walled, hyalodermis absent; ventral merophyte 2 cells wide. Leaf lobes ovate, sometimes falcate, apex rounded to acute, often recurved, margins entire to crenulate to finely denticulate, dorsal leaf surface smooth or finely papillose. Leaf cells convex, smooth or mamillose and with a papilla on the dorsal surface, at the base with 2 transverse rows of larger, elongate cells, the upper, suprabasal row of 2–5 adjacent ocelli, the lower row without ocelli; trigones usually lacking, rarely present. Lobules 1/5–2/5 of leaf lobe, sometimes reduced, inflated, ovoid to

Lejeuneaceae

613

triangular-ovoid, apex with 1–2 teeth, first tooth slightly elongate-falcate, second lobule tooth short and blunt or reduced, teeth often with strongly thickened outer wall, hyaline papilla proximal, keel smooth to roughed with papillae, or finely denticulate. Underleaves small, distant, 1.0–2.5× stem width, bifid, ± orbicular, insertion line almost transverse. Androecia usually on short or elongate branches, bracteoles restricted to the base of the spike. Gynoecia on short or elongate branches, without or with a short, sterile pycnolejeuneoid innovation. Perianths obovate, inflated, sharply 5-keeled, each keel with 2 finely denticulate wings. Sporophytes lejeuneoid. Vegetative reproduction by caducous leaf lobes. An Afro-American genus (5 spp.), 2 species in Colombia and Ecuador. The main features of Xylolejeunea are (1) leaf margins frequently finely crenulate, dorsal leaf surface smooth or rough by mamillose cells crowned by a small papilla; (2) leaf base with 2 transverse rows of larger, elongate cells, the upper row made up of ocelli (2–5); (3) leaf cells without trigones, thin-walled or evenly thick-walled; (4) underleaves small, distant, bifid; (5) gynoecia without or with a short pycnolejeuneoid inovation; and (6) perianths 5-keeled, keels finely toothed and often with paired wings. Literature: He and Grolle (2001) 1. Dorsal leaf surface smooth. Perianth keels winged. . . . . . . . . . . . . . X. crenata 1. Dorsal leaf surface papillose in the upper half. Perianth keels not winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. pellucidissima 1. Xylolejeunea crenata (Nees & Mont.) X.L.He & Grolle  Crossotolejeunea crenata (Nees & Mont.) Lacout. Trachylejeunea pandurantha (Spruce) Steph. Trachylejeunea spruceana Steph.

Fig. 62F, G

On rotten wood in the shaded understory of lowland and submontane rainforests, occasionally on living leaves, 100–1075 m. Colombia: Ama, Caq, Cho, Put, Vau. Ecuador: Na, Or, ZC (Schäfer-Verwimp et  al. 2013a). General distribution: tropical America. Xylolejeunea crenata is a characteristic shade epiphyte in Amazonian rainforests that is recognized by its rather dull greenish color, leaves with a rounded to subacute apex, entire to finely denticulate or crenulate margins and a smooth dorsal surface, leaf cells thin-walled to evenly thick-walled and with 2–5 ocelli in a transverse row near the leaf base (some additional ocelli may occur scattered in the distal portion of leaf lobe), and small, distant underleaves. The gynoecia have no innovation or 1 short innovation, and the perianth keels have paired wings that are finely denticulate. The leaf lobes are frequently caducous. 2. Xylolejeunea pellucidissima (Spruce) Gradst., comb. nov. Lejeunea pellucidissima Spruce, Trans Proc. Bot. Soc. Edinburgh 15: 1884. Type: Venezuela, Amazonas, San Carlos del Río Negro, Spruce s.n. (isolectotype G-00264517!) On trunk bases in Amazonian rainforest, ca. 100–300  m. Ecuador (new): Or (Yasuni National Park, near Yasuni Station, on trunk base in valley, 220 m, 2019,

614

 

Keys and Descriptions

M.  Berdugo et  al. 24VV, QCA). General distribution: Amazonia (Venezuela, Brazil, Ecuador). Xylolejeunea pellucidissima was considered a synonym of X. crenata by He and Grolle (2001) but differs from X. crenata in the papillose dorsal leaf surface (dorsal leaf surface smooth in X. crenata) and the finely denticulate perianth keels without wings (winged in X. crenata) (see also Spruce 1884, p. 185).

Porellaceae A family of two genera, one genus in Colombia and Ecuador. Porella L. Madotheca Dumort. Plants robust, 5–20 cm long and 3–7 mm wide, green to brown, creeping or pendent, (bi)pinnate. Branches Frullania-type. Stems brown, rigid, with a thick-walled cortex; paraphyllia sometimes present. Leaves incubous, divided into a large dorsal lobe and a small ventral lobule, apex of the lobe rounded or mucronate, margins entire or toothed, bases sometimes ciliate-laciniate. Lobules flat and usually distinctly elongate parallel to the stem (or slightly spreading), almost free from the dorsal lobe, margins entire to dentate-ciliate or laciniate, apex broadly rounded. Leaf cells with trigones and smooth cuticle; oil bodies small and homogeneous, numerous per cell. Underleaves large, longer than wide, apex broadly rounded, undivided, margins entire or dentate-ciliate. Rhizoids in bundles from underleaf bases. Dioicous. Gametoecia on short branches. Sporophytes surrounded by a bluntly 3-keeled perianth, foot penetrating into the stem. Setae short and thick. Capsule globose, walls 3–6-layered. Elaters free, not attached to the capsule valves. Spores large, multicellular, germination endosporous. Vegetative reproduction unknown. About 50 species worldwide, 6  in Colombia and Ecuador. The neotropical Porella species have been revised by Swails (1970) and So (2005). The main characters of the genus are (1) plants robust, pinnate, dark green to brown; (2) lobule elongate ± parallel to the stem, flat, not turned into a watersac; (3) gynoecium with numerous archegonia (to 30); (4) perianth without beak; (5) foot of the sporophyte penetrating into the stem; (6) capsule wall 3–6-layered; and (7) elaters free, not attached to the capsule wall. Literature: Swails (1970), So (2005) 1. Ventral margin of leaf lobes undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Ventral margin of leaf lobes not undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Margins of lobules and underleaves strongly toothed to at least the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. brachiata

Porellaceae

615

2. Margins of lobules and underleaves toothed only at the base, otherwise entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. crispata 3. Margins of lobules and underleaves toothed over most of their length. Leaf lobules ligulate, less than 0.5 mm wide. . . . . . . . . . . . . . . . . . . . P. brasiliensis 3. Margins of lobules and underleaves entire or toothed at the base. Leaf lobules ligulate or oblong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Leaf lobules ligulate, 0.2–0.4 wide. Underleaves scarcely wider than the stem, to 0.7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. swartziana 4. Leaf lobules broader, oblong, 0.5–1  mm wide. Underleaves distinctly wider than the stem, usually more than 0.7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Leaf apex recurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. squamulifera 5. Leaf apex plane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. leiboldii 1. Porella brachiata (Taylor) Spruce Madotheca splendida Herzog Pendent from tree trunks and on rock in upper montane rainforest, 2200–3200 m. Colombia: Boy, Cun, Qui, Ris, San, Tol. Ecuador: Lo, Pi. General distribution: Mexico to Bolivia. Porella brachiata is one of the most robust species of Porella in the neotropics, with leafy shoots to 25 cm long and 5 mm wide. The species is distinguished by the undulate ventral margin of the leaf lobes and the strongly ciliate-laciniate margins of lobules and underleaves. Porella brachiata may be confused with P. crispata, but in the latter species, the lobules and underleaves are entire except at the base. 2. Porella brasiliensis (Raddi) Schiffn.

Fig. 63D

On tree trunks and rock in lower montane rainforests, 1000–2000 m. Colombia: Cun, Qui, Ris, San; not known from Ecuador but expected there. General distribution: tropical America. Porella brasiliensis is a common neotropical species that is recognized by the ligulate leaf lobules (only 0.2–0.4 mm wide) with ciliate-laciniate margins and the oblong leaf lobes, which are more than 1.5× longer than wide and entire except at the ventral base. The underleaf margins are ciliate-laciniate. Porella brasiliensis resembles P. swartziana in the shape of leaf lobes and lobules, but in the latter species, the lobules and underleaves are entire except at the base. 3. Porella crispata (Hook.) Trevis. Jungermannia crispata Hook. Madotheca cardoti Steph. Madotheca commutata Gottsche var. lindigiana Gottsche Madotheca piligeri Steph. Porella acanthota Spruce Porella arborea (Taylor) Spruce

Fig. 63C

616

 

Keys and Descriptions

Fig. 63 (A–C) Jubula bogotensis. (A) Habit with gynoecium. (B) Median cells. (C) Leaf. (D) Porella brasiliensis, leaf and underleaf. (E) Porella crispata, habit. (F) Porella leiboldii, leaf and underleaf. (G) Porella squamulifera, leaf. (H) Porella swartziana, habit with gynoecium. (D–H redrawn from Swails 1970; rest from Gradstein et al. 2001). Scale bars: A, D = 500 μm; B = 25 μm; C = 250 μm; E = 1000 μm; F, G = 400 μm; H = 1250 μm

Porellaceae

617

Pendent from tree trunks and on rock in montane rainforest, 1000–3200  m. Colombia: Boy, Cas. Ces, Cun, Hui, Nar, Qui (type), Ris, San, Tol, Val. Ecuador: Az, Ca, Cañ, Ch, Lo, Pi, Tu, ZC.  General distribution: throughout tropical America. Porella crispata is a common neotropical species very similar to P. brachiata but differs from the latter in lobules and underleaves ciliate-laciniate only at the base, otherwise entire. Moreover, the ventral margin of the leaf lobes in P. crispata is more strongly undulate-crispate than in P. brachiata. 4. Porella leiboldii (Lehm.) Trevis. Porella expansa (Lindenb. & Gottsche) Trevis.

Fig. 63F

Pendent from tree trunks and on rock in submontane and lower montane forest, also on soil, 650–3000  m. Colombia: Ant, Boy, Cau, Ces, Cun, Mag, Qui, Tol. Ecuador: EO, Im, Lo (Benitez and Gradstein 2011), Na. General distribution: Mexico to Peru, Greater Antilles. Porella leiboldii is a rather widespread neotropical species distinguished by margins of leaf lobes, lobules, and underleaves ciliate-laciniate at the base, otherwise entire. The leaf margins may be plane or slightly upcurved but the leaf apex is always plane. The species approaches P. squamulifera, but in the latter species, the leaf apex is conspicuously recurved. Both species may produce numerous stem paraphyllia. Porella leiboldii also resembles P. reflexa from subtropical South America, but the latter differs in the reflexed lobule apex and the entire bases of lobules and underleaves. The record of P. reflexa from Magdalena (Winkler 1976 as P. rugulosa Ångstr.) probably refers to P. leiboldii. 5. Porella squamulifera (Taylor) Trevis. Madotheca columbica Taylor Porella rotundifolia Schiffn.

Fig. 63G

On tree trunks and rotten wood in montane forests and páramo, 1200–4000 m. Colombia: Boy, Cau, Ces, Cun, Hui, San. Ecuador: Az, Ca, Es, Im, Na, Pi. General distribution: tropical Andes. Porella squamulifera is close to P. leiboldii in the mostly entire leaf and underleaf margins (some irregularly laciniate teeth are present at the base) but differs in the conspicuously recurved leaf apex. The species often produces stem paraphyllia. 6. Porella swartziana (F.Weber) Trevis. Porella muelleri (Gottsche) Spruce Porella ptilopsis Spruce

Fig. 63H

On trees trunks, rotten logs, rock, and soil in montane forests, 800–3000  m. Colombia: Ces, Cho, Cun, Nar, Qui, Ris, San Tol. Ecuador: Ch, EO, Lo, Tu, ZC. General distribution: throughout tropical and subtropical America; also in South Africa (Hentschel et al. 2007). Porella swartziana is a widespread Afro-American species that is readily distinguished by the ligulate lobules (ca. 0.2–0.4 mm wide) and the narrowly ovate-olong underleaves (scarcely wider than the stem) with margins entire except at the base and often connate with the leaf base. The species resembles P. brasiliensis, but the

618

 

Keys and Descriptions

margins of lobules and underleaves in the latter species are ciliate-laciniate over most of their length. Excluded Record Porella reflexa (Lehm. & Lindenb.) Trevis. (= Porella rugulosa Ångstr.) – Colombia: Mag (Winkler 1976), Qui (Orrego and Uribe 2004). Species endemic to subtropical South America (So 2005); the material from Colombia possibly belongs to P. leiboldii.

Radulaceae A family with a single genus. Radula Dumort. Plants small- to medium-sized, 1–10 cm long, ca. 1–3 mm wide, dull pale green to bright green to brown, creeping, sometimes pendent, (bi)pinnate or dichotomously branched. Branches Radula-type. Stems rigid, without hyalodermis. Leaves incubous, divided into a large dorsal lobe and a small ventral lobule, apex of lobe rounded, occasionally acute, margins usually entire, occasionally crenulate or toothed. Lobules broadly attached to the dorsal lobe by a keel and to the stem, quadrate to oblong, flat or somewhat inflated. Cells thin-walled or with small trigones, cuticle smooth, rarely verruculose; oil bodies very large, brown, 1–2 per cell, rarely more, the additional oil bodies usually smaller. Underleaves lacking. Rhizoids in bundles from the lobules. Dioicous, rarely monoicous. Gynoecium with 2–4 archegonia. Sporophytes surrounded by a narrowly elongate, flattened, 2-keeled perianth with a broad, entire mouth and a thin or fleshy base (when fleshy = perigynium). Foot of the sporophyte not penetrating into the stem. Setae short and thick. Capsules cylindric, walls 2–3-layered. Elaters free, not attached to the capsule valves. Spores unicellular or multicellular, germination exosporous or endosporous. Vegetative reproduction by multicellular gemmae or caducous leaf lobes. Protonema sometimes large and thalloid in epiphyllous species. About 200 species worldwide, 33 in Colombia and Ecuador. Radula is a very distinct genus, recognized by (1) Radula-type branches; (2) large, brown oil bodies, only 1–2 per cell and filling the cell lumen; (3) lack of underleaves; (4) rhizoids in tufts from leaf lobules; and (5) perianth narrowly elongate, flattened above, with a broad, entire mouth. The neotropical species have been studied by Castle (1968) and Yamada (1980, 1981, 1982, 1987, 1988, 1991, 1993a, 1997, 2000, 2003), and these data have been instrumental for the present treatment. A molecular analysis, based on a broad sampling across the entire genus, has shown the existence of seven robust clades in Radula, which were described as subgenera (Devos et al. 2011). At

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least four subgenera are represented among the species of Radula in Colombia and Ecuador, as follows: 1. Radula subg. Cladoradula Spruce (R. gottscheana), characterized by stems with a thick-walled, brown epidermis and subepidermis and by gynoecia without innovations. 2. Radula subg. Metaradula R.M.Schust., characterized by thin-walled stem cells, gynoecia with innovations, and, especially, the presence of a stem perigynium (gynoecial base fleshy, perianth and calyptra multistratose near the base). All neotropical species of this subgenus belong in the section Epiphyllae Grolle (Pócs 2017), distinguished by epiphyllous growth, a very long and narrow trumpet-shaped perianth, and frequent presence of gemmae on leaf margins (R. aguirrei, R. flaccida, R. mammosa, R. stenocalyx, R. yanoella). Two species of this section, R. aguirrei and R. yanoella, have large, persistent thalloid protonemata and show neotenic evolution (Schuster 1991). 3. Radula subg. Radula characterized by stems with a thin-walled epidermis (no subepidermis) and a ± thick-walled medulla, gynoecia with innovations, and the presence or absence of gemmae (R. fendleri, R. javanica, R. quadrata, R. santacruziana, R. tectiloba). 4. Radula subg. Volutoradula Devos et  al. characterized by stems with a thickwalled, brownish epidermis (no subepidermis) and a thin-walled medulla, gynoecia with innovations, and absence of gemmae. A stem perigynium occurs in R. pallens (Schuster 1980a; see also under the latter species). Most neotropical Radula species belong in this section. The subgenera and sections are characterized by gametophytic traits and DNA; synapomorphies are largely lacking. Future taxonomic work should include study of the sporophyte generation, which has been neglected and might yield new characters to support the infrageneric classification (Renner and Braggins 2005). Literature: Yamada (1988, 2003), Schuster (1991), Reiner-Drehwald (1994b), Devos et al. (2011), Patiño et al. (2017), Pócs (2017) Note: lobule characters must be studied on leaves of the main stem, not on branch leaves. 1. Plants growing on living leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Plants not growing on living leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 2. Plants consisting of a small, branched thallus and short leafy shoots arising from thallus margins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Plants without a thallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Thallus irregularly rounded, the lobes radially arranged and dichotomously branched. Keel of leaf lobes straight or slightly concave. Plants of northern South America and Costa Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. yanoella 3. Thallus narrowly elongate, pinnately branched. Keel of leaf lobes strongly convex. Plants of Gorgona Island (Colombia) . . . . . . . . . . . . . . . . . . . . . R. aguirrei 4. Discoid gemmae produced on dorsal leaf margins . . . . . . . . . . . . . . . . . . . . . 5 4. Gemmae lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

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5. Gemmae very large, to 0.7  mm in diameter, base deeply cordate-auriculate. Lobule apex somewhat elongate, obtuse . . . . . . . . . . . . . . . . . . . . . R. flaccida 5. Gemmae much smaller, to 0.25  mm in diameter, base of gemmae almost straight. Lobule apex not elongate, rounded . . . . . . . . . . . . . . . . . R. stenocalyx 6. Keel deeply concave. Lobule strongly inflated along the keel, inflated portion of lobule curved, banana-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. tenera 6. Keel straight or convex. Lobule not strongly inflated along the keel . . . . . . . . 7 7. Rhizoidal area on lobules strongly convex, prominently projecting outward. Lobule base not covering the stem. Leaf lobes not caducous. . . . . R. mammosa 7. Rhizoidal area on lobules not prominently projecting outward. Lobule base covering the stem (1/5–4/5 of stem width). Leaf lobes frequently caducous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. javanica 8. Branching purely dichotomous 8. Branching irregularly or regularly pinnate, sometimes becoming dichotomous in upper stem portions [when in doubt try both leads] . . . . . . . . . . . . . . . . . 10 9. Leaves longer than wide, usually falcate. Lobule base extending almost fully across the stem. Plants very rare (Ecuador). . . . . . . . . . . . . . . . . . R. sinskeana 9. Leaves (sub)orbicular, not falcate. Lobule base only partially extending across the stem. Plants common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. episcia 10. Discoid gemmae produced on leaf margins. . . . . . . . . . . . . . . . . . . . . . . . . . 11 10. Gemmae lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 11. Lobule base not covering the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11. Lobule base covering the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 12. Gemmae very large, to 0.7  mm in diameter, base deeply cordate-auriculate. Lobule apex somewhat elongate, obtuse . . . . . . . . . . . . . . . . . . . . . R. flaccida 12. Gemmae much smaller, to 0.25 mm in diameter, base almost straight. Lobule apex not elongate, rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Plants 1–1.5 mm wide. Keel convex. Rhizoidal area on lobules usually strongly convex, prominently projecting outward. . . . . . . . . . . . . . . . . . . R. stenocalyx 13. Plants larger, ca. 2.5 mm wide. Keel straight. Rhizoidal area not prominently projecting outward. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. santacruziana 14. Lobules densely imbricate, base extending fully across the stem and beyond . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. quadrata 14. Lobules distant to subimbricate, base of most lobules not extending fully across the stem (to be studied on mature stem leaves). . . . . . . . . . . . . . . R. tectiloba 15. Lobules (sub)imbricate. Lobule base of stem leaves auriculate, the auricle coiled (twisted), crossing the stem or not. Leaf cells without trigones . . . . . 16 15. Lobules not imbricate. Lobule base of stem leaves without coiled auricle. Leaf cells with or without trigones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 16. Auricle of lobules longly extending downward, far beyond the keel. Keel very short, less than 1/5 of leaf length. Gynoecia on very short branches without innovations. Stemswith a thick-walled, brown subepidermis. Leaves not caduceus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. gottscheana 16. Auricle not extending downward beyond the keel. Keel longer, 1/4–1/2 of leaf length. Gynoecia terminal on main stems or elongate branches with innovations. Stems without brown subepidermis. Leaves sometimes caducous . . . 17

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17. Coiled auricle large, extending downward and usually reaching the keel, covering almost the whole ventral stem surface. Plants robust, 2.5–4 mm wide, very common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. voluta 17. Coiled auricle small, not extending downward, covering only part of the ventral stem surface. Plants smaller, 2–2.5 mm wide, rare . . . . . . . . . . . . . . R. sinuata 18. Plants with caducous leaf lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18. Plants without caducous leaf lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 19. Caducous leaf lobes produced on small flagelliform shoots at the stem apex, the shoots with very small and densely imbricate lobules. . . . . . . . . . . . R. tenera 19. Caducous leaf lobes not produced on small flagelliform shoots at the stem apex with densely imbricate lobules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Lobules 2.5–4× as long as wide, parallel to the stem, tubularly inflated. Shoots very fragile, fragmenting. Plants of the Sierra Nevada del Cocuy, 4000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. bischlerae 20. Lobules 1–2× as long as wide, obliquely to widely spreading, not tubularly inflated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21. Trigones large, swollen. Cuticle finely papillose. Leaf lobes strongly convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. fendleri 21. Trigones absent or small, not swollen. Cuticle smooth. Leaf lobes convex or plane. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Keel convex at least in part of the leaves. Apicular margin of lobule ± sinuate in the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22. Keel ± straight. Apicular margin of lobule not sinuate . . . . . . . . . . . . . . . . . 26 23. Lobule base covering 1/3–4/5 of stem width . . . . . . . . . . . . . . . . . . . . . . . . . 24 23. Lobule base covering less than 1/3 of stem width (or not covering stem) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24. Plants 2–3 mm wide. Lobule base covering 1/3–1/2 of stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. nudicaulis 24. Plants smaller, less than 2 mm wide. Lobule base covering more than 1/2 of stem width. plants of the Galápagos Islands . . . . . . . . . . . . . . . R. galapagona 25. Leaf cells distinctly mammillose on the dorsal leaf surface and with a thickened wall. Leaf margins ± crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . R. sonsonensis 25. Leaf cells not mammillose, walls thin. Leaf margins not crenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. schaefer-verwimpii 26. Lobule apex narrowly elongate, (sub)acute. Lobule base covering the whole width of the stem. Plants rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. cubensis 26. Lobule apex not or shortly elongate, obtuse. Lobule base not covering the whole width of the stem. Plants very common . . . . . . . . . . . . . . . . . . . . . . R. javanica 27. Lobule apex conspicuously elongate (to be observed on flattened lobules). Plants monoicous or dioicous, rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 27. Lobule apex not or slightly elongate [when in doubt try both leads]. Plants dioicous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 28. Leaf cells with distinct trigones. Lobule base (on main stem) fully extending across the stem. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. angulata 28. Leaf cells ± without trigones. Lobule base (on main stem) not fully extending across the stem, covering only 1/3–1/2 of stem width. Plants monoicous, usually fertile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. mexicana

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2 9. Leaf lobes strongly convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 29. Leaf lobes flat or slightly convex. (Apex of leaf lobes sometimes recurved) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 30. Lobules inflated over the whole surface. . . . . . . . . . . . . . . . . . . . . . . R. decora 30. Lobules inflated along the keel, flattened above . . . . . . . . . . . . . . . . . . . . . . 31 31. Keel deeply concave, inflated carinal portion of lobule curved, ± bananashaped. Lobule base not or scarcely covering the stem. Plants of upper montane forest and páramo, 1900–3500 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. tenera 31. Keel straight or convex, inflated portion not banana-shaped. Lobule base covering 1/4–2/3 of stem width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 32. Plants regularly and densely pinnate. Base of lobule of stem leaves covering 1/2–2/3 of stem width. Trigones large, bulging. . . . . . . . . . . . . . . . . R. eggersii 32. Plants loosely and irregularly pinnate. Base of lobule of stem leaves usually covering less than 1/2 of stem width. Trigones small or large . . . . . . . . . . . . 33 33. Leaf cells with large, bulging trigones. Cuticle finely papillose. Lobules longer than wide, subrectangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. fendleri 33. Leaf cells without or with very small trigones. Cuticle smooth. Lobules about as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. saccatiloba 34. Leaf cells with large, bulging trigones. Lobules strongly inflated along the keel. Keel concave, decurrent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. involvens 34. Leaf cells without or with small trigones. Lobules inflated or not. Keel concave, straight or convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 35. Lobule base in many leaves covering more than 1/2 of stem width. Plants densely 1–2-pinnate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. xalapensis 35. Lobule base mostly covering 1/5–1/2 the width of the main stem or not covering the stem. Plants not densely 1–2-pinnate . . . . . . . . . . . . . . . . . . . . . . . . . 36 36. Keel convex or straight, not decurrent. Lobules conspicuously inflated along the keel (except in R. pusilla) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 36. Keel concave or straight, ± decurrent. Lobules not conspicuously inflated along the keel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 37. Free margin of lobule sinuate-plicate. Lobule base covering 1/4–1/2 of stem width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. nudicaulis 37. Free margin of lobule ± plane. Lobule base not or scarcely covering the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38. Leaf cells strongly mammillose. Keel of stem leaves strongly convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. sonsonensis 38. Leaf cells not or only slightly mammillose. Keel of stem leaves straight or convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39. Lobules strongly inflated along the keel . . . . . . . . . . . . . . . . . . . . R. subinflata 39. Lobules not strongly inflated along the keel . . . . . . . . . . . . . . . . . . . R. pusilla 40. Median leaf cells 20–30 μm, with small trigones. Leaf lobes falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. jamesonii 40. Median leaf cells smaller, 15–20 μm, ± without trigones. Leaf lobes falcate or not falcate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

