Flora of the Venezuelan Guayana, Volume 5, [5] 0915279711, 9780915279715

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Flora of the Venezuelan Guayana

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

Flora of the Venezuelan Guayana VOLUME 5 ERIOCAULACEAE–LENTIBULARIACEAE

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

MISSOURI BOTANICAL GARDEN PRESS St. Louis

Copyright © 1999 by the Missouri Botanical Garden Press All rights reserved. ISBN 0-915279-71-1 Printed in U.S.A. Published on 14 July 1999 by Missouri Botanical Garden Press P.O. Box 299 St. Louis, Missouri 63166-0299, U.S.A.

Contents Preface and Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii Eriocaulaceae (Nancy Hensold) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Erythroxylaceae (Timothy Plowman and Paul E. Berry) . . . . . . . . . . . . . 59 Euphorbiaceae (Grady L. Webster, Paul E. Berry, W. Scott Armbruster, Hans-Joachim Esser, Lynn J. Gillespie, W. John Hayden, Geoffrey A. Levin, Ricardo de S. Secco, and Scott V. Heald) . . . . . . 72 Euphroniaceae (Julian A. Steyermark and Luis Marcano-Berti) . . . . . . 228 Fabaceae (Gerardo A. Aymard C., Nidia L. Cuello A., Paul E. Berry, Velva E. Rudd, Richard S. Cowan, Paul R. Fantz, Richard H. Maxwell, Charles H. Stirton, Hans-Helmut Poppendieck, Haroldo Cavalcante de Lima, Renée H. Fortunato, Basil Stergios, Nereida Xena de Enrich, David A. Neill, Celia Gil, and R. Toby Pennington) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231 Flacourtiaceae (Mark Olson, Paul E. Berry, and Gerardo A. Aymard C.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434 Gentianaceae (Lena Struwe, Paul J. M. Maas, Oliver Pihlar, and Victor A. Albert) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474 Gesneriaceae (Christian Feuillet and Julian A. Steyermark) . . . . . . . . . 542 Gnetaceae (Dennis W. Stevenson) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 573 Haemodoraceae (Paul J. M. Maas and Hiltje Maas) . . . . . . . . . . . . . . . . 576 Haloragaceae (Luther J. Raechal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 581 Heliconiaceae (Lennart L. Andersson) . . . . . . . . . . . . . . . . . . . . . . . . . . . 583 Hernandiaceae (James S. Miller and Paul E. Berry) . . . . . . . . . . . . . . . 592 Hippocrateaceae (Alberta M. W. Mennega and Jennifer P. Hedin) . . . . . 594 Hugoniaceae (Nelson Ramírez, Paul E. Berry, and Antony Jardim) . . . 618 Humiriaceae (José Cuatrecasas and Otto Huber) . . . . . . . . . . . . . . . . . . 623 Hydrocharitaceae (Robert R. Haynes and Lauritz B. Holm-Nielsen) . . . 641

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vi C ONTENTS

Hydrophyllaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Icacinaceae (Richard A. Howard and Rodrigo Duno de Stefano) . . . . . . Iridaceae (Peter Goldblatt and James E. Henrich) . . . . . . . . . . . . . . . . . Ixonanthaceae (Nelson Ramírez and Paul E. Berry) . . . . . . . . . . . . . . . . Juncaceae (Carlos A. Vargas) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Krameriaceae (Lois Brako) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lacistemataceae (Luther J. Raechal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamiaceae (Raymond M. Harley) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lauraceae (Henk van der Werff and Jens G. Rohwer) . . . . . . . . . . . . . . . Lecythidaceae (Scott A. Mori and Ghillean T. Prance) . . . . . . . . . . . . . . Lemnaceae (Lawrence J. Davenport) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lentibulariaceae (Peter G. Taylor) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

644 646 658 664 672 674 676 678 700 750 779 782

Appendix: List of new names published in this volume . . . . . . . . . . . . . 804 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 805

Preface and Acknowledgments This volume continues the taxonomic treatments of all native and naturalized vascular plant families known to occur in the Venezuelan Guayana. It includes 29 families of spermatophytes. The remaining families will be published in alphabetical order in four subsequent volumes. The Flora of the Venezuelan Guayana project has been strongly supported since its inception by the Missouri Botanical Garden and especially by its director, Peter H. Raven. The Herbario Nacional de Venezuela, now part of the Fundación Instituto Botánico de Venezuela, has steadily supported this project in many ways; it was there that Julian Steyermark conceived the idea of the flora and did most of his background research. This volume is based upon work supported by the National Science Foundation under Grants Nos. BSR-8717303, BSR-9045532, and BSR-9201044. The Foundation provides awards for research in the sciences and engineering. The awardee is wholly responsible for the conduct of such research and preparation of the results for publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the authors and do not necessarily reflect the views of the National Science Foundation. The Julian A. Steyermark Fund, established by the late Dr. Steyermark at the Missouri Botanical Garden, provided major funding for the flora. The Lewis B. and Dorothy Cullman Foundation provided funding for the Gentianaceae, in part. The Armand G. Erpf Fund provided financial support for part of the plant illustrations in this volume. Besides the illustrations by Bruno Manara, Bobbi Angell did the drawings of Aldina paulberryi (Fig. 225), Macrocarpaea marahuacae (Fig. 436), Potalia elegans (Fig. 441), P. maguireorum (Fig. 442), P. resinifera (Fig. 443), Rogersonanthus coccineus (Fig. 446), Symbolanthus aureus (Fig. 452), and S. rosmarinifolius (Fig. 453). Sheila M. Hayden did the drawings of Amanoa almerindae (Fig. 95), A. guianensis (Fig. 96), A. cupatensis (Fig. 94), A. oblongifolia (Fig. 93), and A. steyermarkii (Fig. 92). Mark Olson did the drawing of Euceraea rheophytica (Fig. 395). Reviews of certain family treatments in this volume were kindly provided by Rupert Barneby, Kenneth Cameron, Arian Jacobs-Brouwer, Marion J. JansenJacobs, Robert Kral, Paul J. M. Maas, Richard H. Maxwell, James S. Pringle, Gustavo Romero, and Kenneth Wurdack. Dan Nicholson provided nomenclatural corrections to the treatment of Humiriaceae. Information from unpublished revisions and unpublished seed anatomy data for the Gentianaceae was provided by Paul J. M. Maas. Unpublished pollen data was provided for the Gentianaceae by Siwert Nilsson. vii

viii P REFACE AND A CKNOWLEDGMENTS

This project has benefited tremendously from the knowledge and assistance of Otto Huber, the main contributor to Volume 1 and an expert on the flora and vegetation of the Venezuelan Guayana. Numerous institutions and individuals in Venezuela have contributed significantly to different aspects of the flora. These include the Corporación Venezolana de Guayana and its affiliate Electrificación del Caroní, C.A., especially through the assistance of Alfredo Lezama, Hermán Roo, Luis Castro, Gabriel Picón, and Antonio Ahogado; the Ministerio del Ambiente y de los Recursos Naturales Renovables; the Instituto Nacional de Parques and the Dirección General de Parques Nacionales; the Servicio Autónomo para el Desarrollo Ambiental del Estado Amazonas; and the Consejo Nacional de Investigaciones Científicas y Tecnológicas. The Fuerzas Aéreas de Venezuela, Fundación Terramar, Charles Brewer-Carías, and the late Parker Redmond also gave important logistical support on various trips to southern Venezuela. Several Venezuelan herbaria and associated botanists provided considerable assistance over an extended period of time, including the Herbario Nacional de Venezuela, especially through Francisco Delascio, Francisco Guánchez, Gilberto Morillo, and Zoraida Luces de Febres; the Universidad Central de Venezuela, through the Herbario Ovalles at the Facultad de Farmacia and its director Stephen Tillett, and the herbarium of the Facultad de Agronomía in Maracay, particularly with the help of Carmen Emilia Benítez de Rojas; the Universidad Nacional Experimental de los Llanos Ezequiel Zamora in Guanare and its active team of botanists including Gerardo Aymard, Nidia Cuello, and Basil Stergios; the herbarium at Puerto Ayacucho, now associated with the Centro Amazónico de Investigaciones Ambientales Alejandro de Humboldt of the Servicio Autónomo para el Desarrollo Ambiental del Estado Amazonas; and the Universidad de Los Andes in Mérida, mainly through the herbaria of the Facultad de Ciencias Forestales and the Facultad de Farmacia. In the United States, the National Geographic Society provided grants for several explorations associated with this project. Three major institutions, the Missouri Botanical Garden, the New York Botanical Garden, and the Smithsonian Institution, lent their facilities, specimens, and the expertise of their staff to assist in the preparation of numerous floristic treatments. The project has also benefited from the efforts of two postdoctoral researchers, Denis Kearns and John MacDougal. Carlos Vargas worked for a year as a full-time Research Assistant for the project. Other people who worked part time or volunteered to assist on the flora include William Betz, Lois Brako, Germán Carnevali, Kandis Elliot, Luther Raechal, Ronald Liesner, Ivón Ramírez, Erika Rohrbach, and George Yatskievych. A special thanks to George Thornburgh, whose help as a volunteer was instrumental in seeing this volume through to completion.

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

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x INTRODUCTION

FAMILIES INCLUDED IN THIS VOLUME Genera Species Eriocaulaceae Erythroxylaceae Euphorbiaceae Euphroniaceae Fabaceae Flacourtiaceae Gentianaceae Gesneriaceae Gnetaceae Haemodoraceae Haloragaceae Heliconiaceae Hernandiaceae Hippocrateaceae Hugoniaceae

7 1 58 1 67 13 23 17 1 3 1 1 2 11 2

88 30 239 3 368 47 83 58 6 3 1 14 3 38 8

Genera Species Humiriaceae Hydrocharitaceae Hydrophyllaceae Icacinaceae Iridaceae Ixonanthaceae Juncaceae Krameriaceae Lacistemataceae Lamiaceae Lauraceae Lecythidaceae Lemnaceae Lentibulariaceae

6 2 1 7 4 2 1 1 1 9 15 8 4 2 271

24 2 2 16 9 8 1 2 1 30 142 35 4 53 1318

in Volume 5 and their numbers of native or naturalized genera and species in the Venezuelan Guayana are given above. Organization and Scope of the Floristic Treatments The floristic treatments in this flora are organized alphabetically by family, first within the ferns and fern allies, then within the seed plants. The gymnosperms, monocotyledons, and dicotyledons are not treated separately within the seed plant category. The authors for each family treatment are cited below the family heading; in multi-authored families, if different parts were contributed by different authors, the respective authors are cited separately under the parts they were responsible for (such as a particular genus or the key to the genera). Each family treatment contains a complete description of the family and the genera that are present in the flora; these descriptions are worldwide in scope. Within each family, genera are arranged in alphabetical order. At the end of each family or genus description, a separate paragraph lists the overall distribution of the family or genus. This is followed by the author’s estimate of the number of subordinate taxa worldwide, as well as the number of taxa present in the Venezuelan Guayana. For each genus, the estimated number of species in all of Venezuela is also given. All keys used in the flora are dichotomous and bracketed. In each couplet, the number in parenthesis refers to the couplet that led to it. The keys to the genera are designed primarily to cover the species that occur in the Venezuelan Guayana and not necessarily species found elsewhere.

INTRODUCTION

xi

Slightly more than half the species in the flora are illustrated by black-andwhite line drawings. Volume 5 includes 708 figures. The line drawings are consecutively numbered within each volume, and they are usually grouped together by genus near the end of the corresponding species entries to provide easy visual comparisons between species. Since the illustrations and the keys are designed to enable identification of plants from the flora area, the flora does not provide full species descriptions. Species entries are organized alphabetically within their respective genus and begin with the accepted species name, author(s), and place of publication. If the accepted name is a combination, it is immediately followed by its basionym and then by other synonyms based on the same type. Common or vernacular plant names that are known to be used within the Venezuelan Guayana are listed at the end of this paragraph. Synonyms based on a different type from the accepted name are listed in separate paragraphs in chronological order of basionym publication. Synonymy is not intended to be exhaustive, but rather includes synonyms pertinent to the Venezuelan Guayana and adjacent areas. We have attempted to include all names originally based on collections from the flora area. Following the nomenclatural portion of each species entry, a new paragraph indicates the plant’s habit and sometimes includes diagnostic characters of the plant that were not included in the keys. This is followed by a listing of the habitats and the elevational range where the taxon occurs in the flora area only (not worldwide), as well as its geographical distribution within the Venezuelan Guayana. Distribution outside the flora area is listed next, beginning with any states or regions of Venezuela north of the Río Orinoco where the taxon occurs, then any other countries or regions of occurrence. Whenever a taxon is illustrated by a black-and-white line drawing, the symbol “Š” followed by the figure number appears at the end of the paragraph. A separate paragraph is sometimes used to provide taxonomic discussion and/or notes on the uses of the plant in the flora area. Any varieties or subspecies present in the flora area appear below their corresponding species entry, and they contain a similar level of information as species entries. A key is included if more than one subspecies or variety is present. Forms found in the flora area do not have separate entries, but they are discussed under the next higher rank. Throughout the flora, author abbreviations follow Authors of Plant Names (R. K. Brummitt and C. E. Powell, editors. Royal Botanic Gardens, Kew. 1992). The only exception is the use of the abbreviation “H.B.K.” for species described in Nova Genera et Species Plantarum, by A. von Humboldt, A. Bonpland, and K. S. Kunth (Librairie Grecque-Latine-Allemande, Paris. 1815–1825). This series of seven volumes was published in two versions (quarto and folio), each with different page numbers. Page numbers cited in this flora refer to the quarto version, which is more widely available than the folio version, except for volume four, which is the only volume in which the folio edition was distributed before the quarto. Book abbreviations follow Taxonomic Literature: A Selective Guide to Botanical Publications and Collections with Dates, Commentaries and Types, 2nd edition (F. A. Stafleu and R. S. Cowan. Bohn, Scheltema & Holkema, Utrecht, Netherlands. 1976–1988), and journal abbreviations follow Botanico-Periodicum-Huntianum (G. H. M. Lawrence, A. F. G. Buchheim, G. S. Daniels, and H. Dolezal, editors. Hunt Botanical Library, Pittsburgh, Pennsylvania. 1968) or B-P-H/S: Botanico-

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INTRODUCTION

Periodicum-Huntianum/Supplementum (G. D. R. Bridson and E. R. Smith, editors. Hunt Institute for Botanical Documentation, Pittsburgh, Pennsylvania. 1991). When the effective publication date of a reference is different from the imprint date, the effective publication date is placed in brackets following the imprint date. Herbarium abbreviations follow Index Herbariorum, 8th edition (P. K. Holmgren, N. H. Holmgren, and L. C. Barnett, editors. New York Botanical Garden, Bronx, New York. 1990). For each family treated in the flora we have prepared lists of selected voucher specimens of the accepted taxa. These are specimens that have been seen and verified by the authors of the different treatments and are arranged alphabetically by genus, species, and collector. These exsiccatae lists are not included in the flora, but are available upon request from the Missouri Botanical Garden, c/o Flora of the Venezuelan Guayana.

Contributors Victor A. Albert The Lewis B. and Dorothy Cullman Program for Molecular Systematics Studies The New York Botanical Garden Bronx, New York 10458-5126

Lois Brako Graduate School 321 Bascom Hall 500 Lincoln Drive University of Wisconsin–Madison Madison, Wisconsin 53706

Lennart L. Andersson University of Göteborg Department of Systematic Botany Carl Skottsbergs Gata 22 B S-413 19 Göteborg SWEDEN

Richard S. Cowan (deceased) Western Australian Herbarium Department of Conservation and Land Management Como, W.A. 6152 AUSTRALIA

W. Scott Armbruster Department of Botany Norwegian University of Science and Technology Trondheim, N-7491 Dragvoll NORWAY

José Cuatrecasas (deceased) Department of Botany, NHB-166 National Museum of Natural History Smithsonian Institution Washington, DC 20560

Gerardo A. Aymard C. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA Paul E. Berry Botany Department University of Wisconsin–Madison 132 Birge Hall 430 Lincoln Drive Madison, Wisconsin 53706

Nidia L. Cuello A. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA Lawrence J. Davenport Department of Biology Samford University Birmingham, Alabama 35229-2234 Rodrigo Duno de Stefano Herbario Nacional de Venezuela Apartado 2156 Caracas 1010-A VENEZUELA

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CONTRIBUTORS

Hans-Joachim Esser Institut für Allgemeine Botanik und Botanischer Garten Universität Hamburg Ohnhorststasse 18 D-2000 Hamburg 52 GERMANY Paul R. Fantz Dept. of Horticultural Science, Box 7609 Kilgore Hall North Carolina State University Raleigh, North Carolina 27695-7609 Christian Feuillet Department of Botany, NHB-166 National Museum of Natural History Smithsonian Institution Washington, DC 20560 Renée H. Fortunato Secretaria de Agricultura Ganaderia y Pesca, I.N.T.A. Centro de Investigaciones de Recursos Naturales Villa Udaondo 1712, Castelar ARGENTINA Celia Gil Departamento de Biología Universidad Central de Venezuela Apartado 20513 Caracas 1040 VENEZUELA Lynn J. Gillespie Research Division Canadian Museum of Nature P.O. Box 3443, Station D Ottawa, ON K1P 6P4 CANADA Peter Goldblatt Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

Raymond M. Harley Herbarium Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB UNITED KINGDOM W. John Hayden Herbarium Biology Department University of Richmond Richmond, Virginia 23173 Robert R. Haynes Department of Biological Sciences University of Alabama 425 Scientific Collections Building Box 870345 Tuscaloosa, Alabama 35487-0345 Scott V. Heald Institute of Systematic Botany The New York Botanical Garden Bronx, New York 10458-5126 Jennifer P. Hedin Herbarium University of Arizona 113 Schantz Bldg. Tucson, Arizona 85721 James E. Henrich Denver Botanical Garden 909 York Street Denver, Colorado 80206-3799 Nancy Hensold Department of Botany Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Lauritz B. Holm-Nielsen World Bank Education & Social Policies 1818 H Street, NW Washington, DC 20433

CONTRIBUTORS

Richard A. Howard Harvard University Herbaria 22 Divinity Avenue Cambridge, Massachusetts 02138-2020 Otto Huber Apartado 80.405 Caracas 1080-A VENEZUELA Antony Jardim Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Geoffrey A. Levin Herbarium Center for Biodiversity Illinois Natural History Survey 172 Natural Resources Bldg. 607 East Peabody Drive Champaign, Illinois 61820-6970 Haroldo Cavalcante de Lima Herbário, Sistemâtica Jardim Botânico do Rio de Janeiro Rua Pacheco Leão 915 22460 Rio de Janeiro RJ BRAZIL Hiltje Maas Herbarium Institute of Systematic Botany Department of Plant Ecology and Evolutionary Biology State University of Utrecht Heidelberglaan 2 3584 CS Utrecht THE NETHERLANDS Paul J. M. Maas Herbarium Institute of Systematic Botany Department of Plant Ecology and Evolutionary Biology State University of Utrecht Heidelberglaan 2 3584 CS Utrecht THE NETHERLANDS

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Luis Marcano-Berti Facultad de Ciencias Forestales Universidad de Los Andes Mérida 5101 VENEZUELA Richard H. Maxwell Herbarium Indiana University Southeast 4201 Range Line Road New Albany, Indiana 47150 Alberta M. W. Mennega University of Utrecht Department of Plant Ecology and Evolutionary Biology Herbarium Division Heidelberglaan 2 P.O. Box 80.102 3508 TC Utrecht THE NETHERLANDS James S. Miller Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Scott A. Mori Institute of Systematic Botany The New York Botanical Garden Bronx, New York 10458-5126 David Neill Missouri Botanical Garden c/o Herbario Nacional del Ecuador Avenida Río Coca E6-115 Casilla Postal 17-21-1787 Quito ECUADOR Mark Olson Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

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R. Toby Pennington Royal Botanic Garden Edinburgh 20A Inverleith Row Edinburgh EH3 5LR SCOTLAND Oliver Pihlar The Lewis B. and Dorothy Cullman Program for Molecular Systematics Studies The New York Botanical Garden Bronx, New York 10458-5126 Timothy Plowman (deceased) Department of Botany Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Hans-Helmut Poppendieck Institut für Allgemeine Botanik und Botanischer Garten Universität Hamburg Ohnhorststasse 18 D-2000 Hamburg 52 GERMANY Ghillean T. Prance Herbarium Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB ENGLAND, U.K. Luther J. Raechal c/o Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Nelson Ramírez Departamento de Biología Universidad Central de Venezuela Apartado 20513 Caracas 1040 VENEZUELA

Jens G. Rohwer Institut für Spezielle Botanik 55099 Mainz GERMANY Velva Rudd 7307 Newcastle Ave. Reseda, California 91335 Ricardo de S. Secco Herbário, Departamento de Botânica Museu Paraense Emilio Goeldi Caixa Postal 399 66000 Belém, Pará BRAZIL Basil Stergios Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA Dennis W. Stevenson Herbarium The New York Botanical Garden Bronx, New York 10458-5126 Julian A. Steyermark (deceased) Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Charles H. Stirton National Botanic Garden of Wales Middleton Hall Llanarthne, Carmarthen SA32 8HW WALES, U.K. Lena Struwe The Lewis B. and Dorothy Cullman Program for Molecular Systematics Studies The New York Botanical Garden Bronx, New York 10458-5126

CONTRIBUTORS

Peter G. Taylor Herbarium Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB ENGLAND, U.K. Carlos A. Vargas Departamento de Biología Universidad Peruana Cayetano Heredia Apartado 4314 Lima 100 PERU Grady L. Webster Herbarium Section of Plant Biology University of California–Davis Davis, California 95616 Henk van der Werff Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Nereida Xena Departamento de Biología Universidad Central de Venezuela Apartado 20513 Caracas 1040 VENEZUELA

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z 1

ERIOCAULACEAE by Nancy Hensold Annual, perennial, or monocarpic herbs, rosulate to caulescent, sometimes with fertile branches distinct in morphology from the sterile primary stems. Roots fibrous or aerenchymatous (spongy), sometimes diaphragmed (with transverse striations). Leaves spirally arranged, sometimes distichous or verticillate, mostly linear or lanceolate, parallel-veined, sometimes fenestrate (with translucent square window-like lacunae), often with broad sheathing bases; vestiture of filamentous or T-shaped (malpighian) trichomes. Inflorescences involucrate capitula on naked peduncles, either axillary or in terminal umbels, rarely sessile, or flowers solitary, the peduncles enclosed at the base by a tubular or merely open adaxial sheath, mostly 2–12-ribbed, sometimes (not in ours) fasciated and many-capitate; involucral bracts occasionally herbaceous, mostly chaffy, rigid, or hyaline; floral (receptacular) bracts subtending each flower or sometimes absent. Flowers actinomorphic or zygomorphic, shortpedicellate, unisexual (plants monoecious, rarely dioecious, andromonoecious, or hermaphroditic; if monoecious, the two sexes mixed in the same head, though usually occupying distinct whorls). Calyx 2- or 3-merous, the sepals mostly chaffy, free or shortly connate at their bases especially in staminate flowers, sometimes fused into a spathe, often tufted with trichomes, the trichomes acute or often swollen apically and opaque-white due to granular thickening of the internal wall, sometimes tuberculate; corolla 2- or 3-merous, rarely reduced to trichomes or absent, the petals chaffy to membranous, free or variously fused, in staminate flowers often raised above sepals on an intercalary stipe. Stamens twice the number of petals or equal and opposite them; filaments free or adnate basally; anthers with 1 or 2 thecae, introrse, longitudinally dehiscent; pollen spiraperturate, spinulose; staminodia filamentous, scale-like, or absent. Ovary 1–3-locular; ovules solitary, pendulous, orthotropous; style terminal, with 2 or 3 main branches, these vascularized and positioned above the locules (subfamily Eriocauloideae) or nonvascularized, positioned above the septa, and alternating with vascularized nectaries (subfamily Paepalanthoideae); nectaries when present filiform, fusiform, clavate, or funnelform, and variously papillate; stigmas filiform, simple or bifid; pistillodes present in staminate flowers. Fruit usually a loculicidal capsule, rarely 1-seeded and indehiscent. Seeds spheroid or ellipsoid, the testa variously ornamented; endosperm abundant and mealy; embryo small and apical. Pantropics, particularly diverse on the Brazilian and Guayana Shields in South America; 11 genera and ca. 1200 species; 7 genera and 88 species in the flora area. NOTE: The characters that distinguish genera of Eriocaulaceae are technical and sometimes difficult to see and interpret, so the following Key to the Genera is provided mainly for reference purposes. For identification of flora area material, all species are keyed out together in the Key to the Species of Eriocaulaceae. 1

2

E RIOCAULACEAE

Key to the Genera of Eriocaulaceae 1.

1.

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

Stamens 4 or 6, twice as many as the outer perianth segments, rarely 3; petals often with a black gland just below the apex; style branches simple; gynoecial nectaries absent; roots aerenchymatous, diaphragmed ................................................................................................ 1. Eriocaulon Stamens 2 or 3, as many as the outer perianth segments; petals always eglandular; style branches simple to bifid; clavate to filiform nectaries usually inserted on style alternate with the style branches; roots fibrous or aerenchymatous, but rarely diaphragmed ............................. 2 Conspicuous linear staminodes present in pistillate flowers, or flowers staminate and bisexual (plants andromonoecious) ........ 5. Rondonanthus Staminodes tiny and scale-like or lacking; flowers all unisexual or bisexual ..................................................................................................... 3 Petals of pistillate flowers usually connate medially, free below and at apex; T-shaped trichomes often present on vegetative parts ............... 4 Petals of pistillate flowers absent or free; T-shaped trichomes absent .... 5 Anthers 1-thecous ......................................................................... 4. Philodice Anthers 2-thecous ................................................................. 6. Syngonanthus Roots usually aerenchymatous; nectaries inserted on style well below divergence of the style branches; anthers basifixed ................... 2. Leiothrix Roots fibrous; nectaries inserted at or slightly below divergence of style branches; anthers dorsifixed ................................................................. 6 Petals of pistillate flowers present, conspicuous .................. 3. Paepalanthus Petals of pistillate flowers reduced to very small pilose lobes or apparently lacking entirely and replaced by trichomes ........................ 7. Tonina Key to the Species of Eriocaulaceae

1.

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

Plants producing basal rosettes of leaves as well as aerial stems with clusters of peduncles at their apex; aerial stems naked (except for bracts at apex) or foliose and then leaves smaller than rosette leaves, or arranged in verticils .................................................................................. 2 Plants rosulate or caulescent, if caulescent, the basal and cauline leaves not differing in size or arrangement; leaves never in verticils .......... 14 Aerial stems foliose, the cauline leaves few to numerous, spirally arranged .................................................................................................... 3 Aerial stems naked, or if foliose, the cauline leaves arranged in verticils ................................................................................................................ 7 Stems densely leafy, covered with spirally arranged lanceolate leaves > 1 cm long ............................................................................................. 4 Stems with scattered leaves that cover only a fraction of the stem surface, these linear to lance-linear, mostly < 1 cm long ................................... 5 Roots spongy, white; basal leaves bidentate at apex and pellucidmargined ............................................................. Syngonanthus williamsii Roots firm, brownish; basal leaves acute, the margins not pellucid ................................................................................ Paepalanthus formosus Basal leaves 35–130 × 1.5–6 mm, glabrous or nearly so; peduncle

E RIOCAULACEAE

3

sheaths glabrous; peduncles villous, not glandular, 15–40 cm long ................................................................................ Syngonanthus longipes 5. Basal leaves 5–40 × < 1 mm, pubescent; peduncle sheaths and peduncles glandular-hairy; peduncles 3–13 cm long ............................................. 6 6(5). Heads 2.5–3 mm diameter; involucral bracts whitish, hyaline, acuminate; staminate flowers zygomorphic (falcate) .....Syngonanthus bisumbellatus 6. Heads 6–7 mm diameter; involucral bracts cream or often with a greenish or brownish midvein, obtuse; staminate flowers actinomorphic ........... ............................................................................ Syngonanthus fenestratus 7(2). Aerial stem < 2 cm long, much surpassed by the linear, erect leaves; leaves 3.5–5.5 cm long, with 5 or more veins ............... Syngonanthus amapensis 7. Aerial stem > 2 cm long; or, if less, then not hidden by the rosette leaves, these 1–14 cm long and either 1-veined or not erect ............................ 8 8(7). Basal leaves linear, tufted, with a single prominent vein, 1–5 cm long, ca. 1 mm wide or less, involucre and perianths conspicuously pubescent ................................................................................................................ 9 8. Basal leaves linear to spatulate, usually spreading, 3–many-veined, 2–13 cm long, 1.5–9 mm wide or if less, then heads completely glabrous .............................................................................................................. 11 9(8). Heads 1–5 per umbel, 7–9 mm diameter; involucral bracts and flowers coppery red, hyaline; aerial stems always naked ................................... ................................................................... Syngonanthus macrocephalus 9. Heads usually > 5 per umbel, 1.5–6 mm diameter; involucral bracts cream to red-brown; flowers white to buff; aerial stems often bearing verticils of leaves ................................................................................................ 10 10(9). Peduncle sheaths open to the base; peduncles 0.1–1 cm long, sometimes entirely hidden by uppermost verticil of leaves .... Syngonanthus cowanii 10. Peduncle sheaths closed; peduncles 1.8–14 cm long ..... Syngonanthus humboldtii 11(8). Leaves coriaceous, sharp-aristate, 1–1.5 mm wide; heads 7–7.5 mm diameter; involucral bracts reddish when fresh, glabrous ........................ ............................................................................. Syngonanthus ottohuberi 11. Leaves membranous to chartaceous, acute to rounded, 1.5–9 mm wide; heads 4–8 mm diameter; involucral bracts white to gold, glabrous to frequently pilose .................................................................................. 12 12(11). Leaves oblong-elliptic to spatulate, the apex rounded; involucral bracts oblong-obovate, broadly rounded ......................... Syngonanthus oblongus 12. Leaves linear to lance-linear, acute; involucral bracts various .............. 13 13(12). Lower surface of leaves hirsute; involucral bracts hyaline, acuminate ........................................................................... Syngonanthus umbellatus 13. Leaves glabrous or appressed-puberulent; involucral bracts leathery, ovate, rounded ....................................................... Syngonanthus longipes 14(1). Plants in moss-like mats; mature leaves 1–2 mm long; roots spongy; heads sessile, lacking an involucre, reduced to 1 or 2 flowers borne terminally on short unspecialized branches and hidden among the leaves ............................................................................. Syngonanthus acephalus 14. Plants sometimes in branched mats, but the mature leaves at least 3 mm long; roots fibrous or spongy; heads mostly pedunculate, rarely sessile and then many-flowered and subtended by an involucre .................. 15

4

E RIOCAULACEAE

15(14). Surface of head glabrous or nearly so, neither the sepals nor the floral bracts bearded or tufted at apex (though occasionally obscurely ciliate or tufted well below apex; also sometimes with a dense cover of receptacular hairs remaining after abscission of the flowers) ........... 16 15. Surface of head densely whitish pubescent due to the bearded or densely ciliate apices of the sepals and floral bracts ....................................... 41 16(15). Plants caulescent, the stems ca. 8–40 cm long at flowering; heads brown to blackish ............................................................................................ 17 16. Plants of various habit; if caulescent, the heads white .......................... 18 17(16). Leaves capillary-membranous, 3–6 cm long; inflorescences borne in terminal umbels .............................................................. Eriocaulon setaceum 17. Leaves lanceolate, 0.7–2.7 cm long; inflorescences solitary in leaf axils .......................................................................................... Tonina fluviatilis 18(16). Plants rosulate with fenestrate leaves; heads cone-like, i.e., globose to ovoid or cylindric, the floral bracts similar in size and shape to the involucral bracts, appressed against the surface of the head (as in Xyris) enclosing to half-enclosing the flowers ............................................... 19 18. Plants caulescent or rosulate, if rosulate, then not with fenestrate leaves; heads turbinate to globose, the central axis never elongate, the floral bracts when present much smaller than the involucral bracts and visible only by dissection of the head ....................................................... 23 19(18). Roots not diaphragmed; peduncles 3-ribbed; bracts cuspidate, deflexed at apex, usually shedding after seed set starting from the base ............ 20 19. Roots diaphragmed (transversely striate); peduncles 4–6-ribbed; bracts acute to rounded, not cuspidate and deflexed at apex, persistent ..... 21 20(19). Leaves (0.3–)0.4–1 mm wide; flowers with 2 stamens ....................... .......................................................................... Syngonanthus amazonicus 20. Leaves 0.15–0.25 mm wide; flowers with the adaxial stamen only ........... ........................................................................ Syngonanthus trichophyllus 21(19). Floral bracts acute to apiculate, ascending but not tightly appressed; anthers white; gynoecium 3-merous ............................. Eriocaulon cinereum 21. Floral bracts broadly rounded; anthers black; gynoecium 2-merous ..... 22 22(21). Floral bracts completely enclosing flowers; sepals asymmetrically keeled, winged ...................................................................... Eriocaulon guyanense 22. Floral bracts only about half-enclosing flowers; sepals symmetrically conduplicate, not winged ................................... Eriocaulon dimorphopetalum 23(18). Plants submerged rosulate aquatics of flowing water; leaves filiform, terete, 7–30 cm long .............................................. Rondonanthus capillaceus 23. Plants terrestrial to amphibious, rosulate to caulescent; leaves rarely appearing filiform or terete, but if so, then < 7 cm long ........................ 24 24(23). Peduncles distinctly 6-ribbed, usually with silvery, appressed, T-shaped trichomes toward apex; peduncle sheaths papery, pale, the mouth obtuse to truncate, scarious, and usually lacerate ................................. 25 24. Peduncles 3-ribbed, sometimes obscurely so; peduncle sheaths herbaceous, the mouths entire, acute ........................................................... 26 25(24). Involucral bracts triangular-oblong, broadly rounded, surpassing disk by 1–2 mm .................................................................. Syngonanthus tiricensis 25. Involucral bracts lanceolate, acute to acuminate, not surpassing disk

E RIOCAULACEAE 5

.................................................................................. Syngonanthus duidae 26(24). Peduncle sheaths lacking......................................................................... 27 26. Peduncle sheaths present ........................................................................ 28 27(26). Plants caulescent; involucral bracts green, lance-linear, acute ................. ............................................................................. Philodice hoffmannseggii 27. Plants rosulate; involucral bracts gold-brown, ovate, obtuse ..................... ................................................................................ Syngonanthus minutus 28(26). Involucres cup-like or even somewhat constricted at the mouth, membranous to leathery, cream to brown; flowers of outer whorl exserted on long pedicels, surpassing the involucre and eventually abscising; leaves 2–23 cm long ............................................................................. 29 28. Involucres obconic and usually flattening with maturation of head, scarious to membranous, never leathery, white to cream or greenish; flowers never exserted on long pedicels; leaves 0.3–8 cm long ................. 31 29(28). Involucral bracts abruptly aristate and sharply reflexed; roots with a wiry blackish core ............................................................ Syngonanthus reflexus 29. Involucral bracts (at least the lower) ovate to obovate, obtuse to rounded; roots white, spongy or brittle, without a blackish core ...................... 30 30(29). Leaves and involucres coriaceous, bracts mostly of uniform color (cream to dark brown), mostly glabrous, occasionally pubescent abaxially but not at margin; peduncles glabrous at apex ............................................. .................................................................... Syngonanthus xeranthemoides 30. Leaves and involucres membranous to chartaceous, the lower involucral bracts white and obscurely ciliate, the upper gray-brown flecked; peduncles often fringed with trichomes at apex ...... Syngonanthus jenmanii 31(28). Roots dark and wiry; leaves hirsute on lower surface with stiff erect trichomes; leaf upper surface and peduncle sheaths usually covered with white oblong scales ............................................ Syngonanthus kegelianus 31. Roots white to orange, spongy; leaves appressed-pubescent to softly woolly, never hirsute, never scaly ....................................................... 32 32(31). Plants amphibious, caulescent, but sometimes appearing rosulate; leaves usually with several veins, the midvein not prominent; corollas firm, fleshy, not shriveling after anthesis .................................................... 33 32. Plants terrestrial (though often in wet places), rosulate; leaves mostly 1-veined or with midvein more prominent (except sometimes in Syngonanthus nitens and S. tenuis); corollas membranous, shriveling after anthesis ....................................................................................... 34 33(32). Stems brownish, scurfy; involucral bracts usually greenish along the midvein; sepals rounded ..................................... Syngonanthus anomalus 33. Stems with white, arachnoid tomentum; involucral bracts white; sepals sharp-acuminate ................................................ Syngonanthus caulescens 34(32). Upper involucral bracts white, conspicuously surpassing the disk, radiating outward or cupped over it in dry material; lower bracts much smaller, often yellow or brownish ............................. Syngonanthus tenuis 34. Upper involucral bracts white to gold, not or only very slightly surpassing the disk, the lower the same color as the upper ................................. 35 35(34). Roots frequently deep orange; leaves firm, glabrous or appressed-hairy, never pilose or lanate; involucral bracts in 5–8 series ....... Syngonanthus nitens

6

E RIOCAULACEAE

35. 36(35).