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41. Leaves longer than wide, falcate. Lobule base extending almost fully across the stem. Plants very rare (only known from the type) . . . . . . . . . . . . R. sinskeana 41. Leaves (sub)orbicular, not falcate. Lobule base extending only partially across the stem. Plants common . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pallens 1. Radula aguirrei R.M.Schust. On the fronds of a small palm in the understory of dense, wet, undisturbed coastal rainforest, ca. 250  m. Colombia: Cau (Isla Gorgona). General distribution: only known from the island of Gorgona, Colombia. Radula aguirrei (sect. Epiphyllae) is a rare Chocó endemic that is close to R. yanoella and may be a synonym of the latter; differences are shown in the key. The record from “Chocó” in the catalogue of Colombia (Gradstein and Uribe 2016a) refers to the Chocó biogeographic region, not the department. 2. Radula angulata Steph. Fig. 66A Radula bogotensis Steph., syn. nov. Type: Colombia, Bogotá, Azerradero, 2500 m, 1860, Lindig 1738 (lectotype, designated here, G-00043906!; isolectotypes G-00265022!, G-00265923!) Radula korthalsii Steph. On bark, 300–2500  m. Colombia: Cun, Mag, Put. General distribution: West Indies, northern Andes, southeastern Brazil. Radula angulata (subg. Volutoradula) is a little-known neotropical species that is recognized by (1) leaf lobes ovate, slightly convex, not falcate, somewhat fragile but not fully caducous; (2) leaf cells with conspicuous trigones; and (3) lobules distant to (sub)imbricate, variable in size, ca. 1/2 of lobe length, subquadrate, apex distinctly elongate, narrowly rounded to subacute, lobule base in well-developed lobules fully extending across the stem (Yamada 1991). Radula angulata resembles R. mexicana, but the latter species is monoicous and has thin-walled cells without distinct trigones and lobule bases covering only 1/3–1/2 of stem width. Radula angulata may also be confused with R. javanica, but in the latter the leaf lobes are frequently falcate and caducous, tearing off at the base (sometimes broken irregularly but not caducous in R. angulata). Moreover, the lobule apex in R. javanica is less elongate, and the lobule base does not extend fully across the stem. 3. Radula bischlerae Gradst. & Ilk.-Borg., sp. nov.Fig. 64 Type: Colombia, Boyacá, Sierra Nevado del Cocuy, Guicán, Corralitos, 3900 m, on soil in páramo, 10 Jun 1959, H. Bischler 2894 (holotype COL; isotypes MG, PC-0774218) Plants brown, medium-sized, leafy stems ca. 2  mm, all stems and branches highly fragile, fragmenting and mostly denuded by caducous leaf lobes. Leaf lobes highly caducous, suborbicular to ovate, falcate, flat, not extending beyond the stem, insertion line short; lobe apex rounded to bluntly apiculate, margins whitish, irregularly crenulate to subentire, sometimes with a bluntish, lobe-like tooth near the base. Lobe cells small, isodiametrical to slightly elongate, 20–25  ×  15–20  μm, lumen

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Fig. 64  Radula bischlerae. (A) Habit. (B) Lobule. (C) Portion of stem with two lobules, ventral view. (D) Leaf margin, showing initiation of regeneration. (E) Leaf margin with regenerant. (F) Median cells of leaf lobe. (G) Caducous leaf. (All from the type). Scale bars: A, G = 250 μm, B, D = 50 μm; C, E = 100 μm; F = 25 μm

rounded, trigones present, small, triangular, cuticle smooth. Leaf lobules distant to contiguous, suberect, ± tubularly inflated for most of its length, narrowly obtriangularly elongate, 100–125 μm long, 25–50(−70) μm at apex, (2–)2.5–4× longer than wide, narrowest on branches, oriented parallel to the shoots and attached for most of their length, on branches covering much of the branch surface both dorsally and ventrally, lobule base cuneate, free apical part flattened, free margin angular in outline, apex obtuse, free base slightly expanded, covering to 1/3–1/2 of stem width; keel slightly convex, standing at an angle of ca. 25–35° with the stem, smooth. Gametoecia not seen. Vegetative reproduction by caducous leaf lobes, the caducous leaf lobes often with regenerants on leaf margins (Fig. 64D, E). Radula bischlerae (sect. Volutoradula?) is only known from the type from Colombia. It stands out by the very fragile, highly denuded shoots with strongly caducous leaf lobes and by the narrowly elongate, tubularly inflated lobules (ca. 2.5–4× longer than wide), which are positioned subparallel to the stem, with an obtuse apex and the free base covering to 1/3–1/2 of stem width. The new species may be compared with R. javanica and R. nudicaulis with which it shares caducous

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Fig. 65 (A–E) Lejeunea ryszardii. (A) Habit. (B) Leaf. (C) Lobule. (D) Caducous leaf lobe with rhizoids and plantlet. (E) Habit. (F–K) Rectolejeunea halinae. (F) Lobule. (G) Habit. (H) Caducous leaf. (I) Leaf tips. (J) Leaf. (K) Underleaf. (All from the type). Scale bars: A, E, G = 250 μm; B, D, H, J, K = 100 μm; C, F, I = 50 μm

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Keys and Descriptions

Fig. 66 (A) Radula angulata, habit. (B) Radula cubensis, habit. (C) Radula decora, habit. (D) Radula episcia, habit. (E–G) Radula fendleri. (E) Habit. (F) Leaf. (G) Cells. (H and I) Radula flaccida. (H) Habit showing two large gemmae. (I) Gemma. (J) Radula galapagona, habit. (K) Radula gottscheana, habit. (L and M) Radula involvens. (L) Habit. (M) Marginal cells. (N) Radula jamesonii, habit. (O) Radula javanica, habit. (C redrawn from Yamada 1987; (D) redrawn from ReinerDrehwald 1994b; E–G, L, M redrawn from Yamada 1980; J redrawn from Yamada 1991; N redrawn from Yamada 1982; rest from Gradstein and Costa 2003). Scale bars: A, B, D, K, L = 1000 μm; C, J = 650 μm; E = 600 μm; F = 300 μm; G, M = 20 μm; H, O = 500 μm; N = 1500 μm

leaf lobes, an obtuse lobule apex, and the lobule base covering part of the stem. It is immediately separated from the latter two, however, by the narrowly elongate, suberect lobules (Fig.  64A–C). In R. javanica and R. nudicaulis, the lobules are subquadrate.

Radulaceae

4. Radula cubensis K.Yamada

627

Fig. 66B

On bark in montane rainforest, 1800  m. Colombia: Cho (Vasco et  al. 2002). Ecuador: ZC (Parolly et al. 2004). General distribution: Cuba, Ecuador, southeastern Brazil. Radula cubensis (subg. Volutoradula) resembles R. javanica by the falcate, caducous leaf lobes but differs from the latter in the narrowly elongate, subacute lobule apex and the broader lobule base, which extends fully across the stem and sometimes even beyond. By the elongate lobule apex and broad lobule base, R. cubensis is similar to R. angulata, but in the latter species, the leaf lobes are not falcate and not caducous (the leaf lobes in R. angulata may be irregularly broken, but do not drop off entirely). The records from Colombia and Ecuador need verification. 5. Radula decora Steph.  Radula inflata Steph.

Fig. 66C

On bark and rock at low elevation. Ecuador: Es, Su (Arnell 1962). General distribution: Venezuela (type), Ecuador, Chile. Radula decora (subg. Odontoradula?) is distinguished by (1) plants small, 1.2–1.5 mm wide); (2) leaves obliquely spreading, suborbicular, strongly convex; (3) leaf cells very thin-walled; (4) lobules rather large, ca. 1/2× lobe length, subquadrate, fully inflated, apex broadly rounded, margins ± straight and plane, base extending 1/4–1/3 across the stem; and (5) keel slightly convex, not decurrent (Yamada 1987). Radula decora resembles R. fendleri, R. subinflata, and R. tenera but differs in having wholly inflated lobules. The distribution of R. decora includes one locality in Venezuela (type), two in Ecuador (the Galápagos record is erroneous; see Gradstein 2009), and numerous localities in southern temperate regions of Chile where the species was described as R. dusenii Steph. (Castle 1963). Since the Venezuelan and Ecuadorian records are all from low, warm-tropical elevation, the occurrence of the species in temperate Chile is unusual and needs verification. The correct author citation for R. decora is “Gottsche ex Steph.” or, in shortened version, “Steph.” (not “Gottsche”; Renner 2016). 6. Radula eggersii K.Yamada On remnant trees in pasture, 2800–3050 m. Ecuador: Lo, ZC (Yamada 1997; Lüth and Schäfer-Verwimp 2004). General distribution: only known from southern Ecuador. Radula eggersii (subg. Volutoradula) approaches R. fendleri and R. subinflata but differs in the densely and regularly pinnate branching (laxly and irregularly pinnate in R. fendleri and R. subinflata) and the broader lobule base, covering 1/2–2/3 of stem width (maximally 1/3 in R. fendleri and R. subinflata). 7. Radula episcia Spruce Fig. 66D Radula antilleana Castle, syn. nov. Type: Puerto Rico, Mt. Montosa, N. L. Britton & J. F. Cowell 4166 (NY n.v.) Radula cornucopiae Spruce

628

 

Keys and Descriptions

Radula episcia var. opaciuscula Spruce, syn. nov. (Radula opaciuscula [Spruce] Castle). Type: Peru, Mt. Campana, Spruce s.n. (MANCH n.v.), fide Yamada (2000) Loosely creeping or pendent on vertical surfaces of shaded, wet rock faces and on bark in montane rainforests, occasionally on moist soil or growing submerged, 450–3400 m. Colombia: Ant, Boy, Cas, Ces, Cho, Cun, Hui, Mag, Met, Ris, San. Ecuador: Bo, Ch, Co, Ga, Pa, Pi, Tu, ZC. General distribution: widespread in tropical America. Radula episcia (subg. Volutoradula) is a common neotropical-montane species that is readily recognized by (1) plants glossy olive green, 2.5–4 mm wide, purely dichotomously branched; (2) leaf lobes (sub)orbicular, wide-spreading, not falcate; (3) leaf cells without or with small trigones; (4) lobules oblong-rectangular parallel to the stem (being wider than long) or subquadrate, distant, base of lobule variable in width, covering the stem for 1/5–2/3 of stem width; and (5) keel short, 1/4–1/3× lobe length, straight or concave, decurrent. Radula episcia is readily recognized by the purely dichotomous branching, orbicular leaf lobes, and short, wider than long lobules with the base partially extending across the stem. Purely dichotomous branching is also seen in R. sinskeana, but in the latter species, the leaf lobes are clearly longer than wide and usually falcate, and the lobules are subquadrate, with the lobule base fully extending across the stem. Castle (1959) placed R. episcia in sect. Dichotomae Castle and described numerous new species in this section, mostly from the West Indies. Many of them are of doubtful status and probably synonyms of R. episcia. Based on the description and illustration in Castle (1959), R. antilleana, recorded from Ecuador and numerous other neotropical countries, is a synonym of R. episcia. 8. Radula fendleri Steph.

Fig. 66E–G

On bark in lower montane rainforest, to 2000 m. Colombia: Ces (Aguirre and Ruiz 2001), Cho (Winkler 1970). Ecuador: ZC (Nöske et al. 2003). General distribution: West Indies, tropical South America. Radula fendleri (subg. Radula) is an uncommon neotropical species that is distinguished by (1) plants small, 1–1.3 mm wide, irregularly pinnate; (2) leaf lobes strongly convex, obliquely spreading, ovate-orbicular, caducous; (3) leaf cells small, with large bulging trigones and a finely papillose cuticle; (4) lobules distant, longer than wide, subrectangular, ca. 1/2× lobe length, conspicuously inflated along the keel and flattened above, apex obtuse, margins ± straight, base covering maximally 1/2 of stem width; and (5) keel convex, not decurrent (Yamada 1980). Radula fendleri is a misunderstood species and several records are erroneous; the record from the Galápagos Islands is R. subinflata, and that from Magdalena is R. javanica. The correct author citation for R. fendleri is “Gottsche ex Steph.” or, in shortened version, “Steph.” (not “Gottsche”; Renner 2016). 9. Radula flaccida Lindenb. & Gottsche

Fig. 66H, I

On living leaves and bark in lowland rainforest, 30–500 m. Colombia: Cau, Val. Ecuador: Es, Na, Or, Pa. General distribution: tropical America, tropical Africa.

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Radula flaccida (sect. Epiphyllae) is a common Afro-American epiphyllous species that is probably more widely distributed in Colombia and Ecuador than is currently known. The species is characterized by (1) plants small, prostrate, 1–1.5 mm wide, glistening green, irregularly pinnate, forming small, rounded, flat mats on the surface of living leaves or, occasionally, on bark; (2) dorsal leaf margin with a few (1–3) very large discoid gemmae, which are to 0.7 mm in diameter and possess a deeply cordate-auriculate base; the gemmae are quite flat and extend horizontally, in the plane of the leaf; (3) lobules small, distant, subquadrate, with a pronounced protruding rhizoidal area, apex somewhat elongate toward an obtuse tip, base not or very slightly covering the stem; and (4) keel straight or convex, ca. 1/3 length of lobe, not decurrent. Fertile plants are readily distinguished by the narrow-cylindrical, trumpet-shaped perianths, 3–5× longer than wide, arising perpendicularly from the substrate. The unique character of R. flaccida is the huge gemmae with a deeply cordateauriculate base. The strongly bulging rhizoidal area on the lobule surface is shared with R. mammosa and R. stenocalyx and seems to be a characteristic feature of the sect. Epiphyllae. 10. Radula galapagona Steph.

Fig. 66J

Common on rotten bark in moist evergreen Scalesia and Zanthoxylum woodlands and Miconia scrub on the Galápagos Islands, 450–1200  m. Ecuador: Ga (Isabela, Pinta, San Cristóbal, Santa Cruz, Santiago; Yamada 1991; Yamada and Gradstein 1991). General distribution: only known from the Galápagos Islands where it is widespread in the humid belt. Radula galapagona (subg. Volutoradula) is very similar to R. schaefer-verwimpii, and the two might be subspecies of a single species. They differ only in the more strongly expanded lobule base of R. galapagona, covering at least 1/2 of stem width (1/5 or less of stem width in R. schaefer-verwimpii). Radula galapagona also resembles R. javanica in the caducous leaf lobes, but the latter species has a straight keel (strongly convex in R. galapagona). Radula javanica and R. schaefer-verwimpii are not known from the Galápagos Islands. 11. Radula gottscheana Taylor

Fig. 66K

On bark and shaded rock in submontane and lower montane rainforests and in scrub, to 2100 m. Colombia: Boy, Cun, Mag. Ecuador: Ch, Ga, Lo, MS, Pa, Tu, ZC. General distribution: tropical America. Radula gottscheana (subg. Cladoradula) is a characteristic neotropical species that is readily recognized by (1) plants large, irregularly pinnate; (2) stems very rigid, with a thick-walled, brownish cortex of several cell layers; (3) lobules large, widely crossing the stem and with a large auricule that longly extends downward, beyond the base of the very short keel; and (4) gynoecia on very short branches without innovations. Radula gottscheana is the only neotropical member of the subgenus Cladoradula and is closely related to R. boryana (F.Weber) Nees from Africa (Devos et  al. 2011). By its large, auriculate lobule, R. gottscheana resembles

630

 

Keys and Descriptions

R. voluta, but in the latter species, the lobule auricle does not extend downward beyond the base of the keel. 12. Radula involvens Spruce

Fig. 66L, M

On bark in montane rainforest and shrubby páramo, 1000–3000 m. Colombia: Cau, Hui, Ris. Ecuador: Tu, ZC (Reserva Biologica San Francisco, Burghardt 7165, QCA). General distribution: northern Andes, Guyana. Radula involvens (subg. Volutoradula) is an uncommon northern Andean species that is distinguished by (1) reddish-brown plant color; (2) slightly convex, falcate leaves with recurved apex; (3) small lobules that are narrowly inflated along the decurrent keel; and (4) leaf cells with huge trigones. Caducous leaf lobes and gemmae are not produced. By its combination of characters, R. involvens cannot be confused with any other Radula species in the northern Andes. 13. Radula jamesonii Taylor Radula sprucei Steph.

Fig. 66N

On trees, rock, and humic soil in upper montane forest and shrubby páramo, 2200–3900  m. Colombia: Cal, Cun, Tol. Ecuador: Ch, Na, Pi, ZC (páramo of Saraguro and Yacuambi, mossy elfin forest, Benitez et al. 1311, HUTPL). General distribution: tropical Andes. Radula jamesonii (subg. Volutoradula) is an uncommon Andean species that resembles R. pallens but differs from the latter in the falcate leaf lobes and larger leaf cells (20–30 μm in diameter) with trigones. Radula jamesonii also resembles R. xalapensis and R. nudicaulis, but R. xalapensis is more densely bipinnately branched and the lobule base in the latter species extends further across the stem (more than half the stem width) and is sometimes slightly coiled. Radula nudicaulis differs from R. jamesonii in the mostly convex and non-decurrent keel (mostly concave and decurrent in R. jamesonii) and the lobules being distinctly inflated along the keel (not so in R. jamesonii). 14. Radula javanica Gottsche Fig. 66O Radula amazonica Spruce Radula caldana Ångstr. Radula eggersiana Steph. Radula falcifolia Steph. Radula floridana Castle Radula gracilis Mitt. ex Steph., syn. nov. Type: Colombia, “Andes bogotensis,” Weir s.n. (“Weverley”) (G-00042688!) Radula longifolia Steph. Radula macrostachya Lindenb. & Gottsche Radula surinamensis Steph. On bark, dead wood, rock, and living leaves in lowland and montane rainforests and in plantations, in exposed or shaded habitats, 150–2800  m. Colombia: Boy, Cho, Cun, Hui, Mag, Met, Put, Ris, San Vau. Ecuador: Az, EO, Lo, MS, Na, Or, ZC. General distribution: pantropical. Radula javanica (subg. Volutoradula) is a widespread and common pantropical species that has been described under many different names (see Yamada 1979,

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631

2000). The main characters are (1) plants rather small, 1.2–2.5 mm wide, irregularly pinnate; (2) leaf lobes subimbricate, ovate-oblong and ± falcate, frequently caducous; (3) median leaf cells 16–25 μm, trigones minute to medium-sized, never swollen, cuticle smooth; (4) lobules distant to contiguous, subquadrate, inflated along the keel and flattened above with plane margin, apex rounded or obtuse, often slightly elongate, base rounded, variable in width, covering 1/5–4/5 of stem width; and (5) keel straight, 1/3–2/5× lobe length, not decurrent. Radula javanica is the most common neotropical Radula species with caducous leaf lobes. Caducous leaf lobes are also seen in R. cubensis, R. fendleri, R. galapagona, R. schaefer-verwimpii, and R. sonsonensis, but R. cubensis differs in the more narrowly elongate, subacute lobule apex; R. fendleri in the strongly convex leaf lobes, large bulging trigones, and verruculose cuticle; R. galapagona and R. schaefer-verwimpii in the strongly convex keel; and R. sonsonensis in its mammillose leaf cells. Radula lewisii K.Yamada, described from Bolivia and recorded once from Colombia (Risaralda, Mistrato, Gradstein 8544, det. K. Yamada, GOET), is closely similar to R. javanica but lacks caducous leaf lobes (Yamada 1993b, 2000). 15. Radula mammosa Spruce Radula lindigii Castle On living leaves in lowland and montane rainforests, occasionally on bark, from sea level to 2900  m. Colombia: Ama, Ces, Cho, Cun, Mag. Ecuador: Lo, Or, ZC. General distribution: tropical America. Radula mammosa (sect. Epiphyllae) is usually epiphyllous and resembles R. flaccida and R. stenocalyx in the distant lobules with a strongly protruding rhizoidal area, the lobule base not covering the stem, and the very thin-walled leaf cells. It differs from R. flaccida and R. stenocalyx, however, in the absence of gemmae. 16. Radula mexicana Lindenb. & Gottsche On bark in montane rainforest, 1800–2150 m. Colombia: Boy, Cho. Ecuador: ZC (Nöske et al. 2003). General distribution: tropical and subtropical America. Radula mexicana (subg. Volutoradula) is the only monoicous Radula in Colombia and Ecuador and very rare; the species is usually fertile. Radula mexicana resembles R. angulata in the conspicuously elongate lobule apex but differs from the latter in being monoicous and in having thin-walled leaf cells without trigones (Yamada 1993a). Radula mexicana may also be confused with R. pallens, but the lobule apices in the latter species are not elongate and branching is often dichotomous in upper stem portions. Moreover, R. pallens is dioicous. 17. Radula nudicaulis Steph. Radula goebelii Steph.

Fig. 67A

On bark, shaded soil, and rock in montane forests and páramo, ca. 1000–4000 m. Colombia: Ara, Boy, Cal, Cas, Cau, Cho, Cun, Hui, Mag, Qui, Ris, San. Ecuador: Az (Gradstein 12660, QCA), Lo, ZC.  General distribution: tropical South America, Macaronesia (Madeira). Radula nudicaulis (subg. Volutoradula) is a widespread neotropical-Macaronesian species that is recognized by (1) plants yellowish-brown to reddish-brown when dry, 2–3 mm wide, regularly pinnate; (2) leaf lobes slightly convex, widely

Fig. 67 (A) Radula nudicaulis, habit. (B–D) Radula pallens. (B) Habit. (C) Leaf. (D) Leaf cells. (E) Radula quadrata, habit showing gemmae on leaf margins. (F and G) Radula saccatiloba. (F) Habit. (G) Leaf cells. (H) Radula schaefer-verwimpii, habit. (I) Radula sinuata, habit. (J) Radula stenocalyx, habit showing gemmae on leaf margins. (K) Radula tectiloba, habit showing gemmae on leaf margins. (L and M) Radula tenera. (L) Habit. (M) Lobule. (N) Radula voluta, habit. (O) Radula xalapensis, habit. (P) Radula yanoella, habit. (F, G redrawn from Yamada 1987; L, M redrawn from Yamada 1980; O redrawn from Yamada 1982; rest from Gradstein and Costa 2003; Gradstein and Ilkiu-Borges 2009). Scale bars: A, E, I, J, K, N  =  1000  μm; B, H, P  =  500  μm; C = 250 μm; D = 50 μm; F, L = 600 μm; G = 20 μm; M = 200 μm; O = 2000 μm

Radulaceae

633

spreading, ovate, slightly falcate, apex rounded, plane or recurved; (3) median leaf cells 15–25  μm, with minute trigones, cuticle smooth; (4) lobules distant, rather small, subquadrate, narrowly inflated along the keel, apex rounded to obtuse, free margin conspicuously sinuate-plicate (not plane), base covering 1/4–1/2 of stem width; (5) keel obliquely spreading, 1/3–2/5× lobe length, convex or straight, not decurrent; and (6) asexual reproduction by caducous leaf lobes. Radula nudicaulis is only known from sterile material, and reproduction is by caducous leaf lobes. Jans (1979) reported the occurrence of discoid gemmae in the species, but this has not been confirmed; possibly these were regenerants, which develop at the margins of caducous leaf lobes and resemble gemmae when juvenile. In the distant lobules with a sinuate-plicate margin, R. nudicaulis resembles R. jamesonii; for differences see under the latter species. 18. Radula pallens (Sw.) Mont.  Fig. 67B–D Type: Jamaica, Swartz s.n. (S-B28464!) Radula kegelii Steph., syn. nov. Type: Suriname, near Mariepaston, 1846, Kegel 1412 (lectotype, designated here G-00121979!; isolectotypes G-00264270!, G-0026471!) Syntype: Brazil, Santa Catarina, 1847, Pabst 886 (G-00264272!) On bark, rock, and humic soil in lowland and montane rainforests, 200–2000 m. Colombia: Boy, Ces, Mag, San (Gil 2917, 2229, UPTC). Ecuador: EO, Gu, LR, MS, Or, Pa, Su, Tu, ZC. General distribution: tropical America. Radula pallens (subg. Volutoradula) is a widespread neotropical species that is characterized by (1) plants small to large, (1.5–)2–3 mm wide, olive green, irregularly pinnate, often dichotomous in the upper stem portions; (2) leaves widely spreading, (sub)orbicular, not falcate, apex broadly rounded; (3) median leaf cells small, 15–20 μm, without or with minute trigones; (4) lobules small, distant, quadrate to elongate-subrectangular parallel to stem, slightly inflated along keel, apex obtuse, not elongate, free margin often sinuate and incurved, base of lobule covering 1/4–1/2 of the width of the main stem; (5) keel straight or concave, 1/4× lobe length, slightly decurrent, sinus between keel and ventral lobe margin sharp; and (6) gynoecia on long shoots with 1–2 innovations, gynoecial base elaborated into a fleshy stem perigynium, perianth narrowly obtriangular, with a wide mouth, perianth base up to 6 cells thick. Radula pallens has long been considered endemic to Jamaica (Castle 1960), but examination of the types in the Swartz herbarium (S) and in Geneva shows that the species is conspecific with the widespread R. kegelii. The latter species has been described and illustrated by Reiner-Drehwald (1994b), Yamada (2000), and Gradstein and Ilkiu-Borges (2009). Castle (1960) considered R. pallens to be more robust than R. kegelii, but except for shoot length, the types of the two species are fully identical. Radula pallens resembles R. episcia in the shape of lobes and lobules and the small, thin-walled cells but differs from the latter species in the irregularly pinnate to dichotomous branching (purely dichotomous in R. episcia). Dichotomous branches occur in the upper stem portions and are often gynoecial innovations

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Keys and Descriptions

(Reiner-Drehwald 1994b). Radula pallens also approaches R. jamesonii, but the latter species has falcate leaf lobes (not falcate in R. pallens) and larger leaf cells with small trigones. The base of the gynoecium in R. pallens may be elaborated into a short, fleshy stem perigynium (seen in material from Ecuador, Yasuni National Park, abundant on tree trunks in varzea forest near Tiputini Biodiversity Station, Gradstein 12814, MG, QCA). The presence of a stem perigynium in R. pallens was already noted by Schuster (1980a), who placed the species in subg. Metaradula (sect. Pallescentes R.M.Schust.) based on this character. A molecular-phylogenetic study of Radula, however, resolved R. pallens (= R. kegelii) in subg. Volutoradula (Devos et al. 2011). The occurrence of a stem perigynium in subg. Volutoradula has not yet been reported and deserves further study. 19. Radula pusilla Spruce Radula mazarunensis K.Yamada, syn. nov. Type: Guyana, upper Mazaruni, Kamarang, Mt. Latipu, 500 m, 24 Feb 1985, Gradstein 5548 (holotype U!) On bark in submontane and lower montane forest areas, 500–1700 m. Colombia: Qui, Ris. Ecuador: Pa (type). General distribution: tropical Andes, Costa Rica, Guyana, Dominica. Radula pusilla (subg. Volutoradula) is a little-known neotropical species that is distinguished by (1) plants small, ca. 1.5 mm wide, sparsely branched; (2) leaves wide-spreading and sometimes falcate, subimbricate, slightly convex, ovate, apex broadly rounded; (3) median leaf cells small, ca. 20 μm long, without trigones; (4) lobules small, distant, subquadrate-rhombic, little inflated, obliquely spreading, apex rounded to obtuse, free margin plane, lobule base not or little covering the stem (to 1/4 of stem width); and (5) keel convex, 1/3–2/5× lobe length, not decurrent, sinus between keel and ventral lobe margin sharp. Radula pusilla approaches R. pallens, but the latter species is a larger plant with a shorter keel (ca. 1/4 of leaf length), a sinuate free margin of lobule, and the lobule base partly covering the stem. Radula mazarunensis is morphologically identical to R. pusilla and a synonym. 20. Radula quadrata Gottsche Radula andicola Steph. Radula glauca Steph. Radula mollis Lindenb. & Gottsche Radula viridiaurea Spruce

Fig. 67E

On bark, rock, and rotten wood in montane rainforests and plantations up to the páramo, (400–)1200–4000 m. Colombia: Boy, Ces, Cun, Mag, Qui, Ris. Ecuador: Ga, MS, Tu, ZC. General distribution: tropical and subtropical America, extending northward into the USA; also in Africa. Radula quadrata (subg. Radula) is readily recognized by gemmae on leaf margins and the large, imbricate lobules with a widely arched base, covering the entire width of the stem and sometimes extending beyond the stem width. The lobule characters should be studied on mature stem leaves. Radula quadrata is closely similar to R. tectiloba, but the lobules of R. tectiloba are distant or subimbricate (not

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635

densely imbricate) and the lobule base usually covers only part of the stem, rarely the entire stem width. Possibly, R. qudrata and R. tectiloba are conspecific. 21. Radula saccatiloba Steph.

Fig. 67F, G

On bark in montane rainforest, (100–)500–2000  m. Colombia: Cho (Winkler 1970). Ecuador: Es, ZC.  General distribution: scattered in the West Indies, Central America, northern Andes, southeastern Brazil. Radula saccatiloba (subg. Volutoradula) is an uncommon neotropical species that is distinguished by strongly convex leaf lobes, lobules about as long as wide and strongly inflated along the convex keel, and leaf cells flat, thin-walled, with very small trigones and a smooth cuticle. The species resembles R. sonsonensis, but in the latter species, the leaves are less strongly convex and the leaf cells are mammillose (not mammillose in R. saccatiloba). Moreover, R. sonsonensis occurs at higher elevation, above 2500 m. Radula saccatiloba also resembles R. fendleri and was considered a synonym of the latter (Gradstein and Hekking 1989). The latter species clearly differs from R. saccatiloba, however, in having caducous leaf lobes, large bulging trigones, a finely papillose cuticle, and lobules longer than wide. The occurrence of R. saccatiloba on living leaves in lowland rainforest of the Chocó (Winkler 1970) is unlikely; the record must be verified. 22. Radula santacruziana K.Yamada & Gradst. On bark in Miconia scrub, 600–650 m. Ecuador: Ga (Yamada and Gradstein 1991). General distribution: Galápagos Islands, Guianas. Radula santacruziana (subg. Volutoradula) is a rare neotropical species characterized by the rather large plants (2.5–3 mm wide) with broadly ovate leaf lobes, discoid gemmae on leaf margins, leaf cells without or with minute trigones, small subquadrate lobules which are not covering the stem, and straight keels. The species is very similar to R. tectiloba but differs from the latter in larger plant size and distant lobules with a sinuate free margin and the lobule base not covering the stem. 23. Radula schaefer-verwimpii K.Yamada

Fig. 67H

On rotten bark of trees, dead wood, soil, and rock (limestone or sandstone) in montane rainforests and páramo, 700–4000 m. Colombia: Ant, Boy, Cau, Met, Ris, Tol. Ecuador: Lo, ZC.  General distribution: tropical Andes (Colombia to Bolivia), Guyana (Gradstein 5426, GOET), southeastern Brazil. Radula schaefer-verwimpii (subg. Volutoradula) is a rather widespread tropical South American species that is distinguished by (1) plants delicate, 1–1.5 mm wide, irregularly pinnate; (2) leaf lobes strongly caducous, subimbricate, plane to slightly convex, ovate-falcate, margins entire; (3) leaf cells small, ca. 16–24 μm, without or with small trigones, not mammillose, outer wall thin, cuticle smooth; (4) lobules small, distant, ovate-subquadrate, conspicuously inflated along the keel, apex obtuse, base not or very slightly covering the stem (to maximally 1/5 of stem width); and (5) keel strongly convex, not decurrent (Yamada 1990).

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Keys and Descriptions

Radula schaefer-verwimpii resembles R. galapagona, but in the latter species, the lobule base is more strongly expanded, covering more than half the width of the stem. Possibly, the two species should be treated as subspecies. 24. Radula sinskeana K.Yamada On liana, 830 m. Ecuador: Pi (Yamada 1993b; type). General distribution: known only from the type specimen from Ecuador. Radula sinskeana (subg. Volutoradula) is a rare Ecuadorian endemic that is characterized by the ca. 3 mm wide, fragile plants with dichotomous branching (rarely irregularly pinnate according to Yamada 1993b); elongate, ovate-falcate leaf lobes with very small and fully thin-walled cells; and small, distant, subquadrate lobules with a sinuate margin and an expanded base which extends almost fully across the stem. The keel is short, ca. 1/3× lobe length, straight or slightly concave, not decurrent. By its dichotomous to irregularly pinnate branching and small, thin-walled leaf cells, R. sinskeana resembles R. pallens, but in the latter species, the leaf lobes are orbicular and the lobules are shorter, usually wider than long, with the base extending less widely across the stem (to 1/2 of stem width). 25. Radula sinuata Steph. Radula montana Steph. Radula obtusifolia Steph.

Fig. 67I

On bark, rotten wood, and soil in lower montane forest, 900–2000 m. Colombia: Ris (Wolf 1993). Ecuador: EO (Schäfer-Verwimp et al. 2013a). General distribution: scattered in tropical America. Radula sinuata (subg. Volutoradula) is an uncommon neotropical species recognized by the rather regularly (bi)pinnate plants (2–2.5 mm wide) with subimbricate lobules and by the presence of a small, coiled auricle at the lobule base, covering part of the ventral stem surface. Radula sinuata is closely related to R. voluta, but in the latter species, the coiled part of the lobule is much larger, hangs down up to the keel (but not beyond) and covers almost the entire ventral stem surface. Both species may produce caducous leaves (Yamada 1987 as R. montana; Reiner-Drehwald 1994b as R. montana). The record of R. sinuata from the Sierra Nevada de Santa Marta (Winkler 1976) belongs to R. gottscheana. 26. Radula sonsonensis Steph. Type: Colombia, Antioquía, páramo de Sonsón, 1872, G.  Wallis s.n. (G-00043860!) On bark or rock (limestone or sandstone) in upper montane cloud forests and páramo, 2550–4000 m. Colombia: Ant, Boy, Cau, Met, Ris, Tol. General distribution: only known from Colombia. Radula sonsonensis (subg. Volutoradula) is a little-known Colombian endemic that is characterized by (1) leaf lobes occasionally caducous, somewhat convex (but not strongly so), ovate-orbicular, sometimes falcate, apex flat or narrowly recurved, margin ± crenulate; (2) leaf cells conspicuously mammillose on the dorsal surface of the lobe and often with a lens-shaped thickened outer wall, margin cells with a

Radulaceae

637

thickened outer wall, trigones small, with ± concave sides, cuticle smooth; (3) lobules narrowly and often strongly inflated along the keel, apex not elongate, base not covering stem, not or hardly decurrent; and (4) keel convex or almost straight, not or scarcely decurrent (Castle 1963). Radula sonsonensis resembles R. schaefer-verwimpii but differs from the latter in the mammillose leaf cells and the scarcity of caducous leaf lobes. 27. Radula stenocalyx Mont. Radula tenella Gottsche

Fig. 67J

On living leaves and bark in lowland and montane rainforests along the Pacific coast, 50–2300  m. Colombia: Cau, Cho. Ecuador: Es (Schäfer-Verwimp et  al. 2006). General distribution: oceanic regions of tropical South America and tropical Africa. Radula stenocalyx (sect. Epiphyllae) is a widespread tropical-oceanic species that has been found in the Chocó and the Mata Atlântica of Brazil. The species resembles R. flaccida but differs in much smaller gemmae (to 0.25 mm in diameter) with a straight or slightly and shallowly cordate base (not deeply cordate-auriculate) and larger, ovate-triangular lobules without elongate tip. 28. Radula subinflata Lindenb. & Gottsche On trees in montane rainforests up to the páramo, (200–)650–3800 m. Colombia: Cho, Mag. Ecuador: EO, Ga (Yamada and Gradstein 1991), Na, Pi (laguna Mojanda, León-Yánez s.n., QCA), ZC.  General distribution: tropical America, USA. Radula subinflata (subg. Volutoradula) is a widespread but uncommon neotropical species that is distinguished by (1) leaf lobes slightly convex, widely spreading, ovate-orbicular, slightly falcate, apex rounded, plane or recurved; (2) median leaf cells 20–25  μm, not or slightly mammillose, trigones small, cuticle smooth; (3) lobules conspicuously inflated along the keel, free margin weakly concave, base slightly covering the stem; and (4) keel on stem leaves straight or slightly convex (strongly convex on branch leaves), not decurrent. Radula subinflata resembles R. sonsonensis and R. saccatiloba in the inflated carinal region of the lobule, but R. sonsonensis has strongly mammillose cells and R. saccatiloba has strongly convex leaf lobes and a strongly arched keel on stem leaves (Yamada 1987). Radula subinflata also approaches R. nudicaulis, but in the latter species, the margin of the lobule is conspicuously sinuate-plicate and the lobule base extends over 1/4–1/2 of stem width. 29. Radula tectiloba Steph. Radula aurantii Spruce

Fig. 67K

On bark, rock, and rotten wood in montane rainforests, 1000–1900 m. Colombia: Ris (Wolf 1993). Ecuador: ZC (Nöske et al. 2003). General distribution: tropical and subtropical America. Radula tectiloba (subg. Radula) is a rather widespread neotropical species that is distinguished by (1) plants 1.5–2.5 mm wide, irregularly pinnate; (2) leaf lobes with

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marginal gemmae; (3) leaf cells 19–25 μm, trigones small, with concave sides, cuticle smooth; (4) lobules distant to subimbricate, quadrate to subrectangular, usually inflated along the keel, apex angular-obtuse, width of base variable, covering 1/4–4/5 of stem width (rarely the entire stem width); and (5) keel almost straight. The species approaches R. quadrata, but the latter species has larger, densely imbricate lobules with the bases extending fully across and beyond the stem. The lobule characters should be studied on mature stem leaves, not on branch leaves. Radula aurantii is considered a synonym of R. tectiloba following Yamada (1981, 2000, 2003); Reiner-Drehwald (1994b) accepted R. aurantii as a good species. 30. Radula tenera Mitt.

Fig. 67L, M

On twigs and living leaves of trees and shrubs in upper montane forests and páramo, 1900–3500 m. Colombia: Ris (Wolf 1993). Ecuador: Az, Ca, Im, Lo, Na, Pi, ZC. General distribution: Costa Rica, northern Andes, southeastern Brazil. Radula tenera (subg. Volutoradula) is a small, high-montane, ramicolous or epiphyllous neotropical species that is recognized by (1) plants ca. 1 mm wide; (2) leaf lobes strongly convex and with a recurved, rounded to obtuse apex and entire margin; (3) leaf cells small, without trigones, somewhat mammillose but without thickened outer wall, cuticle smooth; (4) lobules subsquadrate, strongly and narrowly inflated along the keel, the inflated portion curved, banana-shaped, keel deeply concave, base of lobule not covering of the stem; and (5) production of small caducous leaf lobes on specialized flagelliform shoots. Radula tenera is somewhat similar to R. involvens; for differences see under the latter species. An overlooked feature of R. tenera, seen in a specimen from southern Ecuador (Parque Nac. Podocarpus, Cajanuma, Gradstein & Mandl 10154, GOET), is the development of caducous leaf lobes on specialized upright, flagelliform shoots. The flagelliform shoots are 2–3 mm long and ca. 0.2 mm wide, arise terminally from leafy stems or branches, and bear small, tightly appressed lobules. The correct author citation for R. tenera is “Mitt. ex Steph.” or, in shortened version, “Steph.” (not “Mitt.”; Renner 2016). 31. Radula voluta Taylor Radula ramulina Taylor Radula ramulina var. microphylla E.Jans

Fig. 67N

Common on bark, rock, rotten wood, humus, and soil in montane forest and páramo, often growing in pendent mats, (450–)1200–4000  m; abundant in moist evergreen Scalesia and Zanthoxylum woodlands on the Galápagos Islands, at 450–1200 m. Colombia: Boy, Cal, Cas, Cau, Ces, Cun, Hui, Mag, Met, Qui, Ris, San, Tol. Ecuador: Az, Ca, Co, Lo, Na, Pa, Pi, Su, Tu, ZC. General distribution: tropical America, Africa, eastern USA, Atlantic coast of western Europe. Radula voluta (subg. Volutoradula) is a widespread montane species that is very common in Colombia and on the Galápagos Islands but is suprisingly known from only few records from the mainland of Ecuador. The species is readily distinguished

Radulaceae

639

by the robust, bipinnate plants (2.5–4 mm wide) with large imbricate lobules that are widely crossing the stem and possess a large auricle, which is circinately coiled across the entire stem surface and reaches downward maximally to the keel (not beyond the keel). The lobule shape must be studied on leaves of the main stem; branch leaves are usually much smaller than stem leaves and may have small, distant lobules that are hardly auriculate. The phenotype with numerous small-leaved branches has been described as R. ramulina var. microphylla. Radula voluta is closely related to R. sinuata; differences between the two are shown in the key. By its large, imbricate lobules of stem leaves, R. voluta may be confused with R. quadrata and R. gottscheana, but R. quadrata has discoid gemmae and lacks a coiled auricle, while in R. gottscheana the lobule auricle extends further downward, well beyond the base of the keel (not extended beyond the base of the keel in R. voluta). Moroever, R. gottscheana has a thick-walled, brownish stem cortex of several cell layers. 32. Radula xalapensis Nees & Mont. Radula frondescens Steph. Radula riparia Spruce

Fig. 67O

On bark, rotten logs, rock, and soil in montane rainforests up to the páramo, also in lowland rainforest, 1500–3800 m. Colombia: Cun, Mag, Qui (Filandia, “Granga Experimental Bengala,” Gradstein s.n., PC), Ris. Ecuador: Lo (Løjtnant & Molau 15051, AAU, PC), (Gradstein 10035, GOET), ZC (Gradstein and Benitez 2017). General distribution: tropical Andes, Costa Rica (Gradstein et al. 2016). Radula xalapensis (subg. Volutoradula) is a characteristic Andean-Costa Rican species that is distinguished by (1) plants rather robust, 3–4  mm wide, regularly 1–2-pinnate; (2) leaf lobes ovate-falcate; (3) median leaf cells 15–25(−30) μm, with small trigones; (4) lobules small, distant, subquadrate, convex, not inflated along the keel, free margin slightly sinuate, plane or somewhat recurved, base rounded and sometimes slightly coiled, usually extending widely across the stem, to ca. 4/5 of stem width or more; and (5) keel concave, decurrent. In the large habit and regularly 1–2-pinnate branching, R. xalapensis resembles R. voluta, but the lobules in the latter species are larger and imbricate and the base of the lobule is much more expanded, extending far across and beyond the stem and being strongly coiled. 33. Radula yanoella R.M.Schust.

Fig. 67P

On angiosperm leaves and fern fronds in lowland and montane rainforest areas, in shaded habitats, 300–2100 m. Ecuador: Na, ZC (Parolly et al. 2004; SchäferVerwimp et al. 2013a); to be expected in Colombia. General distribution: scattered in northern South America (Ecuador, French Guiana, Amazonian Brazil) and in Costa Rica. Radula yanoella (sect. Epiphyllae) is a peculiar, neotenic species that grows in minute patches on living leaves and produces a small thallus, interpreted as a persistent protonema, with short leafy shoots arising from thallus margins (Schuster 1991). The species is known from few localities in the neotropics and has probably

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Keys and Descriptions

been overlooked due to its very small size. The species is close to R. aguirrei; differences are shown in the key. Excluded Records Radula husnotii Castle – Ecuador: Or (Mota de Oliveira and ter Steege 2013). Dubious Caribbean species; the Ecuadorian record should be checked. Radula punctata Steph. (= R. plumosa Steph.)  – Colombia: Ris (Wolf 1993). Patagonian species; the specimen from Colombia is probably misidentified. Váňa and Engel (2013) suggested that the species should be called R. plumosa, but the name R. punctata is slightly older, having been published in September 1884 (Hedwigia 23, fasc. 9: 135), while the name R. plumosa was established in October 1884 (Hedwigia 23, fasc. 10: 154). Radula retroflexa Taylor  – Ecuador: Ga (Evans 1902). Asiatic species; the Galápagos record is considered erroneous (León-Yánez et al. 2006).

HORNWORTS 

(Anthocerotophyta - Anthocerotae)

Plants thalloid, often forming rosettes, growing by a wedge-shaped, four-sided apical cell, sporophyte growing by a basal meristem. Thallus several cells thick at least in the middle, with or without midrib, inner tissue ± undifferentiated, sometimes with large cavities. Nostoc colonies usually present inside the thallus, visible as darkish dots or blue lines. Epidermis cells with or without perforations, with 1(−4) large, plate-like chloroplasts, each sometimes containing a pyrenoid; oil bodies absent. Rhizoids unicellular, smooth. Gametangia immersed in the thallus, originating from an inner thallus cell; antheridia 1 to several in cavities, ± spherical, stalked; archegonia borne single, flask-shaped. Sporophytes projecting from the thallus surface, green, turning black after dehiscence, consisting of foot and capsule, seta lacking. Capsules narrow-cylindrical to long-filiform, surrounded at the base (or up to the apex, in Notothylas) by a tubular involucre, capsule epidermis with or without stomata, spores unicellular or multicellular, with or without trilete mark, elaters (“pseudo-elaters”) usually present, 1–4-celled, with or without spiral band, columella usually present (± absent in Notothylas); capsule dehiscence gradual, from apex downward, by means of 2 valves, rarely by up to 4 valves, with an operculum or rupturing irregularly; spore maturation asynchronous; peristome absent. Protonema very small, thalloid, normally giving rise to only one gametophyte. Contains about 220 species worldwide, in 5 families and 12 genera (Villarreal and Cargill 2016); 20 species in Colombia and Ecuador, in 7 genera. The hornworts of Colombia and Ecuador have been very little studied, and almost nothing is known about the distribution of the species within the countries. To facilitate identification, references to published illustrations are given for species, when available. For

Hornworts

641

practical reasons, the species are treated here in alphabetical order by genus. The classification of the genera of Colombia and Ecuador into families is as follows (Villarreal and Cargill 2016): Anthocerotaceae (Anthoceros). Thallus with globose Nostoc colonies. Sporophytes usually erect. Capsules with columella and stomata. Spore tetrads tetrahedrical, spores ornamented, with a trilete tetrad mark. Elaters without spiral band. On soil in open environments, not inside forests. Dendrocerotaceae (Dendroceros, Nothoceros, Phaeomegaceros). Thallus with globose Nostoc colonies. Sporophytes erect. Capsules with columella, stomata usually absent (present in Phaeomegaceros). Spore tetrads tetrahedrical, spores ornamented, with a trilete tetrad mark. Elaters 1-celled, with spiral band (without spiral band in Phaeomegaceros). In montane forests, usually on bark, rotten wood, rock, or living leaves; rarely on soil in open environments (Phaeomegaceros). Leiosporocerotaceae (Leiosporoceros). Thallus with linearly arranged Nostoc colonies. Sporophytes erect. Columella lacking or short, stomata present. Spore tetrads isobilateral, spores smooth, with a monolete tetrad mark. Elaters without spiral band. On soil in open environments, not inside forests. Notothyladaceae (Notothylas, Phaeoceros). Thallus with globose Nostoc colonies. Sporophytes erect or subhorizontal and lying on the thallus. Columella lacking or short, stomata lacking. Spore tetrads tetrahedrical, spores ornamented, with a trilete tetrad mark. Elaters without spiral band. On soil in open environments, not inside forests. Key to the genera of hornworts of Colombia and Ecuador 1. Thallus usually light green, whole thallus strongly crisped, differentiated into a midrib and unistratose wings. Plants on twigs, branches, or living leaves in humid montane forest areas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendroceros 1. Thallus light green to dark green, whole thallus not strongly crisped, thickened to near the margin, not differentiated into midrib and unistratose wings. Plants on soil, rock, rotten wood, or trunk bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Thallus deep green to blackish-green, elongate, not forming rosettes, margins finely ruffled and ± colorless or subentire and green. Elaters with spiral band, 1-celled. Sporophytes often more than 3  cm long (but sometimes shorter). Spores unicellular or multicellular. Plants on rotten wood, trunk bases, or rock in humid montane forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nothoceros 2. Thallus light green to deep green, usually forming rosettes, margins plane to crispate, green, not finely ruffled. Elaters without spiral band, (1–)2–4-celled. Sporophytes shorter. Spores unicellular. Plants on shaded soil or rock, often on road banks, not inside forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Thallus with internal cavities. Dorsal thallus surface reticulate. Spores brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anthoceros 3. Thallus solid, without cavities. Dorsal thallus surface not reticulate. Spores brown or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

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Keys and Descriptions

4. Thallus margins entire, dorsal thallus surface finely roughened by projecting epidermis cells. Thallus when fresh with long blue lines (= linearly arranged colonies of cyanobacteria inside the thallus). Spores ovoid-oblong, yellow, surface smooth, without trilete mark . . . . . . . . . . . . . . . . . . . . . . . Leiosporoceros 4. Thallus margins entire or dissected, dorsal thallus surface smooth or with lamellate outgrowths, not roughened by projecting epidermis cells. Thallus when fresh without blue lines. Spores rounded-tetrahedral, yellow or brown, surface ornamented, with trilete mark. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Sporophytes lying nearly horizontally on the thallus, less than 5  mm long. Plants mainly in lowlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notothylas 5. Sporophytes erect, more than 5  mm long. Plants montane, up to the páramo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Outer spore surface vermiculate and with one or more rounded depressions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaeomegaceros 6. Outer spore surface with spines or papillae, not vermiculate, without rounded depressions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaeoceros Anthoceros L. Thallus light green, thin, rapidly decaying and becoming blackish in the herbarium, with scattered, globose Nostoc colonies (visible as black dots), thallus usually forming rosettes to 1.5 cm in diameter, without a midrib, margins irregularly dissected into small, often somewhat crispate segments, dorsal surface usually with a reticulate pattern, smooth or covered with crisped or toothed lamellae, cells with 1(−2) chloroplasts with or without pyrenoids; thallus with large internal cavities (cross section). Antheridial chambers with more than 4 antheridia, antheridial jacket of 4 tiers of cells. Sporophytess when mature erect, 1–5 cm long, linear, with stomata and a well-developed columella. Spores brown to dark brown, rounded-tetrahedral, unicellular, outer surface with blunt papillae or sharp spines, inner surface with trilete mark. Elaters (= pseudo-elaters) (1–)2–4-celled, pale blackish-brown, flaccid, walls thin and with irregular bands of thickenings, without spiral band. A subcosmopolitan genus of about 40 species, 5 species in Colombia and Ecuador. The principal characters of Anthoceros are (1) thallus light green, with internal cavities, margins dissected, dorsal surface usually reticulate, smooth or with lamellae; (2) spores dark brown; and (3) elaters (1–)2–4-celled, without spiral band. Based on current knowledge, the neotropical species of Anthoceros can only be identified by mean of spore characters. Literature: Hässel de Menéndez (1990), Gradstein and Costa (2003) 1. Inner spore surface smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Inner spore surface rough, with blunt papillae or tubercles . . . . . . . . . . . . . . . 3 2. Outer spore surface with blunt papillae. . . . . . . . . . . . . . . . . . . . . A. simulans 2. Outer spore surface with sharp spines. . . . . . . . . . . . . . . . . . . . . A. lamellatus

Hornworts

643

3. Trilete mark of spores distinct. Outer spore surface densely covered by sharp spines, ornamentation clearly different from ornamention of inner surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Trilete mark reduced, indistinct. Outer spore surface densely covered by blunt papillae, ornamentation of outer and inner spore surface similar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. granulatus 4. Spores small, 30–35 μm in diameter. Inner spore surface with a large tubercle (much larger than the other ones) in the center . . . . . . . . . . . . . A. tuberculatus 4. Spores larger, 40–60 μm in diameter. Large tubercle lacking . . . . A. punctatus 1. Anthoceros granulatus Gottsche Sphaerosporoceros granulatus (Gottsche) Hässel On soil, 1100–2600  m. Colombia: Cun (Gottsche 1864), Mag (Duckett C8, CBUMAG). General distribution: Mexico, Colombia. Anthoceros granulatus is a rare species that was originally described from Mexico and has been recorded twice from Colombia.The species is characterized by spores densely ornamented by bluntish papillae on outer and inner surface (outer surface not clearly different from inner surface) and trilete mark ± reduced, inconspicuous (Hässel de Menéndez 1988). Illustration: Hässel de Menéndez (1988, Fig. 2A–D) 2. Anthoceros lamellatus Steph.