36. 37(36). 37. 38(37).

38.

39(38).

39.

40(39).

40.

41(15).

41.

42(41).

42.

Roots white to yellowish, never orange; leaf texture and pubescence various; involucral bracts in 2–4 series ..................................................... 36 Pistillate flowers about twice as long as the staminate, the sepals sharpacuminate and conspicuously projecting above surface of the disk ................................................................................ Syngonanthus biformis Pistillate flowers equal or slightly surpassing the staminate, never twice as long; sepals acute to acuminate ...................................................... 37 Leaves filiform, membranous, prostrate or recurved, ca. 0.2 mm wide or less; heads 2–2.5 mm diameter ............................ Syngonanthus setifolius Leaves linear, if 0.2 mm wide or less, then not membranous or prostrate; heads 2–8 mm diameter ...................................................................... 38 Involucres gold, hyaline, contrasting with the paler flowers; leaves with fine T-shaped trichomes evenly on both surfaces, the midvein usually sunken in lower epidermis; flowers lacking nectaries ....... Syngonanthus gracilis Involucres cream to white, the same color as the flowers; leaves pubescent only on upper surface, the midvein prominent on lower surface; flowers with nectaries ....................................................................................... 39 Rosulate or clumping annuals; heads 2–4 mm diameter, often numerous per rosette; leaves never coriaceous or subterete, nor building into a dense cushion; flowers and involucral bracts with a yellowish tinge, lustrous.................................................................... Syngonanthus simplex Perennials; heads 3.5–8 mm diameter, usually produced 1 or 2 at a time per rosette; leaves membranous to coriaceous and subterete and then persistent, forming a thick cushion-like or columnar base; flowers and involucral bracts white, not lustrous .................................................. 40 Leaves 0.3–1 cm long and densely lanate, or to 2 cm but then membranous and glabrate; heads 3.5–5 mm diameter; involucral bracts glabrous with broadly rounded apices; plants of Amazonas and Cerro Guaiquinima ............................................... Syngonanthus albopulvinatus Leaves 1–5 cm long, usually coriaceous and always pilose or lanate; heads 5–8 mm diameter; involucral bracts commonly obscurely hairy abaxially, the uppermost whorl usually reduced, narrow, subacute; plants of the Gran Sabana, Cerro Jaua, and Sierra Pakaraima ............ ..................................................................... Syngonanthus pakaraimensis Plants perennial, forming dichotomously branched clumps or prostrate mats, rarely clumping rosettes branched at ground level; leaves linear, rounded at apex, 0.3–2.5 cm long, 0.3–1.5 mm wide, semisucculent to coriaceous; peduncles obscurely 3-ribbed; involucral bracts hyaline at least at margins; heads 3–6 mm diameter; roots wiry ....................... 42 Plants not as above, rosulate or sparingly branched; heads 1.5–19 mm diameter; if perennial and forming rosulate or erect clumps, then possessing one or more of the following characters: peduncles 4–10-ribbed, leaves chartaceous, roots soft, and/or involucral bracts not semi-hyaline ........................................................................................................ 46 Involucral bracts cream to gold, obtuse, in 5–9 series, shredding early with expansion of the head; peduncles 3.5–11.5 cm long ....................... ......................................................................... Paepalanthus yapacanensis Involucral bracts gold to dark brown, obtuse to acute or aristate in

E RIOCAULACEAE 7

2–4 series, mostly remaining intact; peduncles 0.3–10 cm long ........ 43 43(42). Involucral bracts gold, hyaline, aristate; stems often semiprostrate, ascending only at apices ............................................ Paepalanthus aristatus 43. Involucral bracts brownish, semihyaline, usually acute to obtuse, very rarely aristate; stems usually erect .................................................... 44 44(43). Involucral bracts in 2 or 3 series, glabrous or frequently tufted at apex; leaves with persistent long-spreading cilia especially along lower margins, otherwise glabrous; heads 3–4 mm diameter; peduncles 2–10 cm long ................................................................. Paepalanthus polytrichoides 44. Involucral bracts in 3 or 4 series, glabrous; leaves with silvery, appressed or spreading trichomes along upper margins, and sometimes also abaxially along midvein and adaxially at apex, the long cilia of lower margins, when present, sticky and coherent, not spreading, and only very ephemeral; heads 3–6 mm diameter; peduncles 0.3–5 cm long .............................................................................................................. 45 45(44). Plants clumping rosettes, branched only at or below ground level ............ ................................................................................ Paepalanthus cristatus 45. Plants branching above ground level, forming cushions ................... ............................................................................ Paepalanthus dichotomus 46(41). Heads 1.5–5 mm diameter; leaves < 3 mm wide at midpoint ................ 47 46. Heads 4–19 mm diameter, if < 5.5 mm, then the leaves always at least 3 mm wide at midpoint ........................................................................ 60 47(46). Plants rosulate; leaves filiform, densely tufted, 0.4 mm wide or less .... 48 47. Plants rosulate or caulescent; if rosulate, then leaves at least 0.5 mm wide, never filiform .............................................................................. 49 48(47). Leaves 7–30 cm long, terete; peduncles glabrous, 9–20 cm long; heads 3–5 mm diameter; plants submerged aquatics of rapidly flowing water ........................................................................... Rondonanthus capillaceus 48. Leaves 0.5–4 cm long, flat or subterete; peduncles pubescent, 2.5–10 cm long; heads 2–2.5 mm diameter; plants terrestrial ................................ ............................................................................... Syngonanthus setifolius 49(47). Plants rosulate; roots spongy, unbranched; peduncle sheaths with scarious, often lacerate margins ................................... Eriocaulon tenuifolium 49. Plants rosulate or caulescent; roots fibrous, branched; peduncle sheaths with firm, entire margins or absent .................................................... 50 50(49). Plants perennial; involucral bracts dark brown, opaque, never hyaline, often the basal with the midvein thickened and orange-brown; fruits 1-seeded, indehiscent ........................................................................... 51 50. Plants annual; involucral bracts various, at least the basal membranous or hyaline; fruits 3-locular capsules .................................................... 52 51(50). Peduncles ca. 19–24 cm long; leaves smooth on upper surface .................. ....................................................................... Paepalanthus parvicephalus 51. Peduncles 5–6 cm long; leaves with veins lightly impressed on upper surface .................................................................... Paepalanthus cardonae 52(50). Peduncles pubescent in the basal 1/2, or absent (heads sessile) .............. 53 52. Peduncles always present, glabrous or nearly so, except sometimes with a tuft of trichomes at apex fringing the involucre ................................. 57 53(52). Involucral bracts dark brown or blackish ............................................... 54

8

E RIOCAULACEAE

53. Involucral bracts pale, either green, buff, or hyaline ............................. 55 54(53). Leaves attenuate, acute ....................................... Paepalanthus fasciculoides 54. Leaves with margins slightly rounded toward apex, obtuse ...................... ............................................................................... Paepalanthus lamarckii 55(53). Leaves linear, obtuse to rounded, coriaceous, abruptly widened at base; heads always sessile ......................................... Paepalanthus sessiliflorus 55. Leaves attenuate, acute, membranous, not abruptly widened at base; heads usually pedunculate .................................................................. 56 56(55). Involucral bracts lance-linear, leaf-like, green at first and sometimes turning pale buff; heads globose, 3–4 mm diameter; leaves 1–2.5 mm wide ........................................................................... Paepalanthus bifidus 56. Involucral bracts obtuse to rounded, buff; heads globose to cylindrical, 2–3 mm diameter; leaves 0.5–1 mm wide ........ Paepalanthus fasciculatus 57(52). Leaf apex rounded and abruptly mucronate; base of involucre fringed by long yellowish trichomes of peduncle apex .... Paepalanthus oyapockensis 57. Leaf apex acute to rounded but never mucronate; peduncles glabrous at apex ...................................................................................................... 58 58(57). Stems erect, 0.3–5 cm long; involucres firm, conspicuous even at maturity, the outer bracts hyaline, aristate ............................ Paepalanthus subtilis 58. Stems various; involucres membranous, ephemeral, inconspicuous at maturity, the outer bracts dark, never aristate ....................................... 59 59(58). Plants strictly rosulate; leaves acute; pistillate flowers with sepals uniformly black and petals linear and glabrous or sparsely hairy .............. ............................................................................ Paepalanthus perpusillus 59. Plants caulescent though sometimes apparently rosulate; leaves convex toward apex; pistillate flowers with sepals reddish banded along midveins, and petals spatulate and densely pilose .........Paepalanthus tortilis 60(46). Plants floating aquatics with ribbon-like fenestrate leaves and peduncles ................................................................................... Eriocaulon spongiola 60. Plants terrestrial or submerged aquatics with erect emergent peduncles .............................................................................................................. 61 61(60). Peduncle sheaths deeply buried among leaves and only about 1/4 their length .................................................................................. Leiothrix celiae 61. Peduncle sheaths not hidden, at least 1/2 the length of the leaves and often surpassing them ................................................................................... 62 62(61). Involucral bracts yellow or light reddish brown, the margins rounded toward apex, at least the lower bracts glabrous .................................... 63 62. Involucral bracts mostly dark brown to black, or at least partially so; the apex shape and pubescence various but if bracts a paler brown, then acute or pubescent or both .................................................................. 64 63(62). Leaves firm to membranous; peduncle sheaths 2.5–10 cm long; heads typically globose to ovoid at maturity, fungiform (peduncle apex enclosed by revolute margins of the receptacle), not shattering after anthesis; petals equaling sepals, actinomorphic ....... Eriocaulon tenuifolium 63. Leaves rigid; peduncle sheaths 9–30 cm long; heads typically hemispheric, occasionally subglobose and concave at base but not fungiform, shattering after anthesis; petals strongly surpassing sepals, zygomorphic .......................................................................... Eriocaulon humboldtii

E RIOCAULACEAE 9

64(62). Leaves conspicuously fenestrate, at least at base; roots white, diaphragmed ............................................................................................. 65 64. Leaves not evidently fenestrate, roots black and fibrous to spongy and white, but never diaphragmed ............................................................ 66 65(64). Leaves acute, membranous, fenestrate throughout ... Eriocaulon steyermarkii 65. Leaves obtuse, succulent, fenestrate at base only ........... Eriocaulon jauense 66(64). Roots thick, white, at least 1 mm wide, spongy; leaves with rounded apices; peduncle sheaths subtruncate, herbaceous, shallowly shredding (lacerate) at apex ......................................................... Leiothrix flavescens 66. Roots usually finer, black to white, fibrous or porous and brittle but never spongy; leaves various; peduncle sheaths scarious or herbaceous, mostly splitting along a single suture ................................................. 67 67(66). Involucral bracts lanceolate to narrowly oblong, the upper usually surpassing the disk; sepals blackish distally and strongly contrasting creamy white at base or along midvein .............................................. 68 67. Involucral bracts ovate to lance-oblong, never surpassing the disk; sepals deep to pale brown ............................................................................... 72 68(67). Leaves distichous ..................................................................................... 69 68. Leaves spirally arranged ......................................................................... 70 69(68). Heads 16–19 mm diameter; flowers bisexual and staminate ..................... ....................................................................... Rondonanthus flabelliformis 69. Heads 7.5–14(–15) mm diameter; flowers pistillate and staminate, the pistillate with linear staminodes present ...... Rondonanthus acopanensis 70(68). Involucral bracts of upper series oblong, acuminate, the acumen colorless and tufted ........................................................... Rondonanthus caulescens 70. Involucral bracts of upper series lanceolate, acute, black at apex ......... 71 71(70). Leaves aristate; petals of staminate flowers fused and ciliate at upper margin ...................................................................... Rondonanthus duidae 71. Leaves obtuse to subacute; petals of staminate flowers free, glabrous .............................................................................. Rondonanthus roraimae 72(67). Involucres membranous, the bracts in 2 series, inconspicuous at inflorescence maturity; heads shedding trichomes early; receptacular hairs absent; peduncles 3-ribbed (see couplet 59 for additional characters) ....................................................................................Paepalanthus tortilis 72. Involucres firm, the bracts in 2–4 series; heads persistently pubescent; receptacular hairs present; peduncles mostly 4–6-ribbed .................. 73 73(72). Leaves chartaceous, glabrous or with scattered long cilia, rounded toward the sharp mucronate or aristate apex; fruits various ........................ 74 73. Leaves of various texture and pubescence, never rounded toward a sharp mucronate or aristate apex, sometimes sharp-cuspidate but then always pubescent; fruits 1-seeded, indehiscent ..................................... 75 74(73). Heads 4–5 mm diameter; fruits 3-locular capsules ...... Paepalanthus oyapockensis 74. Heads 6.5–9 mm diameter; fruits 1-seeded, indehiscent ............................ ......................................................................... Paepalanthus chimantensis 75(73). Upper surface of leaves conspicuously pubescent with short to long, spreading or appressed trichomes ....................................................... 76 75. Upper surface of leaves glabrous or pubescent only near the apex (though may be pubescent on margins or lower surface) ................................. 86

10

E RIOCAULACEAE

76(75). 76. 77(76). 77. 78(77).

78.

79(76). 79. 80(79). 80. 81(79).

81.

82(81).

82.

83(81). 83. 84(83).

84.

85(84). 85. 86(75). 86.

Peduncle sheaths surpassing subtending leaves by 2–6 cm ................... 77 Peduncle sheaths surpassing leaves by < 2 cm or not at all................... 79 Leaves sharp-cuspidate .................................................. Paepalanthus major Leaves obtuse to bluntly acute ................................................................ 78 Leaves subacute or ensiform-acute, remaining erect, often twisting at the apex upon drying; involucral bracts lance-ovate, acute to obtuse ......... ....................................................................... Paepalanthus squamuliferus Leaves actually acute but appearing obtuse due to the abruptly deflexed tip, only the uppermost on stem held erect, the lower usually strongly reflexed; involucral bracts ovate, apiculate ........... Paepalanthus fulgidus Leaves densely silky-pubescent on upper surface, the trichomes long, appressed to leaf surface and cohering together .................................... 80 Leaves pubescent on upper surface with short to long hairs, these not silky-appressed, spreading at least at their apices ............................ 81 Leaves 10–14 mm wide, with a blunt coriaceous mucro at apex ............... .................................................................................... Paepalanthus holstii Leaves 1.5–4.5 mm wide, subacute .................. Paepalanthus squamuliferus Leaves sharply acute, rigid, fibrous, especially toward the apex, slightly keeled distally (at least when dry); involucral bracts lance-ovate, dull yellowish to grayish brown, or occasionally blackish brown (fuscous), acute ..................................................................................................... 82 Leaves acute or often rounded toward the apex, membranous to chartaceous, not fibrous-thickened distally, mostly drying flat; involucral bracts ovate to ovate-apiculate, if lance-ovate, then dark fuscous .............................................................................................................. 83 Plants rosulate to subcaulescent, rarely stems to 6 cm long; leaves lancelinear, 2–3 mm wide, ascending, usually slightly falcate; inflorescences 1–4 per stem, often solitary .................................. Paepalanthus fraternus Plants caulescent, the stem at flowering ca. 4–21 cm long; leaves lanceolate, 2–6 mm wide, only the uppermost leaves ascending, the lower reflexed, and densely overlapping; inflorescences 2–9 per stem ........... ............................................................................ Paepalanthus cumbricola Leaves pilose on both surfaces with fine spreading trichomes .................. ............................................................................. Paepalanthus turbinatus Leaves glabrous on lower surface ............................................................ 84 Leaves ligulate, obtuse, 1.5–3 mm wide, pubescent on upper surface with short flat trichomes from margin to margin; anthers brown ................. ....................................................................... Paepalanthus parvicephalus Leaves lance-linear to lanceolate, acute, 3–10(–14) mm wide; pubescent on upper surface with fine spreading trichomes, except for a glabrous, shining submarginal band (the margin itself with scattered cilia); anthers white ........................................................................................... 85 Leaf apex somewhat decurved; leaves 8–10(–14) mm wide; plants of Cerro Jaua ....................................................................... Paepalanthus phelpsiae Leaf apex flat; leaves mostly < 8 mm wide; plants of the Gran Sabana .... ........................................................................ Paepalanthus schomburgkii Lower surface of leaf uniformly soft-hairy ................. Paepalanthus fulgidus Lower surface of leaf glabrous or striate-hairy ....................................... 87

E RIOCAULACEAE 11

87(86). Leaves 5–18 × 1–2.5 mm, the upper surface smooth, without impressed veins ..................................................................................................... 88 87. Leaves mostly 20–75 × 1.5–10 mm, sometimes slightly shorter but then with 1–3 impressed veins above .......................................................... 91 88(87). Leaves 5–6 mm long .............................................. Paepalanthus apacarensis 88. Leaves 7–18 mm long ............................................................................... 89 89(88). Involucral bracts broadly ovate, apiculate, sericeous-villous abaxially and ciliate; anthers brown ................................... Paepalanthus parvicephalus 89. Involucral bracts ovate to lance-ovate, acute or apiculate, ciliate but glabrous abaxially; anthers white ............................................................ 90 90(89). Leaves 1–1.5 mm wide, divergent, rigid, the margins never creasing when dry, upper surface not lustrous, the distal margins glabrous or with scattered long cilia; peduncle sheath with acute to acuminate lamina 4–6 mm long .............................................. Paepalanthus auyantepuiensis 90. Leaves 1–2.5 mm wide, the lower strongly recurved, firm but not rigid, the upper surface lustrous and sometimes creasing horizontally when dry, the distal margins usually very short-ciliate when young; peduncle sheath with obtuse and sharp-cuspidate lamina 1–3 mm long ................................................................................ Paepalanthus venustus 91(87). Leaves 2.5–10 mm wide, triangular-acute, finely striate on both surfaces, drying flat or keeled; involucral bracts ciliate but rarely abaxially pubescent; anthers white ......................................................................... 92 91. Leaves 1.5–5.5 mm wide, the apex ensiform or convex, not truly acute; smooth or with 1–3 veins impressed on the upper surface, finely striate on the lower surface, drying flat or with margins slightly revolute; involucral bracts often pubescent abaxially with silky or subappressed hairs; anthers white to brown ............................................................. 97 92(91). Plants of Roraima-tepui ........................................................................... 93 92. Plants of elsewhere in Bolívar ................................................................. 95 93(92). Leaves more leathery toward apex, the tips often darker and keeled in drying; involucral bracts mostly dark brown, but the lower often reddish or gold and thicker at base, and the upper sometimes mottled ........................................................................... Paepalanthus roraimensis 93. Leaves membranous to firm, not thicker or discolored at tips; involucral bracts of uniform color and thickness ................................................. 94 94(93). Involucral bracts coppery or grayish brown, subhyaline, abruptly apiculate .................................................................. Paepalanthus schomburgkii 94. Involucral bracts dark brown, opaque, acute .... Paepalanthus subcaulescens 95(92). Leaves membranous to somewhat fibrous and then slightly keeled in drying, commonly lustrous with pellucid venation; peduncles 12–35 cm long, rarely fringed at apex; involucral bracts uniform in color and texture or subhyaline along margins, mostly glabrous, occasionally ciliate or pubescent abaxially with subappressed hairs ........ Paepalanthus schomburgkii 95. Leaves fibrous, nearly always conduplicate when dry, dull, never lustrous; peduncles mostly 5–12 cm long, fringed with trichomes at apex; involucral bracts, especially the lower, with a contrasting gold or reddish midrib; all bracts ciliate and mostly hirsute along midvein when young .................................................................................................... 96

12

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96(95). Stems at flowering about 5–6 cm long ...................... Paepalanthus cardonae 96. Stems at flowering ca. 15–60 cm long .................. Paepalanthus scopulorum 97(91). Heads 8–16 mm diameter; peduncle sheaths equaling leaves to surpassing them by 4 cm .................................................................................. 98 97. Heads 5.5–7.5 mm diameter; peduncle sheaths about equaling leaves .............................................................................................................. 99 98(97). Leaves long-ciliate at base and densely and persistently short-ciliate distally .................................................................... Paepalanthus gleasonii 98. Leaves ciliate from base to apex with long, usually coherent, appressed trichomes, or else glabrous ........................... Paepalanthus squamuliferus 99(97). Leaves smooth on upper surface or very rarely striate ........................ 100 99. Leaves always with veins impressed on upper surface ........................ 101 100(99). Flowering stems 8 cm or less in length; leaves glabrous except for the woolly or ciliate base, usually strongly ensiform ................... ......................................................................... Paepalanthus kunhardtii 100. Flowering stems usually 10–40 cm in length; leaves conspicuously ciliate, at least at base, and often also hirsute distally ...................... ................................................................... Paepalanthus parvicephalus 101(99). Leaves with metallic bluish sheen, ciliate only at base; peduncle sheaths glabrous at mouth; involucral bracts obtuse to rounded ..................................................................... Paepalanthus stegolepoides 101. Leaves generally lacking bluish sheen, ciliate to apex; peduncle sheaths ciliate at mouth; involucral bracts acute to acuminate or apiculate ......................................................................................... 102 102(101). Flowering stems to 40 cm long; leaves 2–2.5 mm wide; peduncles 10–14 cm long .................................................... Paepalanthus convexus 102. Flowering stems 15 cm long or less; leaves 2.5–4 mm wide; peduncles 15–25 cm long ..................................................... Paepalanthus sulcatus 1. ERIOCAULON L., Sp. Pl. 87. 1753. Terrestrial or aquatic herbs of simple habit, rosulate or the stem rarely elongate and sparingly branched. Roots white, aerenchymatous (spongy), unbranched, diaphragmed. Leaves membranous to succulent or rigid, usually fenestrate at least at the base, glabrous or obscurely pubescent, the trichomes simple. Inflorescences always in the axils of foliage leaves. Peduncle sheaths usually scarious at apex, splitting down the side or shredding at apex. Flowers 2- or 3-merous, always unisexual (plants monoecious, rarely dioecious). Sepals free or connate into an adaxial spathe (more commonly so in staminate flowers), often densely bearded with opaque, white, clavate or sometimes acute translucent trichomes; prominent, often fleshy, corolla stipe present in staminate flowers, with slight elongation of the floral axis sometimes also occurring in pistillate flowers; petals of both sexes free, the anterior sometimes much larger than the laterals, sometimes reduced or lacking (more commonly so in staminate flowers), often pubescent with opaque or translucent trichomes, usually bearing black multicellular glands adaxially below the apex. Stamens twice as many as the sepals; anthers usually black, sometimes white, with 2 thecae. Ovary 2or 3-locular; style branches simple, positioned above the locules. Fruit a capsule. Pantropics, a few species extending to eastern North America and the British Isles; ca. 400 species, 12 in Venezuela, 9 of these in the flora area.

Eriocaulon 13

Eriocaulon cinereum R. Br., Prodr. 254. 1810. Eriocaulon sieboldianum Siebold & Zucc. ex Steud., Syn. Pl. Glumac. 2: 272. 1855. Membranous rosulate annual; leaves 15– 50 × 0.8–2 mm; peduncles 4–10(–18) cm long; capitula 2.5–4.5 mm diameter, numerous, gold to blackish. Wet openings of sandy savanna, over granitic outcrops, sometimes submerged in shallow water, 50–1200 m; Bolívar (Caicara, Uaipán-tepui), Amazonas (Puerto Ayacucho). Guárico; Pantropics. ŠFig. 1. Eriocaulon dimorphopetalum Mold., Fieldiana, Bot. 28: 116. 1951. Rosulate annual; leaves 20–60 × 1–2 mm, acute; peduncles 5–9 cm long; capitula ca. 5 mm diameter, creamy gold, glabrous; flowers 2-merous; sepals of the pistillate flowers fleshy and conduplicate. Swampy meadows, ca. 1000 m; Bolívar (Río Kukenán south of Hato La Divina Pastora). Endemic. Eriocaulon guyanense Körn. in Mart., Fl. Bras. 3(1): 478. 1863. —Tenahkwa-men (Panare). Rosulate annual; leaves 10–65 × 0.8–2.5 mm, acute; peduncles 5–18 cm long; capitula 3–4.5 mm diameter, creamy gold, glabrous; flowers 2-merous; sepals of the pistillate flowers asymmetrical and winged. Moist areas of savannas, roadsides, 50–100 m; Bolívar (Caicara, lower Río Suapure). Guárico; Panama, Colombia, Guyana, French Guiana, Brazil, Bolivia. Eriocaulon humboldtii Kunth, Enum. Pl. 3: 544. 1841. —Paja vinera. ?Eriocaulon aequinoctiale Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 47. 1903. Robust rosulate herb; leaves 6–53 × 2–6 mm, erect, rigid; peduncles 25–75 cm long; capitula (5–)7–10 mm diameter. Usually terrestrial, occasionally partly submerged, moist meadows or savannas, Mauritia palm swamps, gallery forests, streambanks, 50– 1400 m; Delta Amacuro (lower Río Orinoco), Bolívar (Altiplanicie de Nuria, Ciudad Píar, Gran Sabana, Río Orinoco, Río Paragua), widespread in western and central Amazonas. Anzoátegui, Apure, Guárico, Mona-

gas; Colombia, Guyana, Brazil (Amazonas, Goiás, Mato Grosso). ŠFig. 3. Ruhland distinguished Eriocaulon aequinoctiale, from the lower Río Orinoco, by the leaves entirely rigid and not visibly fenestrate even at the base, a condition which also may occur in E. humboldtii. Eriocaulon jauense Mold., Mem. New York Bot. Gard. 23: 850. 1972. Clumping perennial rosette; leaves 10–15 × 5–7 mm, succulent; peduncles 25–45 cm long; involucre blackish; capitula 9–12 mm diameter. Boggy tepui meadows, 1800–2200 m; Bolívar (summits of Cerro Jaua and Sierra de Maigualida), Amazonas (summits of Cerro Coro Coro, Cerro Yutajé, and Serranía Uasadi). Endemic. Eriocaulon melanocephalum Kunth, Enum. Pl. 3: 549. 1841. Eriocaulon aquaticum Sagot ex Koern in Mart., Fl. Bras. 3(1): 498. 1863. Lasiolepis aquatica Boeck., Flora 56. 91. 1873. Long-stemmed floating aquatic with emergent umbels; leaves (10–)30–60 × 0.25 mm, capillary; peduncles 6–23 cm long; capitula 2.5–3 mm diameter, black. Small streams, 100–1000 m; Bolívar (eastern part of state, Gran Sabana), Amazonas (Puerto Ayacucho). Guárico; Central America, Cuba, Colombia, Guyana, Suriname, French Guiana, central and Amazonian Brazil, Africa. ŠFig. 2. Eriocaulon spongiola Hensold, Ann. Missouri Bot. Gard. 78: 424, figs. 1, 2. 1991. Long-stemmed floating aquatic; leaves ca. 300 × 5–7 mm, ribbon-like, membranous; peduncles 40–50 cm long, ribbon-like, membranous; involucral and floral bracts pale brown with spongy apices; capitula 5–6 mm diameter. Small streams, 100–200 m; Amazonas (4 km south of Macuruco). Colombia (Vaupés). ŠFig. 6. Eriocaulon steyermarkii Mold., Fieldiana, Bot. 28: 117. 1951. Submerged or emergent aquatic rosette; leaves 40–120 × 1–2 mm, tapered; peduncle 10–30 cm long; capitulum 6.5–10 mm diameter, blackish, borne singly. Swamps, shallow

14

E RIOCAULACEAE

streams, muddy places, 500–2700 m; Bolívar (Gran Sabana), Amazonas (Sierra de la Neblina). Carabobo; Guyana, Suriname. ŠFig. 4. Eriocaulon tenuifolium Klotzsch ex Körn. in Mart., Fl. Bras. 3(1): 496. 1863. Eriocaulon brevifolium Klotzsch ex Körn. in Mart., Fl. Bras. 3(1): 496. 1863, non Raf. 1840. Eriocaulon atabapense Mold., Phytologia 2: 132. 1946. Eriocaulon klotzschii Mold., Phytologia 17: 484. 1968.

Perennial rosette herb; leaves 25–150 × (0.3–)1–3.5 mm, texture variable; peduncles 10–35 cm long; capitula 4–8.5 mm diameter, globose, firm, pubescent. Wet sandy savannas, streambanks, granitic outcrops, 50–200 m; Amazonas (Río Atabapo, Río Autana, Río Guainía, upper Río Orinoco, Río Sipapo, Río Ventuari). Colombia (Guainía), Guyana, Brazil (Amazonas, Roraima). ŠFig. 5.

Fig. 1. Eriocaulon cinereum

Fig. 2. Eriocaulon melanocephalum

Eriocaulon 15

Fig. 3. Eriocaulon humboldtii

16

E RIOCAULACEAE

Fig. 4. Eriocaulon steyermarkii

Fig. 5. Eriocaulon tenuifolium

Eriocaulon 17

1 mm

5 mm

6 mm

5 cm

Fig. 6. Eriocaulon spongiola

18

E RIOCAULACEAE

2. LEIOTHRIX Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 225. 1903. Rosulate perennial herbs. Roots usually aerenchymatous, rarely fibrous, never diaphragmed. Leaves needle-like to lanceolate. Inflorescences in axils of foliage leaves; glandular trichomes sometimes present on peduncle and sheath. Involucral bracts ovate to lanceolate, commonly acute to acuminate, pilose, orange- or redbrown; receptacle pilose, very often producing vegetative sprouts from the center; floral bracts always present. Flowers 3-merous, unisexual (plants monoecious). Sepals free in pistillate flowers, variously united in staminate flowers, commonly pilose, the trichomes acute and smooth within; floral axis elongating in staminate flowers of some species, sometimes densely pilose within the sepals or petals of pistillate flowers of some species; petals of pistillate flowers free, of staminate flowers free or fused. Stamens 3, the filaments free; anthers dithecous, white, basifixed. Ovary 3-locular; styles 3-angled, the style branches simple; nectaries usually inserted on style below the level of divergence of the style branches. Fruit a loculicidal capsule. Seeds finely longitudinally striate. South America; ca. 38 species, 2 in Venezuela, both in the flora area. Leiothrix celiae Mold., Mem. New York Bot. Gard. 9: 278. 1957. Rosulate clumping perennial with thick stems; leaves 65 × 3 mm, ciliate, aristate; peduncles 10–13 cm long; capitula ca. 5 mm diameter, numerous, pale brownish. Tepui meadows, ca. 2100 m; Amazonas (Cerro Yutajé). Endemic. ŠFig. 7. Leiothrix flavescens (Bong.) Ruhland, Pflanzenr. IV. 30(Heft 13): 231. 1903. —Eriocaulon flavescens Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 628. 1831. —Paepalanthus flavescens (Bong.) Körn. in Mart., Fl. Bras. 3(1): 423. 1863. Leiothrix umbratilis Mold., Fieldiana, Bot. 28: 119. 1951. Paepalanthus meseticola Mold. & Steyerm., Bol. Soc. Venez. Ci. Nat. 32– 33: 281. 1976. Paepalanthus fraternus var. marahuacensis Mold., Phytologia 49: 385. 1981. Leiothrix flavescens var. chimantensis Mold., Phytologia 54: 66. 1983. Rosulate perennial; leaves obtuse; inflorescences 1–few, pilose, orange- or redbrown. Venezuela, Guyana, Peru, Brazil; 4 varieties, 2 in Venezuela, both in the flora area. This widespread taxon is highly variable in the flora area. Clumping plants with very reduced lustrous leaves no more than 1–2 mm in diameter are found on the Macizo del

Fig. 7. Leiothrix celiae

Leiothrix 19

Fig. 8. Leiothrix flavescens var. flavescens

20

E RIOCAULACEAE

Chimantá, and perhaps merit recognition as Leiothrix flavescens var. chimantensis. This has not been done here because these dwarf forms grade continuously into sympatric forms of typical size. The plants at Cerro Marahuaka, described as Paepalanthus fraternus var. marahuacensis, are distinguished by coriaceous leaves, irregular peduncle length, and firm, deep brown, glabrous involucres and may deserve varietal status. Key to the Varieties of L. flavescens 1. Heads globose; involucral bracts reflexed and hidden at maturity; peduncles 3–14 cm long, ca. 2 times the length of the leaves; flowers (including pedicel) > 5 mm long ............................ var. alpina 1. Heads hemispherical; involucral bracts not reflexed, evident at maturity; peduncles 10–70 cm long, > 3 times the

length of the leaves; flowers < 4 mm long ..................................... var. flavescens L. flavescens var. alpina Mold., Mem. New York Bot. Gard. 9: 279. 1957. Leiothrix umbratilis var. brevipes Mold., Phytologia 32: 47. 1975. Leaves 14–70 mm long. Tepui summit, ca. 2400 m; Bolívar (Auyán-tepui). Endemic. L. flavescens var. flavescens Leaves 30–150 × 1–11 mm; capitula 6–10 mm diameter. Wet meadows and bogs, sandstone outcrops, rarely in tepui forest understories, 1100–2800 m; Bolívar (Cerro Guaiquinima, Cerro Guanay, Cerro Jaua, Gran Sabana including summits of all major tepui formations), Amazonas (summits of Cerro Marahuaka and Cerro Yaví, Sierra de la Neblina). Guyana, Peru, Brazil (Bahia south to Río de Janeiro, and Santa Catarina). ŠFig. 8.