Fig. 70A

On soil, 2960 m. Colombia: Nar (Villarreal 1078 p.p., COL); to be expected in Ecuador. General distribution: Mexico, Panama, Colombia, southeastern Brazil, Argentina (Hässel de Menéndez and Rubies 2009). Anthoceros lamellatus is a widespread neotropical species that is characterized by spores with a smooth inner surface and the outer surface densely covered by sharp spines. Further illustration: Hässel de Menéndez (1990, Fig. 7A–D) 3. Anthoceros punctatus L. Fig. 68A–G Anthoceros hispidus Steph. Anthoceros venosus Lindenb. & Gottsche var. brachyscyphus Spruce (?) Type: Ecuador, Banos, Spruce s.n. (n.v.) On soil in open and shaded, disturbed places, also on flower pots in greenhouses, 650–2700  m. Colombia: Ces, Cun. Ecuador: Ga (Cerro Azul), Lo, Tu, ZC.  General distribution: widely distributed in tropical and warm-temperate regions of the world. Anthoceros punctatus is a subcosmopolitan species that is recognized by the blackish-brown spores densely covered by spines on the outer surface and by blunt papillae on the inner surface. The thallus is typically light green, the margins are irregularly crispate, and dorsal surface is reticulate and smooth or with crisped or toothed lamellae. The species has many synonyms (see Gradstein and Costa 2003). Further illustrations: Hässel de Menéndez (1990, Fig.  3A–D as P. hispidus), Schuster (1992b, Fig. 1046 as Aspiromitus punctatus subsp. punctatus)

644

 

Keys and Descriptions

Fig. 68 (A–G) Anthoceros punctatus. (A) Habit with sporophytes (i = involucre, c = columella, v = capsule valve). (B) External capsule wall with stoma. (C) Spore, outer surface. (D) Spore, inner surface. (E) Elaters. (F) Portion of cross section of thallus, showing cavities and Nostoc colony inside the thallus and lamellae on the dorsal surface. (G) Antheridium. (H–N) Dendroceros crispus. (H) Epidermis cells with a single chloroplast and trigones. (I) Cross section of thallus, showing midrib. (J) Portion of cross section of thallus, showing midrib. (K) Portion of elater. (L) Elaters. (M) Spore, with several nuclei due to endosporous gemination. (N) Habit, dorsal view. (All from Gradstein et al. 2001)

Hornworts

645

4. Anthoceros simulans M.Howe On soil, 650–1600  m. Ecuador: Ga (Isabela, Santa Cruz; Gradstein 2009). General distribution: only known from the Galápagos Islands. Anthoceros simulans is the only species of liverworts or hornworts found above 1500 m on the Galápagos Islands. The species is close to A. lamellatus; differences are shown in the key. Illustration: Hässel de Menéndez (1990, Fig. 7A–D) 5. Anthoceros tuberculatus Lehm. & Lindenb. Anthoceros vegetans M.Howe On earth along roads and trails, at the edge of moist forest, 100–1200 m; on the Galápagos Islands on moist soil over lava rock in pampa vegetation, 500–800 m. Colombia: Cho (Tutunendo, Gradstein 9036, COL), Mag (Gradstein et al. 2016). Ecuador: Ga (Isabela, Santa Cruz, San Cristóbal; Gradstein 2009), Tu (Río Topo, Gradstein 10023, QCA). General distribution: scattered in tropical America. Anthoceros tuberculatus was long known from the Galápagos Islands where it was described as A. vegetans. Recently, it has been recorded from Colombia and the mainland of Ecuador. The species is distinguished by the very small thalli with numerous dorsal lamellae, the small spores (less than 40 μm in diameter), and, especially, the presence of a large tubercle in the center of each of the three facets of the inner spore surface. The outer spore surface is densely covered by sharp, confluent spines. Illustration: Hässel de Menéndez (1990, Fig. 12A–D) Dendroceros Nees Thallus small, yellowish-green, 2–5 mm wide, with scattered, globose Nostoc colonies (visible as black dots), ± pinnately branched or forked, with a midrib and unistratose wings, the wings conspicuously undulate-crispate in neotropical plants and sometimes with large holes (perforations), cells with 1(−2) chloroplasts, epidermis cells with trigones; thallus solid or with cavities in the midrib (subg. Apoceros). Antheridial chambers with 1(−4) antheridia. Sporophytes when mature erect, 1–5  cm long, linear, without stomata, columella present. Spores green, large, ca. 40–80  μm long, rounded, multicellular, outer surface papillose. Elaters long and narrow (to ca. 300 μm long), with spiral band. A pantropical genus of about 30 species, 2 in Colombia and Ecuador. Several further species have been described from the region but their status is doubtful. The principal characters of Dendroceros are (1) thallus differentiated into midrib and unicellular wings, the wings strongly undulate-crispate (neotropical plants), epidermis cells with trigones, midrib solid in neotropical plants; (2) capsule wall without stomata; (3) spores green, multicellular; and (4) elaters long and narrow, with a spiral band. Contrary to other hornwort taxa, the species of Dendroceros are typically ramicolous or epiphyllous, growing on twigs, branches, and living leaves in humid montane forest areas.

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Keys and Descriptions

Literature: Hasegawa (1980), Villarreal et al. (2010a) 1. Thallus wings with large, colorless perforations (holes). Capsule epidermis cells with nodulose thickenings. . . . . . . . . . . . . . . . . . . . . . . . . . . D. crispatus 1. Thallus wings without large perforations. Capsule epidermis cells without nodulose thickenings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. crispus 1. Dendroceros crispatus (Hook.) Nees On palm trunk (Bactris gasipes) in garden and on living leaves, 500–1200 m. Ecuador: Ch (Spruce 1885, as var. simplicior Spruce), Na (Río Payamino, HolmNielsen 837, AAU, PC), ZC (Río Nangaritza, Gradstein 12106, HUTPL). General distribution: tropical America. Dendroceros crispatus is readily recognized by the thallus wings with large, colorless holes (perforations). The record from Chimborazo (R.  Spruce) needs verification. 2. Dendroceros crispus (Sw.) Nees Dendroceros foliatus Spruce (fide Villarreal et al. 2015)

Fig. 68H–N

On branches, twigs, and living leaves of trees and shrubs in humid lowland and montane rainforests, 200–2900 m; on the Galápagos Islands common on twigs and living leaves in moist evergreen woodland and scrub at 450–650  m (Gradstein 2009). Colombia: Hui, Mag, Qui. Ecuador: Ga, Pi (Gradstein 12629, QCA), Or, Tu. General distribution: tropical America. The species of Dendroceros from Colombia and Ecuador have been little studied, and some of the published records of D. crispus might belong to D. crispatus. For differences between the two species, see the key. Further Records Dendroceros allionii Steph. – Ecuador: MS (Stephani 1898–1924). Taxonomic status unclear. Dendroceros breutelii Nees – Ecuador: MS (Stephani 1898–1924 as D. ecuadorensis Steph.). Species accepted by Villarreal et al. (2015). Dendroceros jamesonii Taylor  – Ecuador: Pi (Taylor 1848). Taxonomic status unclear. Leiosporoceros Hässel Thallus deep green, fleshy, with linearly arranged Nostoc colonies (visible as long blue lines in the thallus), thallus forming elongate rosettes to 2  cm in diameter, without midrib, margins entire, flat or somewhat undulate, dorsal surface finely roughened by projecting epidermal cells, not reticulate, cells with 1(−2) chloroplasts; thallus solid (cross section), bi-convex, without cavities. Antheridial chambers with up to 70 antheridia. Sporophytes when mature erect, to 4 cm long, linear, with stomata and a well-developed columella. Spores pale yellow, transparent,

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small, ca. 20  μm long, ovoid-oblong, unicellular, surface smooth, without trilete mark. Elaters long and narrow, to 280 μm long, 1-celled, yellowish-brown, walls thick, without spiral band. A neotropical genus with a single species. The principal characters of Leiosporoceros are (1) thallus rather fleshy, solid, margins entire, dorsal surface finely roughened by projecting epidermis cells; (2) Nostoc colonies linearly arranged inside the thallus; (3) spores pale yellow, ovoid-oblong, small, ca. 20 μm long, surface smooth, without trilete mark; and (4) elaters long and narrow, 1-celled, walls thick, without sporal band. Literature: Hässel de Menéndez (1986), Villarreal and Renzaglia (2006a) 1. Leiosporoceros dussii (Steph.) Hässel Anthoceros dussii Steph. Phaeoceros dussii (Steph.) J.Haseg.

Figs. 69M, N, 70B

On exposed, moist soil, and rock in montane rainforest areas, particularly on roadbanks and along rivers, 1800–3000 m. Colombia: Tol (Villarreal et al. 2010a). Ecuador: Pi, ZC. General distribution: Central America (Costa Rica, Panama), northern Andes, Lesser Antilles. Leiosporoceros dussii resembles Phaeoceros, but the thallus of Leiosporoceros is fleshier and the dorsal surface of the thallus is somewhat velvety in appearance due to projecting epidermal cells. Moreover, the thallus contains unique, linearly arranged colonies of cyanobacteria, which are visible in fresh material as blue lines in the thallus (Villarreal and Renzaglia 2006a). The small, transparent, ovoid-oblong spores of L. dussii with a smooth outer surface and a monolete mark on the inner surface are also very characteristic. Further illustration: Villarreal and Renzaglia (2006a) Nothoceros (R.M.Schust.) J.Haseg. Megaceros Campb. p.p. (non typus) Thallus glistening dark green, 1–5 cm long, 0.3–1.2 cm wide, usually with scattered, globose Nostoc colonies (visible as black dots), forked, pinnate or irregularly branched, usually without a midrib, margins undulate to crispate to finely ruffled and ± colorless, sometimes ± entire, cells with 1–2(−3) chloroplasts, pyrenoid present or lacking; thallus solid (cross section), without cavities. Antheridial chambers with 1(−2) antheridia. Sporophytes when mature erect, often very long (over 5 cm), linear, without stomata, columella present. Spores green to yellowish-brown, usually unicellular (multicellular in N. renzagliensis), rounded to ovoid, outer surface papillose-tuberculate. Elaters long and narrow (to 450 μm long), with spiral band. A widespread, tropical and southern-temperate genus of about ten species, three in Colombia and Ecuador. Nothoceros was recently split from Megaceros on molecular grounds and lacks diagnostic morphological characters separating it from Megaceros (Villarreal et al. 2010a). The two genera share the frequent occurrence

648

 

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of more than one chloroplast in the thallus cells, relatively long sporophytes without stomata and long and narrow elaters with a spiral band. Contrary to other hornwort taxa, the species of Nothoceros and Megaceros typically grow on wet rock, rotten wood, or trunk bases in montane forests. All neotropical species decribed in Megaceros belong to Nothoceros, and most of them are synonyms of N. vincentianus (Villarreal and Renner 2014; Villarreal et al. 2015). The identities of some species remain unclear, and studies using living material are needed (J. C. Villarreal, pers. comm.). Literature: Villarreal et al. (2010a, 2012, 2015), Villarreal and Renner (2014) 1. Plants robust, more than 2 cm long, dark green when fresh, on wet rock, logs or soil, monoicous, frequently with sporophytes. Thallus margins undulate to crispate. Mature sporophytes more than 3 cm long. Epidermis cells with (1–)2–3 chloroplasts, the chloroplasts with or without pyrenoids . . . . . N. vincentianus 1. Plants smaller, less than 2 cm long, on bark, monoicous or dioicous, often sterile. Thallus margins strongly ruffled, sometimes partly entire. Mature sporophytes smaller, less than 3 cm long. Epidermis cells with 1(−2) chloroplasts, without pyrenoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Thallus dark green, ca. 3–8× longer than wide, pinnately branched, margins strongly ruffled throughout. Epidermis cells with one chloroplast. Spores unicellular. Plants dioicous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. aenigmaticus 2. Thallus light green, as wide as long or slightly elongate, 1–2× longer than wide, irregularly branched, margins partly entire. Epidermis cells with 1–2 chloroplasts. Spores multicellular. Plants monoicous . . . . . . . . . . . . N. renzagliensis 1. Nothoceros aenigmaticus (R.M.Schust.) J.C.Villarreal & K.D.McFarland Megaceros aenigmaticus R.M.Schust. On trees trunks in montane forest, growing over a layer of bryophytes, 2000–3230 m. Colombia: Mag (Gradstein et al. 2016), Nar (Villarreal et al. 2012). General distribution: USA, Mexico to Colombia. Characteristics of this species are the small, elongate, pinnately branched, usually dark green thalli with strongly ruffled margins, the presence of only one chloroplast in the epidermis cells (exceptionally 2), and dioicy. The material from Magdalena is sterile and the identification is tentative. Illustration: Schuster (1992b, Figs. 1052 and 1053) 2. Nothoceros renzagliensis J.C.Villarreal, L.V.Campos & Uribe On shrub in upper montane forest, growing over a layer of bryophytes, 2968 m. Colombia: Nar (Villarreal et al. 2012). General distribution: only known from the type from Colombia. Nothoceros renzagliensis the only known species in the genus with multicellular spores. Further characteristic features are the small (usually less than 2 cm long), light green, irregularly branched thalli without midrib, epidermis cells with 1–2 chloroplasts, and monoicous sexuality. The species grows epiphytically on bark attached to a layer of bryophytes, similarly as in N. aenigmaticus. For differences with the latter species, see the key.

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Fig. 69 (A–D) Nothoceros vincentianus. (A) Habit with sporophytes (c = columella, i = involucre, v = capsule valve). (B) Habit (n = Nostoc colony), dorsal view. (C) Spore. (D) Elater. (E–G) Phaeomegaceros squamuligerus. (E) Spore, inner surface. (F) Spore, outer surface showing small rounded depression. (G) Elater. (H–L) Notothylas breutelii. (H) Elaters. (I) Spore, outer surface. (J) Spore, inner surface. (K and L) Habit (c = capsule, i = involucre, n = Nostoc colony). (M and N) Leiosporoceros dussii. (M) Spore. (N) Elater. (All from Gradstein et al. 2001)

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Illustration: Villarreal et al. (2012) 3. Nothoceros vincentianus (Lehm. & Lindenb.) J.C.Villarreal Megaceros vincentianus (Lehm. & Lindenb.) Campb. Megaceros colombianus Steph.

Fig. 69A–D

On rotten wood, logs, dripping wet rock, and soil in montane forests and páramo, sometimes growing submerged in rivulets, 250–3725 m. Colombia: Boy (Gradstein 12715, “cf.”, PC) Cun, Qui, Ris (Aguirre & Gradstein 1439, 1458, GOET), Tol. Ecuador: EO, Lo, MS, Na, Pi, ZC. General distribution: tropical America. Nothoceros vincentianus is the most common Nothoceros species in the tropical Andes and the only one that grows on wet rock and rotten wood; the other local Nothoceros species are smaller and grow on bark. Nothoceros vincentianus occurs normally above 1500 m in the Andes but has been found as low as 250 m in the province of El Oro (Benitez et s.n., HUTPL) and at 500 m in Costa Rica and Panama (J. C. Villarreal, pers. comm.). The species is readily distinguished by the large, dark green thalli (more than 2 cm long) with irregularly undulate to crispate margins, the epidermis cells with (1–)2–3 chloroplasts (with or without pyrenoid), and the long sporophytes, more than 3 cm long. The plants are monoicous and usually possess sporophytes. Notothylas A.Gray Thallus light green to deep green, nearly flat, with scattered, globose Nostoc colonies (visible as black dots), usually forming rosettes about 0.5–3 cm in diameter, without a midrib, irregularly divided, margins entire to crenulate to lobulate, dorsal surface smooth or with a few lamellae, not reticulate, cells with 1(−2) chloroplasts; thallus solid (cross section), without cavities. Antheridial chambers with 1–6 antheridia, antheridial jacket of irregularly arranged cells. Sporophytes when mature very short, to 4 mm long, lying almost horizontally on the thallus and largely enclosed in the involucre, cylindrical or banana-shaped, without epidermal stomata, columella reduced or absent, dehiscence imperfectly by 2(−4) valves or an operculum, or capsule cleistocarpic and rupturing irregularly (N. javanicus). Spores dark brown or yellowish, ca. 40–60 μm in diameter, rounded-tetrahedral, unicellular, the outer surface ornamented, the inner surface with trilete mark. Elaters ± rounded, unicellular, about the same size as spores, pale brown, walls with irregular thickenings, without spiral band; sometimes elaters lacking. A pantropical and northern-temperate genus of 20 species, 4 in Colombia and Ecuador, mostly in open, man-made habitats at low elevation (below 1000 m). The principal characters of Notothylas are (1) thalli usually forming small rosettes, margins entire to lobulate, not ruffled; (2) mature sporophytes short, to 4  mm long, largely enclosed in the involucre, lying horizontally on the thallus or slightly ascending, stomata lacking, columella usually absent or rudimentary; (3) spores yellow or dark brown; and (4) elaters, when present, rounded and without spiral band.

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Literature: Hässel de Menéndez (1976), Schuster (1992b), Gradstein and Costa (2003) 1. Mature spores dark brown. Elaters present . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Mature spores yellow. Elaters present or absent . . . . . . . . . . . . . . . . . . . . . . . 3 2. Thallus rosettes rather large, often more than 2  cm in diameter. Distal spore surface coarsely tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. breutelii 2. Thallus rosettes smaller, 0.5–1 cm in diameter. Distal spore surface vermicular, almost smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. galapagensis 3. Elaters present. Capsule opening by 2–4 longitudinal valves . . . N. orbicularis 3. Elaters lacking. Capsule opening by an operculum, valves lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. vitalii 1. Notothylas breutelii (Gottsche) Gottsche Notothylas amazonica Spruce

Fig. 69H–L

On soil in gardens and cultivated fields, below 1000  m. Colombia: Mag (Gradstein et  al. 2016). General distribution: tropical America, Philippines, Hawaii. Notothylas breutelii is a widespread tropical amphi-Pacific species that is recognized by the rather large thallus, often more than 2 cm in diameter, the brown to blackish spores with a tuberculate outer surface, and the presence of rounded elaters. Further illustration: Schuster (1992b, Fig. 1056) 2. Notothylas galapagensis M.Howe On soil and volcanic rock in rather dry, semi-deciduous woodlands, ca. 100–250 m. Ecuador: Ga (Floreana, Isabela; Tagus Cove, Santiago: James Bay). General distribution: only known from the Galápagos Islands. Notothylas galapagensis is a small plant with thallus rosettes maximally 1 cm in diameter; very short capsules, to 2 mm long; and brown spores with almost smooth outer and inner surface. The rounded elaters are about as large as the spores. The species is endemic to the Galápagos Islands and was originally recorded from the archipelago as N. orbicularis (Evans 1902), but the latter species differs by yellow spores. Illustration: Hässel de Menéndez (1976, Fig. 2A–B) 3. Notothylas orbicularis (Schwein.) Sull. On soil along trails in rather dry forest areas, from sea level to 750 m. Colombia: Mag (Gradstein et al. 2016). Ecuador: EO, Lo (Benitez et al. 2012 as N. vitalii; Schäfer-Verwimp et  al. 2013a as N. vitalii). General distribution: North America, Europe, Japan, tropical Africa, Colombia, Ecuador, eastern Brazil (Pôrto et al. 1999). Notothylas orbicularis is a widespread Holarctic species that is rare in tropical America where it has been recorded from eastern Brazil, Colombia, and Ecuador. The species forms small rosettes, maximally 1 cm in diameter, the spores are yellow, and elaters are present. In the key of Gradstein and Costa (2003), the characters

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of N. orbicularis are interchanged with those of N. vitalii, and as a consequence some records of N. orbicularis have been published under the name N. vitalii. Illustration: Schuster (1992b, Fig. 1054) 4. Notothylas vitalii Udar & D.K.Singh On soil along trails and in gardens, from sea level to 1000 m. Colombia: Mag (obs. J.G. Duckett, Dec 2015). Ecuador (new): MS (Bomboiza village, Pérez et al. s.n., QCA). General distribution: eastern Brazil, Ecuador, Colombia. Notothylas vitalii is a rare South American species that stands out by yellow spores, absence of elaters, and capsules opening by an operculum instead of valves. Several published records from Ecuador (Benitez et  al. 2012; Schäfer-Verwimp et al. 2013a) belong to N. orbicularis (see note under N. orbicularis). After this book went in press, Leandro de Almeida and Peralta (DOI: 10.1007/ s40415-020-00602-x) showed that the correct name for Notothylas vitalii is N. javanicus (Sande Lac.) Gottsche. Illustration: Gradstein & Costa (2003, Fig. 103J–M) Phaeoceros Prosk. Thallus deep green, rather fleshy, with scattered, globose Nostoc colonies (visible as black dots), usually forming rosettes to 2–3 cm in diameter, without a midrib, margins entire or crenate, flat or somewhat crispate, dorsal surface smooth, not reticulate, cells with 1(−2) chloroplasts and usually with pyrenoids inside the chloroplasts; thallus solid (cross section), without cavities. Antheridial chambers with 2–4 antheridia, antheridial jacket of irregularly arranged cells. Sporophytes when mature erect, 1–4(−9) cm long, linear, with epidermal stomata and a well-developed columella. Spores yellow, rounded-tetrahedral, unicellular, outer surface variously ornamented by spines or papillae, inner surface with trilete mark. Elaters (1–)2–4-celled, pale brown, walls thin and with irregular thickenings, without spiral band. A large, subcosmopolitan genus of about 30 species, 4 in Colombia and Ecuador. The principal characters of Phaeoceros are (1) thallus deep green, rather fleshy, solid, margins entire or somewhat dissected, dorsal surface smooth, not reticulate; (2) spores yellow; and (3) elaters (1–)2–4-celled, without spiral band. Based on current knowledge, the neotropical species of Phaeoceros can only be identified by means of spore characters. Literature: Hässel de Menéndez (1989), Schuster (1992b as Anthoceros), Gradstein and Costa (2003) 1. Outer spore surface with small, blunt papillae . . . . . . . . . . . . . . . . . . . P. tenuis 1. Outer spore surface with small, sharp spines . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Inner spore surface with a group of small papillae in each of the three facets (= triangular surface areas bordered by the trilete mark). Plants autoicous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. carolinianus 2. Inner spore surface smooth. Plants dioicous . . . . . . . . . . . . . . . . . . . . . . . . . . 3

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3. Outer spore surface densely covered by spines . . . . . . . . . . . . . . . . . . P. laevis 3. Outer spore surface with few, scattered spines . . . . . . . . . . . . P. pichinchensis 1. Phaeoceros carolinianus (Michx.) Prosk. Phaeoceros laevis subsp. carolinianus (Michx.) J.Haseg. Anthoceros callistictus Spruce (Megaceros callistictus (Spruce) Steph.) (fide Villarreal et al. 2015) Aspiromitus allionii Steph. Type: Ecuador, Gualaquiza, “ad ripam umbrosam rivi Gualaquiza,” 20 Aug 1909, Allioni s.n., Bryotheca Levier 6502 (G-00060895), syn. fide J. Hasegawa in sched. Phaeoceros gualaquizanus (Steph.) Gradst. (Anthoceros gualaquizanus Steph.) Type: Ecuador, Gualaquiza, Bomboiza, Sep 1910, Allioni 561 (G-00061230) On moist soil, logs, and rock at roadsides and along rivers, 500–3500  m. Colombia: Mag (Gradstein et al. 2016), Nar (Villarreal 1079, COL), Qui (Lopez s.n., HUQ). Ecuador: MS (Stephani 1898–1924 as Anthoceros gualaquizanus and Aspiromitus allionii), Na (Holm-Nielsen et al. 26154, AAU, PC), Pa (Spruce 1885 as A. callistictus). General distribution: widely distributed in temperate regions; scattered in the tropics. The distribution of the widespread P. carolinianus in Colombia and Ecuador is very incompletely known. The species has in the past been included in P. laevis, and part of the records of the latter species may belong to P. carolinianus. Differences between the two species are shown in the key. Phaeoceros qualaquizanus is probably a synonym of P. carolininanus based on the description and illustration of Hässel de Menéndez (1989). Illustrations: Hässel de Menéndez (1989, Fig. 10A–D), Gradstein & Costa (2003, Fig. 103N–Q) 2. Phaeoceros laevis (L.) Prosk. Anthoceros laevis L. subsp. laevis On moist soil and rock on roadsides, along rivers and water courses, and near springs, 650–2900 m. Colombia: Cal, Cun, Mag, Nar, Tol. Ecuador: Ga (Santa Cruz), MS.  General distribution: widely distributed in warm-temperate and tropical regions; occurring in warmer climates than P. carolinianus. Phaeoceros laevis is close to P. carolinianus, and some of the records of P. laevis may belong to P. carolinianus. Illustration: Schuster (1992b, Fig. 1043) 3. Phaeoceros pichinchensis (Spruce) Hässel Anthoceros pichinchensis Spruce On rock in river on Mt. Pichincha at high elevation. Ecuador: Pi (Spruce 1885). General distribution: only known from the type from Ecuador. Phaeoceros pichinchensis is a rare rheophilous Ecuadorian endemic close to P. laevis but differs in the outer spore surface with only few, scattered spines (densely spinose in P. laevis). Like Phaeomegaceros squamuligerus, the species is only

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known from the type specimen collected on Mt. Pichincha by Richard Spruce; its continued existence needs confirmation. Illustration: Hässel de Menéndez (1989, Fig. 7A–D) 4. Phaeoceros tenuis (Spruce) Hässel Anthoceros tenuis Spruce On humid soil on steep road banks and along trails, 800–1800 m. Ecuador: MS (road Mácas  – Puyo, Pérez & Gradstein 11614, QCA), ZC (Nöske et  al. 2003). General distribution: scattered in tropical America (Hässel de Menéndez 1989). Phaeoceros tenuis differs from other species of Phaeoceros in Colombia and Ecuador by the outer spore surface with blunt papillae instead of sharp spines. Illustration: Hässel de Menéndez (1989, Fig. 12A–D) Phaeomegaceros R.J.Duff et al. Thallus as in Phaeoceros; chloroplasts without pyrenoids. Antheridial chambers with only 1 antheridium. Sporophytes when mature erect, 1–4(−9) cm long, linear, with stomata and a well-developed columella. Spores yellow, rounded-tetrahedral, unicellular, spore surface vermiculate, with one or more depressions on the outer surface, inner surface with a trilete mark. Elaters (1–)2–4-celled, pale brown, walls thin and with irregular thickenings, without spiral band. A genus of seven species, two in Colombia and Ecuador. The principal characters of Phaeomegaceros are (1) thallus as in Phaeoceros but chloroplasts without pyrenoids; (2) spores yellow, surface vermiculate, with one or more depressions on the outer surface; and (3) elaters (1–)2–4-celled, without spiral band. Literature: Duff et al. (2007), Villarreal et al. (2010b) 1. Dorsal thallus surface smooth or slightly roughened by projecting epidermal cells. Outer spore surface with seven rounded depressions: one central depression surrounded by a ring of six depressions . . . . . . . . . . . . . . . . . P. fimbriatus 1. Dorsal thallus surface with triangular or lamellate outgrowths. Outer spore surface with one rounded depression, ring of six depressions lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. squamuligerus 1. Phaeomegaceros fimbriatus (Gottsche) R.J.Duff et al. Fig. 70C Anthoceros fimbriatus Gottsche (Phaeoceros fimbriatus [Gottsche] Gradst.) Aspiromitus schlimianus (Gottsche) Steph. On moist soil on roadsides and along rivulets in open montane environments and páramo, (600–)2000–4000  m. Colombia: Boy (Gradstein 12715, UPTC), Cun, Mag, Nar. Ecuador: Pi. General distribution: tropical Andes (Venezuela to Bolivia), southern Central America, West Indies (Lavocat Bernard 2018). Phaeomegaceros fimbriatus is a widespread Andean species that is easily distinguished by the presence of seven deep, rounded depressions on the outer spore surface: one central depression and six surrounding ones. The dorsal thallus surface is smooth or slightly roughened by projecting epidermal cells. An undescribed

Hornworts

655

Fig. 70  Spores of hornworts. (A) Anthoceros lamellatus, smooth inner spore surface with tetrad mark (center, left), spinose outer surface (right). (B) Leptosporoceros dussii, smooth inner spore surface with monolete mark. (C) Phaeomegaceros fimbriatus, outer spore surface. (D) Phaeomegaceros squamuligerus, outer spore surface. Photographs by J. C. Villarreal (A–C) and F. Müller (D). Reproduced with permission from Caldasia 40: 268, Fig. 2 (2018)

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species from páramo of Venezuela has spores similarly ornamented as those of P. fimbriatus but larger in size (Villarreal and Renzaglia 2006b). Illustration: Villarreal and Renzaglia (2006b) 2. Phaeomegaceros squamuligerus (Spruce) J.C.Villarreal  Figs. 69E–G, 70D Anthoceros squamuligerus Spruce (Phaeoceros squamuligerus [Spruce] Hässel) Phaeomegaceros squamuliger (Spruce) J.C.Villarreal, orthogr. var. On rock in river on Mt. Pichincha at high elevation. Ecuador: Pi (Spruce 1885). General distribution: Ecuador, central Chile. Phaeomegaceros squamuligerus is a rare rheophilous species that differs from P. fimbriatus by the characters given in the key. Villarreal et al. (2010b) recognized two subspecies, subsp. squamuligerus in Ecuador and subsp. hasseliae in central Chile. Subspecies squamuligerus is only known from the type specimen collected by Richard Spruce on Mt. Pichincha; its continued existence needs confirmation. Villarreal et al. (2010b) and Villarreal and Cargill (2016) changed the spelling of the name of the species to “squamuliger.” However, the original spelling “squamuligerus” is not wrong and should therefore be retained (Art. 60.1 ICN). Illustration: Hässel de Menéndez (1989, Fig.  17A–D as Phaeoceros squamuligerus)

Glossary1

Abaxial  The side of a leaf or branch away from the stem (opposed to adaxial). Acrogynous  Archegonia at the apex of a stem or branch (opposed to anacrogynous). Acuminate  From a broad apex gradually tapered toward a sharp point. Acute  Sharply pointed. Adaxial  The side of a leaf or branch toward the stem (opposed to abaxial). Air chamber A cavity in the thallus of complex thalloid liverworts (Marchantiopsida). Alpine Upper vegetation belt on high mountains, above the upper forest line. Usually without trees. In the Andes the alpine belt is called “páramo“ or “puna.” Amphigastria (sing. amphigastrium)  Underleaves in leafy liverworts. Ampliate In Plagiochila, ventral leaf base expanded, covering the stem. Amphitropical Distributed in the Mediterranean and subtropical regions of the Northern and Southern Hemisphere; absent or very rare in the tropics. Anacrogynous  Archegonia in a lateral position on stem, branch or thallus, not at the apex (opposed to acrogynous). Androecium (plur. androecia) Antheridia and surrounding bracts; the male gametoecium. Angular  Shape or outline with angles, not rounded. Annular thickenings Ring-like thickenings extending over both tangential and radial cell walls in liverwort capsules. Antheridiophore (= male gametangiophore)  Stalked, antheridia-bearing receptacle on the thallus of the Marchantiaceae. Antheridium (plur. antheridia) Male sex organ or gametangium, producing antherozoids (=spermatozoids). Antherozoid (= spermatozoid)  Male gamete.

1  This Glossary is based on the Glossarium Polyglottum Bryologiae (Magill 1990) and the Guide to the Bryophytes of Tropical America (Gradstein et al. 2001), with additions and modifications.

© Springer Nature Switzerland AG 2021 S. R. Gradstein, The Liverworts and Hornworts of Colombia and Ecuador, Memoirs of the New York Botanical Garden 121, https://doi.org/10.1007/978-3-030-49450-6

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658

Glossary

Antical  The leaf margin oriented toward the shoot apex (opposite of postical). Also used for the dorsal surface of a stem. Aphanolejeunea-type branch  Similar to Lejeunea-type branch but without collar at the base. Apical  At the tip. Apical cell  An embryonic cell at the growing tip that divides repeatedly to form new cells. Apiculate  Abruptly ending in a short point (mucronate is shorter pointed; cuspidate is longer and more stoutly pointed). Appendiculate  With short lacinia or auricles at leaf or underleaf bases. Appressed  Lying closely together; leaves lying close or flat against the stem or against each other; plants lying close to the substrate. Aquatic  Of plants, growing permanently submerged or floating, completing the life cycle in water (opposed to rheophytic or terrestrial). Archegoniophore (= female gametangiophore) Stalked, archegonia-bearing receptacle on the thallus of many Marchantiales. Archegonium (plur. archegonia) Female sex organ or gametangium, containing an egg. Areolate (= reticulate)  Surface with a network formed by areoles, e.g., of spores or thallus. Areole  Small angular surface area, the basic unit of a network formed by areoles. Ascending  Pointing or arising obliquely upward, away from the substrate. Athecal  Without a sheath of tissue at the base; said of collarless branches. Attenuate  Slenderly tapering. Auricle  A small, ear-like lobe, often present at leaf base. Auriculate  With auricles. Autoicous (= autoecious)  With archegonia and antheridia on separate branches of the same plant. Axil  Angle between stem and leaf. Axillary  In the leaf axils. Axis  The main stem. Bazzania-type branch  Ventral-intercalary branch (with collar at the base), originating from the axil of underleaves. Beak (= rostrum)  Elongate apex of a lobule, perianth, or operculum (see also lobule beak). Bifid  Divided into two segments or lobes. Bilobed Bifid. Bipolar Distributed in the temperate regions of the Northern and Southern Hemisphere; absent or very rare in the tropics. Biseriate  In two rows, of cells. Bipinnate  Twice pinnately branched. Bisbifid  Twice bifid with the subsequent parts more shallowly bifid. Blunt Obtuse. Bordered  Having margins differentiated from the rest of the structure (leaf, underleaf, perianth, etc.) in cell shape, size, color or thickness.

Glossary

659

Bracteole  A modified underleaf associated with a gametangium. Bracts  Specialized leaves surrounding reproductive organs. Caducous (= deciduous)  Detaching, falling off. Calyptra  A membrane covering the developing sporophyte, derived largely from the archegonial venter. Campanulate Bell-shaped. Canaliculate  Channeled (lengthwise). Capsule (= theca)  The sporangium or spore-bearing portion of the sporophyte. Caulocalyx Protective structure surrounding the sporophyte in Fossombronia, resembling a perianth. Central strand  A small bundle of narrowly elongate cells forming a central axis in stems, thalli or midrib. Chlorophyllose  Containing chlorophyll (=green pigment), generally green unless masked by some other pigments. Cilia (sing. cilium)  Hair-like or thread-like structures, mostly one cell wide and unbranched. Ciliate  With hair-like appendages. Circinate  Curved in a circle. Cladia (sing. cladium) Modified, small caducous branches for vegetative reproduction. Clasping  Closely surrounding or enveloping a structure. Cleistocarpous  Indehiscent; capsule without a regular mechanism for opening. Coalesced  Two or more structures becoming united and losing their identity; e.g., of trigones (compare confluent). Coelocaule A specialized shoot calyptra in certain leafy liverworts, characterized by extreme axial growth and complete penetration of the sporophyte into the shoot apex; frequently associated with the absence of a perianth; e.g., in Trichocolea. Collar  Sheath of tissue around the base of branches in leafy liverworts. Collenchymatous  With cell walls more heavily thickened at the angles (trigone). Columella The central, sterile column of tissue in the capsule of mosses and hornworts. Complanate  Flattened or compressed; e.g., leaves flattened into one plane. Complicate  Folded together. Complicate-bilobed  Having 2 leaf lobes folded against one another. Concave  Hollow (opposed of convex). Conduplicate  Folded longitudinally along the middle. Confluent  Two structures becoming united without losing their identity; e.g., of trigones or margins (compare coalesced). Conical Cone-shaped. Connate  Fused, e.g., the bases of opposite leaves, or underleaves with leaves. Connivent  Directed or pointing together; e.g., tips of leaf segments converging toward one another. Constricted  Abruptly narrowed. Contiguous Adjacent.

660

Glossary

Contorted (= tortuose)  Irregularly curved or twisted. Convex  Bulging outwards (opposed to concave). Convolute  Rolled together. Cordate  Heart-shaped. Of leaves with two large auricles at base, or of trigones with 2 convex sides and one concave side. Cortex  The differentiated outer cell layer(s) of the stem. Cortical  Referring to the outer part of a stem, or cortex. Corticolous  Growing on bark. Cosmopolitan  Occurring in all major floristic zones of the world. Costa (= midrib, nerve) Mid-vein of a leaf or thallus, always more than one cell thick. Costate  With a costa. Crenate  With rounded teeth. Crenulate  With minute, rounded teeth (smaller than crenate). Crisped (= crispate)  Strongly wavy; variously curled, twisted and contorted. Cristate  Crested; bearing a crest-like ridge. Cucullate  Hooded or hood-shaped; used to describe strongly concave leaves that are inflexed at the tips. Cuneate (= cuneiform) Wedge-shaped, narrowed to the base along straight, oblique lines. Cushion  Growth form with stems erect, tightly clustered and somewhat radiating at edges. Cuspidate  Ending abruptly in a stout, rigid point (more longly and stoutly pointed than apiculate or mucronate). Cuticle A wax layer on the outside surface of the epidermis. In bryophytes it is usually very thin. Cylindrical  Narrowly elongate, circular in transverse section. Deciduous (= caducous)  Detaching, falling off. Decolorate  Discolored, colorless. Decorticated  Lacking bark. Decumbent  With stem prostrate but with ascending tips. Decurrent  With leaf bases extending down the stem, beyond the leaf insertion. Decussate  Cross-shaped; of tetrads, spores arranged pair-wise, at a right angle to each other; characteristic of Lejeuneoideae (see Gradstein, 1994, Figs. 6, 7). Dehiscent  Having a distinct opening mechanism. Dendroid  Tree-like; with erect stems that are branched in the upper part. Dentate  With sharp, coarse teeth. Denticulate  Finely toothed. Depauperate  Poorly developed. Diaspore An agent of dispersal; any structure that becomes detached from the parent plant and gives rise to a new individual (seed, spore, gemma, cladium, leaf, etc.) Dimorphic  Occurring in two forms; e.g., with two types of leaves. Dioicous (= dioecious)  With archegonia and antheridia on separate plants. Diploid  Having a double set of chromosomes (2n).

Glossary

661

Disciform  Flat and circular. Discoid  Flat and rounded, shaped like a disc. Disk cell  Large margin cell of the lobule, distal to the apex of the lobule of some species of Lejeunea. Disjunct  Distribution discontinuous, with major gaps. Distal Away from the base or point of attachment; toward the apex of a leaf or stem; the outer, convex face of a spore (opposed to proximal). Distal hyaline papilla See: hyaline papilla. Distant  Well-spaced; e.g., leaves positioned with a space between adjacent leaves. Distichous  Leaves in two rows. Divergent  Turned in opposite directions. Dorsal  The upper surface, away from the substrate (opposed to ventral). Dorsal leaf-free strip  Portion of the dorsal stem surface that is not occupied by the leaf insertion. A leaf-free strip is present when the leaf insertion line does not extend to the dorsal stem-midline, as for example seen in several members of Cephaloziaceae and Zoopsidella. The width of the leaf-free strip is measured as the approximate number of longitudinal stem cell rows between opposite leaf bases. Dorsiventral  Flattened, with distinct upper and lower surfaces. Edentate  Without teeth. Egg  The female gamete, produced in an archegonium. Elater  Long and narrow, diploid, hygroscopic cell with spiral thickenings, derived by mitotic division. Elaterophore  A tuft or brush-like mass of cells to which elaters are attached, at the base or the apex of some liverwort capsules (e.g., in Riccardia, Metzgeria). Elliptical  Oblong with convex sides. Elobulate  Without lobule. Emarginate  Having a broad and shallow notch at the apex (opposed to retuse). Embryo  Young sporophyte, developing from a zygote. Emergent  Partially exposed, referring to capsules or perianths only partly projecting beyond the tips of the bracts. Endemic  Geographic distribution restricted; limited to a single country or floristic area. Endogenous  Originating from internal rather than superficial cells or tissues. Endosporic (= endosporous)  Inside the spore; protonema germinating inside the spore wall, hence spores multicellular (opposed to exosporic). Entally  Toward the inner side, referring to the position of the hyaline papilla of the lobule in Lejeuneaceae. Entire  Without teeth. Epidermis  The outer cell layer of a stem, thallus, capsule or other structure. Epilithic (= saxicolous)  Growing on rock. Epiphyllous (= foliicolous)  Growing on living leaves of other plants. Epiphyte  A plant growing on another living plant. Epiphytic  Growing on another living plant.

662

Glossary

Epistatic In leafy liverworts, lobules of male bracts not overlapping ventrally (opposed to hypostatic). Eplicate  Without keels, referring to the perianth: see also terete. Erect  Upright; leaves directed toward stem apex. Erose  Irregularly notched or ragged. Exosporic (= exosporous)  Outside the spore wall. Explanate (= laminate) In Frullania, lobule that is not rolled inwards forming a sac (opposed to saccate). Exserted Protruding and largely exposed (opposed to immersed; compare also emergent). Exsiccatae (sing. exsiccata)  Dried plants; referring to widely distributed sets of herbarium specimens with printed labels that are used as standards for comparison. Falcate  Curved like the blade of a sickle. Falcate-secund  Strongly curved and turned to one side. Fenestrate  Pierced with openings or perforations. Fertile  In bryophytes, with gametangia or sporophytes (opposed to sterile). Fide  According to. Filamentous Thread-like. Filiform  Very slender and elongate. Flabellate  Fan-shaped, with several branches spreading outwards from almost a single point. Flaccid  Soft, limp. Flagelliform  Whip-like; branches with a gradual attenuation from ordinary leaves at the base to minute, scale-like leaves at the tip. Flagellum (plur. flagella) Whip-like structure; a branch with scale-like or rudimentary leaves (compare stolon). Flexuose  Slightly and irregularly bent or twisted. Foliicolous (= epiphyllous)  Growing on leaves of other plants. Foliose  Leafy or leaf-like. Foot  Basal portion of the bryophyte sporophyte, embedded in the gametophyte and serving as an organ of absorption and attachment. Fragmentation Breaking up; a method of vegetative reproduction in which an organ (or whole plant) is separated into parts capable of producing a new organism. Fringed  With a ciliate margin. Frond  The branched or foliose part of an erect stem, occurring in dendroid liverworts and mosses. Frullania-type branch Terminal branch (without collar at the base) originating from a ventral leaf initial so that it replaces the ventral half of a leaf. The branch is therefore associated with a half-leaf on its dorsal side. Furcate Forked. Furrowed Grooved. Fusiform  Spindle-shaped; narrow (more than 3× as long as wide) and tapered at both ends.

Glossary

663

Gametangiophore  Stalked receptacle bearing gametangia, arising from the thallus in Marchantiales (antheridiophore, archegoniophore). Gametangium  Sex organ (archegonium, antheridium). Gamete  Sexual reproductive cell (spermatozoid, egg). Gametoecium  Gametangia and surrounding bracts (androecium, gynoecium). Gametophyte  The haploid, sexual generation producing gametes (spermatozoids and eggs). In bryophytes, the gametophyte is the actual plant. Gemmae (sing. gemma)  Vegetative diaspores, can be produced on leaves, stems, thalli or rhizoids. Gemma cup  Cup-shaped, gemma-containing structure, produced on the thallus of Marchantia. Gibbous  Swollen on one side. Glabrous Smooth. Glaucous Whitish-blue. Globose  Rounded, spherical. Granular  Composed of small globules, referring to oil bodies (= segmented). Granulose  Roughened with minute, blunt projections. Groove  With a narrow and shallow furrow in the middle. Gynoecium (plur. gynoecia)  Archegonia and the surrounding bracts; the female gametoecium Habit  The general appearance of a plant. Habitat  The local environment of a plant. Haploid  Having a single set of chromosomes (n). Hemiphyll  First leafy appendage at the ventral base of Frullania-type branches in Frullania. Hemiphylls may be bifid or undivided; bifid hemiphylls resemble an underleaf, undivided ones resemble a half-leaf. Heteroicous (= polyoicous)  Used for species that can be monoicous or dioicous. Hexagonal  With 6 angles, usually of cells. Hispid  With short, stiff hairs; bristly. Humus  Decomposing organic material. Hyaline  Colorless or transparent. Hyaline papilla  Thin-walled, secretory cell associated with leaf initial cells in liverworts and persisting in mature leaves, e.g., at the tip of the stylus in Frullania, at the tips of the leaves in Telaranea and Zoopsis, and at the base and the apex of the lobule in the Lejeuneaceae. The papilla at the lobule apex is always positioned at the proximal base of the first lobule tooth (= the distalmost tooth). Sometimes the first tooth is reduced (e.g., Cheilolejeunea); in the latter, case the distalmost visible tooth is the second tooth and the papilla is positioned distal to the base of this tooth. Hyalodermis  Stem epidermis of enlarged cells with thin (not thickened) walls. Hygrophytic  Growing in wet places. Hygroscopic  Moving in response to changes in humidity; e.g., elaters. Hypostatic  In leafy liverworts, lobules of male bracts overlapping ventrally, imbricate (opposed to epistatic).

664

Glossary

Idioblast  Uniquely differentiated cell, distinct from other cells of the same tissue in size, form and/or content; e.g., ocellus, oil-cell. Imbricate  Overlapping, like shingles on a roof. Immersed  Completely below the surface; e.g., said the perianth completed covered by bracts (compare exserted). Inclined  Bent down Incubous An oblique leaf insertion in which the dorsal leaf margin is oriented toward the apex of the stem (when viewed from above; each leaf overlaps the adjacent younger leaf above), e.g., in Calypogeia, Bazzania, and members of Porellales (opposed to succubous). Incurved (= involute) Curved upward (adaxially) and inward (opposed to recurved). Indehiscent  Lacking a distinct opening mechanism (opposed to dehiscent). Inflexed  Bent abruptly upward (adaxially) at more than 90° and inwardly. Inflorescence (= gametoecium)  The term “inflorescence” refers to the flowers of angiosperms and should not be used for bryophytes. Innovation  A branch formed at the base of the perianth. In Lejeuneaceae, innovations can be lejeuneoid (first leafy appendage at the base of the branch is a lateral leaf) or pycnolejeuneoid (first leafy appendage is an underleaf). Inrolled See involute Insertion  The place or line of attachment of a structure. Intercalary  Intermediate in position, somewhere below the apex. Intercalary branch  Branch originating from inside the stem, away from the apex, usually having a collar at the base; the associated leaves are not modified by branch formation (opposed to terminal branch). Involucre A thin structure surrounding the gametangia or the developing sporophyte; also, the bracts (and bracteoles) of the gynoecium (= perichaetium). Involute (= incurved)  Rolled upward (adaxially) and inward (opposed to revolute). Isodiametric  About as broad as long, of cells. Keel Fold. Keeled  Folded along the middle, like the keel of a boat. Laciniae (sing. lacinia)  Appendages that are more than one cell wide. Laciniate  With lacinia. Lamella  Ridge, plate, or filament-like outgrowth of the surface Lamina The undivided portion of the leaf; the thin wings on either side of the midrib of a thallose liverwort. Laminate (= explanate) In Frullania, lobule that is not rolled inwards, not forming a sac (opposed to saccate). Lanceolate  Lance-shaped, narrow and tapered from near the base. Lax Loosely. Lejeunea-type branch  Intercalary branch (with a collar at the base) originating from the lateral side of the stem, from behind a leaf. Lejeuneoid innovation  Innovation in which the first leafy appendage arising at the base of the innovation is a lateral leaf (compare pycnolejeuneoid innovation).

Glossary

665

Lejeuneoid sporophyte Sporophyte of members of Lejeuneaceae subfam. Lejeuneoideae, characterized by slow dehiscence of the capsule, decussate tetrads, elongate spores, and elaters with reduced spirals. Lignicolous  Occurring on dead, decorticated wood. Ligulate  Strap-shaped; long and narrow with parallel sides (sometimes used to mean tongue-shaped or lingulate). Linear  Very long and narrow, with parallel sides; narrower than lingulate. Lingulate  Tongue-shaped, oblong below, somewhat broader toward the apex (cf. ligulate). Lobe  A major segment of a leaf, larger than a tooth (cf. lobule) Lobule  A small lobe; e.g., the smaller segment of a complicate-bilobed leaf with unequal segments. Lobule beak  Downward extension of the sac along the adaxial margin of the laminate part of the lobule, in Frullania subg. Chonanthelia. Longitudinal  Lengthwise, parallel to the long axis; e.g., longitudinal leaf insertion. Lowland  Area between sea level and ca. 500 m elevation Lumen (plur. lumina)  The cavity of a cell, enclosed by the cell wall. Lunulate  With a wide, shallow sinus; moon-shaped. Mamilla  Alternate spelling (British English, Latin) of mammilla. Mammilla (plur. mammillae; American English) Hollow projection from the cell surface, with the cell lumen extending into the protuberance (opposite of papilla). Mammillose With mammillae. Marsupium  A fleshy, cylindrical structure that penetrates downward into the soil and contains the sporophyte. Usually derived from stem tissue. Mat  Growth form with stems horizontal and densely interwoven. Median  Central, middle. Medulla  Internal tissue of stem and seta. Meristem  Embryonic tissue. Merophyte A segment cut off from the apical cell and the tissues and organs derived from it. A mature merophyte consists of one leaf or underleaf and associated stem tissue. Microlepidozia-type branch  Terminal branch (without collar at the base) originating from a dorsal leaf initial so that it replaces the dorsal half of a leaf. The branch is therefore associated with a half-leaf on its ventral side. Micrometer (= micron)  One thousandth of a millimeter (abbreviated μm). Microphyllous  Small-leaved (cf. flagelliform). Midleaf  Middle part of a leaf. Midrib See costa Monochasial  An arrangement of gynoecia in a row, formed by repeatedly innovating branches. Monoicous (= monoecious)  Bisexual, with antheridia and archegonia on the same plant; including autoicous, synoicous, and paroicous. Monolete  In spores, having a single, linear scar on the proximal surface that marks the point of attachment in the tetrad (compare trilete).