3. PAEPALANTHUS Kunth, Enum. Pl. 3: 498. 1841, nom. cons. Paepalanthus Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 17: 13. 1835, nom. rejic. Herbs, acaulescent to caulescent, simple to branched; specialized fertile branches, when present, uniformly leafy; trichomes on vegetative parts simple or rarely T-shaped. Roots firm, cream to brown, the cortex never spongy or diaphragmed. Involucral bracts blackish to cream-colored, frequently pubescent; floral bracts present. Flowers 2- or 3-merous, unisexual (plants monoecious). Sepals free or briefly connate at base, frequently bearded at apex, the trichomes frequently clavate or tuberculate with inner walls ornamented, rarely acute and smooth within; in pistillate flowers of 3-merous species the sepals frequently becoming enlarged, thickened and hygroscopic at maturity, recurving when dry to eject the fruit; corolla stipe commonly present in staminate flowers; petals membranous, those of the staminate flowers connate into a tube, those of the pistillate flowers free. Stamens 2 or 3; filaments mostly free; anthers brown to white. Ovary usually 2- or 3locular, or 1-locular by abortion; style branches simple or usually bifid; nectaries present and inserted at about the level of the style branches. Fruit a loculicidal capsule, or in our area often 1-seeded by abortion and indehiscent. Mexico, Central America, West Indies, tropical and subtropical South America, a few species extending to Africa; ca. 500 species, 50 in Venezuela, 38 of these in the flora area. Paepalanthus apacarensis Mold., Mem. New York Bot. Gard. 9: 408. 1957. Dwarf, clumping perennial; stems to 1.5 cm long, densely foliose; leaves 5–6 × 1–1.5 mm, fleshy; peduncles 1.5–3 cm long; ca-

pitula 6.5 mm diameter. Moist rock ledges, ca. 2500 m; Bolívar (Macizo del Chimantá [Apacará-tepui]). Endemic. This species appears to be a dwarf derivative of Paepalanthus venustus.

Paepalanthus 21

Paepalanthus aristatus Mold., Phytologia 7: 122. 1960. Small delicate plant under 10 cm tall, forming low cushions or prostrate mats; leaves 6–9 × 0.5 mm; peduncles 0.3–2 cm long; capitula 4–5 mm diameter. Sandy banks, savannas, 100–200 m; Amazonas (along Caño Caname of the Río Atabapo, Río Orinoco just below Santa Bárbara). Endemic. ŠFig. 29. This species is very closely related to Paepalanthus dichotomus of the Gran Sabana. It is also related to, and appears to hybridize with, the sympatric P. yapacanensis. An intermediate specimen is Wurdack & Adderley 42874 (F, VEN). Paepalanthus auyantepuiensis Mold., Acta Biol. Venez. 2(7): 47. 1957. Caulescent perennial; stem 5–30 cm long; leaves 7–18 × 1–1.5 mm, linear, divaricate; peduncles 4.5–17 cm long; involucre blackish; capitula 6–9.5 mm diameter. Shaded canyon walls on tepui summit, 2200–2300 m; Bolívar (Auyán-tepui). Endemic. This species is closely related to Paepalanthus venustus, with which it grows in mixed populations. It may prove to be a product of gene exchange with the sympatric species P. squamuliferus. A strongly intermediate specimen is Steyermark 116119 (F). Paepalanthus bifidus (Schrad.) Kunth, Enum. Pl. 3: 512. 1841. —Eriocaulon bifidum Schrad. in Schult., Mant. 2: 468. 1824. Erect, membranous annual; stems 2.5–7 cm long, leafy; leaves 7–25 × 1–2.5 mm; peduncles 1.5–4(–7) cm long; capitula 3–4 mm diameter, buff-colored, in a terminal cluster. Sandy soils of rock outcrops, white-sand savannas, 100–1000 m; Bolívar (Arekuna, Guayaraca, Sabana de Triana in Río Chicanán basin), Amazonas (Cerro Yutajé). Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian and central Brazil. ŠFig. 23. Paepalanthus cardonae Mold., Phytologia 3: 39. 1948. Subcaulescent perennial; stems to 7 cm long; leaves 20 × 2.5–3 mm, lanceolate, acute; peduncles 5–6 cm long; capitula small,

numerous, dark brown. Sandstone outcrops, ca. 2000 m; Bolívar (Macizo del Chimantá [Acopán-tepui]). Endemic. This species is very close to Paepalanthus scopulorum, essentially differing only in the subcaulescent habit, and may only represent a depauperate environmental form or local variety of that species. Paepalanthus chimantensis Hensold, Ann. Missouri Bot. Gard. 78: 428, figs. 3F, 3G, 4. 1991. Rosulate rhizomatous perennial; leaves 30–40 × 3–8 mm, lustrous, aristate; peduncles 13–30 cm long; capitula 6.5–9 mm diameter, gray-brown. Wet areas in open tepui meadows, 2100–2600 m; Bolívar (Macizo del Chimantá [Apacará-tepui]). Endemic. ŠFig. 16. Paepalanthus convexus Gleason, Bull. Torrey Bot. Club 58: 328. 1931. Erect perennial; stem to 40 cm tall; leaves 25–42 × 2–2.5 mm, linear, reflexed; peduncles 10–14 cm long; capitula 6–7.5 mm diameter, numerous, dark brown. Moist bluff faces and ridge tops on tepui summit, 1500– 2000 m; Amazonas (Cerro Duida). Endemic. Paepalanthus cristatus Mold., Brittonia 3: 157. 1939. Small, rosulate, clumping, perennial herb; leaves (7–)10–20 × 0.7–5 mm, leathery, obtuse; peduncles (0.5–)1.2–5 cm long; capitula 4–6 mm diameter. Sandy or organic soil on sandstone outcrops, 1900–2400 m; Bolívar (Auyán-tepui). Endemic. ŠFig. 28. Paepalanthus cumbricola Mold., Mem. New York Bot. Gard. 9: 409. 1957. Robust perennial; stems 4–21 cm, erect, densely foliose; leaves 20–45 × 2–6 mm, tomentose, acute; peduncles 11–28 cm long; capitula 8–11 mm diameter. Sandstone ledges, escarpment faces, shaded tepui slopes, 2000–2500 m; Bolívar (Macizo del Chimantá). Endemic. ŠFig. 9. Paepalanthus dichotomus Klotzsch ex Körn. in Mart., Fl. Bras. 3(1): 348. 1863. Paepalanthus guyanensis Klotzsch ex Körn. in Mart., Fl. Bras. 3(1): 347. 1863.

22

E RIOCAULACEAE

Syngonanthus savannarum Mold., Bull. Torrey Bot. Club 75: 202. 1948. —Paepalanthus savannarum (Mold.) Mold., Phytologia 49: 293. 1981. Paepalanthus steyermarkii Mold., Fieldiana, Bot. 28: 125. 1951. Syngonanthus venezuelensis Mold., Fieldiana, Bot. 28: 128. 1951. Syngonanthus savannarum var. glabrescens Mold., Mem. New York Bot. Gard. 9: 412. 1957. —Paepalanthus savannarum var. glabrescens (Mold.) Mold., Phytologia 49: 293. 1981. Paepalanthus apacarensis var. humilis Mold., Phytologia 54: 66. 1983. Plant low, much-branched, forming cushions or mats, glabrate to densely villous. Venezuela, Guyana, Peru; 3 varieties, 2 in Venezuela, both in the flora area. The lengthy synonymy for this species may be explained by the extreme, probably environmentally induced, variation between local populations, especially with respect to pubescence, stature, leaf length, and peduncle elongation. Key to the Varieties of P. dichotomus 1. Leaves 6–15 mm long (including sheath) × 0.7–1 mm wide ...... var. dichotomus 1. Leaves 2–3 × 0.5 mm ....... var. pumilus P. dichotomus var. dichotomus Cushion plant 3–6 cm tall; peduncles 0.3– 4(–5.5) cm long; capitula 3–5.5(–6.5) mm diameter. Sandy savannas and tepui meadows, 1200–2100 m; Bolívar (Gran Sabana, Macizo del Chimantá). Guyana. ŠFig. 31. P.

dichotomus var. pumilus Mold., Phytologia 49: 385. 1981. Cushion plant ca. 1 cm tall; peduncles 0.5–0.9 cm long; capitula 3–4 mm diameter. White-sand savannas, 1100–1200 m; Bolívar (Kavanayén). Endemic. Paepalanthus fasciculatus (Rottb.) Kunth, Enum. Pl. 3: 506. 1841. —Eriocaulon fasciculatum Rottb., Acta Lit. Univ. Hafn. 271, t. 2, fig. 1. 1778. Paepalanthus congestus Kunth, Enum. Pl. 3: 505. 1841.

Small annual with upright, leafy stems; leaves 5–25 × 0.5–1 mm; peduncles 2–8 cm long; capitula 2–3 mm diameter, globose to cylindrical, buff-colored, in terminal umbels. Wet areas of white-sand savannas, disturbed sites, 50–1000 m; scattered throughout Bolívar and Amazonas, most frequent in lowlands of southwestern Amazonas. Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ŠFig. 21. Paepalanthus fasciculoides Hensold, Ann. Missouri Bot. Gard. 78: 431, figs. 3C–E, 5. 1991. Annual herb; stems to 5 cm, erect, leafy; leaves 5–10 × 0.5–1 mm; peduncles 2–5 cm long; capitula 2–3 mm diameter, dark brown, clustered at the apex of the stem. Rock outcrops, ca. 600 m; Bolívar (Cerro Cotorra east of La Paragua). Brazil (Pará). ŠFig. 25. Paepalanthus fasciculoides is very similar to the common P. fasciculatus but is distinguished by its fuscous (blackish brown) involucral bracts. Paepalanthus formosus Mold., Bol. Soc. Venez. Ci. Nat. 14: 11. 1952. Paepalanthus moldenkeanus R. Schult., Bot. Mus. Leafl. 16: 187, t. 23, 24. 1954. Robust monocarpic herb from basal rosette with flowering stem 0.5–1.5 m tall; basal leaves 130–180 × 8 mm; peduncles 15– 35 cm long; capitula 6–11 mm diameter, these arranged in a single large spherical umbel. Lowlands (ca. 100 m) in swales in sandy savannas, in uplands (1000–2000 m) in shrublands on sandstone plateaus; Bolívar (Cerro Guanay), Amazonas (Canaripó, Cerro Coro Coro, Cerro Duida, Cerro Yutajé, Santa Bárbara). Colombia, Brazil (Goiás, Maranhão, Mato Grosso, Minas Gerais). ŠFig. 10. According to A.M. Giuletti (personal communication), Paepalanthus formosus and P. moldenkeanus R. Schult., described as endemic to Venezuela and Vaupés, Colombia, respectively, are conspecific with the central Brazilian species P. speciosus (Bong.) Körn. However, this name is illegitimate, and the correct name is not yet determined. Paepalanthus fraternus N.E. Br., Trans. Linn. Soc. London, Bot. 6: 69. 1901.

Paepalanthus 23

Rosulate perennial; leaves (15–)22–42 × (1.5–)2–3(–3.5) mm, falcate; inflorescences 1–few per rosette; peduncles (6–)10–28 cm long; capitula 8–11 mm diameter. Locally abundant on rocky open wet sites of tepui summits, 2200–2800 m; Bolívar (Ilú-tepui, Kukenán-tepui, Roraima-tepui). Guyana (Mount Roraima). Paepalanthus fulgidus Mold., Mem. New York Bot. Gard. 9: 279. 1957. Paepalanthus perplexans var. steyermarkii Mold., Phytologia 44: 384. 1979. Robust perennial; stem 7–20 cm tall; leaves (20–)30–53 × (3–)4–6(–7) mm, silveryhairy, obtuse; peduncles (11–)15–30 cm long; capitula (5–)6.5–10 mm diameter. Shrub islands, thickets around boulders, tepui forests, often associated with Bonnetia roraimae, 1900–2700 m; southeastern Bolívar (Macizo del Chimantá, Roraima-tepui area north and west to Sierra de Lema and Kamarkawarai-tepui). Endemic. ŠFig. 19. The plants of the Chimantá massif are irregular and somewhat transitional morphologically with Paepalanthus scopulorum (Abacapá-tepui) and P. schomburgkii (Aprada-tepui). Paepalanthus gleasonii Mold., Phytologia 2: 140. 1946. Paepalanthus robustus Gleason, Bull. Torrey Bot. Club 58: 330. 1931, non Silveira 1908. Paepalanthus convexus var. strigosus Mold., Phytologia 35: 277. 1977. Caulescent perennial; stem 8–40 cm tall; leaves 25–60 × 3–5 mm, linear, ciliate; peduncle 22–47 cm long; capitulum (6–)9–16 mm diameter, robust, solitary on long peduncles. Open tepui meadows and slopes, sandy riverbanks, 1200–2300 m; Amazonas (summits of Cerro Duida and Sierra de la Neblina). Brazil (Amazonas). The type specimen, from Cerro Duida, is an extra robust, fleshy, large-capitate individual, or “gigas” form, common in the Eriocaulaceae. The variation in this species suggests hybridization with one or more sympatric species (probably Paepalanthus major and P. sulcatus), but more study is needed.

Paepalanthus holstii Steyerm., Ann. Missouri Bot. Gard. 75: 311. 1988. Robust caulescent perennial; stems 5–30 cm tall; leaves 40–50 × 10–14 mm, broadly lanceolate, sericeous, mucronate; peduncles 10–16 cm long; capitula 7–10 mm diameter. Shaded sandstone crevices on tepui summit, 2300–2500 m; Bolívar (Murisipán-tepui). Endemic. Paepalanthus kunhardtii Mold., Mem. New York Bot. Gard. 8: 97. 1953. Subcaulescent rhizomatous herb; stem to 8 cm tall; leaves 38–52 × 3.5–5.5 mm, lanceolate to ensiform; peduncles 11–25 cm long; capitula 7.5 mm diameter. Wet mossy hummocks, open riversides, 1500–2100 m; Bolívar (Sierra de Maigualida), Amazonas (Cerro Cuao, Cerro Sipapo). Endemic. ŠFig. 14. Paepalanthus lamarckii Kunth, Enum. Pl. 3: 506. 1841. Eriocaulon fasciculatum Lam., Encycl. 3: 276, t. 50, fig. 3. 1789, non Rottb. 1778. Small unbranched annual, typically pilose, with a cluster of numerous grayish inflorescences terminating the stem; stem to 5 cm tall; leaves 5–35 × 0.5–3 mm; peduncles 1.5–8.5 cm long; capitula 2–3.5 mm diameter. Wet soil of roadcuts, riverbanks, swampy depressions in savannas, thin soil of granitic outcrops, 50–300 m; Bolívar (Altiplanicie de Nuria, lower Río Caroní, Río Orinoco), Amazonas (between Puerto Ayacucho and El Burro). Apure, Guárico, Monagas, Táchira; Central America (north to Belize), West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Africa. ŠFig. 24. Paepalanthus major (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 432. 1991. —Paepalanthus convexus var. major Mold., Phytologia 15: 463. 1968. Paepalanthus fulgidus var. zuloagensis Mold., Phytologia 23: 211. 1972. Subcaulescent perennial; stem 0–17 cm tall; inflorescences robust, solitary; leaves 20–60 × 2–5 mm, silky-pubescent; peduncle (15–)25–54 cm long; capitulum 7–11 mm diameter. Tepui meadows, 1300–2300 m;

24

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Amazonas (Sierra de la Neblina). Brazil (Amazonas: Pico da Neblina). This species is part of a variable complex that includes Paepalanthus gleasonii. They are maintained here as separate species because strongly differentiated populations of the two species have been collected on the Venezuelan part of Sierra de la Neblina. In the Brazilian collections observed, the difference in stem elongation between these two species appears to break down. See Paepalanthus gleasonii. Paepalanthus oyapockensis Herzog, Feddes Repert. Spec. Nov. Regni Veg. 29: 206, pl. 121, figs. f–h. 1931. Paepalanthus leucocyaneus Tutin, J. Bot. 72: 336. 1934. Paepalanthus brunneus Mold., Bull. Torrey Bot. Club 75: 195. 1948. Paepalanthus griseus Mold., Bull. Torrey Bot. Club 75: 197. 1948. Paepalanthus tafelbergensis Mold., Bull. Torrey Bot. Club 75: 199. 1948. Small, lax, membranous herb; stems 6 (–17) cm long; inflorescences numerous; leaves 10–47 × 1–5 mm, ligulate, mucronate; peduncles 5.5–25 cm long; capitula 3–5 mm diameter. Wet or mucky places on granitic outcrops, also reported from sandy banks and pools, 100–500 m; Bolívar (Canaima, Río Parguaza), northwestern Amazonas. Guyana, Suriname, Brazil (Amazonas, Pará, Amapá). ŠFig. 18. Paepalanthus parvicephalus (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 432. 1991. —Paepalanthus convexus var. parvicephalus Mold., Phytologia 52: 19. 1982. Erect or procumbent, long-stemmed perennial with obtuse leaves. Venezuela; 2 varieties, both endemic to Amazonas. Key to the Varieties of P. parvicephalus 1. Leaves usually villous on adaxial tips, at least when young; heads 6.5–8 mm diameter .................. var. parvicephalus 1. Leaves ciliate, but otherwise glabrous; heads 4.5–5 mm diameter ....................... ..................................... var. wurdackii

P. parvicephalus var. parvicephalus Stem 10–40 cm long; leaves (15–)17–30 (–35) × (1.5–)2–3 mm; peduncles 12–20 cm long; capitula 6.5–8 mm diameter. Wet places below rocks and open thickets by forest edge on high tepui summit, 2500–2800 m; Amazonas (Cerro Marahuaka). Endemic. For discussion of possible hybrids, see Rondonanthus duidae. P. parvicephalus var. wurdackii (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 432. 1991. —Paepalanthus perplexans var. wurdackii Mold., Mem. New York Bot. Gard. 8: 98. 1953. Stem ca. 19 cm long; leaves 23–25 × 2.2– 2.5 mm; peduncles 19–24 cm long; capitula 4.5–5 mm diameter. Tepui meadows, ca. 2000 m; Amazonas (Cerro Parú). Endemic. ŠFig. 17. Paepalanthus perpusillus Kunth, Enum. Pl. 3: 503. 1841. Small, rosulate, glabrous annual; leaves 5–15 × 0.5–1.2 mm; peduncles 0.7–4 cm long; capitula 1.5–3 mm diameter, numerous, blackish. Sandy seasonally flooded banks and sandbars, ca. 100 m; Amazonas (near mouth of Río Emoni in Río Casiquiare basin). Apure; Brazil (Amazonas, Minas Gerais, Pará). Paepalanthus phelpsiae Mold., Mem. New York Bot. Gard. 23: 852, fig. 6. 1972, “phelpsae.” Robust perennial; stem to 20 cm tall; leaves 55–82 × 8–10 mm, lanceolate, scabrous; peduncles 15–45 cm long; capitula 7.5–9 mm diameter. Sandstone outcrops, meadows, and streambanks on tepui summit, 1800–2200 m; Bolívar (Cerro Jaua). Endemic. ŠFig. 13. Paepalanthus polytrichoides Kunth, Enum. Pl. 3: 507. 1841. Paepalanthus viscosus Mold., Bull. Torrey Bot. Club 68: 70. 1941. Syngonanthus guianensis Mold., Bull. Torrey Bot. Club 75: 201. 1948. Paepalanthus vigiensis Mold., Phytologia 3: 170. 1949. Paepalanthus villipes Mold., Phytologia 3: 171. 1949.

Paepalanthus 25

Much-branched perennial herb forming wiry cushions or prostrate mats; stem to 5 cm tall; leaves 3–10 × 0.25–1 mm; peduncles 2–10 cm long, filiform; capitula 3–4 mm diameter, globose, gray-white, numerous. White-sand streambanks and savannas, 100–1200 m; scattered throughout Bolívar and Amazonas. Guyana, Suriname, Peru, Brazil (Amapá, Amazonas, Mato Grosso, Pará). ŠFig. 26. Paepalanthus polytrichoides is allied to the widely ranging annual species P. subtilis, and although these are generally distinct taxa, they intergrade in Venezuela, especially at Cerro Guaiquinima. The Guaiquinima plants are intermediate with respect to leaf length, pubescence, and involucral bract form, but are here treated as P. polytrichoides due to branching of the stem above the inflorescences. Paepalanthus roraimensis Mold., Fieldiana, Bot. 28: 121. 1951. Rondonanthus micropetalus Mold., Fieldiana, Bot. 28:126. 1951. Robust perennial; stem 10–40 cm tall; inflorescences numerous; leaves 35–70 × 4.5– 10 mm, lanceolate, with coriaceous apices; peduncles 9–20 cm long; capitula 7–10 mm diameter. Sandstone bluffs, open rocky areas and ravines on tepui summit, 2200–2800 m; Bolívar (summit of Roraima-tepui). Guyana (Mount Roraima). This species may be of hybrid derivation. It displays certain developmental irregularities, such as early necrosis of leaf tips, uneven involucral bract color and irregular form of bracts. The population upon which Rondonanthus micropetalus is based is characterized by a failure of the peduncles to elongate (not uncommon in eriocaul hybrids) and by abortion of the corollas in the staminate flowers. The closest relative of Paepalanthus roraimensis is P. schomburgkii, which ranges widely in the Gran Sabana and adjacent Guyana, but has only been collected once on Roraima-tepui (the type). The differences between these two taxa suggest probable gene exchange with the sympatric P. fraternus (Roraima-tepui). Paepalanthus schomburgkii Klotzsch ex Körn. in Mart., Fl. Bras. 3(1): 375. 1863.

Paepalanthus perplexans Mold., Fieldiana, Bot. 28: 120. 1951. Paepalanthus pendulus Mold., Mem. New York Bot. Gard. 9: 280. 1957. Paepalanthus macrocaulon var. venamensis Mold., Bol. Soc. Venez. Ci. Nat. 26: 411. 1966. Erect or clambering perennial; stem 3–55 cm long; inflorescences several at a node; leaves 25–70(–80) × 3–10(–14) mm, fine-textured, with acute apices; peduncles (5–)12– 35 cm long; capitula 4–8(–10) mm diameter. Open boggy areas, Bonnetia forests, sandstone ledges, talus slopes, along streamsides, in waterfall spray and on dripping bluffs, 500–2400 m; Bolívar (Amaruay-tepui, Aparamán-tepui, Cerro Venamo, Cerro El Sol, Ilú-tepui, Macizo del Chimantá [Chimantátepui], Ptari-tepui, Roraima-tepui). Guyana. ŠFig. 15. Paepalanthus schomburgkii varies considerably in size and habit, with robust subcaulescent plants in open boggy areas of the Roraima formation, large long-stemmed plants usually in shady understory, and small membranous clumps on wet bluffs and ledges. The type of P. pendulus, collected at 2400 m on Ilú-tepui, is atypical, with the large abnormally developing inflorescences similar to those of some specimens of P. roraimensis. It is included here in P. schomburgkii because of its vegetative similarity to collections of that species from Ilú-tepui. Although common on tepui summits in Bolívar, the type (Schomburgk 1026, B) is the only known record from Roraima-tepui, reportedly from the south slope. Paepalanthus scopulorum Mold., Fieldiana, Bot. 28: 122. 1951. Paepalanthus scopulorum var. auyantepuiensis Mold., Acta Bot. Venez. 2(5– 8): 153. 1967. Clambering perennial; stem 15–60 cm long; inflorescences numerous; leaves 20–75 × 3–10 mm, recurved, conduplicate; peduncles 5–12(–14) cm long; capitula 5–8 mm diameter. Commonly hanging from dripping or misty escarpment faces, or in talus cloud forests or on wet meadows, 1400–2400 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Macizo del Chimantá, Ptari-tepui). Endemic. ŠFig. 11.

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This is a fairly uniform species, though plants of Auyán-tepui have heads 7–8 mm in diameter, while at other localities the heads are 5–6 mm. Hybridization may be occurring with Paepalanthus schomburgkii; an intermediate is Wurdack 34275 (VEN), pro parte, from Macizo del Chimantá. Also see P. cardonae. Paepalanthus sessiliflorus Körn. in Mart., Fl. Bras. 3(1): 361. 1863. Diminutive herb with simple leafy stems and glomerules of sessile white heads. Venezuela, Guyana, Brazil; 2 varieties, 1 of these in the flora area. P. sessiliflorus var. venezuelensis Mold., Phytologia 28: 193. 1974. Paepalanthus lilliputianus Mold., Phytologia 3: 115. 1949. Stem to 1.5 cm long; leaves 3–5 × 0.25–0.5 mm; capitula 2–3 mm diameter, sessile. White-sand savannas, rock outcrops, riverbanks, 400–1100 m; eastern Bolívar (Arekuna, Canaima, Salto Kamá in Río Aponguao basin, 15 km north-northwest of Santa Elena de Uairén). Guyana, Brazil (Amazonas). ŠFig. 30. Paepalanthus squamuliferus Mold., Fieldiana, Bot. 28: 124. 1951, “Paepalanthus squamiliferus.” Paepalanthus fraternus var. spathulatus Mold., Phytologia 49: 386. 1981. Subcaulescent perennial bearing 1–few inflorescences per cohort; stem 0–15 cm tall; leaves 20–50 × 1.5–4.5 mm, linear, semisucculent; peduncles (8–)11–40 cm long; capitula 8–12(–14) mm diameter. Moist meadows, sandstone outcrops, rocky streambanks, crevices of tepui summits, 1600–2500 m; Bolívar, (Aparamán-tepui, Auyán-tepui, Macizo del Chimantá, Ptari-tepui). Endemic. Paepalanthus stegolepoides Mold., Mem. New York Bot. Gard. 9: 409. 1957. Paepalanthus fraternus var. chimantensis Mold., Phytologia 54: 66. 1983. Perennial forming low branched clumps; stems to 8 cm long; leaves 18–40 × 1.5–2.5 mm, reflexed, linear, with a bluish metallic sheen; peduncles 6–16 cm long; capitula 5.5– 7.5 mm diameter. Shaded sites, especially

under sandstone ledges on tepui summit, 1800–2500 m; Bolívar (Macizo del Chimantá). Endemic. ŠFig. 20. Paepalanthus subcaulescens N.E. Br., Trans. Linn. Soc. London, Bot. 6: 71. 1901. Clumping perennial; stem to 4 cm long; leaves 20–40 × 4–5 mm, lance-linear, acute; peduncles 6–14 cm long; capitula 4–7 mm diameter. Expected around the base of Roraima-tepui in Bolívar state (currently known only from the Cotinga River valley on the Guyanese side of Mount Roraima). Paepalanthus subtilis Miq., Stirp. Surinam. Select. 221. 1851. Paepalanthus pauperrimus Herzog, Feddes Repert Spec. Nov. Regni Veg. 29: 206. 1931. Erect unbranched annual; stem 0.2–5 cm long; inflorescences numerous, clustered at stem apex, each borne on a minute bracteate short shoot; leaves 5–15 × 0.3–1 mm; peduncles 2.5–6 cm long; capitula 2–3 mm diameter. Sandy soils of scrubby savannas, dry forests, streambanks, roadsides, among rock outcrops, often in depressions or locally moist areas, 50–1200 m; widespread in Bolívar and Amazonas. Guárico; Guyana, Suriname, Brazil (Amapá, Amazonas, Bahia, Maranão, Pará). ŠFig. 27. See discussion of Paepalanthus polytrichoides. Paepalanthus sulcatus Hensold, Ann. Missouri Bot. Gard. 76: 958. 1989. Paepalanthus stegolepoides var. acutalis Mold., Phytologia 15: 463. 1968. Caulescent perennial; stem to 15 cm long; inflorescences several per stem; leaves (30–) 35–55 × 2.5–4 mm, lance-linear, with engraved veins; peduncles 15–25 cm long; capitula 5.5–10 mm diameter. Open or wooded slopes and tepui meadows, 1500–3000 m; Amazonas (summits of Cerro Aracamuni, Cerro Avispa, Cerro Marahuaka, and Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). This species varies significantly between localities. The type material from Brazil differs by its nearly glabrous leaves, apiculate bracts, and larger heads. The plants found

Paepalanthus 27

on Cerro Aracamuni are rather longstemmed. The population on Cerro Marahuaka with dull nearly obtuse leaves intergrades with Paepalanthus parvicephalus, also found on Cerro Marahuaka. Paepalanthus tortilis (Bong.) Körn. in Mart., Fl. Bras. 3(1): 354. 1863. —Eriocaulon tortile Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 624, t. 49. 1831, non Steud. 1855. Paepalanthus salticola Herzog, Feddes Repert. Spec. Nov. Regni Veg. 29: 207, pl. 121, figs. i-l. 1931. Paepalanthus tatei Mold., Brittonia 3: 158. 1939. Paepalanthus killipii Mold., Bull. Torrey Bot. Club 68: 67. 1941. Paepalanthus maguirei Mold., Bull. Torrey Bot. Club 75: 198. 1948. Rosulate to caulescent, soft-textured, weedy herb of extremely variable habit and stature; stem 0–36 cm long; leaves 10–60 × 1–10 mm; peduncles 5–26(–31) cm long; capitula 2.5–6(–8) mm diameter, brittle, blackish. Sandy streambanks, forests, seeps and ledges, 100–2200 m; Bolívar (Gran Sabana), Amazonas (widespread). Lara, Táchira; Colombia, Guyana, Suriname, Brazil (Amazonian and eastern). ŠFig. 12. This common species varies widely in size and habit, with plants ranging from tiny rosettes with leaves 8 × 1 mm and peduncles of 5 cm, to clambering long-stemmed herbs with leaves 60 × 10 mm, and peduncles to 30 cm long. However, the inflorescence size varies by a smaller factor, mostly 2.5– 6(–8) mm diameter. Within Venezuela, plants found on Gran Sabana, regardless of size, are firmer in texture and with more rounded leaf margins than plants of Amazonas or the Andes. Paepalanthus turbinatus (Gleason) Hensold, Ann. Missouri Bot. Gard. 78: 433. 1991. —Leiothrix turbinata Gleason, Bull. Torrey Bot. Club 58: 331. 1931. Clumping rhizomatous herb; stem to 10 cm long; leaves 55–86 × 3–4.5 mm, lanceolate, pilose; peduncles 18–33 cm long; capitula 6–9 mm diameter. Moist meadows on

tepui slopes and summits, 1500–2100 m. Amazonas (Cerro Duida). Endemic. Paepalanthus venustus Mold., Mem. New York Bot. Gard. 9: 281. 1957. Paepalanthus venustoides Mold., Acta Biol. Venez. 2(7): 48. 1957. Lax perennial; stem 5–30 cm long; inflorescences few per stem; leaves 7–17 × 1–2.5 mm, recurved, lustrous; peduncles 8–20 cm long; capitula 7–10 mm diameter. Canyon walls, tepui forest understories and boggy meadows, 1500–2500 m; Bolívar (Auyántepui, Cerro Guaiquinima, Macizo del Chimantá). Endemic. ŠFig. 22. See Paepalanthus apacarensis and P. auyantepuiensis. Paepalanthus yapacanensis (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 433. 1991. —Syngonanthus yapacanensis Mold., Mem. New York Bot. Gard. 8: 102. 1953. Mat-forming perennial to 2.5 cm tall; inflorescences numerous, the heads obconic; leaves 5–24 × 0.5–1.5 mm, linear, firm; peduncles 3.5–11.5 cm long; capitula 3.5–6 mm diameter. White-sand savannas, ca. 100 m. Venezuela; 2 varieties, both in the flora area. Endemic. Key to the Varieties of P. yapacanensis 1. Peduncles soft-villous, usually densely so, with long- spreading or appressed trichomes ............................ var. hirsutus 1. Peduncles glabrous .................................... ............................... var. yapacanensis P. yapacanensis var. hirsutus (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 434. 1991. —Syngonanthus yapacanensis var. hirsutus Mold., Phytologia 36: 51. 1977. Amazonas (upper Río Orinoco and lower Río Ventuari). Endemic. ŠFig. 32. This variety has not been collected from the same localities as var. yapacanensis. P. yapacanensis var. yapacanensis Amazonas (base of Cerro Yapacana, Río Guayapo, Río Siapa and its tributaries). Endemic. See Paepalanthus aristatus.