666

Glossary

Montane  Pertaining to mountains; altitudinal belt between lowlands and upper forest line. In the tropics usually extending from about 500 to 3000(–4000) m. The upper forest line may be lowered due to local climate and human impact and is much lower on islands. Mucro  A short, abrupt point. Mucronate  Ending in a short, abrupt point (apiculate is somewhat longer; cuspidate is even longer and stouter). Multi- Many. n  The haploid (gametophytic) chromosome number of an organism. Neotenic  Mature organism with juvenile characters. Neoteny  Occurrence of juvenile characters in mature organism. Nerve See costa Nodulose  With knob-like wall thickenings. Notch  V-shaped indentation, e.g., at the tip of the thallus. Obcuneate  Wedge-shaped with apex broader than base. Oblique  Slanted; e.g., an oblique leaf insertion is one that is between transverse and longitudinal. Obliquely spreading  Spreading at an angle of 30–60°. Oblong  Rectangular with rounded corners or ends. Obovate  Egg-shaped with apex broader than base. Obpyriform  Pear-shaped with apex narrower than base. Obtuse  Blunt, broadly pointed, at an angle greater than 90°. Ocellus (plur. ocelli)  A specialized cell in the leaves (and perianth) of some members of Frullania and Lejeuneaceae, having one very large oil body and lacking chloroplasts. Ocelli may be light gray, brown or red in color. Oil body  A membrane-bound organelle containing terpenoids, characteristic of liverwort cells. The content of oil bodies may be homogeneous, or segmented and made up of numerous small or large granules. Upon degeneration, homogeneous oil bodies may become segmented while degenerated oil bodies may become homogeneous. Therefore, it is necessary to study oil bodies on fresh material. Oil cell  Colorless cell in the thallus of complex thalloid liverworts (Marchantiopsida), containing a large oil body or oil droplets, and no chloroplasts. Opaque  Not transparent. Operculum  Lid covering the mouth of the capsule, falling away when the spores are mature; found in mosses, some complex thalloid liverworts (Marchantiopsida) and some hornworts (Anthocerotophyta). Orbicular  Rounded, circular. Oval  Widely elliptical Ovate  Egg-shaped, broader at the base than at the apex. Ovoid  Egg-shaped in three dimensions. Ovum Egg. Paired (= twinned)  Associated in groups of two; e.g., hairs, teeth. Palmate  Lobed or branched in a radiating fashion, resembling fingers spreading from the palm of a hand. Pantropical  Occurring throughout the tropics (America, Africa, Asia).

Glossary

667

Papilla (plur. papillae)  A solid projection from the cell surface (opposed to mammilla); papillae can be rounded or elongate in circumference, and rather low and flat (lens-shaped) or highly swollen (tuberculate). Papillose  Roughened by papillae. Páramo  Shrubby or herbaceous vegetation belt above the upper forest line, with harsh, humid climate; in Colombia and Ecuador usually above 3200 m, occasionally at lower elevation; frequently inhabited by large, pachycaulous plants with giant, apical leaf rosettes. Paraphyllia (sing. paraphyllium)  Small, green outgrowths of stems, much smaller than leaves and of various shapes, i.e., filiform, lanceolate, etc. Paraphyses (sing. paraphysis)  Multicellular hairs associated with antheridia and archegonia. Parenchymatic  Isodiametric or rectangular cells with broad, truncate tips (opposite to prosenchymatic). Paroicous (= paroecious)  With antheridia in the axils of bracts just below the perichaetium, on the same shoot. Pegged rhizoids  Rhizoids with papilla-like projections on the inner wall surface. Pellucid  Clear, translucent, transparent, hyaline. Pendent (= pendulous)  Hanging downward. Percurrent  Extending to the apex Peri-Afroamerican  Occurring in tropical America and on the East African Islands but not on the African mainland. Peri-Amazonian  Occurring along the headwaters of the tributaries of the Amazon river, near the margins of the Amazon Basin. Perianth Organ of foliar origin enclosing the developing sporophyte in leafy liverworts. Perichaetium Cluster of modified leaves (bracts, bracteoles) enclosing the archegonia. Periclinal  A plane of cell division parallel to the surface. Perigonium  Cluster of modified leaves (bracts, bracteoles) enclosing the antheridia (in mosses). Perigynium Fleshy structure surrounding and protecting the developing sporophyte, derived from stem cells; a perigynium may or may not be associated with a shoot calyptra. Piliferous  With hair point Pilose  With long hairs. Pinnate  Branched in a feather-like fashion, with numerous branches on opposite sides of the stem. Plane Flat. Plagiochila-type branch  Lateral-intercalary branch (with collar at the base), originating from the axil of lateral leaves. Plica (plur. plicae)  Longitudinal furrow. Plicate  With longitudinal furrows. Plumose  Closely and regularly pinnate, feathery. Pluricellular  Made up of several cells (more than one; compare unicellular).

668

Glossary

Pluriplicate  With many keels or folds (more than 5), referring to the perianth. Polymorphic  Of more than one form, variable. Polyoicous See heteroicous Pore  A small opening. Porose  Having pores. Postical  The leaf margin oriented toward the base of the shoot (opposite of antical). Also used for the ventral surface of a stem. Pre-apical tooth  Tooth arising from the sinus between keel and ventral leaf margin, beyond the first lobule tooth. Seen in Drepanolejeunea and some other genera of Lejeuneaceae. Propagule  Reduced bud, branch, or leaf serving in vegetative reproduction (compare diaspore). Prosenchymatic  Elongate cells with narrow tips (opposite to parenchymatic). Prostrate  Laying flat on the surface; creeping. Protonema (plur. protonemata)  A filamentous or thalloid structure resulting from spore germination and giving rise to one or more gametophytes. Proximal  Near the base or the point of attachment; the internal face of a spore (opposite of distal). Proximal hyaline papilla See: hyaline papilla. Pseudo-elater Unicellular or multicellular sterile cells in the hornwort capsule, lacking spirals. Pseudoperianth  The inner one of two involucres in some thalloid liverworts; usually a thin, tubular structure of thalline origin surrounding the archegonia, the calyptra and the developing sporophyte, and enclosed by the outer involucre. Puna  Dry, herbaceous vegetation belt above the upper forest line, usually above 3200 m; absent in Colombia. Ptychanthoid sporophyte Sporophyte of members of Lejeuneaceae subfam. Ptychanthoideae, characterized by explosive dehiscence of the capsule, tetrahedral tetrads, isodiametric spores, and elaters with well-developed spirals. Pycnolejeuneoid innovation  Innovation in which the first leafy appendage arising at the base of the innovation is an underleaf (compare lejeuneoid innovation). Pyrenoid  Colorless, rounded body in the chloroplasts of algae and hornworts that is involved in carbon fixation and starch formation and storage. Pyriform Pear-shaped. Quadrate  Square or nearly so. Quadrifid  Divided into four lobes or segments. Radiate  Elongate at the angles, of trigones. Radula-type branch  Terminal branch (without collar at the base) originating from the stem epidermis and therefore associated with an unmodified leaf. Receptacle A rounded or elongate structure bearing antheridia or archegonia in thalloid liverworts; found on the thallus, inside the thallus or elevated on a stalk (gametangiophore). Recessed Sunken; perianth beak sunken in a hollow perianth apex, becoming invisible. Rectangular  Longer than wide with straight angles.

Glossary

669

Recurved (= revolute) Curved downward (abaxally) and inward (opposed to incurved). Reflexed  Bent abruptly down (abaxally) at more than 90° and inwardly. Reniform  Kidney-shaped, much wider than long. Reticulate (= areolate) Forming a network; surface covered by a network of areoles. Retuse  Having a narrow, shallow indentation in a broad, rounded apex (opposed to emarginate). Revolute (= recurved) Rolled downward (abaxally) and backward (opposed to involute). Rheophytic (= rheophilous)  Growing partly or periodically submerged in running water (rivers, creeks) and producing reproductive organs on emerged shoots, above the water (opposed to aquatic or terrestrial). Rhizoid disc  A cushion-like swelling at the base of an underleaf, from which rhizoids originate. Rhizoid  A root-like hair that functions in anchorage; in liverworts and hornworts typically 1-celled, in mosses many-celled and with oblique end walls. Rhizome  A slender, creeping, often subterranean stem with reduced leaves, giving rise to erect leafy stems or fronds. Rhombic  Diamond-shaped, oblong-hexagonal. Rhomboidal See rhombic Rosette  In thalloid liverworts, a rounded, segmented thallus with segments radiating from a central point; in leafy liverworts (Frullania, Lejeuneaceae), a small rounded area on the spore surface consisting of radially arranged spines. Rostrum See beak Ruffled  Curled, irregularly crisped; refers to the thallus margin of some hornwort species. Saccate Sac-like. Saxicolous (= epilithic)  Growing on rocks. Scale  A thin, membranous structure; usually present on the ventral surface of the thallus of complex thalloid liverworts (Marchantiopsida). Secondary rhizoid disc  Disc of fused rhizoids in epiphyllous liverworts, firmly fixing the plant to the leaf surface. Secund  Turned to one side Segmented  Divided in parts, e.g., lobed leaves, granular oil bodies. Seriate  In a row, e.g., uni-, bi-, tri-, multiseriate; applied to number of adjacent rows of cells. Serrulate  Finely toothed. Seta (plur. setae)  Stalk-like portion of the sporophyte, between capsule and foot. Sheathing  Surrounding and clasping the stem or the perianth. Shoot calyptra A somewhat fleshy calyptra derived from the archegonial venter and from axial or thalline tissue peripheral to the archegonium. Shouldered In Plagiochila, leaf margin rather longly stretched upwards before curving outwards. Sinuate (= sinuose) Wavy.

670

Glossary

Sinus  The depression separating two segments or lobes. Slime hair  A hair-like, mucilage-secreting outgrowth of the epidermis. Slime papilla  A mucilage-secreting cell. Spathulate  Tapering to the base from a broad, rounded apex. Spermatozoid (= antherozoid)  The male gamete, produced in an antheridium. Spinose  With long and sharp teeth. Spore  In bryophytes, a reproductive unit produced in the capsule as a result of meiosis, upon germination giving rise to a protonema. Sporeling  Protonema and juvenile gametophore arising from a spore. Sporophyte The diploid, asexual generation producing (meio)spores, in bryophytes normally consisting of a foot, seta and capsule and permanently attached to the gametophyte. Spreading  Forming an angle of 30–90°; e.g., leaves spreading from the stem. Squarrose  Spreading at almost right angles. Stellate Star-shaped. Sterile  In bryophytes, without gametangia or sporophytes (opposed to fertile). Stipitate  Having a stalk-like base. Stolon  Slender branch or shoot without leaves. Stoloniferous  With stolons. Stoma (plur. stomata)  Pore bordered by two guard cells, occurring in the capsule wall of many mosses and hornworts. -Stratose  In layers; e.g., uni-, bi-, multistratose. Striate  With fine ridges or lines. Stylus (plur. styli)  A minute, subulate to triangular structure usually crowned by a slime papilla, found between the lobule and the stem in some leafy liverworts, especially Frullania. Sub-  Under, below; almost. Suberect  Almost upright; almost parallel to the stem; Spreading at an angle of less than 30°. Subfloral innovation A branch that arises just below the perianth in leafy liverworts. Subpáramo  Lowest páramo belt, directly above the forest line, characterized by shrubby vegetation. Subspecies  Taxon characterized by one or two morphological characters and by a different geographic distribution (= geographic race, allopatric race). Subula  A long, slender point. Subulate  Slenderly long-acuminate. Superpáramo  The upper páramo belt, above 4000 m, characterized by very harsh climate and low rocky, discontinuous vegetation. Succubous Oblique leaf insertion in which the dorsal leaf margin is oriented toward the base of the stem (when viewed from above, each leaf overlaps the adjacent younger leaf underneath), e.g., in Lophocolea and Plagiochila (opposed to incubous). Synoicous (= synoecious) With antheridia and archegonia mixed in the same gametoecium.

Glossary

671

Terete  Smooth-cylindrical, rounded in cross section. Terminal  At the apex or distal end (opposite of basal). Terminal branch  A branch formed at the stem apex; the branch is often associated with a half-leaf or -underleaf and has no collar at its base (opposed to intercalary branch). Terrestrial (= terricolous)  Growing on soil, not submerged. Tetrad  Group of four spores produced by meiotic division of a spore mother cell, often tightly pressed together so that the inner (= proximal) face shows the mark of the other spores. Tetrads in bryophytes are usually tetrahedral in shape, with three spores in one plane and one centrally over the other three. The presence of a trilete scar on the inner spore surface marks the tetrahedral position of the spores in the tetrad. Some bryophytes, e.g., members of the subfamily Lejeuneoideae, have decussate tetrads, characterized by the spores being arranged pair-wise in the tetrad, at a right angle to each other (see Gradstein, 1994, Figs. 6, 7). Tetrahedral  Four-sided; e.g., shape of the spore tetrad of most bryophytes (see above); shape of the apical cell of leafy liverworts (Jungermanniidae). Thalloid (= thallose)  Thallus-like; flat and ribbon- or rosette-like. Thallus (plur. thalli) A flattened plant body that is not differentiated into stems and leaves. Theca (plur. thecae) See capsule Tomentose Woolly. Tooth (plur. teeth)  Sharp, projecting cells on the margin or surface of a leaf. Tortuose  Irregularly bent or twisted. Transverse  Across, perpendicular to the axis (opposed to oblique and longitudinal). Trifid  Divided into three segments. Trigone  A three-angled thickening at corners of cells (compare collenchymatous). Trigonous  Three-angled; applied to a three-keeled perianth. Trilete mark  Triradiate scar on the inner (= proximal) surface of spores, marking the tetrahedral position of the spores in the tetrad. Truncate  Abruptly cut off at the apex. Tubercle  Wart, large papilla. Tuberculate  With warts (coarser than papillae). Tubular  Cylindrical and hollow. Tuft  Small cushion; growth form with stems erect but radiating at the edges. Twinned (= paired)  Associated in groups of two. Underleaves (= amphigastria) Leaves originating from the ventral surface of the stem. Undulate Wavy Unicellular  Made up of only one cell (compare pluricellular). Uniseriate  In one row, of cells. Unistratose  One-layered, of a single layer of cells. Utricle The strongly enlarged, bladder-like lobule on branch leaves of Ceratolejeunea. Valve One of the segments into which the capsule separates upon dehiscence; small lamina regulating the opening of the lobule sac in Colura and Pleurozia.

672

Glossary

Variety  Taxon characterized by a single diagnostic morphological character but not differing in geographic distribution. Ventrad  Directed toward the ventral side of the plant. Ventral  The lower surface or side, next to the substrate (compare dorsal). Verrucose See papillose Verruculose  Slightly roughened by small papillae. Vitta  In liverworts, band of elongate cells in a leaf, resembling a nerve but only one cell layer thick. Wart  A solid projection from the surface; large papilla. Wide-spreading  Spreading at an angle of 60–90°. Wing  A thin, flat expansion or appendage. Xerophytic  Growing in dry places.

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Index

A Acanthocoleus, 395, 396 aberrans var. aberrans, 396 aberrans var. laevis, 396 Acrobolbaceae, 130–136 Acrobolbus, 130–135 antillanus, 133 azoricus, 133 bispinosus, 295 caducifolius, 131 cuneifolius, 131 diversifolius, 134 laceratus, 316 laxus, 133 madeirensis, 133 mittenii, 134 ruwenzorensis, 133 setaceus, 134 wilsonii, 134 wilsonii var. andinus, 135 Acrolejeunea, 396, 398 emergens, 398 torulosa, 398 Adelanthaceae, 136–142 subfam. Jamesoniellaceae, 191 Adelanthus, 137–139 aureocinctus, 141 aureomarginatus, 138 carabayensis, 138 corticolus, 139 crossii, 141 decipiens, 141 decipiens subsp. aureocinctus, 141 decurvus, 138 integerrimus, 139

pittieri, 139 Aitonia lanigera, 45 subplana, 45 Allorgella, 398, 399 schnellii, 399 Alobiella, 161 biloba, 166 campanensis, 161 husnotii, 161 Alobiellopsis acroscypha, 163 dominicensis, 163 Amblyolejeunea, 502 fulfordiae, 531 Amphilejeunea, 502 catinulifera, 521 patellifera, 521 reflexistipula, 545 viridissima, 521 Anastrophyllaceae, 324–338 Anastrophyllum, 326–329 adulterinum, 329 auritum, 327 austroamericanum, 336 calocystum, 327 conforme, 329 crebrifolium, 327 gemmiferum, 337 gradsteinii, 329 hians, 327 leptodictyon, 329 leucocephalum, 327 leucostomum, 327 minutirete, 327

© Springer Nature Switzerland AG 2021 S. R. Gradstein, The Liverworts and Hornworts of Colombia and Ecuador, Memoirs of the New York Botanical Garden 121, https://doi.org/10.1007/978-3-030-49450-6

699

Index

700 Anastrophyllum (cont.) minutum, 337 nigrescens, 327 paramicola, 327 pearcei, 329 piligerum, 327, 336 pusillum, 329 stellatum, 329 subg. Crossocalyx, 336 tubulosum, 329 Andrewsianthus achrous, 331 jamesonii, 331 Androcryphia porphyrorhiza, 76 Aneura, 88, 90 allionii, 101 latissima, 88 pectinata, 99 pinguis, 88 pseudopinguis, 88 Aneuraceae, 87–105 Anomoclada, 168–172 mucosa, 171 portoricensis, 171 Anoplolejeunea, 399, 400 conferta, 399, 400 Anthelia, 142, 143 juratzkana, 142, 143 Antheliaceae, 142–143 Anthoceros, 642 callistictus, 653 dussii, 647 fimbriatus, 654 granulatus, 643 hispidus, 643 laevis subsp. laevis, 653 lamellatus, 643 pichinchensis, 653 punctatus, 643 simulans, 645 squamuligerus, 656 tenuis, 654 tuberculatus, 645 vegetans, 645 venosus var. brachyscyphus, 643 Anthocerotaceae, 641 Anthocerotophyta, 640–646 Aphanolejeunea, 440 angustissima, 451 azorica, 445 camillii, 446 cingens, 446 crenata, 447 cyathiphylla, 451

ensifolia, 446 ephemeroides, 450 gracilis, 447 heterophylla, 447 kunertiana, 451 liliputana, 446 longifolia, 445 microscopica, 450 minuta, 451 misiones, 446 papilliloba, 451 pterocolea, 452 pustulosa, 451 sicifolia, 453 subsphaeroidea, 454 truncatifolia, 447 verrucosa, 450 Apotomanthus, 338 succulentus, 339 Arachniopsis, 238–241 coactilis, 240 diacantha, 239 pecten, 240 Archilejeunea, 400–404 auberiana, 467 badia, 401 bischlerana, 467 crispistipula, 401 cruegeri, 467 florentissima, 467 fuscescens, 401 juliformis, 401, 403 leprieurii, 467 ludoviciana, 403 ludoviciana subsp. ludoviciana, 403 ludoviciana subsp. porelloides, 403 nebeliana, 404 parviflora, 467 spruceana, 401 subg. Dibrachiella, 466 viridissima, 467 Arnelliaceae, 143–145 Aspiromitus punctatus subsp. punctatus, 643 schlimianus, 654 Asterella, 43 lateralis, 44 lindenbergiana, 45 macropoda, 44 pringlei, 44 saccata, 45 venosa, 45 Athalamia, 46, 47 andina, 47 Aureolejeunea, 421

Index aurifera, 427 fulva, 438 paramicola, 436 paramoensis, 436 quinquecarinata, 436 tonduzana, 438 Austrofossombronia, 70–77 peruviana, 75 Aytoniaceae, 42–46 B Balantiopsidaceae, 145–152 Bazzania, 205–219 acanthostipa, 208, 209 acuminata, 208 affinis, 209 ancistrodes, 212 angustifalcata, 215 arcuata, 213 arcuata var. mamillosa, 214 aurescens, 209 bidens, 210 bidens var. dissodonta, 217 boliviana, 212 breuteliana, 213 canelensis, 210 chilensis, 210 chimantaensis, 207 chimborazensis, 213 crassidentata, 211 creberrima, 217 cuervi, 213 cuneistipula, 211 decidua, 215 denticulata, 211 denticulifera, 211 diversicuspis, 212 eggersiana, 212 elongata, 212 falcata, 212, 213 flavicans, 213 glaziovii, 213 gottscheana, 213 gracilis, 213 heteroclada, 213 heterostipa, 219 hookeri var. hookeri, 213, 214 hookeri var. mamillosa, 214, 215 jamaicensis, 215 latidens, 213 lechleri, 217 leptostipa, 218 liebmaniana, 218 lindigii, 209

701 longa, 213, 219 longistipula, 215 macrostipula, 212 pallidevirens, 217 peruviana, 217 phyllobola, 217 placophylla, 218 planiuscula, 211 praerupta, 219 quadricrenata, 209 robusta, 212 roraimensis, 218 rusbyi, 211 schlimiana, 212, 213 serrata, 218 skottsbergii, 217 speciosum, 213 spruceana, 218 stolonifera, 218, 219 taleana, 209 tayloriana, 219 teretiuscula, 213 tricrenata, 219 tricuspidata, 215 vincentina, 218 viridissima, 213 Bazzania fendleri, 215 Blepharolejeunea, 404–406 harlingii, 405 incongrua, 405 saccata subsp. harlingii comb. nov., 405 saccata subsp. saccata, 405 securifolia, 406 Blepharostoma, 321, 322 trichophyllum, 321 Blepharostomataceae, 320–324 Bonneria, 236, 237 granatensis, 236 Brachiolejeunea, 406, 407 asplundii, 406 bicolor, 493 chinantlana, 496 columbica, 493 corticalis, 493 densifolia, 493 grandidentata, 495 hans-meyeri, 406 laciniatiflora, 495 laxifolia, 407 leiboldiana, 407 liebmaniana, 495 nitidiuscula, 405 paramicola, 436 phyllorhiza, 407 rupestris, 493

Index

702 Brachiolejeunea (cont.) succisa, 493 tristis, 496 wrightii, 496 Bromeliophila, 407, 408 helenae, 408 Bryolejeunea, 408 Bryopteris, 408, 409, 411 diffusa, 409 filicina, 409 fruticulosa, 409 galapagana, 409 liebmaniana, 409 longispica, 409 tenuicaulis, 409 trinitensis, 409 C Calobryum, 39 andinum, 39 Calypogeia, 152–158 afrocoerulea, 156 amazonica, 156 andicola, 153 biapiculata, 156 caespitosa, 158 colombica, 153 crenulata, 159 cyclostipa, 159 densifolia, 155 euthemona, 136 granatensis, 144 grandistipula, 155 heterophylla, 156 laxa, 155 lechleri, 156 lechleri var. densifolia, 155 mastigophora, 159 miquelii, 156 muscicola, 156 nephrostipa, 159 oblata, 156 parallelogramma, 160 peruviana, 156 puiggarii, 155 rhombifolia, 157 rhombifolia var. colombiana, 157 subg. Mnioloma, 158–160 subintegra, 157 tenax, 157 uncinulatula, 157 Calypogeiaceae, 152–160 Calyptrocolea, 137–139

aureomarginata, 138 carabayensis, 138 lindenbergiana, 139 pittieri, 139 Campanocolea, 260–264 fragmentissima, 263 Caudalejeunea, 411 harpaphylla, 411 harpaphylla fo. longiloba, 411 lehmanniana, 411 Cephalozia, 164 bicuspidata, 164 bischlerae, 164 caribbeania, 165 colombica, 164 crassifolia, 165 crossii, 164 dussii, 164 granatensis, 174 grandifolia, 164 infuscata, 166 patagonica, 165 pleniceps, 165 subforficata, 165 subg. Macrocephalozia, 165 venezuelana, 164 Cephaloziaceae, 160–172 Cephaloziella, 173–176 andina, 174 antillana, 176 divaricata, 174 fragillima, 174 granatensis, 174 hampeana, 176 pygmaea, 174 Cephaloziellaceae, 172–179 Cephaloziopsis, 176, 177 intertexta, 176 schistophila, 176 Ceratolejeunea, 412–420 amazonica, 416 andicola, 415 belangeriana, 416 brasiliensis, 417 brevinervis, 414 caducifolia, 418 calabariensis, 415, 416 ceratantha, 414 coarina, 414, 415 commixta, 416 confusa, 415 corniculata, 417 cornuta, 415 cubensis, 416

Index dentatocornuta, 585 diversicornua, 414 dussiana, 585 ecuadoriensis, 416 fallax, 417 fastigiata, 417 filaria, 417 globulifera, 417 granatensis, 416 grandiloba, 417 guianensis, 418 hylophila, 415 laetefusca, 418 litoralis, 415 maritima, 415 mascarena, 416 megalophysa, 415 microrhegma, 415 monticola, 414 oxygonia, 418 parisii, 416 panamensis, 418 patentissima, 420 platyceras, 420 plumula, 414 pungens, 420 rubiginosa, 420 sacculosa, 417 spinosa, 420 tenuicornuta, 416 valida, 415 variabilis, 415 wallisii, 420 Ceratolejeunea subg. Ceratolejeunea, 412 Ceratolejeunea subg. Ceratophora, 412 Chaetocolea, 322 palmata, 322 Cheilolejeunea, 421–440 acutangula, 425, 426 adnata, 426 aneogyna, 427 asperiflora, 440 asperrima, 427 aurifera, 427 beyrichii, 428 choachina, 428 compacta, 431 curvatiloba, 440 decidua, 426 decurviloba, 594 discoidea, 425, 440 duriuscula, 437 filiformis, 429, 430 filiformis var. platycoleus, 429

703 filiformis var. wallisii, 429, 430 fragrantissima, 430 holostipa, 430, 431 inflexa, 431 intertexta, 431 invaginata, 428 jackii, 431, 432 laevicalyx, 432 lineata, 432, 434 lobulata, 434 myriantha, 440 neblinensis, 434, 435 norisiae, 423 oncophylla, 434 ornata, 435 ovalis, 435 paramicola, 436 polyantha, 437 quinquecarinata, 436 revoluta, 436, 437 rigidula, 437 rupestris, 428 schiavoneana, 437, 438 serpentina, 437 tonduzana, 438 trifaria, 438 unciloba, 439 urubuensis, 439 valenciae, 424 xanthocarpa, 439, 440 Chiloscyphus bomboizanus, 249 purpureus, 255 quadridentatus, 258 Clasmatocolea, 244, 245 fragillima, 257 heterostipa, 245 vermicularis, 245 Clevea, 46, 47 andina, 47 spathysii, 47 Cleveaceae, 46–47 Cololejeunea, 440–455 antillana, 445 azorica, 445 bekkerae, 445 bischleriana, 445, 446 camillii, 446 cardiocarpa, 446 cingens, 446, 447 clavatopapillata, 442 conchifolia, 455 contractiloba, 442 cornuta, 453

Index

704 Cololejeunea (cont.) cremersii, 455 crenata, 447 dauphinii, 579 diaphana, 447 ecuadorensis, 447 ensifolia, 446 ensifolia var. pygmaea, 447 ephemeroides, 450 fefeana, 455 gracilis, 449 hildebrandii, 449 jamesii, 442 kuciana, 449 lanceolata, 446 lanciloba, 452 latilobula, 449, 450 linopteroides, 450 manaosensis, 452 megalandra, 450 microscopica, 450 minuscula, 451 minutissima, 578, 579 obliqua, 451 ochyrana Pócs, 457 papilliloba, 451 papillosa, 451 planifolia, 452 planissima, 452 planiuscula, 452 platyneura, 452 pterocolea, 452, 453 scabrifolia, 451 sicifolia, 453 sicifolia subsp. jamaicensis, 453 sintenisii, 450 stotleriana, 453 subcardiocarpa, 454 subscariosa, 452 subsphaeroidea, 454 surinamensis, 454 tixieri, 579 truncatifolia, 447 turbinifera, 578 verwimpii, 454 vitaliana, 455 winkleri, 455 yanoanae, 454 Colura, 455–462 andoi, 457, 458 calyptrifolia, 458 clavigera, 461 cylindrica, 458 greig-smithii, 458

herzogii, 462 irrorata, 458, 460 kilimanjarica, 460 lyrata, 461 naumannii, 460 ornithocephala, 460, 461 patagonica, 460 sagittistipula, 461 stotleri, 461 tenuicornis, 461 tortifolia, 461, 462 ulei, 462 Colurolejeunea, 455 Corsiniaceae, 48 Cronisia, 48 weddellii, 48 Crossotolejeunea, 502 angulata, 524 bogotensis, 512 caulicalyx, 532 cavifolia, 534 crenata, 613 cristatella, 512 inflexiloba, 534 intricata, 534 lindeniana, 524 pustuloso-alata, 534 Cryptochila, 191 grandiflora, 199 Cryptogynolejeunea, 502 reflexistipula, 545 Cryptolophocolea, 245, 246 connata, 246 guadalupensis, 246 martiana var. martiana, 247 martiana var. perissodonta, 247 Cyathodiaceae, 48–51 Cyathodium, 48, 50, 51 cavernarum, 50 foetidissimum, 50, 51 Cyclolejeunea, 462–466 accedens, 464 affixa, 464 angulata, 465 angulistipa, 465 chitonia, 464 convexistipa, 464 ecuadorensis, 464 foliorum, 465 glimeana, 465 grandistipula, 464 luteola, 465 ocellata, 466 papillata, 464

Index peruviana, 466 subintegerrima, 464 triocellata, 464 truncatulata, 464 Cylindrocolea, 177 obtusifolia, 177 planifolia, 177 rhizantha, 177 Cyrtolejeunea, 421 holostipa, 430 Cystolejeunea, 421 lineata, 432 D Dendroceros, 645, 646 allionii, 646 breutelii, 646 crispatus, 646 crispatus var. simplicior, 646 crispus, 646 ecuadorensis, 646 foliatus, 646 jamesonii, 646 Dendrocerotaceae, 641 Dendrolejeunea, 607 Dibrachiella, 466, 467, 469 auberiana, 467 bischleriana, 467 parviflora, 467, 469 Dicladolejeunea, 502 anomala, 547 reflexistipula, 545 rotundifolia, 547 saccatiloba, 547 Dicranolejeunea, 469, 470 axillaris, 470 ciliatiflora, 470 cipaconea, 567 circinnata, 567 grossiloba, 583 hypoacantha, 470 loxensis, 396 pulcherrima, 470 renauldii, 396 Diplasiolejeunea, 470–478 alata, 472, 473 armatiloba, 478 brachyclada, 473 brunnea, 473 caribea, 475 cavifolia, 473 columbica, 473, 474 erostrata, 474

705 galloana, 473 grandirostrata, 474 involuta subsp. andicola, 474 johnsonii, 474 lanceolata, 474, 475 lanciloba, 473 mayaykuensis, 475 papilionacea, 475 pauckertii, 475 pellucida, 475 pluridentata, 476 pocsii, 476 reflexiloba, 475 replicata, 476 rudolphiana, 476, 478 Diplophyllum, 329, 330 obtusatum, 330 Drepanolejeunea, 478–491 andina, 481, 482 anoplantha, 482 araurcariae, 482, 483 aurita, 483 bidens, 483 biocellata, 483, 484 campanulata, 484 capitulata, 485 crassiretis, 484, 485 crucianella, 485 cutervoensis, 485 evansii, 486 fragilis, 486 granatensis, 486 grandistipula, 481 hamulata, 488 hamulata var. espinulosa, 488 inchoata, 486, 487 inchoata var. biocellata, 486 inchoata var. roraimae, 486 infundibulata, 486, 487 integribracteata, 484 lancifolia, 492 lichenicola, 487 longirostris, 492 mosenii, 487, 488 navicularis, 485 orthophylla, 488, 489 orthophylla subsp. saxicola, 489 palmifolia, 490 polyrhiza, 490 pterocalyx, 490 pungens, 492 ramentiflora, 490, 491 sabaliana, 487 spinosa, 491

Index

706 Drepanolejeunea (cont.) squarrosula, 483 subdissitifolia, 492 submuricata, 484 subvittata, 491 urceolata, 491 Dumortiera, 51, 52 hirsuta, 51 hirsuta var. irrigua, 51 hirsuta var. trichopus, 51 Dumortieraceae, 51–52 E Echinocolea, 502 asperrima, 516 prionocalyx, 557 subspathulata, 552 Eopleurozia, 122, 123 paradoxa, 123 Eulejeunea, 502 monimiae, 551 Euosmolejeunea, 421 beyrichii, 428 duriuscula, 437 fragrantissima, 430 fragrantissima var. columbica, 430 laxiuscula, 438 neglecta, 437 opaca, 438 F Fimbriaria, 43 bakeri, 44 canelensis, 44 elegans, 44 macropoda, 44 Fossombronia, 70–77 brasiliensis, 76 columbica, 73 crassifolia, 73 fernandeziensis, 73 herzogii, 75 jostii, 73 lamellata, 75 paranapanema, 77 peruviana, 75 porphyrorhiza, 76 ptychophylla, 75 pusilla, 76 tuberifera, 75 Fossombroniaceae, 70–77 Frullania, 350

aculeata, 358 acuminata, 358 albertii, 380 ambronnii, 358 apiculata, 380, 381 apollinarii, 372 arecae, 375 arietina, 378 armata, 383 arnellii, 380 arsenii, 380 atrata, 358, 360, 377 atrata β subtilissima, 380 atrosanguinea, 377 bakeri, 377 beyrichiana, 360 beyrichii, 360 bicornistipula, 361 blepharozia, 361 bogotensis, 361 bomboizana, 362 brachyclada, 382 brasiliensis, 361 breuteliana, 360 bullatiflora, 372 campanensis, 375 caroliniana, 374 clandestina, 362 closterantha, 369 coalita, 360 columbica, 361 compsotera, 374 confertiloba, 362 confertiramea, 380 convoluta, 362, 364 crenulifolia, 380 crispiloba, 383 cuencensis, 364 cylindrica, 361 darwinii, 364 decidua, 364 dulimensis, 365 dusenii, 365 ecklonii, 365 ericoides, 365, 366 exilis, 380 flammea, 364 flexicaulis, 375 formosa, 366 galapagona, 374 gibbosa, 367 glomerata, 367 gradsteinii, 367 granatensis, 361

Index grandifolia, 369 guadalupensis, 361 gymnotis, 374 haeckeriana, 377 haematocysta, 369 harpantha, 358 heinrichsii, 353 hians, 375 hirtiflora, 367 holostipula, 354 intumescens, 369, 370 intumescens var. sabanetica comb. nov., 370 involuta, 382 jelskii, 370 kunzei, 371 latilobula, 378 laxiflora, 371 laxiflora var. crossii, 371 lechleri, 358 leptomitra, 378 lindenbergii, 371, 372 lobatohastata, 372 longipinna, 362 macrocephala, 372, 373 mathanii, 377 megalostipa, 379 meridana, 373 mesophila, 367 mirabilis, 373 montagnei, 373, 374 moritziana, 383 mucronata, 361, 372 neesii, 370 nodulosa, 374 obcordata, 374 obscura, 375 ocanniensis, 366 pachoensis, 369 paraensis, 380 parasitica, 370, 374 pendulostyla, 373 peruviana, 377 phalangiflora, 377 pittieri, 377 planifolia, 377, 378 pluricarinata, 378 remotifolia, 377 repanda, 372 replicata, 374 ringens, 378 riojaneirensis, 378, 379 rio-janeirensis var. megalostipa comb. nov., 379

707 robusta, 361 semiconnata, 381 serrata, 383 setigera, 379 sphaerocephala, 379 spruceana, 361 squamulifera, 367 squarrosa, 366 standaertii, 380 stenostipa, 380 strobilantha, 372 subaculeata, 358 subalpina, 377 subg. Chonanthelia, 353–355 subg. Diastaloba, 353 subg. Meteoriopsis sect. Intumescentes, 356–358 subg. Meteoriopsis sect. Meteoriopsis, 355, 356 subtilissima, 380, 381 tamarisci, 383 taxodiocola, 380 tenera, 378 tenuis, 361 tetraptera, 381 tolimana, 367 transfalcicula, 365 trollii, 375 tubiflora, 372 tunguraguana, 382 uleana, 382 vaginata, 383 vanderhammenii, 382 villosa, 372 vimicola, 365 winteri var. vanderhammenii, 382 Frullaniaceae, 350–383 Frullanoides corticalis, 493 densifolia subsp. densifolia, 493, 495 densifolia subsp. grandidentata, 495 laciniatiflora, 495 liebmaniana, 495, 496 tristis, 496 Fulfordianthus, 496 pterobryoides, 496, 497 Funicularia bischlerana, 48 Fuscocephaloziopsis, 165 biloba, 166 crassifolia, 165 pleniceps, 165 pulvinata, 166 subintegra, 166, 167

Index

708 G Gongylanthus, 143–145 granatensis, 144 innovans, 145 liebmanianus, 144 limbatus, 145 muelleri, 145 oniscoides, 144 Gymnocoleopsis, 177, 178 cylindriformis, 178 multiflora, 178 Gymnomitriaceae, 179–183 Gymnomitrion, 180, 181 andinum, 181 atrofilum, 181 bolivianum, 181 setaceum, 181 truncatoapiculatum, 181 H Haplolejeunea amazonica, 394 Haplomitriaceae, 39–42 Haplomitriopsida, 38, 39 Haplomitrium, 39 blumei, 39 Haplozia carnosa, 144 cucculata, 144 limbata, 145 rorida, 201 Harpalejeunea, 497–502 acanthogona, 512 acuta, 498 ancistrodes, 498 aspera, 500 buenaventurae, 499 cinchonae, 498, 499 cinchonae var. strigulosa, 498 diaphana, 557 galeata, 531 granatensis, 486 grandis, 499 grandistipula, 498 harpaphylla, 502 heterodonta, 500 longilobula, 420 oxyphylla, 499, 500 pellucida, 544 pilifera, 499 puelensis, 425 scabra, 500 stricta, 500 subvittata, 491 tridens, 500, 502 tuberculata, 498

verrucosa, 434 Herbertaceae, 183–191 Herbertus, 184, 185 acanthelius, 185 angustevittatus, 185 bivittatus, 185 colombianus, 187 commutatus, 187 divergens, 185 grandifolius, 187 grossispinus, 187 inaequalis, 185 juniperoideus, 187 juniperoideus subsp. acanthelius, 185 juniperoideus subsp. bivittatus, 185 juniperoideus subsp. pensilis, 187 limbatus, 185 obtusifolius, 187 pensilis, 187 sendtneri, 187, 188 serratus, 185 subdentatus, 187 subnivalis, 187 Heterogemma, 337 capitata, 337 patagonica, 337 Heteroscyphus, 247–251 arnellii, 258 contortuplicatus, 248 marginatus, 249 polyblepharis, 249 polychaetus, 249 thraustus, 251 Homalolejeunea, 569 Hornworts, 640–656 Hygrolejeunea, 502 fulva, 528 grossereticulata, 558 obtusiserrulata, 562 osculatiana, 538 pallida, 530 paramicola, 526 pastasensis, 561 planiloba, 561 punctata, 561 sullivantii, 562 Hymenophyton, 70 I Isopaches, 330, 331 bicrenatus, 330 Isotachis, 146–150 allionii, 150 aubertii, 149 ecuadorensis, 149

Index haematodes, 149 lacustris, 149 lindigiana, 149 lopezii, 147 madida, 149 mascula, 149 obtusa, 149 parva, 147 piliflora, 149 ripensis, 189 serrulata, 149 tenax, 147 woronowii, 149 J Jamesoniella, 191 autumnalis, 194 boliviana, 136 dependula, 289 paramicola, 200 rotundifolia, 198 rubricaulis, 198 undata, 200 Jamesoniellaceae, 191–200 Jensenia, 78–79 erythropus, 79 erythropus var. nobandae, 79 florschuetzii, 78, 79 spinosa, 79 Jubula, 383, 384 bogotensis, 384 hutchinsiae subsp. bogotensis, 384 pennsylvanica subsp. bogotensis, 384 Jubulaceae, 383–384 Jungermannia, 201, 203 acutangula, 425 amoena, 341 brachiata, 571 callithrix, 341 cordifolia var. longifolia, 201 crispata, 615 decolor, 341 filiformis, 429 heteracria, 151 hyalina, 342 laetevirens, 341 linguifolia, 341 ovatotrigona, 201, 203 picta, 341 sphaerocarpa, 342 Jungermanniaceae, 201–203 Jungermanniales, 124–130 Jungermanniidae, 123 Jungermanniopsida, 68, 69

709 K Kantia, 152, 158–160 Kingiolejeunea ornata, 559 Kurzia, 219–221 brasiliensis, 220 capillaris, 220 capillaris subsp. páramicola, 221 flagellifera, 221 Kymatocalyx, 178, 179 cubensis, 178 dominicensis, 178 rhizomaticus, 179 stoloniferus, 178 L Leiomitra, 346–349 argentea, 349 elliotii, 347 flaccida, 347 paraphyllina, 347 robusta, 348 sphagnoides, 348 tomentosa, 348 Leioscyphus, 251–260 campanulata, 257 chamissonis, 245 ecuadorensis, 339 fragilifolius, 257 galipanus, 253 obcordatus, 256 schizostomus, 257 Leiosporoceros, 646, 647 dussii, 647 Leiosporocerotaceae, 641 Lejeunea, 502–558 acanthogona, 512, 513 adpressa, 513, 514 albida, 530 ambigua, 558 ancistrodes, 498 angusta, 514 anisophylla, 513 aphanes, 515, 516, 555 apiculata, 557 asthenica, 516, 517 autoica, 515 bermudiana, 517 biformis, 557 blepharogona, 512 bombonasensis, 517, 518 bongardiana, 571 boryana, 518 caespitosa, 513, 519 cancellata, 518, 519

Index

710 Lejeunea (cont.) capensis, 519, 520 capillaris, 576 caracensis, 558 caripensis, 520, 521 carolensis, 513 catinulifera, 521 cauapunensis, 513 caulicalyx, 532 cephalandra, 540 cerina, 521, 522 chimborazensis, 522, 523 cladiophora, 518 cladobola, 536 cladogyna, 558 comans, 428 concinnula, 523, 524 controversa, 524 corynantha, 524 crispistipula, 401 cuspidata, 561 cyathophora, 524 debilis, 526 deplanata, 526, 527 diaphana, 557 dictyocalyx, 557 dimorphophylla, 576 diversicuspis, 558 drehwaldii, 527 drymophila, 513 elliottii, 557 elongella, 528 filipes, 515 flaccida, 528, 529 flagellaris, 529, 530 flava, 530, 531 flavicans, 538 florida, 528 fulfordiae, 531 fulva, 528 fusagasugana, 557 galeata, 531 gaudichaudii, 530 geophila, 535 glaucophylla, 536 globosiflora, 537 granatensis, 530 hahnii, 558 hapalochroa, 583 hebetata, 526 herminieri, 533 herpestica, 399 heterocheila, 526 huctumalcensis, 585 hyalina, 557

hygrophila, 557 inflexilobula, 534 intricata, 534 inundata, 601 juruana, 554 killipii comb. nov., 535 kunthiana, 425 laeta, 535, 536 laetevirens, 536 lancifolia, 492 languida, 573 lepida, 536 leptalea, 529 leucophaea, 530 longifissa, 513 longiuscula, 521 lucens, 536 lusoria, 528 macrorhycha, 557 magnoliae, 513 maxonii, 526 megalostipa, 526 meridensis, 536 minutiloba, 513 mojandae sp. nov., 537, 538 monimiae, 551 montana, 526 nemoralis, 438 obtusangula, 528 orbicularis, 513 osculatiana, 538 ovalis, 435 palaeflora, 573 pallescens, 538 pallida, 530 parviloba, 553 paucidentata, 506 paucispina, 524 paucispina var. rufescens, 524 pellucidissima, 614 pentotantha, 559 phyllobola, 540, 541 pililoba, 554 planiuscula, 486 plicata, 610 polyantha, 554 polycephala, 540 prionodes, 541, 542 prominula, 538 prorepens, 554 pterobryoides, 496 puiggariana, 542, 543 pulverulenta, 514, 515, 543, 544 quinqueumbonata, 544 raddiana, 544, 545

Index radicans, 526 ramulosa, 545 recurva, 509 recurvans, 401 reflexistipula, 545, 546 remotifolia, 558 rionegrensis, 509, 553 robusta, 573 rotundifolia, 547 ryszardii, 547 schultesii sp. nov., 548, 549 sect. Echinocolea, 503 semiscabrida, 557 seriata, 551 serpillifolioides, 550 setiloba, 550 silvatica, 530 smaragdina, 558 sporadica, 557 stachyclada, 464 subhyalina, 536 subrufula, 536 subsessilis, 551 subsimplex, 521 subspathulata, 552 suffruticola, 552 sulphurea, 552 surinamensis, 464 symphoreta, 530 tamasii, 511 tapajosensis, 546, 553 tarapotensis, 532 tarapotensis var. microstipa, 532 topoensis, 554 torticuspis, 562 trigonostipa, 566 trinitensis, 554, 555 ulicina subsp. bullata, 575 viridis, 518 xiphophylla, 555 yasuniensis sp. nov., 555 Lejeuneaceae, 384–614 Lepicolea, 203, 204 pruinosa, 203, 204 Lepicoleaceae, 203–204 Lepidolejeunea, 558–562 auriculata, 559 cordifissa, 559 cuspidata, 561 eluta, 561 grossepapulosa, 562 involuta, 562 ornata, 559 punctata, 561

711 spongia, 562 sullivantii, 562 Lepidozia, 221–228 alstonii, 223 andicola, 223 appendiculata, 223 armata, 226 auriculata, 223 bogotensis, 223 brasiliensis, 223 caespitosa, 223 columbica, 203 commutata, 226 cupressina, 223 dendritica, 225 heterophylla, 223 inaequalis, 226 incurvata, 226 karstenii var. standleyi, 226 lindigiana, 223 macrocolea, 226 moritziana, 225 muenchiana, 225 patens, 226 peruviensis, 225 pinnaticruris, 227 quitensis, 225 reptans, 227 serpens, 228 squarrosa, 227 subdichotoma, 228 tenuicola, 226 tenuicuspis, 226 tunguraguae, 225 wallisiana, 225 Lepidoziaceae, 204–242 Leptolejeunea, 562–566 convexistipa, 566 elliptica, 564 exocellata, 565 jamaicensis, 565 maculata, 565, 566 obfuscata, 565 obovata, 565 pterocalyx, 490 radicosa, 565 serratifolia, 566 stenophylla, 488 tridentata, 566 trigonostipa, 566 unguiculata, 488 Leptoscyphus, 251–260 amphibolius, 253 autoicus, 253, 254

Index

712 Leptoscyphus (cont.) cleefii, 254 cuneifolius subsp. cuneifolius, 254 cuneifolius subsp. fragilis, 254 gibbosus, 255 hexagonus, 255 intermedius, 255 jackii, 256 leoniae, 256 liebmanianus, 257 obcordatus, 256 physocalyx, 257 porphyrius, 257 porphyrius subsp. azoricus, 257 sotiauxii, 258 spectabilis, 258 trapezoïdes, 258, 260 Lethocolea, 135, 136 glossophylla, 136 repens, 136, 144 Leucolejeunea, 421 ecuadoriensis, 403 unciloba, 439 xanthocarpa, 439 Leucosarmentum, 236, 237 bifidum, 237 portoricense, 237 Lindigianthus, 566, 567 cipaconeus, 567 Lindigina granatensis, 144 Liverworts, 37, 38 Lobatiriccardia, 90 oberwinkleri, 90 verdoornioides, 90 Lophocolea, 260–264 aberrans, 262 bidentata, 262 blepharoptera, 262 brachydonta, 248 brasiliensis, 265 breutelii, 258 canelensis, 246 coadunata, 262 columbica, 258 connata, 246 contortuplicata, 248 cuspidata, 262 diaphana, 262 effusidens, 258 elata, 245 erosa, 264 foliicola, 265 fragmentissima, 263 granatensis, 246 guadalupensis, 246

hirta, 264 irrigata, 248 latifolia, 262 leptantha, 265 liebmanniana, 263 lindmannii, 265 longiflora, 262 mandonii, 262 martiana, 247 martiana complex, 245, 246 mascula, 262 muricata, 264 orbigniana, 258 pertusa, 247 polychaeta, 249 pycnorrhiza, 253 quadridentata, 258 sect. Connatae, 245, 246 subtrilobata, 262 tenerrima, 264 trapezoïdes, 258 widgrenii, 265 Lophocoleaceae, 243–266 Lopholejeunea, 567–568 andersonii, 568 cocosensis, 568 eulopha, 568 muelleriana, 568 nigricans, 568 quelchii, 568 sagreana, 568 saxatilis, 571 subfusca, 568 Lophonardia, 331 caespitosa, 332 jamesonii, 331, 332 laxifolia, 332 Lophozia bicrenata, 330 capitata, 337 debilis, 135 incisa, 335 laxifolia, 332 opacula, 332 patagonica, 337 stolonifera, 331 subg. Hypolophozia, 331 subg. Isopaches, 330 subg. Schistochilopsis, 335 subinflata, 332 Lophoziaceae, 324–338 Lunularia, 52 cruciata, 52 Lunulariaceae, 52 Luteolejeunea, 569 herzogii, 569

Index M Macrocolura, 455 sagittistipula, 461 Macrolejeunea, 502 cerina, 521 lancifolia, 538 pallescens, 538 subsimplex, 521 Madotheca, 614 cardoti, 615 columbica, 617 commutata var. lindigiana, 615 piligeri, 615 splendida, 615 Marchantia, 53–57 berteroana, 54 breviloba, 55 chenopoda, 55 columbica, 55 discoidea, 54 ecuadorensis, 55 hirsuta, 57 inflexa, 55 paleacea, 55, 57 papillata, 57 plicata, 57 polymorpha, 57 subandina, 57 Marchantiaceae, 53–57 Marchantiophyta, 37, 38 Marchantiopsida, 41, 42 Marchesinia, 569–573 aquatica, 571 bongardiana, 571 brachiata var. brachiata, 571, 573 brachiata var. robusta, 573 extensa, 401 galapagona, 571 guilleminiana, 571 languida, 573 longirostris, 571 longistipula, 403 nigrescens, 571 robusta, 573 Marsupella, 182 andina, 182 emarginata, 182 lacerata, 181 miniata, 182 pusilla, 327 subhyalina, 181 trollii, 181 truncatoapiculata, 181 Marsupidium, 130–135 Mastigobryum, 205