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Fig. 9. Paepalanthus cumbricola

Fig. 10. Paepalanthus formosus

Paepalanthus 29

Fig. 11. Paepalanthus scopulorum

Fig. 12. Paepalanthus tortilis

30

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Fig. 13. Paepalanthus phelpsiae

Fig. 14. Paepalanthus kunhardtii

Paepalanthus 31

5 mm

Fig. 15. Paepalanthus schomburgkii

Fig. 16. Paepalanthus chimantensis

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Fig. 17. Paepalanthus parvicephalus var. wurdackii

Fig. 19. Paepalanthus fulgidus

Fig. 18. Paepalanthus oyapockensis

Fig. 20. Paepalanthus stegolepoides

Paepalanthus 33

Fig. 21. Paepalanthus fasciculatus

Fig. 22. Paepalanthus venustus

Fig. 23. Paepalanthus bifidus

34

E RIOCAULACEAE

Fig. 24. Paepalanthus lamarckii

Fig. 25. Paepalanthus fasciculoides

3 mm

Fig. 26. Paepalanthus polytrichoides

Fig. 27. Paepalanthus subtilis

Fig. 28. Paepalanthus cristatus

Philodice 35

Fig. 29. Paepalanthus aristatus

Fig. 31. Paepalanthus dichotomus var. dichotomus

Fig. 30. Paepalanthus sessiliflorus var. venezuelensis

Fig. 32. Paepalanthus yapacanensis var. hirsutus

4. PHILODICE Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. 17(1): 16, t. 3. 1835. Small herbs, possibly annuals, with erect elongate densely foliose stems. Roots aerenchymatous, not diaphragmed. Leaves narrowly linear. Inflorescences clustered in upper leaf axils; peduncle sheaths lacking; involucral bracts lance-linear, acute, surpassing or equaling the disk, the floral bracts similar but narrower. Flowers 3-merous, unisexual (plants monoecious). Sepals lance-ovate, acuminate, white, glabrous; short corolla stipe present in staminate flowers; petals in staminate flowers connate into a low, urceolate tube, in pistillate flowers connate medially, free at base and apex, in both sexes acuminate, white, glabrous, firm, non-involute. Sta-

36

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mens 3; filaments free; anthers with 1 theca, purplish. Ovary 3-locular; style branches simple; nectaries present and inserted at level of divergence of style branches. Fruit a loculicidal capsule. Colombia, Venezuela, Guyana, Suriname, central and Amazonian Brazil, Bolivia; 2 species, 1 of these in the flora area. Philodice hoffmannseggii Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. 17(1): 17, t. 3. 1835. Small annual with clusters of capitula terminating stem; stem to 10 cm long, erect, foliose; leaves 5–15 × 0.25 mm; peduncles 0.1–1.2 cm long; capitula 2.5–3.5 mm diameter, lustrous white. Wet sandy savannas, floodplains, granitic outcrops, Mauritia palm swamps, 50–200 m; western Bolívar, Amazonas (Río Samariapo). Apure, Cojedes, Guárico; Colombia, Guyana, Suriname, Brazil, Bolivia. ŠFig. 33.

Fig. 33. Philodice hoffmannseggii

5. RONDONANTHUS Herzog, Feddes Repert. Spec. Nov. Regni Veg. 29: 210. 1931. Wurdackia Mold., Mem. New York Bot. Gard. 9: 413. 1957. Cespitose rosulate or caulescent perennial herbs. Roots white to pale brown, aerenchymatous, brittle. Leaves spirally arranged or distichous, filiform to ligulate, the bases lanate, the distal portions glabrous or pubescent with T-shaped trichomes. Inflorescences axillary; peduncle sheaths scarious at the upper margin, often lacerate; peduncles glabrous or pubescent with T-shaped trichomes; involucral bracts (or at least their exposed apices) lanceolate, acute to subacute, gold to black or variegated, variously pubescent; floral bracts present. Flowers 3-merous, bisexual and staminate (andromonoecious) or more commonly all unisexual. Staminate flowers: sepals free, pubescent abaxially at apex; petals free or connate into a 3-lobed tube and then sometimes separating after anthesis, glabrous or ciliate at upper margin; stamens with filaments often adnate to corolla at base; anthers dorsifixed, whitish. Pistillate and bisexual flowers: sepals as in staminate flowers, but thickening and expanding with the maturation of the fruit; petals free or connate distally, glabrous or pilose; linear staminodes, or more rarely fully developed stamens present opposite petals. Ovary usually subtended by hairs at base; style branches strongly bifid, inserted at the same level as or slightly above the papillose nectaries. Fruit a loculicidal capsule. Endemic to the Guayana Shield of Venezuela, Guyana, and Brazil; 6 species, all in the flora area. Rondonanthus acopanensis (Mold.) Hensold & Giul., Ann. Missouri Bot. Gard. 78: 447. 1991. —Syngonanthus acopanensis Mold., Phytologia 3: 41, fig. 1. 1948. Perennial herb; stem 1–4 cm long; leaves 30–130 × 1–4(–6) mm, distichous, leathery, with persistent woolly bases; peduncles 12– 45(–60) cm long; involucral bracts blackish

with hyaline apices; capitula 7.5–12(–15) mm diameter. Venezuela; 2 varieties endemic to the flora area. This species occurs in two discrete size classes, sympatric and about equally common, differing by leaf width, diameter of heads, size of flowers, and possibly habitat preferences. They are here treated as varieties.

Rondonanthus 37

Key to the Varieties of R. acopanensis 1. Leaves 1–2 mm wide; heads 7.5–10 mm in diameter; involucral bracts mostly about 3-seriate ................... var. acopanensis 1. Leaves (2–)2.8–6 mm wide; heads 10–14 (–15) mm in diameter; involucral bracts mostly about 4- or 5-seriate .................... .................................. var. obtusifolius R. acopanensis var. acopanensis Leaves 1–2 mm wide; capitula 7.5–10 mm diameter. Locally abundant on wet sandy depressions and banks on shallow soils, in the open or in scrub forests, 1900–2600 m; Bolívar (Angasima-tepui, Macizo del Chimantá). Endemic. ŠFig. 37. R. acopanensis var. obtusifolius (Mold.) Hensold & Giul., Ann. Missouri Bot. Gard. 78: 448. 1991. —Syngonanthus obtusifolius Mold., Mem. New York Bot. Gard. 9: 410. 1957. Leaves (2–)2.8–6 mm wide; capitula 10– 15 mm diameter. Locally abundant in open wet savannas and among sandstone outcrops, 1800–2200 m; Bolívar (Macizo del Chimantá). Endemic. One exceptionally large form, represented by Huber 11552 (MO), differs from other material by the leaves arranged spirally in a rosette rather than distichously. Rondonanthus capillaceus (Klotzsch ex Körn.) Hensold & Giul., Ann. Missouri Bot. Gard. 78: 448, fig. 2. 1991. —Paepalanthus capillaceus Klotzsch ex Körn. in Mart., Fl. Bras. 3(1): 415, t. 53, fig. II. 1863. Paepalanthus capillaceus var. proliferus Gleason, Bull. Torrey Bot. Club 58: 328. 1931. —Paepalanthus capillaceus f. proliferus (Gleason) Mold., Phytologia 44: 384. 1979. Submerged aquatic of flowing water, with coarse, much-branched roots; leaves 70–300 × 0.2–0.4 mm, filiform; peduncles 5.5–20 cm long, emergent; capitula 3–5 mm diameter. Adhering to rocks of streambeds, 100–2400 m; Bolívar (Gran Sabana), Amazonas (Cerro Duida, Cerro Marahuaka, Río Mawarinuma, Río Siapa). Guyana, Brazil (Amapá, Amazonas). ŠFig. 35.

Rondonanthus caulescens (Mold.) Hensold & Giul., Ann. Missouri Bot. Gard. 78: 451. 1991. —Paepalanthus jauensis var. caulescens Mold., Phytologia 44: 215. 1979. Caulescent perennial; stem 4–6 cm long; leaves 30–40 × 0.6–1 mm, linear, densely woolly at base; peduncles 15–35 cm long; involucral bracts with hyaline apices; capitula 7.5–10 mm diameter. Open areas on tepui summit, ca. 2500 m; Bolívar (Aprada-tepui). Endemic. Rondonanthus duidae (Gleason) Hensold & Giul., Ann. Missouri Bot. Gard. 78: 453, fig. 3. 1991. —Paepalanthus duidae Gleason, Bull. Torrey Bot. Club 58: 329. 1931. Syngonanthus phelpsiae var. pilosus Mold., Mem. New York Bot. Gard. 9: 282. 1957. Syngonanthus phelpsiae var. viridis Mold., Phytologia 22: 126. 1971. Paepalanthus jauensis Mold., Mem. New York Bot. Gard. 23: 850, fig. 5. 1972. Paepalanthus duidae var. parvifolius Mold., Phytologia 54: 121. 1983. Rosulate or unbranched short-stemmed clumping herb; stems to 10 cm long; leaves 15–150 × 0.3–1.5 mm, linear, aristate; peduncles (6–)11–39 cm long; involucral bracts long-lanceolate, black or black and cream; capitula 6–14 mm diameter. Shallow, rocky, wet soils of open tepui summits, 1600–2800 m; Bolívar (Aprada-tepui, Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Duida, Cerro Marahuaka, Sierra de la Neblina). Endemic. ŠFig. 36. The type of Paepalanthus fraternus var. radiatus (Phytologia 49: 383. 1981; Steyermark 124407, MO, NY, VEN), from Cerro Marahuaka, probably represents a hybrid between this species and Paepalanthus parvicephalus. Rondonanthus flabelliformis (Mold.) Hensold & Giul., Ann. Missouri Bot. Gard. 78: 454, fig. 4. 1991. —Wurdackia flabelliformis Mold., Mem. New York Bot. Gard. 9: 413. 1957. Perennial; stems to 16 cm long; leaves 90– 100 × 5–8 mm, distichous, ligulate, leathery; peduncles 13–16 cm long; capitula 15–19 mm diameter. Moist places at bases of rocks

38

E RIOCAULACEAE

Fig. 34. Rondonanthus roraimae

5 cm

and dripping crevices on tepui summit, ca. 2500 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Endemic. ŠFig. 38. Rondonanthus roraimae (Oliv.) Herzog, Feddes Repert. Spec. Nov. Regni Veg. 29: 210. 1931. —Paepalanthus roraimae Oliv., Trans. Linn. Soc. London, Bot. 2: 286. 1887. Rosulate or unbranched short-stemmed herb in dense clumps; stems to 10 cm long; leaves 15–40 × 0.5–1.3 mm; peduncles 5–28 cm long; involucral bracts radiating, dark brown; capitula 8–13 mm diameter. Open rocky or boggy areas over thin soils on tepui summits, 2300–2800 m; Bolívar (Ilú-tepui, Kukenán-tepui, Roraima-tepui, Yuruanítepui). Guyana (Mount Roraima). ŠFig. 34. This species is very similar to Rondonanthus duidae, from which it is firmly distinguished only by the early separating glabrous petals of the staminate corolla. Differences in flower and trichome dimensions suggest that R. roraimae may be a polyploid relative to R. duidae.

Fig. 35. Rondonanthus capillaceus

Rondonanthus 39

6 mm

Fig. 36. Rondonanthus duidae

Fig. 37. Rondonanthus acopanensis var. acopanensis

40

E RIOCAULACEAE

Fig. 38. Rondonanthus flabelliformis

Syngonanthus 41

6. SYNGONANTHUS Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 242. 1903. Comanthera L.B. Sm., Contr. Gray Herb. ser. 2, 117: 38, pl. 2. 1937. Carptotepala Mold., Mem. New York Bot. Gard. 9: 278. 1957. Annual or perennial herbs, rosulate or caulescent and branched, sometimes with specialized fertile branches. Roots white, aerenchymatous, usually spongy, or sometimes sloughing the cortex early and then dark and ligneous. Leaves various; pubescence of T-shaped trichomes common. Inflorescences axillary or in terminal umbels; peduncles and sheaths sometimes glandular-hairy; involucral bracts usually pale and hyaline, sometimes dark; floral bracts present or often lacking. Flowers usually 3-merous, rarely 2-merous, unisexual (plants monoecious) or rarely bisexual. Sepals free or briefly connate, often glabrous, sometimes pilose but then rarely tufted apically, more often ciliate along lower margins or pilose below the apex, the trichomes acute and smooth within; corolla in staminate flowers stipitate or not, usually connate into a tube, sometimes secondarily free; petals of pistillate flowers usually connate at medial margins, free at base and apex, rarely free entirely. Stamens as many as the corolla lobes; filaments free or adnate to corolla; anthers with 2 thecae, usually white. Ovary 2- or 3-locular, often subtended by a ring of hairs; style terete, style branches simple; nectaries present and inserted at level of divergence of style branches. Fruit a loculicidal capsule. Southeastern U.S.A., Mexico, Central America, West Indies, South America, Africa; ca. 200 species, 32 in Venezuela, 31 in the flora area. The type of Syngonanthus phelpsiae var. cardonae Mold. (Bol. Soc. Venez. Ci. Nat. 23: 100. 1962), Cardona 1990 (VEN), was reported from Aprada-tepui, Bolívar state, but was not seen for this treatment. Syngonanthus phelpsiae (= Syngonanthus duidae) is not otherwise reported from this area. I suspect that var. cardonae, said to be characterized by very long stems, probably represents either Rondonanthus caulescens, Syngonanthus duidae, or one of the varieties of S. pakaraimensis. Syngonanthus acephalus Hensold, Ann. Missouri Bot. Gard. 78: 434, figs. 3H, I, 6. 1991. Much-branched, dense, moss-like cushions ca. 1 cm high with abundant spongy white roots; leaves 1–2 mm × 0.2 mm; flowers solitary or paired, 1–1.3 mm long, sessile, terminal on leafy stems. White-sand scrub (bana), secondary forests, 50–200 m; southwestern Amazonas (basins of Río Negro and Río Guainía). Endemic. ŠFig. 42. Syngonanthus albopulvinatus (Mold.) Mold., Phytologia 20: 243. 1970. —Paepalanthus albopulvinatus Mold., Bol. Soc. Venez. Ci. Nat. 14: 10. 1952. Clumping perennial herb, the rosettes usually densely woolly in the center; leaves 3–20 × 0.5–1 mm, recurved; peduncles 6.5– 25 cm long; capitula 3–5 mm diameter. Savannas, thin woods, streambanks, 700–2200 m; Bolívar (Cerro Guaiquinima), northern Amazonas (Cerro Autana, Cerro Cuao, Cerro

Duida, Cerro Guanay, Cerro Parú, Cerro Yutajé). Endemic. ŠFig. 41. This species includes both a hairy, leathery-leaved form very like a dwarf Syngonanthus pakaraimensis (Cerro Guaiquinima, Cerro Guanay, Cerro Parú, Cerro Yutajé, and at 1750 m on Cerro Duida), and a membranous-leaved glabrescent form (Cerro Autana, Cerro Cuao, and at 750 m on Cerro Duida). Because of intermediacy and insufficient material, these have not been distinguished here as varieties. Plants of Syngonanthus albopulvinatus from Cerro Guaiquinima are atypical, lacking the involucral characters of the Amazonas plants, and may be more closely allied to the related species S. pakaraimensis of the Gran Sabana. Hybrids occur at Río Coro Coro, with a hybrid swarm population well-collected by Holst & Liesner (Liesner & Holst 21292, 21795, Holst & Liesner 3183, 3222, all at MO), and probably representing gene ex-

42

E RIOCAULACEAE

change with Syngonanthus biformis or S. gracilis, both of which occur in the area. Hybrids probably also occur with S. simplex at Cerro Duida (Steyermark 57804, F, MO; Steyermark 126134, LL, VEN). Syngonanthus amapensis Mold., Phytologia 5: 90. 1954. Perennial rosette with capitula borne singly or in pairs at the apex of 0.3–2.5 cm long, bracteate fertile stems hidden by the leaves; leaves 35–80 × 0.6–3.5 mm, erect, arising from a small corm-like base; peduncles 9.5– 29 cm long; capitula 4–6.5 mm diameter. White-sand savannas, 100–200 m; Amazonas (basins of upper Río Orinoco and lower Río Ventuari). Colombia, Brazil (Amapá, Pará). ŠFig. 45. Syngonanthus amazonicus Mold., Phytologia 3: 42. 1948. —Yuwije (Baré). Cespitose herb; stems to 3 cm long; leaves 12–65 × (0.3–)0.4–1 mm, membranous; peduncles 3–12 cm long; capitula 2–4.5 mm diameter at widest point, conical, pale. Sandy banks, wet sandy or rocky depressions, 100– 800 m; Bolívar (Cerro Guaiquinima), Amazonas (base of Cerro Duida, Río Casiquiare, Río Yatúa). Colombia (Guainía), Brazil (Amazonas). Syngonanthus anomalus (Körn.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 267. 1903. —Paepalanthus anomalus Körn. in Mart., Fl. Bras. 3(1): 458. 1863. Syngonanthus huberi Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 266. 1903. Syngonanthus macrocaulon Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 269. 1903. Syngonanthus paraensis Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 265. 1903. Syngonanthus brevifolius Gleason, Bull. Torrey Bot. Club 56: 14. 1929. Syngonanthus baldwinii Mold., Phytologia 3: 174. 1949. Aquatic or subaquatic, membranous herb in low branched mats or floating, with capitula emergent and solitary from nodes; stems 1.5–22 cm long; leaves 4–26 × 0.5–2 mm; peduncles 6–50 cm long; capitula 2–5 mm diameter. Sandy riverbanks, lake margins, frequently in forested areas, occasionally in savannas, 100–800 m; Bolívar (Río

Caroní, Río Hacha), Amazonas (widespread). Apure; Colombia, Guyana, Brazil (Amapá, Amazonas, Pará). ŠFig. 62. Syngonanthus biformis (N.E. Br.) Gleason, Bull. Torrey Bot. Club 56: 394. 1929. —Paepalanthus biformis N.E. Br., Trans. Linn. Soc. London, Bot. 6: 71. 1901. Syngonanthus simplex var. appendiculifera Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 248. 1903. Syngonanthus densus var. pumilus Mold., Phytologia 3: 277. 1950. Delicate rosulate annual, with numerous capitula; pistillate flowers about twice as long as staminate ones and projecting with sharp tips; leaves 5–20 × 0.5–1.5 mm; peduncles 3–13 cm long; capitula 3–4.5 mm diameter, whitish. Wet sandy soil of streambanks and savannas, depressions and seepages over rock, white-sand savannas, 50–1300 m; widespread in Bolívar (Cerro Guaiquinima, Gran Sabana, Río Parguaza basin, Sabana de Triana in Río Chicanán basin) and Amazonas (Cerro Aracamuni, Río Coro Coro, Río Cuao, Río Guayapo, upper Río Orinoco, and Río Sanariapo, Tobogán de la Selva near Coromoto). Colombia (Vaupés), Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 53. This species is related to and probably hybridizes with members of the Syngonanthus gracilis complex. See S. albopulvinatus, S. simplex, S. pakaraimensis, and S. tenuis. Syngonanthus bisumbellatus (Steud.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 263. 1903. —Eriocaulon bisumbellatum Steud., Syn. Pl. Glumac. 2: 275. 1855. Paepalanthus steudelianus Körn. in Mart., Fl. Bras. 3(1): 450, t. 58, fig. I. 1863. Delicate rosette bearing reddish, sparingly leafy fertile stems topped by umbels of capitula; stems 3–6.5 cm long; leaves 5–12 × 0.3–0.5 mm; peduncles 3–7 cm long; capitula 2.5–3 mm diameter, white. Open swampy areas on tepui slope, ca. 1000 m; Bolívar (Guayaraca). Brazil (Pará, Piauí). Syngonanthus caulescens (Poir.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 267. 1903. —Eriocaulon caulescens Poir.,

Syngonanthus

Encycl. suppl. 3: 162. 1813. —Paepalanthus caulescens (Poir.) Kunth, Enum. Pl. 3: 537. 1841. —Bola de fuego, Guanak, Poma-taca (Yekwana). Paepalanthus heteropeplus Körn. in Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 238. 1867. —Syngonanthus heteropeplus (Körn.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 248. 1903 Syngonanthus glandulosus Gleason, Bull. Torrey Bot. Club 56: 394. 1929. Syngonanthus glandulosus var. epapillosus Mold., Phytologia 26: 177. 1973. —Syngonanthus glandulosus f. epapillosus (Mold.) Mold., Phytologia 44: 384. 1979. Syngonanthus caulescens var. hirsutus Mold., Phytologia 53: 367. 1983. Weedy terrestrial to subaquatic herb, caulescent or apparently rosulate, with capitula clustered at stem apex; stems 0.5–18 cm long; leaves 10–25 × 1–5 mm; peduncles 4–15 cm long; capitula 3–6 mm diameter, shiny white. Marshes, wet savannas, streambanks, pool margins, Mauritia palm swamps, wet cliff faces, rocky sandstone or granitic seeps, 100–1500 m; common throughout Bolívar, much less common in northern Amazonas (bases of Cerro Duida, Cerro Yapacana, and Cerro Yutajé, Puerto Ayacucho). Anzoátegui, Apure, Guárico, Monagas, Portuguesa; Neotropics from Mexico south to Peru and Argentina. ŠFig. 63. Syngonanthus glandulosus is a name commonly applied to rosulate individuals of S. caulescens, but intergrading forms exist, and these habit differences are probably due to habitat. The rosulate forms usually are collected from streambanks, and the upright robust long-stemmed plants from stable soils not subject to flooding. Syngonanthus cowanii Mold., Mem. New York Bot. Gard. 8: 99. 1953. Syngonanthus cowanii var. longipedunculatus Mold., Mem. New York Bot. Gard. 9: 282. 1957. Syngonanthus cowanii var. tabulatus Mold., Phytologia 44: 215. 1979. Syngonanthus cowanii var. involucratus Mold., Phytologia 51: 302. 1982. Syngonanthus cowanii var. simplex Mold., Phytologia 51: 245. 1982. Herb from rosette, with naked or verticillately leafy aerial stems with dense,

43

flat-topped, buff-colored umbels of capitula; stems 1–5.5 cm long; basal leaves 2–14 × 0.5 mm; peduncles 0.1–1 cm long; capitula 1.5–3 mm diameter. White-sand savannas, 100– 200 m; Amazonas, (Canaripó, Caño Yagua, Cerro Yapacana, Río Atabapo, Río Guayapo, Río Siapa). Adjacent Colombia. ŠFig. 59. The characters used to distinguish the numerous varieties listed above in synonymy (peduncle length, presence of leaf verticils on stem) are highly variable, even within populations. Syngonanthus duidae Mold., Fieldiana, Bot. 28: 127. 1951. Syngonanthus phelpsiae Mold., Bol. Soc. Venez. Ci. Nat. 14: 12. 1952, “phelpsae.” Syngonanthus phelpsiae var. elongatus Mold., Mem. New York Bot. Gard. 8: 102. 1953. Leiothrix marahuacensis Mold., Phytologia 55: 113, pl. 1. 1984. Robust perennial, usually a low dense circular cushion; inflorescences solitary or nearly so; leaves 4–40 × 0.4–1 mm, erect, linear; peduncle 2–35 cm long; capitulum 4–8 mm diameter. Open rocky plateaus, open or often shrubby savannas, sandy riverbanks on tepui slope and summits, 1000–2600 m. Venezuela; 2 varieties, both restricted to the flora area. Key to the Varieties of S. duidae 1. Leaves mostly 10–40 mm long; peduncle to 35 cm long, though occasionally also dwarfed ............................ var. duidae 1. Leaves 4–6 mm long; peduncle 2–7 cm long .................................. var. humilis S. duidae var. duidae Stems to 5 cm long; leaves 4–40 × 0.4–1 mm; peduncle 2–35 cm long; capitulum 4–8 mm diameter. 1000–2600 m; Bolívar (Auyántepui, Cerro Jaua, Cerro Guaiquinima, Ptari-tepui), Amazonas (Cerro Aracamuni, Cerro Autana, Cerro Cuao, Cerro Duida, Cerro Guanay, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé). Endemic. ŠFig. 43. See discussion of Syngonanthus pakaraimensis. S. duidae var. humilis Hensold, Ann. Missouri Bot. Gard. 78: 434. 1991.

44

E RIOCAULACEAE

Stems to 1 cm long; leaves 4–6 × 0.5 mm; peduncle 2–7 cm long; capitulum 3–6.5 mm diameter. 2100–2600 m; Bolívar (Cerro El Sol, Cerro Kukenán, Roraima-tepui). Endemic. This variety is expected to be found on the Guyanese portion of the summit of Mount Roraima. Syngonanthus fenestratus Hensold, Ann. Missouri Bot. Gard. 78: 434, figs. 3K–J, 7. 1991. Herb with basal rosette and sparingly leafy aerial stems with terminal umbels; leaves 15–40 × 0.5–0.7 mm, fenestrate; peduncles 7–13 cm long; capitula 6.5–7 mm diameter. Along large waterfalls, ca. 400 m; Bolívar (Canaima). Guyana, Brazil (Amazonas). ŠFig. 61. Syngonanthus gracilis (Bong.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 249. 1903. —Eriocaulon gracile Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 634. 1831. —Paepalanthus gracilis (Bong.) Körn. in Mart., Fl. Bras. 3(1): 460, t. 59, fig. 1. 1863. Solitary rosulate annual; leaves 5–20 × 0.2–0.6 mm, linear, olivaceous; peduncles 7– 15 cm long; involucral bracts lustrous, redbrown, hyaline; capitula 3–5 mm diameter. Moist parts of savannas, sometimes over granite, 50–300 m; Bolívar (Río Orinoco basin), Amazonas (Cerro Morrocoy, Puerto Ayacucho). Anzoátegui, Apure, Guárico, Portuguesa, Sucre; Colombia, Guyana, Suriname, Brazil, Bolivia, Paraguay. ŠFig. 54. Ruhland [Pflanzenr. IV. 30(Heft 13): 249– 252. 1903] described 13 varieties of Syngonanthus gracilis, two of these have been reported from Venezuela, both from the lower Río Orinoco. The types of neither were seen in preparation of this treatment so their identity cannot be ascertained with certainty. Syngonanthus gracilis var. pallida Ruhland, possibly refers to S. nitens. Syngonanthus gracilis var. recurvifolius Ruhland seems to refer to the material here being recognized as S. gracilis. Workers since Ruhland have tended to use the name for nearly any small rosulate Syngonanthus with hyaline bracts and several to many inflorescences. However, in the the flora area very diverse forms occur with this suite of

characters, and typical S. gracilis as it occurs in central Brazil and as here treated is certainly a distinct species not identical with the bulk of the Guayanan material, here recognized as Syngonanthus simplex. Syngonanthus humboldtii (Kunth) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 262. 1903. —Paepalanthus humboldtii Kunth, Enum. Pl. 3: 535. 1841. Eriocaulon umbellatum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 252. 1815 [1816], non Lam. 1789. Eriocaulon bonplandianum Steud., Syn. Pl. Glumac. 2: 275. 1855. Paepalanthus fertilis Körn. in Mart., Fl. Bras. 3(1): 448, t. 58, fig. 1. 1863. —Syngonanthus fertilis (Körn.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 263. 1903. Syngonanthus humboldtii var. glandulosus Gleason, Bull. Torrey Bot. Club 58: 327. 1931. —Syngonanthus fertilis var. glandulosus (Gleason) Mold., Phytologia 55: 104. 1984. Syngonanthus humboldtii var. elongatus Mold., Mem. New York Bot. Gard. 8: 101. 1953. Syngonanthus humboldtii var. orinocensis Mold., Mem. New York Bot. Gard. 8: 102. 1953. —Syngonanthus fertilis var. orinocensis (Mold.) Mold., Phytologia 55: 105. 1984. Syngonanthus humboldtii var. glabrescens Mold., Phytologia 46: 155. 1980. Rosulate plant with erect fertile stems bearing 0–many verticils of leaves along their length and an umbel of pilose heads at the apex. Venezuela, Colombia, Guyana, Suriname, Brazil; 3 varieties, all in Venezuela, 2 of these in the flora area. Key to the Varieties of S. humboldtii 1. Plants usually > 10 cm tall, fertile stem interrupted by 1–many verticils of leaves ........................ var. humboldtii 1. Plants < 10 cm tall, fertile stem naked except at apex ...................... var. parvus S. humboldtii var. humboldtii Stems 3–20 cm long; leaves 10–50 × 0.5–1 mm; peduncles 3–14 cm long; capitula (3–)4– 6 mm diameter. Common in wet sandy savannas, disturbed sites, streambanks, flood-

Syngonanthus 45

plains, Mauritia palm swamps, occasionally on shallow soils on granitic outcrops, 50– 1300 m; Bolívar (southern bases of Auyántepui and Cerro Ikabarú, Río Caroní, Río Paragua), Amazonas (northwestern part of state, also Cerro Duida and Cerro Vinilla south to Río Guainía). Apure; Colombia, Guyana, Suriname, Brazil (Amazonas, Goiás, Mato Grosso, Pará, Piauí, Rondônia, Roraima). ŠFig. 64. Note: this taxon may actually include two distinct but possibly hybridizing species, one of which is distinguished by the presence of floral bracts. Further study is needed.

—Comanthera kegeliana (Körn.) Mold., Phytologia 13: 218. 1966. Comanthera linderi L.B. Sm., Contr. Gray Herb. ser. 2, 117: 38, pl. 2. 1937. Syngonanthus akurimensis Mold., Phytologia 2: 371. 1947. Rosulate perennial with dark and wiry roots; leaves 4–15 × 0.4–1.5 mm, hirsute with stiff upright trichomes and appressed oblong scales; peduncles 7–14 cm long; capitula 3–4 mm diameter. Sandy savannas, ca. 900 m; Bolívar (Santa Elena de Uairén area). Guyana, Suriname, Brazil (Amazonas, Pará). ŠFig. 51.

S. humboldtii var. parvus (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 436. 1991. —Syngonanthus allenii var. parvus Mold., Mem. New York Bot. Gard. 8: 99. 1953, "alleni." Stems 2–6 cm long; leaves 2–15 × 0.2–0.7 mm; peduncles 1.8–4.4 cm long; capitula 0.25–4 mm diameter. Wet savannas and granitic outcrops, 50–200 m; northwestern Bolívar, Amazonas (along the Río Orinoco from San Pedro south to the Río Parguaza and Río Suapure). Possibly Colombia (specimens too poor for positive identification).

Syngonanthus longipes Gleason, Bull. Torrey Bot. Club 56: 15. 1929. Large rosulate perennial bearing clusters of long-pedunculate capitula on naked, lax, fertile branches; stems 10–42 cm long; leaves 35–130 × 1.5–6 mm; peduncles 15–41 cm long; capitula 5–8 mm diameter. Wet sandy soils of open or shrubby gallery forests, savannas, Mauritia palm swamps, 100–1500 m; Bolívar (Cerro Guaiquinima, Gran Sabana), Amazonas (widespread). Colombia, Guyana, Suriname, Brazil (Amazonas, Mato Grosso, Pará, Rondônia). ŠFig. 60.

Syngonanthus jenmanii (Gleason) Giul. & Hensold, Ann. Missouri Bot. Gard. 78: 462. 1991. —Paepalanthus jenmanii Gleason, Bull. Torrey Bot. Club 56: 14. 1929. —Carptotepala jenmanii (Gleason) Mold., Mem. New York Bot. Gard. 9: 278. 1957. Carptotepala insolita Mold., Fieldiana, Bot. 28: 114. 1951. Terrestrial to subaquatic herb; stems to 9 cm long; leaves 70–120 × 1–1.5 mm, linear, erect, soft; peduncles 13–15 cm long; involucre deep-cupulate; capitula 6–7 mm diameter. Riverbanks, moist rock outcrops, swampy savannas or meadows, 400–2600 m; Bolívar (northern Gran Sabana, Ilú-tepui, Macizo del Chimantá). Guyana. ŠFig. 47. This species is closely related to Syngonanthus xeranthemoides, and possibly hybridizes with it. An intermediate is Davidse 4865 (LL, MO) from the Gran Sabana. Syngonanthus kegelianus (Körn.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 273. 1903. —Paepalanthus kegelianus Körn. in Mart., Fl. Bras. 3(1): 438. 1863.

Syngonanthus macrocephalus (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 436. 1991. —Syngonanthus humboldtii var. macrocephalus Mold., Mem. New York Bot. Gard. 8: 101. 1953. Perennial from rosette; stems 4–6.5 cm long; inflorescence borne singly at apex of short naked fertile branches, the heads hirsute and coppery red; leaves 10–50 × 0.5–1 mm; peduncle 6.5–11.2 cm long; capitulum 6–9 mm diameter. Bogs and shallow soil over rock of tepui summit, 1500–1800 m; Amazonas (Cerro Sipapo). Endemic. Syngonanthus minutus (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 436. 1991. —Paepalanthus minutus Mold., Brittonia 3: 158. 1939. Tiny short-stemmed herb; heads dull gold, hyaline; flowers 2-merous; leaves 0.6– 0.8 cm long, filiform, acicular; peduncles 5– 15 cm long; capitula 1–2 mm diameter. Tepui meadows, ca. 2200 m; Bolívar (Auyán-tepui). Endemic. Syngonanthus nitens (Bong.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 254.

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1903. —Eriocaulon nitens Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 633, t. 55. 1831. —Paepalanthus nitens (Bong.) Kunth, Enum. pl. 3: 531. 1841. Syngonanthus flavipes Mold., Mem. New York Bot. Gard. 8: 100. 1953. Robust rosulate perennial with spongy orange roots; leaves 12–80 × 0.8–1.5 mm; peduncles 16–45 cm long; capitula obconic, 4.5– 6.5 mm diameter. Moist parts of savannas, Mauritia palm swamps, 100–900 m; Bolívar (eastern part of state, Santa Elena de Uairén), Amazonas (Raudal de Atures, basins of Río Atabapo, upper Río Orinoco, and lower Río Ventuari). Colombia, Brazil (mostly central plateau), Bolivia, Paraguay. ŠFig. 46. Syngonanthus oblongus (Körn.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 260. 1903. —Paepalanthus oblongus Körn. in Mart., Fl. Bras. 3(1): 446. 1863. Perennial with basal rosettes of spatulate leaves and naked fertile stems; inflorescences umbels of whitish heads. Venezuela, Colombia, Brazil (Amazonas, Maranhão, Pará, Piauí); 2 varieties, 1 of these in the flora area. S. oblongus var. aequinoctialis Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 260. 1903. Stems 1.5–18 cm long; leaves 22–80 × 4–9 mm; peduncles 3.5–8 cm long; capitula 4–5 mm diameter. Streambanks in gallery forests or otherwise shaded sites, 1000–1300 m; Bolívar (Cerro Marutani), Amazonas (Cerro Duida, Cerro Marahuaka). Colombia (Vaupés), Brazil (Amazonas). ŠFig. 55. Syngonanthus ottohuberi Hensold, Ann. Missouri Bot. Gard. 78: 437. 1991. Syngonanthus similis var. venezuelensis Mold., Phytologia 45: 209. 1980. Syngonanthus densifolius var. venezuelensis Mold., Phytologia 48: 291. 1981. Robust perennial from leathery rosettes; inflorescences umbels of heads on black leafless erect aerial stems 6–12 cm long; leaves 22–45 × 1–1.5 mm; peduncles 20–27 cm long; capitula 7–7.5 mm diameter. White-sand savannas, 100–200 m; Amazonas (Canaripó,

Caño Yagua, Cerro Yapacana). Endemic. ŠFig. 57. Syngonanthus pakaraimensis Mold., Mem. New York Bot. Gard. 9: 282. 1957. Leiothrix steyermarkii Mold., Fieldiana, Bot. 28: 118. 1951. Syngonanthus lanatus var. alpinus Mold., Act. Bot. Venez. 2: 153. 1967. Syngonanthus duidae var. longifolius Mold., Phytologia 51: 245. 1982. Clumping perennial with villous linear leaves in dense cushion-like rosettes. Venezuela, Guyana; 2 varieties, both in the flora area. Key to the Varieties of S. pakaraimensis 1. Involucral bracts cream, hyaline, remaining appressed to head; floral bracts absent ...................... var. pakaraimensis 1. Involucral bracts brownish, firm, becoming strongly reflexed with maturation of head; floral bracts present ...................... ....................................... var. rivularis S. pakaraimensis var. pakaraimensis ?Paepalanthus venetifolius Mold. & Steyerm., Bol. Soc. Venez. Ci. Nat., 32– 33: 284. 1976. Rosulate herb; leaves 10–35(–50) × 0.5– 1.5 mm; peduncles 5–39 cm long; capitula (4–)5–8 mm diameter. Wet open or shrubby savannas, rocky plateaus, sandy streambanks, 800–2400 m; Bolívar (Auyán-tepui, Gran Sabana, Ilú-tepui, Ptari-tepui, Sierra Pakaraima). Guyana. ŠFig. 39. The type of Paepalanthus venetifolius, with its dwarf peduncles, upright leaves, and long narrow involucral bracts, appears to be a hybrid between this variety and Syngonanthus duidae at Cerro Jaua. Gene exchange probably occurs between S. pakaraimensis and the two related annual species S. simplex and S. biformis, where they occur together at the Gran Sabana and Cerro Marutani. These intermediate collections are listed separately in the exsiccatae. Also see discussion of S. albopulvinatus. S. pakaraimensis var. rivularis (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 437. 1991. —Syngonanthus rivularis Mold., Mem. New York Bot. Gard. 9: 411. 1957.

Syngonanthus 47

Stems to 4 cm long; leaves 20–40 × 0.5 mm; peduncles 19–21 cm long; capitula 9 mm diameter. Sandy openings and along riverbanks, 1900–2200 m; Bolívar (Macizo de Chimantá [Apacará-tepui]). Endemic. This variety with its erect leaves, rather large, firm, reflexing bracts, and floral bracts, appears intermediate between Syngonanthus pakaraimensis and S. tiricensis, and should be investigated as a possible hybrid. Syngonanthus reflexus Gleason, Bull. Torrey Bot. Club 58: 327. 1931. Leiothrix echinulata Mold., Bull. Torrey Bot. Club 77: 390. 1950. Syngonanthus reflexus var. longifolius Mold., Phytologia 46: 155. 1981. Rosulate herb; leaves 35–210 × 1.5–3.5 mm, coriaceous; peduncles 19–46 cm long; involucre cupulate, bristly; capitula 7.5–10 mm diameter. Locally abundant in wet areas in swampy sandy savannas, 100–200 m; central and western Amazonas. Colombia, Brazil (Amazonas, Mato Grosso, Pará). ŠFig. 44. Syngonanthus setifolius Hensold, Ann. Missouri Bot. Gard. 78: 438, figs. 3A, 3B, 8. 1991. Small cespitose perennial; leaves 5–40 × 0.1–0.2 mm, filiform; peduncles 2.5–10 cm long; capitula 2–2.5 mm diameter, whitish. White-sand savannas, streambanks, 100– 1200 m; western and central Amazonas. Brazil (Amazonas). ŠFig. 50. Syngonanthus simplex (Miq.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 248. 1903. —Paepalanthus simplex Miq., Stirp. Surinam. Select. 222. 1851. Paepalanthus eriophyllus Körn. in Mart., Fl. Bras. 3(1): 463. 1863. —Syngonanthus eriophyllus (Körn.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 249. 1903. Paepalanthus filipes Mold., Bull. Torrey Bot. Club 75: 196. 1948. Syngonanthus vareschii Mold., Acta Biol. Venez. 2(7): 50. 1957, pro parte (species based on mixed collection; the vegetative parts belong to a moss). Habit highly variable; leaves 3–20(–32) × 0.5–1 mm; peduncles 3–10 cm long; capitula

2–4 mm diameter. Sandy banks or depressions, sandstone crevices, forest borders, wet savannas, 100–1500 m; Bolívar (Arekuna, Cerro Guaiquinima, Gran Sabana, Sabana de Triana in Río Chicanán basin), Amazonas (slopes of Cerro Duida, Cerro Marahuaka, and Sierra de la Neblina). Guyana, Suriname, Brazil (Amazonas eastward and south to restingas of Bahia). Together with Syngonanthus gracilis, this species belongs to a difficult complex, where species boundaries need to be resolved. In terms of vegetative habit, S. simplex is highly variable, with solitary to clumping, and woolly to glabrous forms predominating in different local areas. “Gigas” types also occur, as represented by Steyermark 59305 (F, NY) and Davidse 4826 (MO). For a discussion of hybrids, see Syngonanthus albopulvinatus and S. pakaraimensis. Although presumably more difficult to detect, hybridization is also likely with S. biformis. In areas where these two species occur together, variability is found in the relative length of the staminate and pistillate flowers in S. simplex. Syngonanthus tenuis (H.B.K.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 253. 1903. —Eriocaulon tenue H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 253. 1815 [1816]. —Paepalanthus tenuis (H.B.K.) Kunth, Enum. Pl. 3: 282. 1855. ?Syngonanthus tenuis var. minor Mold., Phytologia 37: 275. 1977 (type not seen). Syngonanthus drouetii var. parviceps Mold., Phytologia 54: 68. 1983. Rosulate herb frequently with woolly buds in upper leaf axils; involucre showy, white, glabrous, surpassing the capitula. Southern Venezuela, Colombia, Brazil; 2 varieties, both in the flora area. The variation in the Venezuelan material is complex, and the species is here provisionally divided into two imperfectly delimited varieties. The varieties are roughly sympatric, but only occasionally collected at the same site, and var. tenuis unlike var. bulbifer does not appear to occur south of Venezuela. Hybridization with sympatric species in the Syngonanthus gracilis complex should be investigated as a possible basis for this variation pattern.

48

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Key to the Varieties of S. tenuis 1. Leaves 12–50 × 0.5–2 mm; peduncle sheaths 1.6–3.8 cm long; peduncles 10– 27.5 cm long, ca. 1–15 per rosette; capitula 4–5.5 mm diameter; upper involucral bracts waxy white, oblanceolaterounded, often held erect or slightly radiating in dry material; flowers ca. 1.4–1.8 mm long ............ var. bulbifer 1. Leaves 0.4–3.8 cm long, 0.2–1 mm wide; peduncle sheaths 0.8–2(–3) cm long; peduncles 4–14 cm long, ca. (4–)8–60 per rosette; capitula 2.5–4(–4.5) mm diameter; upper involucral bracts hyaline or with waxy white streaks, obovate to broadly oblanceolate, always cupped over the disk of the capitulum in dry material; flowers ca. 1(–1.4) mm long ........................................... var. tenuis S. tenuis var. bulbifer (Huber) Hensold, Ann. Missouri Bot. Gard. 78: 440. 1991. —Paepalanthus bulbifer Huber, Bol. Mus. Paraense Hist. Nat. 2: 499. 1898. —Syngonanthus bulbifer (Huber) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 272. 1903. Syngonanthus drouetii L.B. Sm., Contr. Gray Herb. 117: 34, pl. 2, figs. 41–43. 1937. Syngonanthus vaupesanus Mold., Phytologia 2: 6. 1941. White-sand savannas and meadows, 100– 1200 m; Bolívar (Cerro Guaiquinima, upper Río Paragua), Amazonas (Río Atabapo, Río Autana, Río Guainía, Río Orinoco, Río Sipapo). Colombia (Vaupés), Brazil (Amazonas, Pará, Mato Grosso, Goiás). ŠFig. 48. Hybridization has been observed with Syngonanthus biformis on the upper Río Ventuari (Infante s.n., VEN 44017). S. tenuis var. tenuis. —Aribái-panáru-kusí (Pemón). White-sand savannas, 100–900 m; Bolívar (Río Yuruaní), Amazonas (Cerro Yapacana, Río Atabapo, Río Autana, Río Guainía, upper Río Orinoco, Río Parú, Río Sipapo). Endemic. The isolated collections at Río Yuruaní are intermediate in size range between the two varieties, but lack the bract characters of var. bulbifer.

Syngonanthus tiricensis Mold., Mem. New York Bot. Gard. 9: 412. 1957. Perennial forming low dense circular mats; leaves 7–30 × 0.5–0.9 mm; peduncles 8–25.5 cm long; capitula 7–11 mm diameter, showy, white, large-bracteate, borne singly or in pairs. Locally common in wet tepui meadows, 2100–2500 m; Bolívar (Apradatepui, Macizo del Chimantá). Endemic. ŠFig. 40. See discussion of Syngonanthus pakaraimensis var. rivularis. Syngonanthus trichophyllus Mold., Phytologia 6: 329. 1958. Paepalanthus saxicola var. conicus Mold., Bol. Soc. Venez. Ci. Nat. 23: 300. 1963. Cespitose herb; stems to 3 cm long; leaves 7–30 × 0.5–0.9 mm, filiform, membranous; peduncles 8–25.5 cm long; capitula conical, 7–11 mm diameter at widest point, pale. Wet sandy soil of streambanks, depressions in savannas, often semi-submerged in pools or small streams, 50–1200 m; Bolívar (Gran Sabana), Amazonas (scattered). Guyana, Brazil (Amazonas). ŠFig. 52. Syngonanthus umbellatus (Lam.) Ruhland in Urb., Symb. Antill. 1: 488. 1900. —Eriocaulon umbellatum Lam., Encycl. 3: 277. 1789; Encycl. Recueil 1: pl. 50, fig. 4. 1823. —Paepalanthus umbellatus (Lam.) Kunth, Enum. Pl. 3: 537. 1841. Syngonanthus umbellatus f. proliferens Mold., Phytologia 39: 161. 1978. Syngonanthus humboldtii var. simplex Mold., Phytologia 51: 245. 1982. Rosulate perennial with naked fertile stems bearing umbels of capitula; stems 4– 17 cm long; leaves basal, (14–)25–65 × 1.5–4 mm, hirsute; peduncles 5–23 cm long; capitula globose, 5–7 mm diameter, white. Wet savannas, Mauritia palm swamps, 100–500 m; scattered in Bolívar (Amaruay-tepui, Canaima, Río Chicanán, Río Paragua), Amazonas (La Esmeralda). Colombia (Vaupés), Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, Minas Gerais, Roraima, Pará). ŠFig. 56. Syngonanthus williamsii (Mold.) Hensold, Ann. Missouri Bot. Gard. 78: 440. 1991. —Paepalanthus williamsii Mold., Phytologia 2: 367. 1947.

Syngonanthus 49

Robust, rosulate herb with umbels of capitula at the apex of uniformly leafy stems; stems 5–45 cm long; leaves 50–150 × 5–10 mm; peduncles 10–27 cm long; capitula 7– 9.5 mm diameter. White-sand savannas and adjacent forests, 100–300 m; southwestern Amazonas (basins of Río Atabapo, Río Guainía, and upper Río Orinoco). Colombia (Guainía, Vaupés), Brazil (Amazonas, Pará, Roraima). ŠFig. 58. Syngonanthus xeranthemoides (Bong.) Ruhland in Engl., Pflanzenr. IV. 30(Heft 13): 276. 1903. —Eriocaulon xeranthemoides Bong., Mém. Acad. Imp. St.Pétersbourg, Sér. 6, Sci. Math. 1: 635. 1831. —Aribái-panáru-kusí-yek (Arekuna). Syngonanthus tricostatus Gleason, Bull. Torrey Bot. Club 56: 16. 1929. —Syngonanthus xeranthemoides var. tricostatus (Gleason) Mold., Phytologia 26: 179. 1973. Syngonanthus xeranthemoides f. brevifolius Mold., Phytologia 26: 178. 1973. Syngonanthus xeranthemoides var. angustifolius Mold., Phytologia 51: 302. 1982. Syngonanthus xeranthemoides var. alpinus Mold., Phytologia 55: 268. 1984. Rosulate perennial; leaves 20–230 × 1–5 mm, coriaceous; peduncles 2.5–50 cm long; involucre cupulate, leathery; capitula 4–9 mm diameter. Sandy, periodically flooded savannas, thin saturated soils of sandstone outcrops and streambanks, 50–2200 m; Bolívar (Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá), Amazonas (Cerro Parú, Cerro Yutajé, Puerto Ayacucho, Río Atabapo). Apure, Guárico; Colombia, Guyana, Brazil, Bolivia. ŠFig. 49. The plants of Macizo del Chimantá tend to form dwarf clumps, with small falcate leaves, short peduncles, brownish involucral bracts, and a high percentage of abortive inflorescences. Because of the abnormal developmental features, it is suspected that these may be derived from hybridization with Rondonanthus acopanensis. Most of the Gran Sabana plants have irregularly twisted, short thick leaves, and have been recognized by Moldenke as var. tricostatus. Narrow, straight-leaved forms mostly occur in southern Amazonas and have been de-

scribed as var. angustifolius. In northern Amazonas, eastern Bolívar, and Apure very robust forms with leaves to 35 cm long occur. Intermediates exist between these varieties, so that they are not here recognized. See also S. jenmanii.

Fig. 39. Syngonanthus pakaraimensis var. pakaraimensis

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Fig. 40. Syngonanthus tiricensis

2 mm

2 cm

Fig. 41. Syngonanthus albopulvinatus

Fig. 42. Syngonanthus acephalus

Fig. 43. Syngonanthus duidae var. duidae

Syngonanthus 51

Fig. 44. Syngonanthus reflexus

Fig. 45. Syngonanthus amapensis

Fig. 46. Syngonanthus nitens

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Fig. 48. Syngonanthus tenuis var. bulbifer

Fig. 47. Syngonanthus jenmanii

Fig. 49. Syngonanthus xeranthemoides

Syngonanthus 53

Fig. 50. Syngonanthus setifolius

Fig. 51. Syngonanthus kegelianus

2.5 mm

Fig. 52. Syngonanthus trichophyllus

Fig. 53. Syngonanthus biformis

Fig. 54. Syngonanthus gracilis

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Fig. 55. Syngonanthus oblongus var. aequinoctialis

Fig. 56. Syngonanthus umbellatus

Syngonanthus 55

Fig. 57. Syngonanthus ottohuberi

Fig. 58. Syngonanthus williamsii

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Fig. 59. Syngonanthus cowanii

Fig. 60. Syngonanthus longipes

Fig. 61. Syngonanthus fenestratus

Syngonanthus 57

Fig. 62. Syngonanthus anomalus

Fig. 63. Syngonanthus caulescens

Fig. 64. Syngonanthus humboldtii var. humboldtii

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7. TONINA Aubl., Hist. Pl. Guiane 857, pl. 330. 1775. Weedy amphibious herbs with long, floating or procumbent, mat-forming stems. Roots fibrous, brownish; stems glabrate. Leaves lanceolate, membranous. Inflorescences in leaf axils, very short-pedunculate; peduncle sheaths leaf-like, open, often exceeding peduncle; peduncles 2-ribbed; involucral bracts brown, ciliate with tuberculate trichomes. Flowers 3-merous, unisexual (plants monoecious). Sepals brown, ovate-acuminate, free, in staminate flowers the tips incurved to enclose the corolla; fleshy obconic corolla stipe present in staminate flowers; petals of pistillate flowers very reduced, appearing as densely pilose lobes, those of staminate flowers connate into a short tube. Stamens 3; filaments free; anthers with 1 theca. Ovary 3-locular; style branches bifid; nectaries inserted at level of style branches. Fruit a capsule. Mexico, Central America, West Indies, South America; 1 species. Tonina fluviatilis Aubl., Hist. Pl. Guiane 857, pl. 330. 1775. Amphibious weed; stems 8–35 cm long, lax, with axillary capitula; leaves 7–27 × 0.8– 4 mm, short-ligulate; peduncles 0.1–1.2 cm long; capitula 5–6 mm diameter, brownish. Streambanks, riversides, roadside ditches, 0–400 m; Delta Amacuro (Caño Acoimito near Río Orocoima), northern Bolívar, widespread in Amazonas. Apure, Guárico, Monagas, Táchira, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian and northeastern Brazil. ŠFig. 65.

Fig. 65. Tonina fluviatilis

Erythroxylum 59

ERYTHROXYLACEAE by Timothy Plowman and Paul E. Berry Glabrous shrubs or small trees. Leaves usually alternate (opposite in Aneulophus), stipulate, petiolate, entire; stipules persistent on twigs or caducous. Flowers actinomorphic, pedicellate, often heterostylous, solitary or fascicled in the axils, sometimes short-pedunculate, arising from small, scarious bracteoles. Calyx persistent, sepals 5, valvate, united below; petals 5, free, alternate with the sepals, imbricate in bud, caducous, usually appendaged on the adaxial surface. Stamens usually 10; filaments united at the base usually forming a short tube; anthers 2-locular, longitudinally dehiscent. Ovary superior, (2)3-locular, usually only 1 locule ovuliferous; ovules solitary, rarely 2 (in Nectaropetalum), axile, pendulous, epitropous; styles (2)3, free or connate from the base; stigmas capitellate, rarely subsessile. Fruit small, drupaceous, 1-seeded, rarely capsular and 2- or 3-seeded (in Aneulophus); embryo straight; endosperm present or absent. Neotropics, Asia, tropical Africa, Madagascar, Australia; 4 genera and ca. 250 species, 1 genus and 30 species in the flora area. 1. ERYTHROXYLUM P. Browne, Civ. Nat. Hist. Jamaica 278. 1756. Glabrous shrubs or small trees, evergreen or deciduous, rarely dioecious or subdioecious, twigs compressed at apex, often bearing persistent, distichous, imbricated cataphylls and stipules. Leaves alternate, often distichous, simple, stipulate, petiolate, entire, with involute venation that sometimes imprints 2 parallel lines and/or a distinct, discolorous, central panel on lower surface. Stipules intrapetiolar, appearing as a single organ, dorsally bicostate, often 2- or 3-setulose at apex, persistent or caducous, often leaving an obliquely transverse scar. Flowers actinomorphic, small, heterostylous, axillary, solitary or fascicled, rarely short-pedunculate, arising from small, scarious, often persistent bracteoles, pedicellate. Calyx persistent, sepals 5, valvate, united below; petals 5, free, caducous, imbricate in bud, appendaged on the adaxial surface with a 2-lobed ligule. Stamens 10 in 2 whorls of 5, the outermost alternate with the petals; filaments united at base and forming a short tube surrounding the ovary, persistent; anthers 2-locular, longitudinally dehiscent. Ovary 3-locular but with only 1 locule ovuliferous; ovule solitary in the fertile locule, pendulous from axile placenta, anatropous, epitropous; styles 3, free or partly connate from base; stigmas capitellate. Fruit a small, fleshy, red or purplish drupe; endocarp 1- or 3-locular but with only 1 fertile locule. Seed 1, with or without endosperm; embryo straight. Pantropical, in the New World in Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay, and Argentina; ca. 230 species, 41 in Venezuela, 30 of these in the flora area. To effectively identify neotropical species of Erythroxylum, it is often necessary to have complete material at hand. Among vegetative characters, the foliar stipules are often diagnostic; it is important that newly formed and undamaged (usually subterminal) stipules be examined whenever possible. Mature leaves should be sought, especially in deciduous species. Leafy specimens should never be preserved with

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ethanol or other preservatives, since subtle but useful characters of coloration are destroyed. Mature endocarps are also important. The key is based primarily on specimens from the Venezuelan Guayana and is tailored to identify material from there. Both vegetative and reproductive characters are used in the key, and easily observed, macroscopic characters are emphasized. Key to the Species of Erythroxylum 1. 1.

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

6.

7(2).

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

9.

Stipules of mature leaves conspicuously longitudinally striately veined .... 2 Stipules of mature leaves smooth, not conspicuously striately veined (but stipules of younger leaves of E. vernicosum and E. squamatum may appear striate) ...................................................................................... 12 Leaves rounded, retuse, or obtuse at apex; evergreen or deciduous species ................................................................................................................ 3 Leaves acute or acuminate at apex, the tip itself often obtuse; evergreen species .................................................................................................... 7 Stipules 5–15 mm long, caducous or variably persistent; evergreen species ................................................. E. macrophyllum var. savannarum Stipules 1–4 mm long, mostly caducous; deciduous species ..................... 4 Branchlets stout, the bark becoming thick-corky; leaves coriaceous; stipules persistent, turning dark brown or black with age ........ E. suberosum Branchlets slender, never corky; leaves membranous or chartaceous; stipules not conspicuous on older twigs ................................................ 5 Flowers produced at successive nodes on last season’s twigs, often prior to leaf flush; pedicels 9–20 mm long ................................................ E. rufum Flowers produced near apex of twigs or short shoots with the new leaves; pedicels 3.5–6 mm long .......................................................................... 6 Petioles 1–2 mm long; leaves membranous, translucent; venation prominulous on upper surface of leaf when dried; fruits slightly curved at maturity; the endocarp falcate-ellipsoid, laterally compressed, prominently 3-ribbed, 3-locular with 1 fertile and 2 empty locules ....... E. steyermarkii Petioles 3–7 mm long; leaves chartaceous, opaque; venation inconspicuous on upper surface of leaf when dried; fruits radially symmetric at maturity; the endocarp ovoid-ellipsoid, terete, 1-locular ..... E. lindemanii Stipules persistent, ≤ 3 mm long (not including setae), with 2 conspicuously long apical setae, often fimbriate and recurving, appearing to peel off abaxial side of stipule; only known from one collection in the flora area (Sierra de Lema) .................................................. E. fimbriatum Stipules persistent or caducous, 4–35 mm long, the setae if present short, filamentous, often minute and evanescent ........................................... 8 Stipules apically obtuse or rounded, briefly 2- or 3- setulose ................... 9 Stipules apically acute or tapered to a long point .................................. 10 Leaves broadly elliptic, elliptic-oblong, or obovate, subcoriaceous; stipules usually persisting, turning dark brown with age; pedicel apically thickened into calyx; developing fruits oblong-obovoid, mature fruits 12–14 × 5–6 mm .................................................................. E. amazonicum Leaves lanceolate or oblong-lanceolate, chartaceous; stipules usually caducous; pedicel slender, not thickened toward apex; developing fruit oblong-ellipsoid, mature fruits 7–10 × 3–4 mm .................... E. citrifolium

Erythroxylum

10(8).

10. 11(10). 11. 12(1).

12. 13(12).

13. 14(13). 14. 15(14). 15. 16(14). 16. 17(16). 17. 18(17). 18.

19(17). 19. 20(19).

20.

21(19).

21.

61

Leaves 10–25 cm long; calyx 3.5–5 mm long, the lobes broadly ovate, elliptic, or obovate, abruptly acuminate at apex, the margins overlapping .......................................................................................... E. macrophyllum Leaves 3–10 cm long; calyx 1–3 mm long, the lobes triangular-ovate, oblong or lanceolate, acute at apex, the margins not touching ............. 11 Stipules usually caducous, or if persisting, then not fraying at apex; pedicel slender, not thickened into calyx ........................ E. macrophyllum Stipules usually persisting, fraying at apex (marcescent), becoming paleaceous; pedicel gradually thickened into calyx ........... E. mucronatum Stipules with 2 or 3 conspicuous, long, recurved fimbriate setae at apex; only known from one collection in the flora area (east of Upata) .......... ..................................................................................... E. aff. subrotundum Stipules not setulose at apex, or the setae short, filamentous, inconspicuous, often evanescent ........................................................................... 13 Lower surface of leaf with 2 distinct parallel lines running parallel to midrib, or, if lacking lines, with a discolorous central panel (often quite faint in E. spruceanum) ....................................................................... 14 Lower surface of leaf uniform, without 2 parallel lines nor with a discernible central panel ................................................................................. 22 Flowers produced at apex of new shoots on leafy shoots; endocarp terete .............................................................................................................. 15 Flowers produced on last season’s shoots, proximal to the leaves; endocarp sulcate ...................................................................................... 16 Leaves narrowly lanceolate, 3–6 × 0.8–1.1 cm; growing in waterfall spray ............................................................................................ E. lenticellosum Leaves elliptic, > 2 cm wide; riparian forests .......................... E. spruceanum Leaves mostly acute or acuminate at apex ................................. E. gracilipes Leaves rounded, retuse, or obtuse at apex .............................................. 17 Lower surface of leaf usually with a discernible, discolorous central panel, but without definite lines .................................................................... 18 Lower surface of leaf usually with 2 distinct parallel lines .................... 19 Flowers borne on leafless stems prior to leaf flush; branchlets without distinct lenticels; calyx lobes usually recurving in fruit .............. E. foetidum Flowers usually produced on leafy branches of current season’s shoots; branchlets with distinct punctate or elongate lenticels; calyx lobes erect or spreading in fruit ....................................................... E. gracilipes Fruiting pedicels thickened, 1–5 mm long .............................................. 20 Fruiting pedicels slender, not thickened apically, > 5 mm long ............. 21 Mature leaves coriaceous, often white-waxy on lower surface; branchlets 2–4 mm diameter, lenticels lacking or sparse; flowering pedicels < l mm long ....................................................................................... E. hypoleucum Mature leaves chartaceous or subcoriaceous, pale green or silvery on lower surface but not white-waxy; branchlets 1.5–2 mm diameter, covered with rounded or elongate lenticels; flowering pedicels 1.5–5 mm long ......................................................................................... E. lineolatum Twigs appearing knobby at nodes from persistent bracteoles; leaf blades 25–50 mm wide, often greenish when dry; flowering pedicels 3–6 mm long, in fruit 5–11 mm long ..................................................... E. gracilipes Twigs not conspicuously knobby at nodes; leaf blades 15–25 mm wide, dark brown when dry; flowering pedicels 1–4 mm long, in fruit 5–8 mm

62

E RYTHROXYLACEAE

long .................................................................................... E. cataractarum 22(13). Flowers produced on scaly lateral short shoots or in axils of persistent distichous imbricated cataphylls crowded on tips of twigs; some stipules occasionally with faint longitudinal veins .......................................... 23 22. Flowers produced in axils of leaves or cataphylls on new shoots; imbricate stipules not covering the branchlets; stipules never longitudinally veined ................................................................................................... 25 23(22). Leaves chartaceous, not shiny, 10–17 × 5–6.5 cm, distinctly acuminate at apex; petiole 4–7 mm long; pedicels 4–6 mm long .............. E. squamatum 23. Leaves coriaceous, shiny, mostly < 10 cm long, abruptly acuminate, acute, or obtuse at apex; petiole 1–3 mm long; pedicels 6–11 mm long ....... 24 24(23). Most leaves < 4 cm long, obovate to orbicular; endocarp terete; on tepuis > 1200 m elevation ................................................................ E. oreophilum 24. Most leaves > 5 cm long, elliptic to oblong or elliptic-obovate; endocarp sulcate; at elevations below 800 m ....................................... E. vernicosum 25(22). At least some of leaves acute or acuminate at apex, the tip itself sometimes obtuse ......................................................................................... 26 25. Leaves rounded, retuse, or obtuse at apex .............................................. 29 26(25). Pedicels 1–2 mm long; secondary leaf venation obscure ....... E. divaricatum 26. Pedicels > 2 mm long ............................................................................... 27 27(26). Leaves subcoriaceous, broadly elliptic, the apex rounded-retuse; petioles 4–7 mm long; pedicels 8–12 mm long ...................................... E. roraimae 27. Leaves membranous or chartaceous, narrowly elliptic, the apex acute to rounded; petioles 2–4 mm long; pedicels 3–6 mm long ...................... 28 28(27). Stems usually lenticellate; leaves usually > 7 cm long, not glaucous on lower surface, apex acute to acuminate .......................... E. kapplerianum 28. Stems not lenticellate; leaves usually < 7 cm long, glaucous on lower surface, apex acute to rounded ................................................. E. ligustrinum 29(25). Pedicels 1–2 mm long ............................................................................... 30 29. Pedicels > 2 mm long ............................................................................... 31 30(29). Leaves obovate to suborbicular, 8–23 mm long, deciduous; on granitic lajas in northwestern Amazonas state and adjacent parts of Bolívar ................................................................................................. E. williamsii 30. Leaves oblong-elliptic, 25–55 mm long, evergreen; rare, known only from one collection near Roraima-tepui ................................... E. schomburgkii 31(29). Flowers produced on leafless stems before the leaves; mature endocarp sulcate ..................................................................................... E. havanense 31. Flowers produced with the leaves; endocarp sulcate or terete .............. 32 32(31). Stipules and cataphylls subcoriaceous, black when dry; endocarps sulcate or terete ................................................................................................ 33 32. Stipules membranous, paleaceous when dry; endocarps terete ............. 34 33(32). Petiole 2–5 mm long; stipule 1.5–3.5 mm long; pedicel 5–7 mm long; drupe 4.5–5.5 mm long; endocarp sulcate ....................................... E. guanchezii 33. Petiole 6–9 mm long; stipule 3–6 mm long; pedicel 8–12 mm long; drupe 12–13 mm long; endocarp terete .............................................. E. roraimae 34(32). Mature leaves ovate to elliptic, rather shiny on both surfaces, the margins plane; calyx lobes with distinctly pale margins ..................... E. orinocense 34. Mature leaves elliptic to obovate, shiny or matte on upper surface, matte on lower surface, often revolute at margins; calyx lobes without pale margins .................................................................................. E. impressum

Erythroxylum

63

Erythroxylum amazonicum Peyr. in Mart., Fl. Bras. 12(1): 167. 1878. Evergreen shrub or medium-sized tree 3– 10 m tall. Moist forests and forest edges, often on sandy or rocky soil, in both lowland and montane forests, 50–1000 m; Bolívar (Altiplanicie de Nuria, middle Río Aponguao), Amazonas (Cañón Grande of Sierra de la Neblina, Pimichín, San Carlos de Río Negro, Sierra Parima, Yavita). Anzoátegui, Distrito Federal, Falcón, Miranda, Monagas, Yaracuy; Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 66. This poorly understood species grades into both Erythroxylum citrifolium and E. mucronatum, and some specimens are difficult to determine with certainty. Specimens from cloud forests outside the flora area are fairly distinct from the Amazonian collections.

m; widely scattered in Bolívar and Amazonas. Falcón, Lara, Mérida, Táchira, Zulia; Nicaragua to Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

Erythroxylum cataractarum Spruce ex Peyr. in Mart., Fl. Bras. 12(1): 149. 1878. Leafy evergreen shrub or treelet to 4 m tall. Moist forests, partially deciduous gallery forests, river margins, often growing in flooded sandy or rocky soils and on rocky river banks, also in marshy mixed forest of Mauritia palms, 50–200 m; Amazonas (Río Atabapo, Río Casiquiare, Río Emoni, Río Guayapo, Río Pasimoni, Río Sipapo, Río Ventuari). Apure, Guárico, Miranda; eastern Colombia, Brazil (Amazonas: upper Rio Negro basin).

Erythroxylum fimbriatum Peyr. in Mart., Fl. Bras. 12(1): 162. 1878. Evergreen shrub or treelet to 6 m tall. Moist lower montane forests, ca. 600 m; Bolívar (Sierra de Lema at La Escalera). Costa Rica, Colombia, French Guiana, Ecuador, Peru, Brazil (Acre, Amazonas, Rondônia), Bolivia. This species is known in Venezuela only from a single fruiting collection, Foldats 2853 (F, VEN).

Erythroxylum citrifolium A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 2: 67[94]. 1829. Erythroxylum gomphioides Planch. & Linden ex Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 341. 1862. Erythroxylum acutifolium Steud. ex Peyr. in Mart., Fl. Bras. 12(1): 166. 1878. Erythroxylum micranthum Bong. ex Peyr. in Mart., Fl. Bras. 12(1): 164, t. 30, fig. 1. 1878. Erythroxylum paraense Peyr. in Mart., Fl. Bras. 12(1): 164, t. 30, fig. 3. 1878. Erythroxylum duckei Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 416. 1909. Evergreen shrub or small tree 3–10 m tall. Widespread in moist gallery forests, secondary forests, and forest margins, 50–1300

Erythroxylum divaricatum Peyr. in Mart., Fl. Bras. 12(1): 146. 1878. —Jobo amarillo. Slender evergreen shrub or treelet 2–7 m tall, with straight, divaricating branches. Occasional in seasonally flooded riparian forests and savannas on sandy soils, 50–500 m; Bolívar (Río Paragua at Raudal de Guaiquinima and San Pedro de las Dos Bocas, Río Parguaza), Amazonas (Caño Majagua, scattered along Río Orinoco from Puerto Ayacucho south to Santa Bárbara and also at Raudal de los Guaharibos, Río Ventuari, San Carlos de Río Negro). Guyana, northern Brazil (Amazonas, Pará, Roraima). ŠFig. 67.