713 brevifolium, 211 columbicum, 215 Mastigolejeunea, 607 auriculata, 609 innovans, 609 pittieri, 609 plicatiflora, 610 Megaceros, 647 aenigmaticus, 648 callistictus, 653 colombianus, 650 vincentianus, 650 Metalejeunea, 573, 574 cucullata, 574 Metzgeria, 105–121 agnewiae, 108 albinea, 109 allionii, 113 atramentaria, 121 attenuata, 112 aurantiaca, 121 australis, 109, 117 bialata, 114 bischlerae, 109 bracteata, 120 bracteata var. robusta, 120 chilensis, 111 ciliata, 111 claviflora, 111 cleefii, 112 conjugata, 112 consanguinea, 112 convoluta, 121 crassipilis, 113 crenata, 120 crenatiformis, 120 cylindra, 113 decipiens, 111 dorsipara, 113 ecuadorensis, 117 filicina, 114 fruticola, 114 fruticulosa, 121 furcata, 115 furcata var. decipiens, 111 furcata var. extensa, 117 gigantea, 111 gracilimascula, 120 grandiflora, 115 hamata, 117 herminieri, 120 heteroramea, 121 holzii, 115, 116 hygrophila, 111 inflata, 116

Index

714 Metzgeria (cont.) jackii, 121 jamesonii, 116 lechleri, 116 leptomitra, 118 leptomitra var. angustifrons, 118 leptomitra var. longifurca, 118 leptoneura, 117 liebmaniana, 117 lindbergii, 117, 118 metaensis, 113 mexicana, 118 minor var. cyanogutta, 112 myriopoda, 118, 119 neotropica, 112 papulosa, 117 parviinvolucrata, 119 pilosa, 117 polytricha, 119 procera, 119 procera var. columbica, 119 pulvinata, 120 rufula, 120 saccata, 115, 116 sandei, 117 saxbyi, 117, 118 scyphigera, 120 temperata, 112 thomeensis, 109 tonsa, 111 uleana, 114 uncigera, 121 violacea, 121 violacea var. dorsipara, 113 vivipara, 116 wallisiana, 117 Metzgeriaceae, 105–122 Metzgeriidae, 86, 87 Microlejeunea, 574–577 acutifolia, 575 aphanella, 612 bullata, 575 capillaris, 576 cardotii, 577 colombiana, 576 concinnula, 523 diversiloba, 576 epiphylla, 576 globosa, 577 grandistipula, 558 laetevirens, 536 monoica, 574 perpusilla, 577 stricta, 576 triangulata, 575

Microlepidozia, 219 Micropterygium, 228–232 campanense, 229 carinatum, 229 duidae, 230 exalatum, 229 lechleri, 230 leiophyllum, 230 longicellulatum, 230 parvistipulum, 231 pterygophyllum, 231 pterygophyllum var. gracile, 231 reimersianum, 231 trachyphyllum, 231 Mnioloma, 158–160 caespitosum, 158 cellulosum, 159 crenulatum, 159 cyclostipum, 159 nephrostipum, 159 parallelogrammum, 160 venezuelanum, 159 Monoclea, 58 gottschei subsp. australis, 58 gottschei subsp. elongata, 58, 59 gottschei subsp. gottschei, 58, 59 Monocleaceae, 58–59 Monodactylopsis, 232 minima, 232 monodactyla, 232 Mylia fragilis, 254 minuta, 257 quitoensis, 257 Myriocolea, 455 irrorata, 458 Myriocoleopsis, 577–579 gymnocolea, 578 minutissima, 578, 579 riparia, 578 sect. Myriocoleopsis, 578 sect. Protocolea, 578 tixieri comb. nov., 579 Mytilopsis, 232, 234 albifrons, 234 N Nanomarsupella, 183 xenophylla, 183 Nardia, 338, 339 succulenta, 339 tortistipula, 339 Neesioscyphus, 150–151 allionii, 150

Index argillaceus, 151 carneus, 151 Neopotamolejeunea, 502 Neoprasanthus, 135, 136 granatensis, 136 Neurolejeunea, 580–581 breutelii, 580, 581 sastreana, 581 sect. Aneurolejeunea, 580 seminervis, 580 Noteroclada, 85–87 confluens, 86 leucorhiza, 86 Nothoceros, 647, 648, 650 aenigmaticus, 648 renzagliensis, 648 vincentianus, 650 Notoscyphus carneus, 151 lindmannii, 245 Notothyladaceae, 641 Notothylas, 650–652 amazonica, 651 breutelii, 651 galapagensis, 651 orbicularis, 651 vitalii, 652 Nowellia, 167–168 caribbeania, 168 curvifolia, 167 evansii, 168 O Odontolejeunea, 581–584 calcarata, 583 chaerophylla, 583 decemdentata, 583 ecuadoriensis, 583 grandiloba, 584 longispica, 583 lunulata, 583, 584 nigrescens, 583 obversilobula, 584 peruviana, 466 rhomalea, 584 sieberiana, 583 spiniloba, 583 Odontoschisma, 168–172 atropurpureum, 172 bifidum, 168 cleefii, 169 cordifolium, 171 denudatum, 171 denudatum var. andinum, 172

715 engelii, 171 falcifolium, 172 jishibae, 168 longiflorum, 171 portoricense, 171 soratamum, 172 sphagni var. amazonicum, 172 steyermarkii, 168 variabile, 172 Olgantha, 188 eophylla, 189 Omphalanthus, 421 apiculatus, 557 cuervi, 557 filiformis, 429 filiformis var. laxus, 429 geminiflorus, 429 grandistipulus, 429 jackii, 431 ovalis, 435 platycoleus, 429 wallisii, 430 Omphalolejeunea, 421 Oryzolejeunea, 502 antillana, 533 saccatiloba, 533 Otigoniolejeunea, 584, 585 acanthotis, 585 huctumalcensis, 585 quinqueumbonata, 544 reducta, 532 schizotis, 585 Otolejeunea schnellii, 399 subg. Allorgella, 398 P Pallavicinia, 79, 80 lyellii, 80 Pallavicinia sp., 80 Paracromastigum, 236, 237 bifidum, 237 granatense, 236 pachyrhizum, 237 Paramomitrion paradoxum, 126 Pelliaceae, 85–87 Pelliidae, 69, 70 Peltolejeunea, 421 acuminata, 470 galapagona, 470 jackii, 431 rotundistipula, 435 Perilejeunea granatensis, 561

716 Phaeoceros, 652–654 carolinianus, 653 dussii, 647 fimbriatus, 654 gualaquizanus, 653 laevis, 653 laevis subsp. carolinianus, 653 pichinchensis, 653 squamuligerus, 656 tenuis, 654 Phaeomegaceros, 654, 656 fimbriatus, 654 squamuliger, 656 squamuligerus, 656 Phycolepidoziaceae, 172–176 Physotheca, 251–260 autoica, 253 Physotium paradoxum, 123 Pictolejeunea, 586 picta, 586 sprucei, 586 Plagiochasma, 45, 46 rupestre, 45 Plagiochila, 266–319 abscedens, 302 acanthoda, 314 adiantoides, 277 aerea, 277, 278 aliena, 311 alpina, 277 alternans, 278, 279 amazonica, 315 amicta, 279 anderssonii, 311 andicola, 288 apicidens, 320 arcta, 319 arcuata, 279, 280 arnelliana, 280 arrecta, 280 asperifolia Steph. var. columbica, 314 aspleniformis, 320 atrovirens, 280 azuayensis, 294 bancroftii, 300 barutana, 307 beskeana, 307 bicaudata, 288 biceps, 280 bicuspidata, 280 bidens, 317 bifaria, 280, 282, 283 binominis, 300

Index blepharobasis, 289 bogotensis, 304 boissieri, 300 breuteliana, 283 breuteliana var. acutifolia, 283 breuteliana var. novo-granatensis, 283 bryhnii, 283, 284 bryopterioides, 288 buchtiniana, 284 bursata, 277 bursata var. andina, 277 caespiticia, 306 calomelanos, 278 canelensis, 284 centrifuga, 280 chimborazensis, 287 chinantlana, 310 choachina, 310 cipaconensis, 301 cleefii, 285 cobana, 313 cobana fo. linearis, 277 columbica, 278 compressula, 280 confundens, 313 connivens, 311 conspicua, 316 contigua, 301 contingens, 303 cordilliera, 305 corniculata, 295 crispabilis, 320 crispatodecurrens, 311 cristata, 285 cuatrecasii, 286 cucullifolia var. anomala, 286 cucullifolia var. cucullifolia, 286 cuervina, 305 cultrifolia, 316 cuneata, 287 cuneata var. loriloba, 302 deflexa, 287, 288 deflexirama, 288, 289 demissa, 298 densa, 318 densispina, 277 dependula, 289 depressa, 319 dichotoma, 307, 320 diffusa, 292 dilatata, 288 dimorpha var. ecuadorica, 293 discreta, 307 disticha, 289, 291, 292

Index distinctifolia, 292 divaricata, 313 diversifolia, 292 diversispina, 319 dominicensis, 293 dotensis, 292 douinii, 279 dubia, 307 echinella, 280 ecuadorensis, 287 ecuadorica, 293, 294 eggersii, 294 elegantula, 314 ensiformis, 294, 295 erronea, 313 esmeraldana, 277 exesa, 280 exigua, 295, 296 eximia, 277 falcatoserrata, 291 fallax, 307 fastigiata, 296 fendleri, 313 filicaulis, 310 filicina, 288 fimbristipula, 291 flabellifrons, 301 flaccida, 303 flavescens fo. ampliata, 319 florida, 304 fragilis, 282 friabilis, 284 frontinensis, 316 funckiana, 311 fuscolutea, 296, 298 galapagona, 292 geniculata, 282 glaucescens, 292 glomerulifera, 277 gradsteinii, 282 granatensis, 318 grandicrista, 298 grandifolia, 278 grateloupii, 299 gualaquizana, 311 guevarii, 298 guilleminiana, 311 gymnocalycina, 298, 299 gymnocalycina var. surinamensis, 299 gymnostoma, 296 hans-meyeri, 318 harlingii, 315 heteromalla, 299, 300 heteromalla var. densifolia, 300

717 heteromalla var. latifolia Spruce, 300 heterophylla, 302 homochroma, 305 horrida, 282 hortorum, 291 humboldtiana, 319 husnotii, 300 hylacoetis, 319 hypantra, 292 hypnoides, 304 hystrix, 282 impluviata, 301 increscentifolia, 282 inouei, 280 insularis, 319 interjecta, 307 intricata, 307 irmscheri, 320 jamesonii, 294 jamesonii var. secundifolia, 303 jaramilloi, 282 jelskii, 296 jovoensis, 320 juruensis, 291 keckiana, 277 kegeliana, 315 laetevirens, 300, 301 leptodictyon, 315 leptophylla, 303 lindigiana, 292 linearis, 313 lingua, 312 longaeva, 319 longiramea, 320 longispina, 301 loriloba, 302 ludoviciana, 311 macra, 302, 303 macrifolia, 303 macrifolia var. taylorii, 303 macrostachya, 303 macvicarii, 277 magellanica, 139 martiana, 307 matanga, 301 meridana, 303 meridana var. funkii, 303 micropteryx, 301 minutidens, 305 miqueliana, 292 montagnei, 304 montana, 307 multispina, 305 mutila, 287

Index

718 Plagiochila (cont.) notidophila, 277 nova, 304 nudiuscula, 280 oblita, 278 orbigniana, 291 oresitropha, 299 ovata, 305 ovato-obconica, 318 ovifolia, 299 oxyphylla, 314 pachoensis, 301 pachyloma, 305, 306 paludosa, 299 panamensis, 282 papillifolia, 306 paramicola, 310 paraphyllina, 306 pastasensis, 301 patentissima, 320 patriciae, 307 patula, 307, 309 patzschkei, 310 paucispinula, 283 pautaphila, 309 pendula, 289 pensilis, 291 perbella, 311 permista, 318 perrottetiana, 313 pichinchensis, 316 pichinchensis var. basidentata, 293 pinnata, 282 pinnatidens, 282 pinnatispina, 294 plicata, 311 polopolensis, 299 potrerillana, 296 praetermissa, 292 priceana, 310 procera, 277 pseudopatula, 301 punctata, 310, 311 pungens, 277 quitensis, 277 raddiana, 311, 312 rara, 287 reclinata, 283 regeliana, 313 relicta, 288 revolvens, 312 rhizophila, 311 rigidula, 307 rosariensis, 278

rudischusteri, 312 rufoviridis, 320 rutilans, 313 rutilans fo. foliicola, 320 rutilans var. aequatorialis, 277 rutilans var. angustifolia, 299 sacramenti, 303 saettonii, 278 saladona, 277 saltuensis, 318 scabrifolia, 317 schlimiana, 311 scopulosa, 296 semidentata, 278 serrata, 291 silvatica, 311 simplex, 313, 314 soratensis, 305 spinifera, 282 sprucei, 320 stolonifera, 310 stricta, 314 subaequalis, 279 subcristata, 311 subdeflexiramea, 288 subedentata, 292 subplana, 315 subplana var. scoposa, 315 subrara, 310 subrotundifolia, 282 subsimplex, 311 subtrinitensis, 287 subundulata, 315 superba, 316 sylvicultrix, 299 tabinensis, 316, 317 tamariscina, 317 tenuis, 317 thyoides, 320 thysanotis, 291 tocarema, 301 tovarina var. trianae, 287 trianae, 288 triangulifolia, 296 trichostoma, 318 tridenticulata, 295 trifida, 282 trollii, 288 truncatella, 307 tunguraguensis, 302 turgida, 319 umbrosa, 313 undulifolia, 311 vincentina, 319

Index virens, 293 wallisiana, 305 wolfii, 299 wolframii, 296 xanthochroma, 307 zygophylla, 303 Plagiochilaceae, 266–320 Plagiochilion, 266 bryhnii, 283 heteromallum, 299 reclinata, 283 Platycaulis, 265, 266 renifolia, 265 Platylejeunea, 603 Plectocolea subamoena, 341 Pleurozia, 122, 123 heterophylla, 123 paradoxa, 123 Pleuroziaceae, 122–123 Pleuroziidae, 121 Porella, 614–618 acanthota, 615 arborea, 615 brachiata, 615 brasiliensis, 615 crispata, 615, 617 expansa, 617 leiboldii, 617 muelleri, 617 ptilopsis, 617 reflexa, 618 rotundifolia, 617 rugulosa, 618 squamulifera, 617 swartziana, 617 Porellaceae, 614–618 Porellales, 349–350 Potamolejeunea, 421 riparia, 427 sprucei, 601 Prionocolea, 502 viridissima, 518 Prionolejeunea, 587–592 aemula, 588, 589 ampliretis, 589 arguta, 589 circinulata, 590 cuatrecasii, 586 decora, 589 denticulata, 589, 590 fabroniaefolia, 592 gemmata, 588 helleri, 589

719 leptocardia, 464 luxurians, 464 macrocardia, 589 magnistipula, 590 microdonta, 589 mucronata, 590 muricatoserrulata, 590 prionodes, 588 recurvula, 590 scaberula, 592 schlimiana, 592 trachyodes, 592 Pseudocephalozia, 237 quadriloba, 237 Pseudolepicoleaceae, 320–324 Pseudomarsupidium, 141, 142 aureocinctum, 141 decipiens, 141 Pteropsiella, 238 frondifrons, 238 metzgeriiformis, 238 serrulata, 238 Ptychanthus theobromae, 602 Ptychocoleus polycarpus, 600 Pycnolejeunea, 592–595 callosa, 595 chocoensis, 593 contigua, 594 decurviloba, 594 densistipula, 594 flagellifera, 598 granatensis, 561 hyalina, 557 macroloba, 595 revoluta, 436 spruceana, 594 surinamensis, 427 R Radula, 618–640 aguirrei, 623 amazonica, 630 andicola, 634 angulata, 623 antilleana, 627 aurantii, 637 bischlerae sp. nov., 623, 626 bogotensis, 623 caldana, 630 cornucopiae, 627 cubensis, 627

Index

720 Radula (cont.) decora, 627 eggersiana, 630 eggersii, 627 episcia, 627, 628, 636 episcia var. opaciuscula, 628 falcifolia, 630 fendleri, 628 flaccida, 628, 629 floridana, 630 frondescens, 639 galapagona, 629 glauca, 634 goebelii, 631 gottscheana, 629, 630 gracilis, 630 husnotii, 640 inflata, 627 involvens, 630 jamesonii, 630 javanica, 630, 631 kegelii, 633 korthalsii, 623 lewisii, 631 lindigii, 631 longifolia, 630 macrostachya, 630 mammosa, 631 mazarunensis, 634 mexicana, 631 mollis, 634 montana, 636 nudicaulis, 631, 633 obtusifolia, 636 pallens, 633, 634 plumosa, 640 punctata, 640 pusilla, 634 quadrata, 634, 635 ramulina, 638 ramulina var. microphylla, 638 retroflexa, 640 riparia, 639 saccatiloba, 635 santacruziana, 635 schaefer-verwimpii, 635 sect. Dichotomae Castle, 628 sect. Epiphyllae, 619 sinuata, 636 sonsonensis, 636 sprucei, 630 stenocalyx, 637 subg. Cladoradula, 619 subg. Metaradula, 619

subg. Radula, 619 subg. Volutoradula, 619 subinflata, 637 surinamensis, 630 tectiloba, 637 tenella, 637 tenera, 638 viridiaurea, 634 voluta, 638, 639 xalapensis, 639 yanoella, 639, 640 Radulaceae, 618–640 Rectolejeunea, 595–598 berteroana, 598 brittoniae, 540 colombiana, 540 emarginuliflora, 596 flagelliformis, 596, 598 halinae, 598 heteroclada, 427 intermedia, 585 longiloba, 526 maxonii, 526 phyllobola, 540 versifolia, 598 Regredicaulis, 232 monodactylus, 232 serrus, 242 Reinerantha, 599 foliicola, 599 Rhaphidolejeunea polyrhiza, 490 Riccardia, 91–105 aberrans, 94 algoides, 102, 103 amazonica, 103 andina, 98 bogotensis, 98 boliviensis, 101 calcarea, 97, 98 capillacea, 101 capillacea var. dentata, 101 cataractarum, 96 cervicornis, 102, 103 chamedryfolia, 96, 97 ciliolata, 97 columbica, 99 crassa, 94 crassicaulis, 98 digitiloba, 97, 98 fucoidea, 98 gradsteinii, 92 hans-meyeri, 99 hans-meyeri var. dentata, 99

Index herzogiana, 101 hymenophytoides, 99 judithae, 104 latifrons subsp. parasitans, 100, 101 lepidomitra, 98 leptophylla, 101 metzgeriiformis, 97, 98 multifida, 105 pallida, 101 palmata, 105 papillata, 98 paramorum, 96 parasitans, 100, 101 pinguis, 88 pinnatifida, 96, 97 plumiformis, 98 poeppigiana, 102, 103 reginae, 104 regnellii, 103 sinuata, 96, 97 smaragdina, 104 sprucei, 99 tenuicula, 101 trichomanoides, 101 verticillata, 103 wallisii, 104 Riccia, 59–67 albopunctata, 61 amazonica, 65 atromarginata subsp. iodocheila, 64 australis, 61 crassifrons, 62 echinatispora, 65 howellii, 62 huebeneriana, 64 ianthina, 64 indusiata, 65 iodocheila, 64 lamellosa, 65 limicola, 65 membranacea, 65 planobiconvexa, 65, 66 ridleyi, 66 sorocarpa, 60 stenophylla, 62 vitalii, 66 weinonis, 66 Ricciaceae, 59–67 Ricciocarpos, 67 natans, 67 Ruizanthus, 151, 152 lopezii, 147 venezuelanus, 151

721 S Sauteria, 47 chilensis, 47 berteroana, 47 Scalia andina, 39 Scapania, 332, 334, 335 cuspiduligera, 334 geppii, 334 portoricensis, 334 portoricensis var. roraimensis, 334 splendida, 334 Scapaniaceae, 324–338 Schiffneriolejeunea, 599, 600 amazonica, 600 polycarpa, 600 Schisma, 184 columbiae, 187 granatense, 187 striolatum, 187 wallisianum, 185 Schistochilopsis, 335 incisa, 335 Schusterolejeunea, 601 andina, 453 inundata, 601 Sendtnera, 184 aequabilis, 203 Solenostoma, 339–342 amoena, 341 amplexifolium, 341 callithrix, 341 decolor, 341 hyalinum, 342 sphaerocarpa, 342 Solenostomataceae, 338–342 Sphaerolejeunea, 502 umbilicata, 527 Sphaerosporoceros granulatus, 643 Sphenolobus, 336, 337 austroamericanus, 336 minutus, 337 Spruceanthus, 601, 602 theobromae, 602 Steereochila, 266 ecuadorica, 293 Stenorrhipis, 178 Stephaniella, 343, 345 paraphyllina, 343 rostrata, 345 uncifolia, 345 Stephaniellaceae, 342–346

Index

722 Stephaniellidium, 345, 346 sleumeri, 346 Stictolejeunea, 602, 603 balfourii, 603 herzogii, 569 kunzeana, 603 squamata, 603 Strepsilejeunea, 421 acutangula, 425 choachina, 428 kunthiana, 425 laevicalyx, 432 Stylolejeunea pililoba, 554 Subfamily Lejeuneoideae, 386 Subfamily Ptychanthoideae, 386 Symbiezidium, 603–640 barbiflorum, 605, 606 dentatum, 606 hobsonianum, 607 pogonopterum, 605 setosum, 605 subrotundum, 606 taeniopsis, 607 transversale, 606, 607 transversale subsp. dentatum, 606 vincentinum, 607 Symphyogyna, 80–85 apiculispina, 82 aspera, 82 bogotensis, 84 brasiliensis, 82 brongniartii, 82 canaliculata, 82 circinata, 82 digitisquama, 81 hymenophyllum var. bogotensis, 84 hymenophyllum var. heterogena, 84 leptothelia, 84 marginata, 84 podophylla, 84 rubescens, 85 trivittata, 85 Symphyomitra glossophylla, 136 Syzygiella, 191–196, 198–200 anomala, 193, 194 autumnalis, 194 bilobata, 194 burghardtii, 194, 195 campanulata, 195 ciliata, 195 colombiana, 196

concreta, 195, 196 contigua, 196 geminifolia, 196 grollei, 195 integerrima, 196 liberata, 200 linguifolia, 198 manca, 196 oppositifolia, 198 pectiniformis, 196 perfoliata, 198 plagiochiloides, 196 renifolia, 195 riclefii, 195 rubricaulis, 198 setulosa, 198, 199 sonderi, 199 tonduzana, 199 trigonifolia, 200 uleana, 200 undata, 200 Szweykowskia, 266 cucullifolia, 286 T Targionia, 67, 68 hypophylla, 68 robusta, 47 stellaris, 68 Targioniaceae, 67–70 Taxilejeunea, 502 apiculata, 557 asperrima, 427 asthenica, 516, 517 auriculata, 547 bombonasensis, 517 caracasensis, 558 caracensis, 558 caripensis, 520 chimborazensis, 522 crebriflora, 517 cuervi, 557 debilis, 526 densiflora, 542 dictyocalyx, 557 dissitifolia, 558 florida, 528 galapagensis, 529 herzogiana, 558 implexa, 517 isocalycina, 550 killipii, 535

Index lancifolia, 538 leioscypha, 538, 540 lusoria, 517, 528 macroloba, 529 macrorhyncha, 557 obtusangula, 528 peruviana, 540 prominula, 538 pterigonia, 541, 542 renistipula, 552, 553 serpillifolioides, 550 speciosa, 522 steyermarkii, 543 sulphurea, 552 tenax, 517 tenuiplica, 530 xiphophylla, 555 Telaranea, 234, 236, 238–241 bicruris, 240 chaetophylla, 240 coactilis, 240 diacantha, 239 nematodes, 240 pecten, 240 rectangularis, 234 sejuncta, 240 Temnoma, 324 chaetophyllum, 324 Thysananthus, 607–610 amazonicus, 608, 609 auriculatus, 609 innovans, 609 mexicanus, 407 olivaceus, 409 plicatiflorus, 610 Thysanolejeunea, 607 Trachylejeunea, 421 aneogyna, 427 asperiflora, 440 decurviloba, 594 didrichsenii, 465 dominicensis, 434 inflexa, 431 pandurantha, 613 spruceana, 613 Triandrophyllum, 188, 189, 191 eophyllum, 189 maegdefrauii, 149 subtrifidum, 189, 191 subtrifidum var. trifidum, 189 Tribe Brachiolejeuneeae, 386

723 Tribe Lejeuneeae, 386 Tribe Symbiezidiae, 386 Trichocolea, 346 allionii, 348 argentea, 349 difficilis, 347 elliottii, 347 filicaulis, 349 flaccida, 347 floccosa, 348 herzogii, 347 paraphyllina, 347 robusta, 348 sprucei, 348 tomentosa, 348 uleana, 348 Trichocoleaceae, 346–349 Tritomaria, 337, 338 capitata, 337 Tuyamaella, 599 Tylimanthus, 130–135 azoricus, 133 bispinosus, 295 cuneifolius, 131 laxus, 133 madeirensis, 133 ruwenzorensis, 133 setaceo ciliatus, 133 V Verdoornianthus, 611 griffinii, 611 marsupiifolius, 611 Vitalianthus, 612 aphanellus, 612 urubuensis, 439 X Xylolejeunea, 612–614 crenata, 613, 614 pellucidissima comb. nov., 614 Z Zoopsidella, 241–242 dichotoma, 242 integrifolia, 242 macella, 242 serra, 242