Erythroxylum foetidum Plowman, Brittonia 40: 256, t. 1. 1988. Completely deciduous shrub to 3 m tall, with erect-ascending branches; leaves glaucous on lower surface. Shrub islands on granitic outcrops, 50–200 m; Amazonas (near Puerto Ayacucho). Apure. Erythroxylum gracilipes Peyr. in Mart., Fl. Bras. 12(1): 159. 1878. —Fruto de paloma. Erythroxylum novogranatense var. macrophyllum O.E. Schulz in Engl., Pflanzenr. IV. 134(Heft 29): 87. 1907. Erythroxylum recurrens Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 417. 1909. Erythroxylum cuatrecasasii W.A. Gentner, J. Wash. Acad. Sci. 47: 6, t. 1. 1957. Variable evergreen or partly deciduous shrub or small tree to 5 m. Wet to seasonally

64

E RYTHROXYLACEAE

dry forests on terra firme and along riverbanks, often on sandy or rocky substrate, 0–700 m; Delta Amacuro (middle Río Grande, Sacupana), Bolívar (Cerro Arimagua near Represa Guri, Río Canaracuni, upper Río Caura, Río Cuyuní, Serranía de Imataca, Serranía de los Pijiguaos), Amazonas (Río Atacavi, Río Baría, Salto Yureba in lower Río Ventuari basin, San Carlos de Río Negro, Yavita to Maroa, Yutajé). Apure, Barinas, Distrito Federal, Guárico, Mérida, Miranda, Táchira, Zulia; widely distributed in Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 71. Erythroxylum guanchezii Plowman, Brittonia 40: 261, t. 4. 1988. Evergreen treelet 5–6 m tall; leaves elliptic-obovate, retuse. Shrublands on white sand, 100–200 m; Amazonas (La Esmeralda, Río Guayapo south of Raudal Moriche). Endemic. Erythroxylum havanense Jacq., Enum. Syst. Pl. 21. 1760. —Cabeza de negro, Escoba, Jayo, Fruto de paloma. Erythroxylum ovatum Cav., Diss. 404, t. 233. 1789. Erythroxylum obtusum DC., Prodr. 1: 574. 1824. Erythroxylum havanense var. continentis O.E. Schulz in Engl., Pflanzenr. IV. 134(Heft 29): 92. 1907. Deciduous shrub or treelet to 2–6 m tall. Deciduous forests especially in outcrop areas and bordering lajas, gallery forests, shrublands, 100–300 m; Bolívar, (Ciudad Bolívar, Puerto Ordaz, Represa Guri, middle Río Caura, Serranía de Imataca). Anzoátegui, Aragua, Carabobo, Distrito Federal, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Zulia; Mexico, Central America, Cuba, Lesser Antilles, Colombia, Trinidad. Erythroxylum hypoleucum Plowman, Bot. Mus. Leafl. 29: 280, t. 30. 1983. —Cachito, Palo de máguari, Palo gallineta, Turí. Evergreen slender tree to 8 m tall, with erect-ascending branches. White-sand shrub savannas (bana), 100–200 m; Amazonas (San Carlos de Río Negro, between Yavita

and Maroa). Northern Brazil (Amazonas: Rio Uaupés). Erythroxylum impressum O.E. Schulz in Urb., Symb. Antill. 5: 202. 1907. —Escobo negro, Jayo, Tepa pa meh (Panare). Erythroxylum squamatum var. orinocense Kuntze, Revis. Gen. Pl. 1: 86. 1891. Deciduous shrub or treelet to 6 m tall. Low semideciduous forests and shrubby savannas over granitic outcrops, often on rocky slopes and along rocky streams, 50–400 m; northern Bolívar (Altiplanicie de Nuria, 23 km southwest of Canaima, Cerro Médano near Caicara, west of El Manteco, Maniapure, Represa Guri, Serranía de Imataca, Serranía de los Pijiguaos, west of Upata), Amazonas (Puerto Ayacucho north to Galipero). Colombia (opposite Puerto Ayacucho), Trinidad, Guyana, Brazil (Amazonas). ŠFig. 69. Erythroxylum kapplerianum Peyr. in Mart., Fl. Bras. 12(1): 159. 1878. Erythroxylum ligustrinum var. grandifolium Sagot, Ann. Sci. Nat. Bot. sér. 6, 11: 179. 1881. Erythroxylum mapuerae Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 421. 1909. Evergreen shrub or tree 3–10 m tall. Forested granitic, often flooded, outcrop areas, sandy soils along black-water rivers, 100– 400 m; Bolívar (Río Asa, 25 km southwest of Maripa, Reserva Forestal La Paragua, middle and lower Río Caura, Río Erebato, Salto Pará), Amazonas (lower Río Ventuari, Río Yureba). Colombia (Vaupés), Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, Pará, Rondônia). ŠFig. 73. Erythroxylum lenticellosum Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 420. 1909. Evergreen shrub 1–2 m tall; flowers subapical, few or solitary; pedicels 4–5 mm long. On rocks in spray of waterfall, 100–200 m; Amazonas (Salto Gallineta on Río Siapa). Brazil (Pará). Erythroxylum ligustrinum DC., Prodr. 1: 574. 1824.

Erythroxylum

Deciduous shrub or small tree to 12 m tall. Seasonally dry, semievergreen forests on rocky or sandy soil, often in association with igneous outcrops, bordering white-sand savannas and along streams, 50–500 m; Amazonas (Garcitas between Puerto Ayacucho and Samariapo, Puerto Ayacucho to Gavilán, Río Coromoto). Suriname, French Guiana, Brazil (Amapá, Maranhão, eastern Pará). The Venezuelan specimens differ from the eastern Amazonian material mainly in having fewer (8–12 versus 12–20) pairs of lateral veins in the leaves. Erythroxylum lindemanii Plowman, Phytologia 58: 172, t. 1. 1985. Deciduous shrub or treelet 2–7 m tall. Restricted to shrub islands on granitic outcrops, 100–200 m; Amazonas (Puerto Ayacucho north to the border with Bolívar). Colombia (opposite Puerto Ayacucho), Guyana, Suriname. Erythroxylum lineolatum DC., Prodr. 1: 575. 1824. Evergreen shrub or treelet to 4 m tall. Riparian forest formations and low woods on rocky slopes, cloud forests, savanna margins, 300–800(–1400) m; Bolívar (summit of Cerro Guaiquinima, Río Tonoro north of Cerro Guaiquinima, upper Río Trueno 35 km west of Chiguao), Amazonas (Río Autana). Aragua, Barinas, Carabobo, Sucre; Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana. Outside the Venezuelan Guayana, this uncommon species grows mainly in dry deciduous forests or savanna margins. The few and incomplete collections of Erythroxylum lineolatum from the Venezuelan Guayana approach E. hypoleucum or even depauperate forms of E. gracilipes, and some hybridization may have occurred among these species. Erythroxylum macrophyllum Cav., Diss. 401, t. 227. 1789. Erythroxylum lucidum H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 179. 1821 [1822]. Erythroxylum floribundum Mart., Beitr. Erythroxylon 118. 1840 [preprint from Abh. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. 3: 398. 1841].

65

Erythroxylum laurinum Planch. & Linden ex Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 341. 1862. Erythroxylum filipes Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 415. 1909. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 4 varieties, 2 in Venezuela, both in the flora area. The two varieties of Erythroxylum macrophyllum that occur in the Venezuelan Guayana are quite distinct, morphologically and ecologically. In our area, var. macrophyllum matches well with the type from French Guiana. However, some confusion may be expected when var. savannarum is compared with material from other areas of the range of this complex group. Key to the Varieties of E. macrophyllum 1. Leaves 10–23 cm long, usually acuminate (sometimes abruptly so) at apex; stipules and cataphylls caducous or lasting just one season; calyx lobes foliose, broadly ovate or obovate, overlapping at the margins; fruits ca. 12 mm long ........ ............................. var. macrophyllum 1. Leaves 3–10 cm long, acute, obtuse, or rounded at apex; stipules and cataphylls sometimes persisting more than one season, giving branchlets a scaly appearance; calyx lobes narrowly ovate, oblong or lanceolate, not overlapping at margins; fruits 7–10 mm long ................ ................................ var. savannarum E. macrophyllum var. macrophyllum Evergreen shrub or small tree to 15 m tall. Evergreen lowland to lower montane forests on both sandy and clay soils, sometimes on rocky slopes, 100–800 m; Bolívar (Amaruay-tepui, Perai-tepui west of Santa Elena, Represa Guri, headwaters of Río Túriba), Amazonas (upper Río Baría, Río Coromoto, upper Río Ventuari, San Juan de Manapiare). Aragua, Barinas, Carabobo, Mérida, Miranda, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 75.

66

E RYTHROXYLACEAE

E. macrophyllum var. savannarum Plowman, Brittonia 40: 263, t. 5. 1988. —Escobo, Miel de pajarito, Pilón negro, Socoroco negro. Evergreen shrub or treelet to 12 m tall. Moist forests adjacent to or surrounded by savannas, in tree islands, on rocky outcrops, gallery forests, on well-drained, often rocky soils, 50–1000 m; Bolívar (Río Paragua basin, Serranía de Imataca), Amazonas (Caño Yapacana, base of Cerro Moriche, Río Orinoco near Puerto Ayacucho, upper Río Parú). Apure; eastern Colombia. Erythroxylum mucronatum Benth., London J. Bot. 2: 372. 1843. —Kawadadek (Arekuna), Kawudadek (Arekuna), Perumatá. Erythroxylum comosum O.E. Schulz in Engl., Pflanzenr. IV. 134(Heft 29): 32, t. 8. 1907. Erythroxylum kirkianum O.E. Schulz, Repert. Spec. Nov. Regni Veg. 30: 179. 1932. Erythroxylum venezuelense Steyerm., Fieldiana, Bot. 28: 271. 1952. Erythroxylum albertianum Kuhlm. & W.A. Rodrigues, Publ. Inst. Nac. Pesq. Amazônia Bot. 5: 3. 1957. Evergreen shrub or treelet 1–8 m tall. Evergreen lowland to montane forests, often along rivers and rocky streams, tepui slopes, commonly in sandy or rocky soils, 100–1200 m; Bolívar (widely scattered in the Gran Sabana, Sierra de Lema), widely scattered in Amazonas. Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ŠFig. 72. Erythroxylum oreophilum (O.E. Schulz ex Pilg.) Steyerm. & Maguire, Mem. New York Bot. Gard. 17: 445. 1967. —Erythroxylum vernicosum var. oreophilum O.E. Schulz ex Pilg., Notizbl. Königl. Bot. Gart. Berlin 5: 142. 1914. Evergreen shrub or small tree 2–8 m tall. Elfin tepui forests, edges of meadows, upper tepui slopes (on both sandstone and granitic substrates), 1200–2200 m; Bolívar (Auyán-tepui, Cerro El Sol on Venezuela-Brazil border, Cerro Guaiquinima, Cerro Jaua, Macizo del Chimantá), Amazonas (Sierra de Maigualida). Brazil (Roraima). ŠFig. 68.

This is the only species of Erythroxylum endemic to the tepuis, and it represents the highest altitudinal record for the genus. It is closely related to, and probably derived from, E. vernicosum, a species that occurs at lower elevations. Erythroxylum orinocense H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 178, t. 453. 1821 [1822]. —Fruta de paloma. Deciduous shrub 1–3 m tall. Savanna margins and shrub islands, 50–100 m; Bolívar (Cerro San Borja 50 km southwest of Caicara, Túriba), Amazonas (Puerto Ayacucho and vicinity). Apure, Barinas, Cojedes, Distrito Federal, Guárico, Miranda, Portuguesa, Zulia; llanos of Colombia. ŠFig. 76. Erythroxylum roraimae Klotzsch ex O.E. Schulz in Engl., Pflanzenr. IV. 134(Heft 29): 104. 1907. Evergreen tree to 10 m tall. Lower montane forests, 300–1200 m; Bolívar (near Cerro Venado, headwaters of Río Túriba 45 km east of Los Pijiguaos, southern slope of Roraima-tepui), Amazonas (Reserva Forestal El Sipapo). Colombia (Amazonas), Guyana, Suriname, Brazil (Amazonas, Roraima, possibly Amapá). In the populations from Guyana, Suriname, French Guiana, and eastern Bolívar state in the flora area, the leaf apices are rounded or obtuse. In the western part of the range, the apices are abruptly acuminate or acute. Erythroxylum rufum Cav., Diss. 404, t. 232. 1789. —Jaillito. Deciduous shrub or tree to 5 m tall. Deciduous forest on rocky slopes, shrub islands on granitic outcrops and bordering savannas, 50–200 m; Bolívar (Caicara, Cerro Médano, islands in Represa Guri, Río Asa, Serranía de los Pijiguaos), Amazonas (near Puerto Ayacucho). Apure, Guárico, Monagas, Zulia; widely distributed in Greater Antilles, Guyana, French Guiana, northern Brazil. Erythroxylum schomburgkii Peyr. in Mart., Fl. Bras. 12(1): 148. 1878. Evergreen shrub or treelet to 5 m tall. Granitic outcrops, savanna margins on ex-

Erythroxylum

posed rocks, ca. 1000 m(?); Bolívar (base of Roraima-tepui). Guyana, Brazil (Roraima). Erythroxylum spruceanum Peyr. in Mart., Fl. Bras. 12(1): 160, t. 30, fig. 2. 1878. Evergreen tree 3–7(–15) m tall. Seasonally flooded forests along black-water rivers, 100–200 m; Amazonas (Caño Tamatama, Río Guainía, Río Yatúa). Barinas; Amazonian Colombia, Peru (Loreto), Brazil (Amazonas). A specimen from the upper slopes of Sierra de la Neblina, Maguire et al. 37217 (F), at 1700–1800 m elevation, is referred here, but occurs at much higher elevations than is normal for the species. Erythroxylum squamatum Sw., Prodr. 75. 1788. Erythroxylum aristigerum Peyr. in Mart., Fl. Bras. 12(1): 157. 1878. Erythroxylum bahiense Peyr. in Mart., Fl. Bras. 12(1): 160. 1878. —Erythroxylum aristigerum var. bahiense (Peyr.) O.E. Schulz in Engl., Pflanzenr. IV. 4(Heft 134): 103. 1907. Erythroxylum squamatum var. microcarpum O.E. Schulz in Urb., Symb. Antill. 5: 192. 1907. Erythroxylum trinerve Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 417. 1909. Evergreen shrub or small tree to 10 m tall. Riparian forests, 100–200 m; Bolívar (Río Erebato), Amazonas (Río Guainía near Maroa). Miranda, Sucre; Lesser Antilles, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Erythroxylum steyermarkii Plowman, Brittonia 34: 442, t. 1. 1982. —Jaillito. Deciduous shrub to 5 m tall. Deciduous forests or scrub, often on rocky igneous slopes and ravines, 50–200 m; Bolívar (Caicara and 35 km to the southwest). Anzoátegui, Nueva Esparta, Sucre, Zulia. ŠFig. 70. Erythroxylum suberosum A. St.-Hil., Pl. Usuel. Bras. pl. 69, fig. A. 1824. —Atónyek (Arekuna). Erythroxylum areolatum Vell., Fl. Flumin. 194. 1825 [1829], non L. 1758. Erythroxylum testaceum Peyr. in Mart., Fl. Bras. 12(1): 170. 1878.

67

Gnarled evergreen or semideciduous shrub 1–2 m tall; bark thick, corky. Open rocky or grassy savannas, on rocky slopes and on mounds in moist savannas, 800–1000 m; Bolívar (Cerro Akurimá and San Antonio del Morichal near Santa Elena de Uairén, Gran Sabana). Guyana, French Guiana, Brazil, Bolivia, Paraguay. ŠFig. 74. Erythroxylum aff. subrotundum A. St.Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 2: 71[99]. 1829. —Erythroxylum christii O.E. Schulz, Repert. Spec. Nov. Regni Veg. 30: 179. 1932, nom. nud. Deciduous shrub or spreading tree 3–7 m tall. Deciduous forests on manganese outcrops, 300–400 m; Bolívar (Cerro Los Bloques and Cerro Paja [=Guacuripia] 25 km east of Upata). Guárico; eastern and central Brazil, Bolivia. The identity of the Venezuelan material remains questionable, but it most closely resembles Erythroxylum subrotundum, based on the distinctive long-setulose, fimbriate stipules. A similar disjunct distribution from Guayana to the Brazilian Shield is also seen in the species pair E. steyermarkii Plowman and E. laetevirens O.E. Schulz. Erythroxylum vernicosum O.E. Schulz in Engl., Pflanzenr. IV. 134(Heft 29): 106. 1907. Erythroxylum squamatum var. emarginatum Peyr. in Mart., Fl. Bras. 12(1): 158. 1878. Slender evergreen shrub or treelet to 6 m tall. Rocky outcrops, white-sand savannas, open forests on sandy soil, river banks on rocky soil, 100–800 m; Bolívar (Canaima, slopes of Cerro Bolívar, Río Guaniamito), Amazonas (upper Río Orinoco at Salto Salas). Guyana, Brazil (Roraima). Erythroxylum williamsii Standl. ex Plowman, Brittonia 34: 453, t. 6. 1982. —Punteral. Diffusely branched deciduous shrub or small tree to 6 m tall. Shrub islands on exposed granitic outcrops, 50–100 m; Bolívar (Agua Amena along Caicara to Puerto Ayacucho road, Río Parguaza), Amazonas (Puerto Ayacucho to Galipero). Guárico; Colombia (Vichada). ŠFig. 77.

68

E RYTHROXYLACEAE

Fig. 66. Erythroxylum amazonicum

Fig. 67. Erythroxylum divaricatum

Fig. 68. Erythroxylum oreophilum

Erythroxylum

Fig. 69. Erythroxylum impressum

Fig. 70. Erythroxylum steyermarkii

Fig. 71. Erythroxylum gracilipes

69

70

E RYTHROXYLACEAE

Fig. 72. Erythroxylum mucronatum

Fig. 74. Erythroxylum suberosum

Fig. 73. Erythroxylum kapplerianum

Erythroxylum

Fig. 75. Erythroxylum macrophyllum var. macrophyllum

Fig. 76. Erythroxylum orinocense

Fig. 77. Erythroxylum williamsii

71

72

E UPHORBIACEAE

EUPHORBIACEAE by Grady L. Webster, Paul E. Berry, W. Scott Armbruster, Hans-Joachim Esser, Lynn J. Gillespie, W. John Hayden, Geoffrey A. Levin, Ricardo de S. Secco, and Scott V. Heald Trees, shrubs, herbs, or vines, sometimes succulent. Leaves alternate or less commonly opposite (rarely whorled), simple or palmately lobed to compound, glabrous or pubescent with simple or branched trichomes, sometimes with glands at base of blade; margins entire or dentate; stipules persistent or deciduous, sometimes absent. Inflorescences cymose, racemose, spicate, or pseudanthial, or flowers solitary; bracts persistent or deciduous, sometimes enlarged or glandular. Flowers unisexual (plants monoecious or dioecious); perianth biseriate, uniseriate, or rudimentary to absent. Sepals imbricate or valvate, free or basally connate; petals showy to reduced or absent; floral disk entire to dissected, or absent. Stamens (1)2–many, free or united; anthers mostly 2-locular and dehiscing longitudinally. Ovary superior, mostly 3- or 4-locular (rarely 1-locular or multilocular); placentation axile; ovules 1 or 2 per locule, anatropous or less commonly hemitropous, usually inserted beneath an obturator (a placentary tissue from the funicle that covers the micropyle and serves to conduct the sperm tubes towards the embryo sac); styles free or united, often bifid or multifid. Fruit usually capsular and separating into 3 cocci, sometimes baccate or drupaceous. Seeds with dry or fleshy coat, often carunculate, usually with endosperm; embryo large. Tropical and subtropical regions of both hemispheres; ca. 320 genera and 8100 species, 58 genera and 239 species in the flora area. Chonocentrum cyathophorum (Müll. Arg.) Pierre ex Pax & K. Hoffm., the sole member of the genus, is known only from the type collection, Spruce 3781 (BM, GDC, K, NY, P). The Kew specimen lists the locality as an island in the Rio Negro at the mouth of the Rio Marania, in Brazil (A. Radcliffe-Smith, personal communication), not the mouth of the Río Casiquiare in Amazonas state, Venezuela, as reported by Pittier et al. (Catálogo de la Flora Venezolana 2: 69. 1947).

Key to the Genera of Euphorbiaceae by Paul E. Berry, Grady L. Webster, and Hans-Joachim Esser 1. 1. 2(1).

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

Vines or lianas ............................................................................................ 2 Herbs, shrubs, or trees ............................................................................... 6 Inflorescences long-pedunculate (in Venezuelan species), usually bracteate pseudanthia (bearing 3 pistillate and 6 or more staminate flowers); minute stinging trichomes present ................................. 19. Dalechampia Inflorescences not bracteate pseudanthia ................................................. 3 Woody lianas, stems with reddish exudate or copious white latex; stamens either 2 or numerous ............................................................................. 4 Twining vines or lianas, stems lacking exudate; stamens 3 or more ....... 5 Stems with reddish exudate; leaves (broadly) elliptic-ovate, with palmate venation and 2 large petiolar glands; inflorescence paniculate; stamens 2, connate, anther connectives greatly enlarged; fruit a schizocarpous capsule or berry, 3-lobed or globose, sometimes with a conical beak,

EUPHORBIACEAE 73

4.

5(3). 5. 6(1). 6. 7(6). 7. 8(6). 8. 9(8). 9. 10(9). 10. 11(8). 11. 12(11).

12.

13(12).

13.

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

15.

8–12 cm diameter .................................................................. 40. Omphalea Stems with copious white latex; leaves (narrowly) elliptic, with pinnateparallel venation, and lacking petiolar glands; inflorescence a cylindrical “bottlebrush” thyrse; stamens numerous; fruit capsular, velvetypubescent, much smaller than above ......................................... 34. Mabea Stinging hairs absent; leaf blades glandular near base; stamens > 10; styles connate into a long column; fruits > 1 cm diameter ............ 46. Plukenetia Stinging hairs present; leaf blades not glandular near base; stamens < 10; styles basally connate; fruits < 1 cm diameter ................. 58. Tragia Leaves opposite or verticellate, at least in upper part of stems; flowers in cyathia with connate bracts simulating a calyx ................................... 7 Leaves entirely alternate (sometimes clustered); flowers variously arranged, sometimes in cyathia ................................................................ 8 Herbs; leaves opposite, asymmetrical at base ...................... 15. Chamaesyce Shrubs or small trees; leaves verticellate, symmetrical at base ................ ...................................................................... 24. Euphorbia (E. cotinifolia) Leaves palmately compound ...................................................................... 9 Leaves simple, although sometimes deeply lobed .................................. 11 Shrubs; stems with white latex; perianth petaloid ..................... 35. Manihot Trees; stems with or without white latex; perianth not petaloid ........... 10 Latex present; leaflets stalked (petiolule > 5 mm long); inflorescence paniculate ..................................................................................... 29. Hevea Latex absent; leaflets subsessile (petiolule < 5 mm long); inflorescence spicate ..................................................................................... 45. Piranhea Leaves palmately lobed ............................................................................ 12 Leaves not palmately lobed (but may be palmately veined at the base) .............................................................................................................. 16 Weedy annual herbs usually < 0.5 m tall, or trees; stellate trichomes present, sometimes mixed with simple trichomes; inflorescences terminal racemes ........................................................................ 18. Croton Weedy or nonweedy, usually perennial large herbs or shrubs generally > 0.5 m tall; stellate trichomes absent; inflorescences axillary or terminal, dichasial or racemose ................................................................... 13 Stinging or glandular trichomes present (except Jatropha curcas); perianth present and petaloid, white; stamens 8–10, largely connate; inflorescences terminal dichasia ................................................................. 14 Stinging or glandular trichomes absent; perianth absent, or, if present, then yellowish to purplish; stamens either 10 and free or 50–many; inflorescences terminal racemes or panicles .......................................... 15 Petals absent in pistillate flowers, the sepals petaloid; trichomes strongly stinging ................................................................................ 16. Cnidoscolus Petals present in pistillate flowers; trichomes glandular (or absent), not stinging .................................................................................... 33. Jatropha Stems without latex; leaves markedly serrate and peltate; perianth not petaloid; stamens 50–many, connate into irregular fascicles; ovary and fruit muricate ............................................................................. 50. Ricinus Stems with white latex; leaves neither markedly serrate, nor peltate; perianth petaloid; stamens 10, free, filaments not branched; ovary and fruit smooth or ribbed, not muricate ....................................... 35. Manihot

74

E UPHORBIACEAE

16(11). 16. 17(16). 17. 18(17).

18. 19(18).

19. 20(19).

20. 21(20).

21.

22(17).

22.

23(22).

23. 24(23).

24.

25(24). 25. 26(25). 26.

Leaf margins serrate, crenulate, or dentate ........................................... 17 Leaf margins entire .................................................................................. 42 Herbs or subshrubs (< 1 m tall) ............................................................... 18 Shrubs or trees ......................................................................................... 22 Pistillate inflorescences densely spicate, resembling a bushy tail, with the flowers subtended by conspicuous toothed bracts, the bracts enlarging in fruit; style branches much divided and laciniate ................ 1. Acalypha Inflorescences different from above ........................................................ 19 Leaves with densely, shortly serrate margins (teeth < 0.5 mm apart), with simple (multicellular) trichomes; staminate flowers in spikes opposite the leaves; pistillate flowers borne on the stems between nodes; fruits with triangular spur-like projections ............................... 39. Microstachys Leaves less densely and more coarsely serrate ....................................... 20 Plants with abundant white latex; the staminate and pistillate flowers reduced to a single stamen and ovary, respectively, grouped together in a cyathium .................................................. 24. Euphorbia (E. heterophylla) Plants lacking white latex; flowers in raceme- or spike-like inforescences, flowers not reduced as above ............................................................... 21 Plants usually found in wet or swampy areas, lacking stellate trichomes; stems usually hollow; venation pinnate with 8–20 pairs of secondary veins ...................................................................................... 12. Caperonia Plants usually not found in swampy areas, with stellate trichomes; stems solid; venation palmate-pinnate, with fewer secondary veins than above ..................................................................... 18. Croton (C. trinitatus) Trunk densely covered by woody, conical spines; staminate flowers many in a fleshy, reddish cone; pistillate flowers solitary, carpels mostly 10–15, stylar column terminating in an expanded disk with lobed margins; fruit a woody capsule to 8 cm diameter (explodes with great force) ........................................................................................................ 30. Hura Trunk without dense conical spines (occasional slender spines present in Adelia); flowers and inflorescences different from above; fruit smaller than above, with fewer carpels ............................................................ 23 Stellate or peltate trichomes present on some part of the plant, usually on the lower surface of the leaves, sometimes the stellate trichomes minute .................................................................................................. 24 Plants glabrous or else trichomes simple or branched ........................... 27 Trichomes stellate or peltate; stamens mostly 8–30, incurved in bud; petioles often with a pair of glands at the junction with the leaf blade; inflorescences terminal, racemoid .............................................. 18. Croton Trichomes stellate, never peltate; stamens 3–ca. 60, straight in bud; with or without glands at the junction of the petiole and leaf blade; inflorescences various ...................................................................................... 25 Inflorescences axillary racemes or thyrses; styles 2, long and slender, unbranched; stamens 8; fruits 2-locular; seeds tuberculate ....... 6. Alchornea Inflorescences robust terminal racemes; styles 2 or 3, bifid; stamens 15 or more; fruits 2- or 3-locular; seeds smooth ........................................... 26 Styles 3, branches blunt; pistillate sepals free; anthers blunt ............. .............................................................................................. 17. Conceveiba Styles 2 (rarely 3), branches acute; pistillate sepals connate; anthers apiculate ............................................................................... 25. Gavarretia

E UPHORBIACEAE 75

27(23). Leaf blades widest above the middle and gradually narrowed toward the base ....................................................................................................... 28 27. Leaf blades elliptic or lanceolate, widest near the base or middle, ± obovate or oblanceolate in Acidoton, Astrococcus, and Richeria ........................ 29 28(27). Leaves with secondary veins < 8 per side; petiole < 2 cm long .................. .......................................................................................... 5. Adenophaedra 28. Leaves with secondary veins > 10 per side; petiole > 3 cm long ................ .............................................................................................. 41. Pausandra 29(27). Leaves with a pair of stipel-like projections at the base on the upper surface and round glands in vein axils on the lower surface, covered by very short, straight trichomes; staminate flowers with 3–5 stamens .......................................................................................... 9. Aparisthmium 29. Leaves without the combination of stipels and glands described above, glabrous, or with simple, T-shaped, dendritic, or stinging trichomes; staminate flowers covering a wider range of stamen number ........... 30 30(29). Flowers in pseudanthia with pink or rarely yellowish involucral bracts at anthesis; inflorescence glands yellow or greenish, the surface papillate, producing spicy fragrance ................. 19. Dalechampia (D. magnoliifolia) 30. Flowers non-pseudanthial, without colorful bracts; flowers not fragranceproducing and lacking inflorescence glands ....................................... 31 31(30). Leaf blades with teeth only in the upper 1/2–2/ 3 of the margin, with 3–7 prominent raised secondary veins paralleling the midrib; stipules persistent; stinging trichomes present on the gynoecium and apex of the anthers ................................................................................. 2. Acidoton 31. Leaf blades with teeth or crenations ± evenly distributed along the margin, without the venation described above; stipules usually caducous; stinging trichomes absent ................................................................... 32 32(31). Petioles with raised glands below the junction of the leaf blade ............ 33 32. Petioles without raised glands below the leaf base ................................ 34 33(32). Plants glabrous, with white latex; flower groups sessile on the rachis; staminate flowers with 2 stamens, pistillate flowers with simple, unbranched stigmas ...................................................................... 53. Sapium 33. Plants with an indumentum, lacking latex; flower groups on short (1–3 mm long) lateral peduncles; staminate flowers with 3 stamens, pistillate flowers with bifid stigmas .............................................. 57. Tetrorchidium 34(32). Plants with milky latex; inflorescence dense, like a bottlebrush; flowers long-pedicellate; floral bracts with (0)1 pair of elliptic-cylindrical glands in variable position to the axis; fruits velvety pubescent or tomentose .................................................................................... 34. Mabea 34. Plants without milky latex; inflorescence narrow, with short-pedicellate flowers; bracts lacking the kind of glands mentioned above; fruits not velvety or tomentose ............................................................................ 35 35(34). Style branches much divided and laciniate; staminate flower buds minute, < 1 mm diameter; fruit usually subtended by a broad bract ................................................................................................... 1. Acalypha 35. Style branches simple to multifid, not laciniate; staminate flower buds 1–3 mm diameter; fruits not subtended by a broad bract .................. 36 36(35). Flowers with well-developed petals; stamens 10, filaments connate; small shrub 1–2 m tall ..........................................................................22. Ditaxis 36. Flowers apetalous; stamens 3–7(–20), free to connate; small to large

76

E UPHORBIACEAE

shrubs or small trees ........................................................................... 37 37(36). Fruits drupaceous; plants dioecious; leaves coriaceous and usually with asymmetrical bases; pistillate flowers with 1 or 2 carpels; stigmas sessile, dilated .......................................................................... 23. Drypetes 37. Fruits capsular; plants monoecious or dioecious; leaves membranaceous to coriaceous, the bases usually symmetrical; pistillate flowers with 3 carpels; styles present, variously shaped, free or connate .............. 38 38(37). Shrubs or small trees, mostly 1–4 m tall; stamens 4, filaments connate; capsules dry, horned or not .................................................................. 39 38. Small to large shrubs or trees; stamens 3–8, free; capsules dry or fleshy, not horned ............................................................................................ 40 39(38). Staminate flowers with a disk; pistillate flowers with an ovoid-urceolate style; carpels horned in fruit .............................................. 10. Astrococcus 39. Staminate flowers without a disk, but the filaments highly dilated and fused at the base; pistillate flowers with styles connate into a massive, elongated, cup-shaped, hollow-centered column, truncate and papillose at apex; carpels not horned in fruit ............................. 28. Haematostemon 40(38). Leaves tripliveined; stamens 4–8, inserted on a massive pubescent disk; anthers with connective enlarged and glandular; capsules dry, 3-lobed ............................................................................................7. Alchorneopsis 40. Leaves pinnately veined; stamens 3–6(–20), alternating with the disk glands or disk absent; anthers without glandular connectives.......... 41 41(40). Small shrubs 1–2 m tall; leaves pubescent on lower surface, margins finely serrate; capsules dry, globose ...................................... 11. Bernardia 41. Trees > 4 m tall; leaves glabrous on both surfaces, margins crenate; capsules fleshy, oblong .................................................................. 49. Richeria 42(16). Plants with stellate or peltate trichomes on leaves and inflorescences .............................................................................................................. 43 42. Plants glabrous or with simple, T-shaped, or dendritic trichomes on leaves and inflorescences ................................................................................ 45 43(42). Inflorescences racemose, without side branches; petals present, at least in staminate flowers; stamens inflexed in bud, 8–50 per flower; fruit capsular ............................................................................................. 18. Croton 43. Inflorescences paniculate or else flowers in axillary clusters; petals absent; stamens erect in bud, 3–6 per flower; fruit drupaceous or capsular (schizocarp) .......................................................................................... 44 44(43). Inflorescence pseudanthial, in short axillary clusters surrounded by a pair of fused bracts; stamens 3 or 4; fruit a schizocarp ................. 43. Pera 44. Inflorescence paniculate, not pseudanthial; stamens 3–6; fruit a drupe .............................................................................................. 31. Hyeronima 45(42). Leaves tripliveined, with glands near the base of the blade .................. 46 45. Leaves pinnately veined, with or without glands near the base of the blade, or, if appearing tripliveined, then without glands near the base of the blade ........................................................................................... 49 46(45). Leaf blade with a pair of glands outside the axils of the basal pair of lateral veins; pit domatia often present in the vein axils; styles and carpels 3; seeds with outer coat fleshy, bright red; inflorescence spicate ............................................................................................7. Alchorneopsis 46. Leaf blade with glands either at the petiole apex or inside the basalmost

EUPHORBIACEAE 77

47(46).

47.

48(47).

48.

49(45). 49.

50(49).

50.

51(49).

51.

52(51).

52. 53(52). 53. 54(53).

54.

55(54).

pair of lateral veins; styles and carpels 2 (3 in Actinostemon schomburgkii); seeds with outer coat dry, not red; inflorescence paniculate or spicate .................................................................................................. 47 Leaves with a pair of conspicuous sunken glands at the apex of the petiole; stigmas sessile, bifid, contorted; stamens 25–30; fruit drupaceous, fusiform, ca. 4–6 cm long; seeds crustaceous .................. 26. Glycydendron Leaves glandular but without conspicuous sunken glands at the apex of the petiole; styles 2 or 3; stamens fewer than above; fruit capsular, smaller than above; seeds tuberculate or smooth .............................. 48 Leaves with glands in the axils of the basal pair of lateral veins; styles 2, long and slender, undivided; stamens 8; capsules 2-locular ............... .................................................................................................. 6. Alchornea Leaves with embedded foliar glands; styles 3, partially connate, free and recurved distally, undivided; stamens 4–20 or more; capsules 3-locular .............................................................. 3. Actinostemon (A. schomburgkii) Raised glands present on the petiole at or below the junction of the leaf blade ..................................................................................................... 50 Petioles without raised glands below the junction of the leaf blade (glands may be present at the junction with the leaf blade or on the blade itself) .............................................................................................................. 51 Plants glabrous, with white latex; flower groups sessile on the rachis; staminate flowers with 2 stamens, pistillate flowers with simple, unbranched stigmas ...................................................................... 53. Sapium Plants with an indumentum, lacking latex; flower groups on short (1–3 mm long), lateral peduncles; staminate flowers with 3 stamens, pistillate flowers with bifid stigmas ............................... 57. Tetrorchidium Pistillate flowers and fruits in fascicles, pedicellate and solitary in the leaf axils, or flowers in a colored bracteate pseudanthium or reduced to mere stamens and pistils and grouped in a cyathium ....................... 52 Pistillate flowers and fruits in multiflowered (often bisexual) racemes, spikes, panicles, or thyrses (neither in cyathia or pseudanthia nor in fascicles, sometimes solitary but then either pedicellate or sessile on the internodes) ..................................................................................... 64 Fruits drupaceous; dioecious shrubs or trees; leaves coriaceous and usually with an asymmetrical bases; staminate flowers with a central disk; pistillate flowers with 1 or 2 carpels; stigmas sessile, dilated; flowers apetalous .................................................................................. 23. Drypetes Fruits capsular; monoecious or dioecious herbs to trees; leaf bases symmetrical or not; flowers with or without petals .................................. 53 Weedy herbs to subshrubs; flowers apetalous ........................................ 54 Shrubs or trees; flowers with or without petals ...................................... 56 Plants with abundant white latex; the staminate and pistillate flowers reduced to a single stamen and ovary, respectively, grouped together in a cup-shaped cyathium ......................................................... 24. Euphorbia Plants lacking latex or else latex very scanty; the staminate and pistillate flowers separate, with perianth and more parts per flower, axillary, spicate, on internodes, or in colorful bracteate pseudanthia .................. 55 Leaves lanceolate to ovate, pubescent; staminate flowers in spikes opposite the leaves; pistillate flowers borne on the stems between nodes;

78

E UPHORBIACEAE

55. 56(53).

56. 57(56). 57. 58(57).

58. 59(57).

59.

60(59).

60.

61(60).

61.

62(61). 62.

63(62).

63.

fruits with triangular spur-like projections ..................... 39. Microstachys Leaves variously shaped, usually glabrous; flowers in axillary clusters; fruits without any spur-like projections ........................... 44. Phyllanthus Succulent shrubs, the stems green and zigzag; flowers reduced to a mere stamen and ovary, both sexes appearing together in a red, asymmetrical, slipper-shaped cyathium ............................................. 42. Pedilanthus Plants not as above .................................................................................. 57 Staminate and pistillate flowers with petals; stamens 4 or 5 ................ 58 Flowers lacking petals; stamens 2–30 ..................................................... 59 Leaves acute to acuminate, without a prominent submarginal vein; filaments connate; styles dilated, ovary pubescent; fruits muricate or pubescent ................................................................................ 21. Discocarpus Leaves acuminate, with a prominent submarginal vein; filaments free; styles slender, ovary glabrous; fruits glabrous ............................ 54. Savia Leaf blades pellucid-punctate (with more than a single gland), usually obovate, with small domatia in vein axils; trunk often with long spines; pistillate flowers long-pedicellate, sepals reflexed at anthesis; staminate flowers sessile in axillary pincushions; stamens 6–30 ......... 4. Adelia Leaf blades not pellucid-punctate (sometimes with a single subapical gland in Phyllanthus), lacking domatia; trunk without spines; pistillate flowers sessile to pedicellate, sepals usually not reflexed at anthesis; staminate flowers usually not in obvious pincushions; stamens 2–15 ...................................................................................................... 60 Young leaves with leafy, asymmetrical stipules; stamens 8–15; ovary muricate, the styles nearly free, bifid, coarse-papillose; ovules 1 per locule; fruit spiny; seeds smooth, black, shiny, carunculate ...... 14. Chaetocarpus Leaves without leafy or asymmetrical stipules; stamens 2–6; ovary usually ± smooth, the styles not as above; ovules 2 per locule (and usually 2 seeds per locule as well); fruit generally smooth, not spiny; seeds not as above ................................................................................................ 61 Inflorescences terminal, usually bisexual, of 1–4 solitary, pedicellate, pistillate flowers at basal nodes, the staminate flowers densely aggregated in a strobiform mass at the end of the rachis, separated from the pistillate part by an elongated internode; leaves with embedded leaf glands; ovules 1 per locule ................................................... 36. Maprounea Inflorescences axillary, unisexual or bisexual, not as above; leaves eglandular or with either a single apical gland or 2 basal glands; ovules 2 per locule ........................................................................................... 62 Semideciduous dioecious trees; sepals and stamens 4; fruits 4- or 5-carpellate, the outer seed coat fleshy and metallic-blue ...........37. Margaritaria Monoecious or occasionally dioecious evergreen shrubs or small trees; sepals 5 or 6, stamens 2–6; fruits 3-carpellate, the seeds dry or, if fleshy, not metallic blue .................................................................................. 63 Leaves narrowly elliptic, narrowly acute at both ends, subsessile, with small basal laminar glands; flowers subsessile; stamens 5, free; seed coat fleshy ............................................................................ 32. Jablonskia Leaves variously shaped, never with basal glands but sometimes with a single subapical large gland, or the leafy branchlets pseudocompound;

E UPHORBIACEAE 79

64(51). 64. 65(64).

65.

66(65).

66.

67(66).

67.

68(66).

68.

69(64). 69. 70(69). 70. 71(69). 71. 72(71).

flowers usually pedicellate; stamens 2–6, filaments free or basally connate; seed coat dry ............................................................. 44. Phyllanthus Leaves usually with conspicuously different-length petioles and/or with flexed or thickened, pulvinate petiole apices ...................................... 65 Leaves with ± uniform-length petioles; petiole apices not noticeably thickened, flexed, or pulvinate .................................................................... 69 Inflorescences axillary, racemes or dense panicles 2–5 cm long, flowers short-pedicellate; petals 5 in flowers of both sexes, with silky trichomes; stamens 5 in staminate flowers ......................... 48. Pogonophora Inflorescences terminal, or, if axillary, then > 5 cm long, either branched or with clearly pedicellate flowers; petals absent, or, if present, with numerous stamens in staminate flowers ............................................ 66 Inflorescences obviously branched, generally > 6 cm long; staminate flowers without petals; pistillate flowers with glabrous ovaries; stems with white latex; leaves usually glandular at the base .............................. 67 Inflorescences unbranched, raceme-like, generally < 6 cm long; staminate flowers with petals; pistillate flowers with pubescent ovaries; stems without latex; leaves without glands at the base ............................... 68 Leaves usually scattered along twigs, not in successive clusters; blades usually with two discrete round glands on the upper surface at the base; terminal stipules generally conspicuous and leaving a scar similar to Ficus species; stilt roots often present; latex copious .......... 38. Micrandra Leaves clustered at intervals along twigs; blades with two ± diffuse glands at the base on the lower surface; stipules not leaving a noticeable Ficus-like scar; stilt roots absent; latex scanty .................. 55. Senefeldera Pistillate flowers apetalous, with 5 free, narrowly lanceolate sepals that remain persistent in fruit; styles branches short, papillose; staminate flowers with a 5- or 6-parted, imbricate calyx, 5 free petals; leaves not abruptly cuspidate ..................................................................... 51. Sagotia Pistillate flowers with 5 minute petals; sepals either 5 and free or 3 and partly connate, not conspicuously persistent in fruit; style branches elongate, thick or slender; staminate flowers with a 2-laciniate calyx, valvate in bud, and 3 or 4 imbricate petals; leaves abruptly and sharply cuspidate ............................................................................. 52. Sandwithia Inflorescences all axillary ........................................................................ 70 Inflorescences all terminal or a combination of axillary and terminal positions ...................................................................................................... 71 Leaves elliptic to ovate, the base acute to rounded or cordate; fruit a dry to fleshy capsule < 5 mm diameter ........................................ 44. Phyllanthus Leaves elliptic-obovate, the base attenuate; fruit a fleshy capsule ca. 1–2 cm diameter ...................................................................... 49. Richeria Plants with copious white latex; leaves with or without a prominent pair of basal leaf glands .............................................................................. 72 Plants without latex or latex very scanty; leaves without a prominent pair of basal glands ...................................................................................... 75 Leaf blades lacking glands at or near the base; dendritic trichomes present; leaves often with marginal glands; staminate and pistillate flowers interspersed in a bottlebrush-like inflorescence; pistillate flow-

80

E UPHORBIACEAE

72.

73(72).

73.

74(73).

74.

75(71).

75.

76(75). 76. 77(76).

77.

78(77).

78.

ers more numerous, the calyx 6-lobed and not accrescent in fruit; styles connate into a conspicuous and often elongate column; fruits velvety pubescent or tomentose .............................................................. 34. Mabea Leaf blades with or without prominent glands at or near the base; inflorescences more openly branched than above; pistillate calyx < 6-lobed; fruits glabrous or pubescent ................................................................ 73 Terminal stipule usually enveloping the terminal bud, leaving a prominent leaf scar; flowers pedicellate, sepals 5, ± cup-shaped in pistillate flowers, stigmas sessile; stamens 5–10 in staminate flowers; stilt roots often present ......................................................................... 38. Micrandra Stipules small or lacking, not enveloping terminal bud nor leaving a prominent leaf scar; flowers sessile to shortly pedicellate, perianth 2- or 3-lobed in both sexes, inconspicuous; stamens 2–5 in staminate flowers; pistillate flowers with styles connate into a short column; stilt roots not present .................................................................................. 74 Leaves eglandular on upper surface; bract glands cup-shaped, sometimes multiple; inflorescences with dendritic trichomes; staminate flowers with 2 or 3 fused stamens; fruits < 8 mm diameter, dry outside ............................................................................................ 20. Dendrothrix Leaves usually with glands at the base on the upper surface; bract glands flattened-elliptic, in a single pair; inflorescences without dendritic trichomes; stamens free, 3–5 in staminate flowers; fruits > 10 mm diameter, fleshy outside ........................................................ 56. Senefelderopsis Inflorescences terminal, usually bisexual, of 1–4 solitary, pedicellate, pistillate flowers at basal nodes, the staminate flowers densely aggregated in a strobiliform mass at the end of the rachis, separated from the pistillate part by an elongated internode; leaves with petioles usually > 1/4 the length of the blade .......................................... 36. Maprounea Inflorescences axillary or terminal, bisexual or unisexual, without the aggregation of staminate flowers described above; petioles usually shorter in relation to the blade ........................................................... 76 Inflorescences spicate; flowers with small petals; stipules intrapetiolar ..................................................................................................... 8. Amanoa Inflorescences racemose or paniculate; flowers apetalous; stipules lateral .............................................................................................................. 77 Plants glabrous; leaves without glands; staminate inflorescences unisexual, racemose or paniculate; pistillate flowers solitary and longpedicellate, the calyx accrescent in fruit and 10–15 mm long; tepui summits at 1000–1500 m elevation ....................................... 13. Celianella Plants glabrous or with pubescence, especially in the young inflorescences; leaves with or without glands on the lower surface; inflorescences bisexual or unisexual; calyx not accrescent and much smaller than above; lowland areas < 500 m elevation ..................................... 78 Plants dioecious, without latex; leaves eglandular; pistillate flowers with sericeous ovary; columella persistent after fruit dehiscence ................. ................................................................................................ 47. Podocalyx Plants monoecious, with scanty latex or latex not noticeable; leaves glandular on the lower surface; pistillate flowers with ± glabrous ovary; columella persistent or not in fruit .......................................................... 79

Acalypha 81

79(78). Leaves glabrous and green on the underside; inflorescences with protective strobiliform (scale-like) bracts enclosing the buds; stamens 4–16 in staminate flowers ............................................................... 3. Actinostemon 79. Leaves whitish-farinose on the underside; inflorescences without protective scales in bud; stamens (2)3–6 in staminate flowers ......... 27. Gymnanthes 1. ACALYPHA L., Sp. Pl. 1003. 1753. by Geoffrey A. Levin Annual or perennial herbs, shrubs, or small trees, without latex, plants monoecious or less often dioecious. Leaves alternate, simple; margins usually toothed; stipules mostly small. Inflorescences axillary or terminal, spicate, racemose, or paniculate, unisexual, or if bisexual, then usually pistillate below and staminate above; bracts subtending the pistillate flowers usually enlarging following flowering. Staminate flowers very small, subsessile, several in the axil of a minute bract; calyx 4-lobed, petals and disk lacking; stamens mostly 8, the anthers pendent, elongated, and twisted at maturity; gynoecium none. Pistillate flowers 1–3 in the axil of a bract; calyx lobes 3–5, petals and disk lacking; stamens none; pistil with (1–)3 carpels, (1–) 3-locular; ovules 1 per locule; styles usually dissected into several to many threadlike segments; distal pistillate flower(s) sometimes differing from others (allomorphic), the ovary 1–3-lobed and the style subbasal. Fruit a capsule with (1–)3 locules, each segment splitting and falling from the persistent columella; fruit of allomorphic flowers a deeply lobed schizocarp or indehiscent. Seeds 1 per locule; testa usually carunculate, variously sculptured. Widespread in the Americas, Africa, and Asia, chiefly in tropical and subtropical regions; ca. 400 species, ca. 16 in Venezuela, 9 of these in the flora area. Acalypha amentacea Roxb. subsp. wilkesiana (Müll. Arg.) Fosberg is a variegated-leaf plant that is locally cultivated (Caicara). Acalypha hispida Burm. is another widely cultivated species, grown for its furry red inflorescences. Key to the Species of Acalypha 1.

1.

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

3.

Annual herbs, generally < 90 cm tall; pistillate inflorescences ovoid, with many tightly overlapping bracts, these hispid and with linear lobes > 1/2 the length of the bracts (linear in A. setosa) ................................. 2 Shrubs or small trees, generally > 1 m tall; pistillate inflorescences cylindrical to linear, or if ovoid then with few bracts, the bracts loosely overlapping if at all, not hispid, the bract lobes broader and no more than 1/4 the length of the bract ....................................................................... 4 Pistillate inflorescence laxly flowered, narrow, the rachis easily visible ........................................................................................................ A. setosa Pistillate inflorescence densely flowered, ellipsoid to cylindrical, the rachis not visible ........................................................................................ 3 Plants with gland-tipped trichomes on the stems or petioles in addition to the inflorescences of both sexes; pistillate inflorescences strictly terminal, on peduncles no more than 1 cm long; styles unbranched ........................................................................................... A. alopecuroides Plants lacking gland-tipped trichomes except on the pistillate inflorescences; pistillate inflorescences axillary, on peduncles 0.4–3 cm long;

82

4(1). 4.

5(4).

5.

6(5). 6.

7(4).

7.

8(7).

8.

E UPHORBIACEAE

styles with 3–5 branches ........................................................... A. arvensis Leaves with pinnate venation, the basal pair of secondary veins not much different from the other secondary veins in thickness or course ......... 5 Leaves with palmate or subpalmate venation, the basal pair of secondary veins prominent, extending to near or above the middle of the leaf blade ....................................................................................................... 7 Pistillate bracts solitary or few, at the base of bisexual inflorescences or in unisexual inflorescences < 2 cm long, each bract subtending 2 or 3 flowers; capsules < 3 mm diameter; petioles mostly < 2 cm long; leaves drying brownish .......................................................................... A. diversifolia Pistillate bracts numerous in unisexual inflorescences > 7 cm long, each bract subtending 1 flower; capsules 4–5 mm diameter; petioles mostly 3–8 cm long; leaves drying greenish ..................................................... 6 Leaf blades obovate, the bases subcuneate; pistillate inflorescences 7–15 cm long; capsules 4–5 mm diameter ................................. A. cuneata Leaf blades ovate or lanceolate, the bases rounded or subcordate; pistillate inflorescences 25–40 cm long, capsules ca. 2.5 mm diameter ................................................................................................... A. scandens Pistillate inflorescences racemose or sometimes paniculate, the flowers pedicellate; peduncles and rachises 0.2–0.3 mm thick; pistillate bracts not enlarging in fruit; lower surface of leaves with minute resinous dots ................................................................................................ A. villosa Pistillate inflorescences spicate, the flowers sessile; peduncles and rachises > 0.7 mm thick; pistillate bracts enlarging in fruit; lower surface of leaves without resinous dots ............................................................. 8 Pistillate inflorescences terminal, 5–10 cm long in fruit, ± congested; pistillate bracts with long stipitate glands; stipules with a filiform awn 7–10 mm long arising from a base 1 mm long and wide, not glandular ................................................................................................ A. schiedeana Pistillate inflorescences axillary, 15–35 cm long in fruit, ± lax; pistillate bracts with inconspicuous sessile glands; stipules ovate to ovate-lanceolate, often abruptly tapering to a long acuminate apex, the base > 2.5 mm long and wide, the margins glandular ............. A. macrostachya

Acalypha alopecuroides Jacq., Collectanea 3: 196. 1789 [1791]. Annual, weedy herb to 60(–90) cm tall. Disturbed places, 50–100 m; Bolívar (Moitaco), Amazonas (Puerto Ayacucho). Scattered in northern Venezuela; southern U.S.A. (introduced), Mexico to Panama, West Indies, Colombia, Trinidad. Acalypha arvensis Poepp., Nov. Gen. Sp. Pl. 3: 21. 1845 [1841]. Annual, weedy herb, sometimes becoming woody at base, 20–70 cm tall. Disturbed places, 50–100 m; Amazonas (Puerto Ayacucho). Scattered in Venezuela; Mexico, Central America, West Indies, Colombia, Trinidad, Tobago, Guyana, Suri-

name, French Guiana, Ecuador, Peru, Brazil, Bolivia. Acalypha cuneata Poepp., Nov. Gen. Sp. Pl. 3: 22. 1845 [1841]. Acalypha obovata Benth., Bot. Voy. Sulphur 163, t. 53. 1846. —Acalypha cuneata var. obovata (Benth.) Müll. Arg., Linnaea 34: 11. 1865. Shrub or small tree 2–5(–8) m tall; staminate inflorescences 5–15 cm long. Evergreen lowland forests, 100–300 m; Bolívar (upper Río Caura), Amazonas (Río Manapiare, Río Matacuni). Northern and central Venezuela; Panama, Colombia, Ecuador, Peru. Acalypha diversifolia Jacq., Pl. Hort. Schoenbr. 2: 63, t. 244. 1797.

Acalypha 83

Acalypha leptostachya H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 96. 1817. —Acalypha diversifolia var. leptostachya (H.B.K.) Müll. Arg. in A. DC., Prodr. 15(2): 854. 1866. Acalypha samydifolia Poepp., Nov. Gen. Sp. Pl. 3: 21. 1845 [1841]. Acalypha leptostachya var. carpinifolia Müll. Arg., Linnaea 34: 35. 1865. —Acalypha diversifolia var. carpinifolia (Müll. Arg.) Müll. Arg. in A. DC., Prodr. 15(2): 854. 1866. Shrub to small tree 1.5–5 m tall, branches arching; staminate and androgynous inflorescences 2–10 cm long. Primary and secondary evergreen nonflooded and riparian forests, 100–300 m; Bolívar (lower Río Caura), Amazonas (Raudal Arata on Río Ocamo). Widespread in northern Venezuela; Mexico to Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 80. As with several other shrubby Acalypha species, A. diversifolia has been divided into several varieties based on stem and leaf pubescence. These forms are found throughout the range of the species, and distinguishing them is neither easy nor useful. Acalypha macrostachya Jacq., Pl. Hort. Schoenbr. 2: 63, t. 245. 1797. Acalypha hirsutissima Willd., Sp. Pl. 4: 528. 1805. —Acalypha macrostachya var. hirsutissima (Willd.) Müll. Arg., Linnaea 34: 11. 1865. Acalypha macrophylla H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 96. 1817. —Acalypha macrostachya var. macrophylla (H.B.K.) Müll. Arg. in Mart., Fl. Bras. 11(2): 345. 1874. Acalypha tristis Poepp., Nov. Gen. Sp. Pl. 3: 22. 1845 [1841]. —Acalypha macrostachya var. tristis (Poepp.) Müll. Arg. in Mart., Fl. Bras. 11(2): 345. 1874. Small tree or large shrub 2–6 m tall, ± erect; leaves usually broadly ovate; staminate inflorescences 5–20 cm long. Semideciduous or evergreen lowland to montane forests, secondary forests, 50–1200 m; Delta Amacuro (Río Aguirre, Serranía de Imataca), Bolívar (widespread), Amazonas (Cerro Yutajé, headwaters of Río Orinoco). Widespread in northern Venezuela; Mexico to Panama, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ŠFig. 78.

As with Acalypha diversifolia, the several varieties, which are based on pubescence differences, are all widespread, scarcely distinct, and do not warrant recognition. Acalypha scandens Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 329. 1854. Shrub or weak tree 1–6 m tall, often scandent; leaves usually narrowly ovate to lanceolate; staminate inflorescences 5–20 cm long. Evergreen lowland forests, generally riparian, 50–300 m; Delta Amacuro (Río Acure, Río Cuyubini), Bolívar (5 km north of El Manteco, Río Botanamo, Río Cuyuní basin, Río Toro). Northeastern Venezuela; Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. This species resembles Acalypha macrostachya but is more delicate; both species show similar variation in pubescence. The leaves of A. scandens tend to be narrower, especially near the base, than those of A. macrostachya, and the stipules of A. scandens are narrower and taper evenly. Leaf venation is the most reliable difference between these species. Acalypha schiedeana Schltdl., Linnaea 7: 384. 1832. Acalypha schiedeana f. angustifolia Müll. Arg., Linnaea 34: 20. 1865. —Acalypha schiedeana var. angustifolia (Müll. Arg.) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 85): 55. 1924. Shrub 1–3 m tall; staminate inflorescences 3–8 cm long. Semideciduous lowland forests, 50–300 m; Bolívar (La Vergareña, lower Río Caroní, islands in Lago Guri, San Félix). Widespread in northern Venezuela; Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Colombia. ŠFig. 79. Acalypha setosa A. Rich., Hist. Fis. Cuba, Fanerogamia 11: 204. 1850. Weedy herb 0.5–0.9 m tall. Associated with cultivated areas or towns, 100–300 m; Amazonas (San Juan de Manapiare, Yutajé). Aragua, Distrito Federal, Portuguesa, Yaracuy; Mexico, West Indies, Colombia. Acalypha villosa Jacq., Enum. Syst. Pl. 32. 1760. Acalypha villosa var. intermedia Müll. Arg., Linnaea 34: 8. 1865.

84

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Fig. 78. Acalypha macrostachya

Acalypha villosa var. tomentosa Müll. Arg., Linnaea 34: 8. 1865. Acalypha villosa var. paniculata Müll. Arg. in A. DC., Prodr. 15(2): 802. 1866. Acalypha karsteniana Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 85): 19. 1924. Shrub to small tree 1–5 m tall; staminate inflorescences 3–12 cm long. Semideciduous to evergreen lowland, often secondary forests, 50–500 m; northwestern Bolívar, Amazonas (headwaters of Río Orinoco). Widespread in northern Venezuela; Mexico to Panama, West Indies (Grenada), Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 81. Acalypha villosa is a variable species, particularly in leaf shape and pubescence, hence the large number of varieties that have been described. These forms are widespread and indistinct, and do not warrant recognition. Though plants with racemose pistillate inflorescences are most common, individuals differing only in having paniculate inflorescences are found throughout the range of the species. Occasional plants bear both types of inflorescences.

Acalypha 85

Fig. 79. Acalypha schiedeana

Fig. 80. Acalypha diversifolia

Fig. 81. Acalypha villosa

86

E UPHORBIACEAE

2. ACIDOTON Sw., Prodr. 84. 1788. Gitara Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 85): 187. 1924. by Lynn J. Gillespie Dioecious shrubs or small trees. Leaves alternate, simple, stipulate, shortly petiolate, eglandular; blades chartaceous to coriaceous, pinnately veined; margins entire to coarsely serrate. Inflorescences axillary, fasciculate or racemose; bracts small, triangular. Staminate flowers pedicellate; sepals 3–5, valvate; petals and disk absent; stamens 20–60; filaments slender and free; anther connective with apical tuft of stinging trichomes; pistillode absent. Pistillate flowers pedicellate; sepals 5 or 6; petals and disk absent; ovary 3-locular, covered with stinging trichomes, locules each with a single ovule; styles partly connate, arms slender, undivided, spreading, with inner surface papillose. Fruit a 3-seeded, 3-lobed schizocarpous capsule, sparsely to moderately densely covered with stinging trichomes. Seeds subglobose, ecarunculate. Central America, Cuba, Jamaica, Hispaniola, Colombia, Venezuela, Ecuador, Peru, Amazonian Brazil; 6 species, 1 in Venezuela.

Fig. 82. Acidoton nicaraguensis

Actinostemon 87

Acidoton nicaraguensis (Hemsl.) G.L. Webster, Ann. Missouri Bot. Gard. 54: 191. 1967. —Cleidion? nicaraguense Hemsl., Biol. Cent.-Amer., Bot. 3: 130. 1883. Gitara venezolana Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 85): 187. 1924. —Acidoton venezolanus (Pax & K. Hoffm.) G.L. Webster, Ann. Missouri Bot. Gard. 54: 191. 1967.

Shrub to 3(–5) m tall; leaves oblanceolate, 10–20 × 3–8 cm, acuminate to caudate at apex, thinly chartaceous, margin moderately to coarsely serrate above the middle; inflorescence racemose, 1–6 cm long. Semideciduous to evergreen lowland forests, 100– 600 m; north-central and northeastern Bolívar. Carabobo, Miranda, Táchira, Yaracuy; Central America, Colombia, Ecuador, Peru, Amazonian Brazil. ŠFig. 82.

3. ACTINOSTEMON Klotzsch, Arch. Naturgesch. 7: 184. 1841. Dactylostemon Klotzsch, Arch. Naturgesch. 7: 181. 1841. by Paul E. Berry and Hans-Joachim Esser Monoecious shrubs or trees, sometimes with scanty white latex. Leaves alternate to verticillate, simple, with embedded foliar glands, pinnately veined or tripliveined at the base; margins entire; short-petiolate; stipules lateral. Inflorescences terminal or axillary, bisexual or unisexual, racemose, 1–3 per node, the buds enclosed by a tight series of imbricate, stipule-like or cone-like bud scales; bracts reduced or absent. Staminate flowers usually 2 or 3 per bract, each on a thin pedicel, minute, with 1–3 very small sepals; disk lacking; stamens (2–)4–16 or more; filaments free; anthers erect, ovoid. Pistillate flowers single or 2 or 3 and proximal on the rachis, borne on long pedicels, small, calyx absent or represented by 1–3 minute lobes; corolla, disk, and staminodes absent; ovary 3-locular, smooth or tuberculate; ovules 1 per locule; styles partially connate, free and recurved distally, undivided. Fruits capsular, columella persistent. Seeds subglobose, carunculate. Nicaragua, Costa Rica, West Indies, Colombia(?), Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Paraguay, Argentina, Uruguay; ca. 14 species, 3 in Venezuela, 2 of these in the flora area. Actinostemon is sometimes included within Gymnanthes, but can be differentiated by having 4–16 or more stamens (3–6 or rarely 2 in Gymnanthes) and embedded foliar glands and protective strobiliform bracts enclosing the young inflorescences (versus marginal laminar glands and unprotected young inflorescences in Gymnanthes). Key to the Species of Actinostemon 1.

1.

Leaves 9–25 × 4–8 cm, pinnately veined; staminate inflorescence > 2 cm long at anthesis, sepals of staminate flowers pubescent; bud scales densely pubescent; fruits smooth, pubescent ..................... A. amazonicus Leaves 3–8 × 1.5–3 cm, tripliveined at the base; staminate inflorescence < 2 cm long at anthesis, sepals of staminate flowers glabrous; bud scales glabrous; fruits muricate, glabrous or slightly pubescent ........................................................................................... A. schomburgkii

Actinostemon amazonicus Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 63. 1912. Shrub or tree 3–15 m tall; leaf blade narrowly elliptic, base attenuate; inflorescence

bud scales tan, striate on the outside, tomentose on the inside. Seasonally to barely flooded riparian vegetation and river banks, 50–200 m; Amazonas (30 km southeast of La Esmeralda, Río Orinoco at Isla Hormiga be-

88

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Fig. 83. Actinostemon schomburgkii

tween Caño Grulla and Caño Ucata, Río Orinoco near junction with Río Casiquiare, Santa Bárbara del Orinoco, Santa Rosa de Ucata, 10 km east of Tamatama). Colombia(?), Peru, Brazil. ŠFig. 84. Actinostemon schomburgkii (Klotzsch) Hochr., Bull. New York Bot. Gard. 6: 278. 1910. —Dactylostemon schomburgkii Klotzsch, London J. Bot. 2: 45. 1843. Actinostemon depauperatus Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 68) Addit. 6: 58. 1919. Actinostemon parvifolius Pittier, Bol. Soc. Venez. Ci. Nat. 5: 306. 1938. Shrub 0.5–4 m tall. Semideciduous to evergreen lowland to lower montane forests, rocky slopes, 50–400 m; northern and eastern Bolívar, northwestern Amazonas. Guyana, Suriname, French Guiana, Brazil, Bolivia. ŠFig. 83.

Fig. 84. Actinostemon amazonicus

Adelia 89

4. ADELIA L., Syst. Nat. ed. 10, 2: 1298. 1759. Ricinella Müll. Arg., Linnaea 34: 153. 1866. by Paul E. Berry Dioecious shrubs or small trees; branchlets often with spines derived from leafless short shoots. Leaves simple, alternate, petiolate; blades pinnately veined; margins entire, membranous to chartaceous, pellucid-punctate, with tufts of hairs (domatia) in the vein axils on lower surface. Inflorescences axillary, staminate flowers fasciculate on cushions in reduced short shoots, pistillate flowers paired in the axils and long-pedicellate. Staminate flowers with 4- or 5-parted, valvate calyx; petals absent; disk extrastaminal and annular, adnate to the calyx; stamens 6–30; filaments mostly free; anthers versatile, dorsifixed. Pistillate flowers with 5–7 narrow sepals, reflexed at anthesis, petals and staminodes absent, disk annular and pubescent; ovary 3-lobed and 3-locular, ovule 1 per locule; styles 3, free, laciniate. Fruits capsular, 3-lobed. Seeds carunculate or not. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Brazil, Bolivia(?), Paraguay; 10–12 species, 1 in Venezuela. Adelia triloba (Müll. Arg.) Hemsl., Biol. Cent.-Amer., Bot. 3: 130. 1883. —Ricinella triloba Müll. Arg., Linnaea 34: 153. 1866. Shrub or small tree, sometimes with spiny stems; leaf blades pellucid-punctate, with small domatia in vein axils; flowers

white, the staminate ones fasciculate. Riparian vegetation near semideciduous forests and savannas, 200–300 m; Amazonas (Río Ocamo near Raudal Arata). Widespread in dry areas of northern Venezuela; Nicaragua to Panama, Colombia, Ecuador. ŠFig. 85.

Fig. 85. Adelia triloba

90

E UPHORBIACEAE

5. ADENOPHAEDRA (Müll. Arg.) Müll. Arg. in Mart., Fl. Bras. 11(2): 385. 1874. —Bernardia sect. Adenophaedra Müll. Arg., Linnaea 34: 172. 1865. by Grady L. Webster Dioecious trees or shrubs. Leaves alternate, pinnately veined, eglandular, dentate; stipules deciduous. Flowers apetalous, in axillary or terminal spiciform thyrses, the staminate ones often compound; bracts eglandular. Staminate flowers pedicellate; sepals 3, valvate; disk absent; stamens 2 or 3; filaments free, short; anthers introrse, connective enlarged with a large gland; pistillode absent. Pistillate flowers pedicellate; sepals 6, imbricate, entire; disk 3-lobed; ovary 3-locular; styles stigmatiform; ovules 1 per locule. Fruit capsular, prominently 3-lobed. Seeds globose, 1 per locule; caruncle absent. Costa Rica, Panama, Colombia, Venezuela, Guyana, French Guiana, Brazil; 3 species, 1 in Venezuela.

Fig. 86. Adenophaedra grandifolia

Adenophaedra grandifolia (Klotzsch) Müll Arg. in Mart., Fl. Bras. 11(2): 386. 1874. —Tragia grandifolia Klotzsch, London J. Bot. 2: 46. 1843. —Maikanwakuyek (Arekuna), Warama-ana-chiyek (Arekuna). Shrub or small tree 2–8 m tall; leaf margins dentate and gland-tipped; petioles short and thick; flowers minute on unisexual, spiciform inflorescences. Montane forests along streams, 900–1300 m; Bolívar (Río Kapuy, Río Karaurín at Río Asadón, base of Roraima-tepui, base of Sororopán-tepui). Costa Rica, Panama, Colombia, Guyana, French Guiana, Brazil. ŠFig. 86.

Alchornea 91

6. ALCHORNEA Sw., Prodr. 98. 1788. by Grady L. Webster Dioecious trees or shrubs. Leaves alternate, pinnately or palmately veined, ± pubescent with minute stellate trichomes on lower surface, commonly with 2 or more flat laminar glands near the base, entire to dentate, sometimes with domatia in vein axils; stipules inconspicuous. Flowers apetalous, in simple or compound spiciform thyrses; bracts eglandular. Staminate calyx splitting into 2–5 valvate sepals; disk central; stamens usually 8; filaments free; anthers introrse; pistillode absent. Pistillate flowers sessile or short-pedicellate; sepals 4, imbricate; ovary 2-locular; styles 2(3 or 4), free, elongated; ovules 1 per locule. Fruit capsular, splitting into two 2-valved cocci; columella persistent. Seeds tuberculate, ecarunculate. Pantropics, mostly in the Neotropics but with secondary centers in Africa/ Madagascar and southeast Asia; ca. 80 species, 7 in Venezuela, 5 of these in the flora area. Key to the Species of Alchornea 1. 1. 2(1). 2. 3(1).

3.

4(3). 4.

Leaves pinnately veined ............................................................................ 2 Leaves palmately veined ........................................................................... 3 Leaves linear-lanceolate, > 5 times as long as wide, remotely and sharply serrate; pistillate spikes unbranched ................................. A. castaneifolia Leaves oblong-lanceolate, < 5 times as long as wide, bluntly dentate or crenate ......................................................................................... A. discolor Styles 3–7 mm long, thick; branches glabrous or subglabrous at tip; leaves coriaceous, rigid, glabrous on lower surface, except sparsely pubescent along veins, coarsely dentate; spikes axillary...................... A. grandiflora Styles 5–20 mm long, slender; branches closely pubescent at tips; leaves chartaceous to subcoriaceous, subentire to dentate (but not coarsely so); spikes axillary or borne on the trunk ............................................. 4 Leaves chartaceous, the lower surface with 5–10 glands; lateral veins ca. 5–8; spikes mostly axillary ................................................... A. glandulosa Leaves rigid, the lower surface with 2–4 glands; lateral veins fewer; spikes mostly cauliflorous .................................................... A. triplinervia

Alchornea castaneifolia (Willd.) A. Juss., Euphorb. Gen. 42. 1824. —Hermesia castaneifolia Willd., Sp. Pl. 4: 809. 1806. —Aliso blanco, Mangle. Shrub or small tree 3–5(–10) m tall. Seasonally flooded riparian forests, river banks, near sea level to 200 m; Delta Amacuro (Caño Mánamo), Bolívar (below La Urbana, Río Caura below Las Trincheras, along Río Orinoco), Amazonas (Isla Ratón, Puerto Ayacucho, along Río Orinoco near mouth of Río Parhueña). Anzoátegui, Apure, Cojedes, Guárico, Lara, Zulia; Colombia, Peru, Brazil, Bolivia, Paraguay. ŠFig. 90.

Alchornea discolor Poepp., Nov. Gen. Sp. 3: 19. 1841. —Reventillo. Alchornea schomburgkii Klotzsch, London J. Bot. 2: 46. 1843. Shrub or tree 2–20 m tall. Open areas, riparian forests, granitic outcrops, edges of Mauritia palm swamps, 50–600 m; Bolívar (6 km from Maniapure, Río Tonoro), Amazonas (Caño Yagua, 2 km west of La Esmeralda, Ocamo, surroundings of Puerto Ayacucho, Río Pasimoni, Salto Yureba, San Carlos de Río Negro, Santa Bárbara del Orinoco, Yutajé). Apure, Anzoátegui; Colombia, Guyana, Suriname, French Guiana, Ec-

92

E UPHORBIACEAE

uador, Peru, Brazil, Bolivia, Paraguay. ŠFig. 88. The form of this species with branched pistillate inflorescences has often been recognized as a distinct species, Alchornea schomburgkii.

Tree 5–10(–18) m tall. Evergreen lowland and swamp forests, near sea level to 100 m; Delta Amacuro (Caño Arature, Caño Cupurito in Caño Capure basin). Widespread in montane areas of much of northern Venezuela; Panama, Colombia, Ecuador, Peru.

Alchornea glandulosa Poepp., Nov. Gen. Sp. 3: 18, t. 221. 1841. —Algodoncillo. Tree 3–20 m tall; leaves ovate, chartaceous, with ca. 6 small glands embedded in the blade near the base; young stems stellate-pubescent. Lower montane to montane (occasionally lowland) forests, (100–)400– 1400 m; Bolívar (Cerro Venamo, northern Gran Sabana, Serranía de Maigualida), Amazonas (Cerro Yutajé, Salto Yureba off lower Río Ventuari). Barinas, Mérida, Portuguesa, Táchira, Trujillo, Zulia; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil (Amazonas), Bolivia. ŠFig. 87.

Alchornea triplinervia (Spreng.) Müll. Arg. in A. DC., Prodr. 15(2): 909. 1866. —Antidesma triplinervia Spreng., Neue Entd. 2: 116. 1821. —Ajumma (Yekwana), Tibure, Tiburi, Tiu. Tree 5–20 m tall. Evergreen lowland to montane forests, often in disturbed areas, sometimes on white sands or granitic outcrops, 50–900 m; Bolívar (El Paují, Río Tabaro in Río Nichare basin, Santa Elena de Uairén, Santa María de Erebato), Amazonas (Capibara, Río Casiquiare, Río Mawarinuma, Río Pasimoni, San Carlos de Río Negro to Solano). Aragua, Apure, Lara, Miranda, Trujillo, Zulia; Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 89.

Alchornea grandiflora Müll. Arg., Linnaea 34: 170. 1865. —Algodón, Cañoflote, Pata de paloma.

Fig. 87. Alchornea glandulosa

Alchornea 93

Fig. 88. Alchornea discolor

Fig. 89. Alchornea triplinervia

94

E UPHORBIACEAE

Fig. 90. Alchornea castaneifolia

7. ALCHORNEOPSIS Müll. Arg., Linnaea 34: 156. 1865. by Grady L. Webster Dioecious trees. Leaves alternate, simple, tripliveined, biglandular at base, nearly glabrous (veins with minute simple trichomes); stipules absent; margins entire or crenate. Flowers apetalous, in axillary spiciform thyrses; bracts eglandular. Staminate calyx splitting into 3 or 4 lobes; stamens 4–8, free, inserted on a massive pubescent disk; anthers with connective enlarged and glandular; pistillode 3-lobed. Pistillate flowers pedicellate; sepals 4 or 5, slightly imbricate; disk annular, pubescent; ovary 3-locular; styles free, short, unlobed, papillose; ovules 1 per locule. Fruit capsular; columella persistent, massive. Seeds flattened, ecarunculate, with fleshy coat; endosperm copious. Costa Rica, Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 3 closely related species that should probably be regarded as subspecies of a single species, 1 in Venezuela.

Amanoa 95

Alchorneopsis floribunda (Benth.) Müll. Arg., Linnaea 34: 156. 1865. —Alchornea glandulosa var. floribunda Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 331. 1854. Tree 10–35 m tall. Lower montane forests, 200–900 m; Bolívar (Icabarú), Amazonas (upper Río Cuao, base of Sierra de la Neblina). Mérida; Honduras, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 91.

Fig. 91. Alchorneopsis floribunda

8. AMANOA Aubl., Hist. Pl. Guiane 256, t. 101. 1775. by W. John Hayden Monoecious or dioecious trees or shrubs, latex absent. Leaves alternate, distichous, evergreen, simple, coriaceous, glabrous; stipules intrapetiolar, paired, or confluent across the leaf axil; margins entire; venation pinnate. Inflorescence axillary and/or terminal, of densely bracteate clusters (reduced cymules), in the axils of ordinary foliage leaves, in nonleafy pseudoterminal aggregates that revert to vegetative growth, or (in neotropical species) in the axils of alternate, reduced, crescentiform stipular bracts of determinate deciduous spiciform axes borne in groups of 1–several per branch apex; axes straight or sinuous; floral bracts minute, deltate, with abaxially pubescent midribs. Staminate flowers sessile or pedicellate, regular; perianth biseriate; sepals 5, free; petals 5, minute, scale-like, obovate or (in neotropical species) clawed with a reniform limb. Disk annular, extrastaminal. Stamens 5, receptacular or elevated on a short androphore; filaments free; anthers basifixed, bilocular; pistillode columnar. Pistillate flowers short-pedicellate, regular; perianth fugaceous; otherwise as in staminate flowers. Disk hypogynous, annular. Ovary 3-locular, globose; ovules 2 per locule; stigmas sessile, entire or lobed. Fruit an explosively dehiscent globose capsule, the bony endocarp twisting and separating from the remainder of the pericarp. Seeds globose to subcylindric; caruncle absent;

96

E UPHORBIACEAE

testa chartaceous to stony; endosperm present; embryo straight, the cotyledons large, oblong, the radicle short, superior. Widespread in eastern Central America, northern South America, and tropical west Africa; 15 species, 6 in Venezuela, all in the flora area. Key to the Species of Amanoa 1. 1. 2(1).

2.

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

4.

5(4).

5.

Leaves finely puncticulate abaxially with uniformly scattered cork warts ................................................................................................................ 2 Leaves without abaxial cork warts ............................................................ 3 Leaves spreading, margins flat; staminate flowers sessile, ca. 5 mm diameter; fruiting pedicels < 5 mm long; growing at elevations of 50–300 m .............................................................................................. A. oblongifolia Leaves stiffly erect, margins revolute; staminate flowers pedicellate, ca. 10 mm diameter; fruiting pedicels 8–15 mm long; growing at elevations of 1500–2100 m ................................................................... A. steyermarkii Shrubs mostly < 2 m tall; leaf apex round or retuse ................. A. cupatensis Trees or shrubs, > 2 m tall; leaf apex acute to acuminate ........................ 4 Inflorescence axis and pedicels of pistillate flowers densely hirtellous; flowers ca. 5 mm diameter; stamens elevated on androphore 1 mm long; fruits 9–13 mm long, on pedicels 2–3 mm long; pericarp ca. 1.5 mm thick; shrubs or trees to 12 m tall ................................................... A. almerindae Inflorescence axis and pedicels of pistillate flowers glabrous; flowers 7–10 mm diameter; stamens inserted on receptacle; fruits 20–40 mm long, on pedicles 3–10 mm long; pericarp 3–6 mm thick; trees to 35 m tall ................................................................................................................ 5 Leaves oblong, margins revolute; plants dioecious; staminate flowers pedicellate; fruits 4–5 cm diameter, with woody exocarp 5–7 mm thick ............................................................................................. A. glaucophylla Leaves elliptic, ovate, or obovate, margins flat or weakly revolute; plants monoecious; staminate flowers sessile; fruits smaller than above, with thinner fruit wall ................................................................... A. guianensis

Amanoa almerindae Leal, Arch. Jard. Bot. Rio de Janeiro 11: 68, pl. 8, 2, fig. 2. 1951. —Reventillo. Amanoa pubescens Steyerm., Fieldiana, Bot. 28: 304. 1952. Shrub or tree to 12 m tall. Periodically flooded savannas and borders of streams, 100–300 m; Amazonas (Río Atabapo, Río Atacavi, Río Casiquiare, Río Guainía, Río Sipapo, Río Temi). Brazil (Amazonas: Rio Negro). ŠFig. 95. Amanoa cupatensis Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 7: 296. 1913. Low shrub 0.5–2 m tall, flowers unknown. Savannas, said to be locally common but sel-

dom collected, 100–200 m; Amazonas (between La Esmeralda and Cerro Duida, Sabana Hechimoni near mouth of Río Siapa). Brazil (Amazonas). ŠFig. 94. Amanoa glaucophylla Müll. Arg. in Mart., Fl. Bras. 11(2): 11. 1873. Tree to 15 m tall. Low forests (bana) on white sand, river banks of black-water rivers, 50–200 m; Amazonas (Caño San Miguel, Río Sipapo). Colombia (Amazonas), Brazil (Amazonas, Goias, Mato Grosso). This is a rare species, known from few collections. Amanoa guianensis Aubl., Hist. Pl. Guiane 256, t. 101. 1775. —Guayabo rebalsero,

Amanoa 97

Fig. 92. Amanoa steyermarkii

Fig. 93. Amanoa oblongifolia

98

E UPHORBIACEAE

Fig. 94. Amanoa cupatensis

Fig. 96. Amanoa guianensis

Fig. 95. Amanoa almerindae

Aparisthmium 99

Hicaquillo, Icaco, Jubey de Arau, Suruauray, Temora-urai. Amanoa guianensis var. grandiflora Müll. Arg. in A. DC., Prodr. 15(2): 219. 1866. —Amanoa grandiflora (Müll. Arg.) Müll. Arg., Flora 55: 2. 1872. Tree to 35 m tall. Evergreen lowland forests, near sea level to 500 m (rarely higher); common throughout Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Guárico; widespread in lowlands of eastern Central America and northern South America. ŠFig. 96. Amanoa oblongifolia Müll. Arg., Linnaea 32: 77. 1863. —Reventillo de bocón.

Tree to 20 m tall. Periodically flooded forests and borders of streams, 100–300 m; southern Amazonas (Río Baría, Río Emoni off Río Casiquiare, Río Pasimoni, ca. 30 km south of San Carlos de Río Negro). Widespread in southeastern Colombia, eastern Peru, Brazil (Amazonas, Pará, Rondônia, Roraima), northwestern Bolivia. ŠFig. 93. Amanoa steyermarkii Jabl., Acta Bot. Venez. 2: 237. 1967. Tree to 20 m tall. Highland riparian and tepui forests, 1500–2100 m; Bolívar (Auyántepui, Cerro Jaua, Macizo del Chimantá [Amurí-tepui]), Amazonas (Cerro Yutajé, Cerro Marahuaka). Endemic. ŠFig. 92.

9. APARISTHMIUM Endl., Gen. Pl. 1112. 1840. by Grady L. Webster Dioecious trees or shrubs; stems without latex. Leaves alternate, ovate, pinnately veined but tripliveined, with paired laminar (stipel-like) glands near the base, remotely dentate, with sparse simple trichomes; stipules small, deciduous. Flowers apetalous, without disk, staminate flowers in panicles (compound thyrses), pistillate flowers in simple or compound racemes. Staminate calyx splitting into 2 or

Fig. 97. Aparisthmium cordatum

100

E UPHORBIACEAE

3 valvate sepals; stamens 3–5; filaments connate below; anther connective not enlarged; pistillode absent. Pistillate flowers pedicellate; sepals 4–6, entire; ovary 3locular; styles 3, nearly free, dilated, bilobed, papillose; ovules 1 per locule. Fruit capsular. Seeds 1 per locule, ecarunculate. Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 1 species. Aparisthmium cordatum (A. Juss.) Baill., Adansonia 5: 307. 1865. —Conceveiba cordatum A. Juss., Euphorb. Gen. 43, pl. 13, fig. 422. 1824. —Caruana, Hoja de danta, Mapagarro, Moradita, Onotillo, Reventillo, Tabaliz morado, Tibure, Momishi (Yekwana). Alchornea orinocensis Croizat, J. Arnold Arbor. 26: 191. 1945.

Shrub or small tree 3–10 m tall. Edges of forests, secondary vegetation, evergreen lowland to lower montane forests, 50–1000 m; widespread in Bolívar and Amazonas. Apure, Barinas, Carabobo, Mérida, Táchira, Zulia; Costa Rica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 97.

10. ASTROCOCCUS Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 327. 1854. by Lynn J. Gillespie Monoecious shrubs or small trees. Leaves alternate, simple, stipulate, very shortly petiolate, eglandular; blades oblanceolate to narrowly elliptic, acuminate at apex, acute at base, chartaceous, pinnately veined; margins serrulate. Inflorescences axillary, spicate, bisexual; lower 2–8 nodes with solitary pistillate flowers, upper nodes with highly condensed few-flowered staminate cymules, terminal flower pistillate; bracts linear-lanceolate, eglandular. Staminate flowers short-pedicellate; sepals 4, valvate; petals absent; disk 4-lobed, extrastaminal, each lobe oppo-

Fig. 98. Astrococcus cornutus

Bernardia 101

site and partly enclosing a stamen; stamens 4; pistillode absent. Pistillate flowers short-pedicellate; sepals 4 (or more in terminal flower); petals and disk absent; ovary 3-locular, distinctly 3-lobed with lobes horned, ovules 1 per locule; styles entirely connate into a massive ovoid-urceolate hollow-centered column. Fruit a 3seeded schizocarpous capsule; carpels 3, triangular, divergent, truncate and horned at apex with 2 large and several smaller horns, joined only at narrowed base. Seeds broadly turbinate, truncate and slightly convex at apex, ecarunculate. Endemic to the Guayana Shield from the upper Río Negro region of Venezuela and Brazil; 1 species. Astrococcus cornutus Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 327. 1854. Shrub or slender tree to 5 m tall; leaves to 17 × 5 cm; inflorescences 6–12 cm long; capsule 0.4 × 1.5 cm, horns reddish. Black-wa-

ter, seasonally flooded lowland forests, 100– 200 m; Amazonas (near Piedra Cocuy, San Carlos de Río Negro). Brazil (Amazonas: São Gabriel da Cachoeira). ŠFig. 98.

11. BERNARDIA Houst. ex Mill., Gard. Dict. ed. 4, 28. 1754. by Grady L. Webster Monoecious or dioecious shrubs; pubescence of simple or stellate trichomes. Leaves alternate, pinnately veined or tripliveined, often with basal laminar glands, ± dentate; stipules persistent. Flowers apetalous, in spiciform axillary or pseudoterminal thyrses; bracts eglandular. Staminate calyx splitting into 4 segments; disk interstaminal or absent; stamens 3–20, free; pistillode rudimentary or absent. Pistillate flowers sessile; sepals 4–6, imbricate, entire; disk annular or dissected; ovary 3locular; styles free, bifid or bipartite; ovules 1 per locule. Fruit capsular; columella persistent. Seeds rounded to prismatic, ecarunculate. American tropics and subtropics, from U.S.A. (California, Texas) south to Argentina and Uruguay; ca. 40 species, 7 in Venezuela, 1 of these in the flora area. Bernardia amazonica Croizat, J. Arnold Arbor. 24: 166. 1943. Shrub 1–2 m tall; leaves pubescent, glandular on lower surface. Semideciduous and lower montane forests, granitic outcrops, Mauritia palm swamps, 50–600 m; northwestern Bolívar (35 km southwest of Caicara, Cerro Borja, Cerro La Guacamaya east of Los Pijiguaos and 43 km south of Los Pijiguaos, along Río Orinoco at Cerro Gavilán near Piedra Marimare), Amazonas (along Río Orinoco). Endemic. ŠFig. 99. This species has been collected only in Venezuela so far, but undoubtedly is present in adjacent Colombia (Vichada).

Fig. 99. Bernardia amazonica

102

E UPHORBIACEAE

12. CAPERONIA A. St.-Hil., Hist. Pl. Remarq. Brésil 244. 1824. by Grady L. Webster Annual or perennial, monoecious or dioecious herbs or subshrubs. Leaves alternate, pinnately or palmately veined, with appressed simple trichomes and sometimes glandular trichomes, serrate; stipules persistent. Flowers in axillary racemiform or spiciform thyrses; bracts eglandular. Staminate flowers with articulate pedicels; sepals 5, valvate; petals 5, often unequal, adnate to staminal column; disk absent; stamens 10; filaments connate; anthers in 2 whorls; pistillode present. Pistillate flowers sessile or short-pedicellate; sepals 5, sometimes with supernumerary lobes; disk obsolete; ovary 3-locular, muricate; styles free, deeply 3–7-lobed or laciniate; ovules 1 per locule. Fruit capsular, 3-lobed, echinate to hispid or verrucose; columella persistent. Seeds spheroidal, minutely foveolate, ecarunculate. Southern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay, Argentina, Africa; ca. 35 species, 5 in Venezuela, 1 of these in the flora area. Caperonia castaneifolia (L.) A. St.-Hil., Hist. Pl. Remarq. Brésil 245. 1824. —Croton castaneifolius L., Sp. Pl. 1004. 1753.

Fig. 100. Caperonia castaneifolia

Annual herb 0.5–1 m tall. Swamps, lagoons, savannas, and other poorly drained areas, near sea level to 500 m; Delta Amacuro (Caño Mánamo, Capure, Tucupita),

Celianella 103

Bolívar (5 km north of Ciudad Piar, 16 km north of El Manteco, Kamarata. base of Serranía de los Pijiguaos), Amazonas (San Juan de Manapiare). Apure, Cojedes, Guá-

rico, Monagas, Portuguesa; widespread in tropical America from U.S.A. (Florida) south to Paraguay and Argentina. ŠFig. 100.

13. CELIANELLA Jabl., Mem. New York Bot. Gard. 12(2): 176. 1965. by Paul E. Berry Subshrubs or shrubs 0.5–5 m tall, apparently dioecious. Leaves subsucculent, alternate, entire, pinnately veined, obovate to elliptic, base long-cuneate, central vein prominent; stipules narrowly lanceolate, 1–2 cm long, leaving prominent scars when falling off. Staminate inflorescences racemose, flowers with 5 orbicular sepals; stamens 5; anthers 2-locular, introrse; central disk 5-lobed. Pistillate inflorescence subterminal, 1–3-flowered, or solitary at apex of rachis; flowers apetalous, sepals 10–13 mm long; ovary 3-locular; styles 3, basally connate, apically bifid; ovules 2 per locule. Fruit capsular, with accrescent sepals to 1.8–3 × 0.4–1.2 cm, reticulateveined. Seeds elongate-fusiform, with a small caruncle. Endemic to tepuis of southern Venezuela; 1 species. Celianella montana Jabl., Mem. New York Bot. Gard. 12(2): 178. 1965. Tepui thickets and meadows, 1000–1500

Staminate inflorescence

Fig. 101. Celianella montana

m; Bolívar (Cerro Sarisariñama), Amazonas (Cerro Duida, Cerro Parú, Cerro Yutajé). Endemic. ŠFig. 101.

104

E UPHORBIACEAE

14. CHAETOCARPUS Thwaites, Hooker’s J. Bot. Kew Gard. Misc. 6: 300. 1854. by Grady L. Webster and Paul E. Berry Dioecious shrubs or trees, glabrescent. Leaves alternate, pinnately veined, entire, coriaceous, without laminar or petiolar glands; stipules strongly asymmetric, sometimes leafy, caducous. Inflorescence dense axillary glomerules of apetalous flowers; bracts eglandular. Staminate sepals 4, imbricate; disk urceolate; stamens 8–15; filaments usually united into an androphore, hirsute; anthers introrse; pistillode 2- or 3-lobed. Pistillate flowers pedicellate; sepals 4 or 5, imbricate, entire; disk annular; ovary muricate, 3-locular; styles nearly free, bifid, coarsely papillose; ovules 1 per locule. Fruit capsular, spiny. Seeds smooth, black, shiny, covered in upper third by a thin aril. Greater Antilles, Venezuela, Guyana, Suriname, French Guiana, Brazil, western Africa, Madagascar, Sri Lanka and southeast Asia to Borneo; ca. 15 species, 1 in Venezuela.

Fig. 102. Chaetocarpus schomburgkianus

Chaemaesyce 105

Chaetocarpus schomburgkianus (Kuntze) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 10. 1912. —Gaedawakka schomburgkianus Kuntze, Revis. Gen. Pl. 2: 606. 1891. —Cacho, Cobija, Yaraya-yek (Arekuna). Drypetes spruceana Müll. Arg. in A. DC., Prodr. 15(2): 454. 1866. Chaetocarpus stipularis Gleason, Bull. Torrey Bot. Club 56: 397. 1929. Discocarpus mazarunensis Croizat, Bull. Torrey Bot. Club 75: 400. 1948. Drypetes maguireana Monach., Phytologia 3: 33. 1948. Chaetocarpus williamsii Steyerm., Fieldiana, Bot. 28: 306. 1952. Tree 6–30 m tall. Semideciduous to ev-

ergreen lowland and gallery forests, near sea level to 900 m; Delta Amacuro (Serranía de Imataca), Bolívar (El Manteco to San Pedro de Las Dos Bocas, widespread in the Gran Sabana, Serranía de Imataca, Urimán), Amazonas (between Maroa and Yavita, Río Casiquiare, Río Guainía in front of Maroa, 44 km southeast of Santa Bárbara del Orinoco). Monagas; Colombia, Guyana, Suriname, French Guiana. ŠFig. 102. Chaetocarpus williamsii from Maroa, Amazonas, is known only from the fruiting type collection and appears to differ only in its relatively larger fruits and seeds; it is here considered conspecific with C. schomburgkianus.

15. CHAMAESYCE Raf., Amer. Monthly Mag. & Crit. Rev. 2: 119. 1817. by Paul E. Berry Annual or perennial herbs or subshrubs, prostrate, ascending, or erect, with milky latex; stems often reddish. Leaves opposite, simple, short-petiolate or subsessile; stipules separate or interpetiolar, often lacerate and persisting; blades usually asymmetric at the base; margin entire to serrate, chlorophyll-bearing cells mostly in sheaths around the veins with colorless areas in between. Inflorescences terminal or axillary, in cymose clusters or glomerules of 1–many cyathia. Cyathia pseudanthial, the involucre resembling a calyx with 5 lobes alternating with 4(5) sessile glands, these simple or with petal-like appendages. Staminate flowers few to many, each consisting of a single, stipitate stamen; anther with 2 divergent thecae. Pistillate flowers terminal, solitary, consisting of a 3-locular ovary; ovules 1 per locule; styles 3, free or united at the base, distally bifid. Fruit a schizocarpous capsule, usually exserted from the cyathium by elongation of the stipe. Seeds ovoid, angled, or smooth, ecarunculate. Cosmopolitan, most species in Neotropics and subtropics; ca. 250 species, ca. 10 species in Venezuela, 6 of these in the flora area. Key to the Species of Chamaesyce 1. 1. 2(1).

2. 3(1).

Capsules glabrous ...................................................................................... 2 Capsules puberulent .................................................................................. 3 Cyathia in nearly leafless cymules; capsules 0.9–1.5 mm long, usually broadest at the middle; stipules conspicuous, usually 1–1.5 mm long ............................................................................................. C. hypericifolia Cyathia in leafy cymules; capsules 1.5–2 mm long, usually broadest below the middle; stipules inconspicuous, ca. 0.5 mm long .......... C. hyssopifolia Cyathia in leafless, capitate glomerules on short-pedunculate shoots branching only near the base; larger leaves 15–40 mm long; plants with yellow trichomes ............................................................................. C. hirta

106

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3.

4(3).

4.

5(4). 5.

Cyathia usually subtended by leaves (sometimes reduced), not in congested pedunculate glomerules; larger leaves 5–20 mm long; leaves glabrous or with colorless trichomes ..................................................... 4 Capsules usually exserted and readily visible on short peduncles, not closely subtended by cyathium and leaves; distal leaf axils and inflorescences not obscured by cotton-like trichomes .......................... C. prostrata Capsules sessile on the cyathium and closely subtended by leaves; distal leaf axils and base of inflorescences with cotton-like, whitish trichomes ................................................................................................................ 5 Capsules splitting the cyathial involucre at maturity; petaloid appendages small or obscure, equal in size ....................................... C. thymifolia Capsules closely subtended by the cyathia and not splitting them at maturity; petaloid appendages conspicuous and unequal ................ C. dioeca

Chamaesyce dioeca (H.B.K.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 384. 1914, “dioica.” —Euphorbia dioeca H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 53. 1817. Mat-forming, densely branched herb; internodes 2–16 mm long; leaf blades 3–15 × 1.5–6 mm. Beaches, open areas, 50–100 m; Bolívar (La Paragua, Puerto Ordaz, Río Orinoco at Piedra Marimare). Widespread in northern Venezuela; Mexico, Central America, Colombia, Guyana, Ecuador, Brazil.

Fig. 103. Chamaesyce hirta

Chamaesyce hirta (L.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 303. 1909. —Euphorbia hirta L., Sp. Pl. 454. 1753. Euphorbia globulifera H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 56. 1817. Decumbent or erect weedy herb with multicellular yellow trichomes; internodes 0.5–5 cm long; leaf blades 6–45 × 5–15 mm; cymes in pedunculate clusters. Open sites, near sea level to 900 m; Delta Amacuro (Isla Paloma in lower Río Orinoco), Bolívar (El Paují), Amazonas (Puerto Ayacucho). Scattered in

Fig. 104. Chamaesyce prostrata

Cnidoscolus 107

northern Venezuela; southern U.S.A. and West Indies to Argentina, also a widespread weed in the Old World tropics. ŠFig. 103. Chamaesyce hypericifolia (L.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 302. 1909. —Euphorbia hypericifolia L., Sp. Pl. 454. 1753. Prostrate or erect herb, usually fewbranched with internodes 1–7 cm long; leaf blades 8–35 × 3–15 mm. Open areas, 50–100 m; Bolívar (74 km southwest of Caicara). Scattered in northern half of Venezuela; southern U.S.A. and West Indies to Argentina. Chamaesyce hyssopifolia (L.) Small, Bull. New York Bot. Gard. 3: 429. 1905. —Euphorbia hyssopifolia L., Syst. Nat. ed. 10: 1048. 1759. Euphorbia brasiliensis Lam., Encycl. 2: 423. 1786 [1788]. Spreading or erect herb; internodes of main stems 1–3 cm long; leaf blades 6–30 × 3–15 mm. Open areas, near sea level to 500 m; Delta Amacuro (Pedernales, Tucupita to Los Guires), Bolívar (Ciudad Piar, El Pao), Amazonas (Puerto Ayacucho, San Juan de Manapiare). Scattered in northern half of Ven-

ezuela; southern U.S.A. and West Indies to Argentina, adventive in the Old World tropics. Chamaesyce prostrata (Aiton) Small, Fl. S.E. U.S. 713, 1333. 1903. —Euphorbia prostrata Aiton, Hort. Kew 2: 139. 1789. Euphorbia tenella H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 53. 1817. Prostrate herb; internodes 1–10 mm long; leaf blades 2–9 × 1–5 mm. Open sunny areas, river banks, grassy savannas, 50–100 m; Bolívar (savannas near Maripa), Amazonas (Río Orinoco near Maypures and Carichana). Scattered in northern Venezuela; southern U.S.A. and West Indies to Argentina, Old World tropics. ŠFig. 104. Chamaesyce thymifolia (L.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 413. 1915. —Euphorbia thymifolia L., Sp. Pl. 454. 1753. —Golondrina. Prostrate, mat-forming, weedy herb; internodes 2–12 mm long; leaf blades 3–10 × 1–5 mm. Sunny, open areas, river edges, 50– 500 m; widespread in northern Bolívar and Amazonas. Widespread elsewhere in Venezuela; Mexico and West Indies to Argentina, Old World tropics.

16. CNIDOSCOLUS Pohl, Pl. Bras. Icon. Descr. 1: 56. 1827. by Grady L. Webster Monoecious trees, shrubs, or herbs; stems and leaves with milky latex, armed with stinging hairs. Leaves alternate; stipules persistent, often dissected; petioles glandular at juncture with blade; blades palmately veined, usually palmately lobed. Flowers apetalous, in terminal dichasial cymes, pistillate flowers at lower nodes; bracts eglandular. Staminate calyx white, gamosepalous, lobes imbricate in bud; disk annular; stamens mostly 8–10; inner and sometimes outer filaments connate into a column. Pistillate flowers pedicellate; calyx yellowish, greenish, or purplish, the sepals usually deciduous; disk annular; ovary 3-locular; styles free, usually several times bifid or laciniate; ovules 1 per locule. Fruit capsular; columella persistent. Seeds carunculate; endosperm copious. Southern U.S.A. to Argentina, most species in arid regions of Mexico and Brazil; ca. 60 species, 2 in Venezuela, 1 of these in the flora area. Cnidoscolus urens (L.) Arthur, Torreya 21: 11. 1921. —Jatropha urens L., Sp. Pl. 1007. 1753. —Guaritoto. Perennial herb, sometimes somewhat woody at base, 0.5–2 m tall; trichomes stinging. Deciduous forests, disturbed or rocky areas, near sea level to 500 m; Delta Amacuro

(between Caño Guire and La Horqueta), northern Bolívar, Amazonas (Puerto Ayacucho). Scattered throughout most of northern Venezuela; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 105.

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E UPHORBIACEAE

Fig. 105. Cnidoscolus urens

17. CONCEVEIBA Aubl., Hist. Pl. Guiane 923. 1775. Conceveibastrum (Müll. Arg.) Pax & K. Hoffm., Pflanzenr. IV. 147. (Heft 63): 217. 1914. —Alchornea section Conceveibastrum Müll. Arg., in Mart., Fl. Bras. 11(2): 375. 1874. by Grady L. Webster Dioecious trees. Leaves alternate, long-petiolate, pinnately or palmately veined, often biglandular and stipellate at base, crenate-denticulate, the lower surface ± pubescent with stellate and sometimes also simple trichomes; stipules deciduous or persistent. Flowers apetalous, without disk, in terminal simple or compound thyrses; bracts biglandular. Staminate calyx splitting into 3 or 4 segments; stamens 10–50, free, the inner stamens sometimes infertile; anther connective sometimes enlarged; pistillode absent. Pistillate flowers pedicellate; sepals 5–8, imbricate, glandular at base; ovary 3-locular; styles free or nearly so, apically bilobed to deeply bipartite; ovules 1 per locule. Fruit capsular, walls massive. Seeds smooth, carunculate; endosperm copious. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Africa; 12 species, 6 in Venezuela, 4 of these in the flora area. This genus is closely allied to Gavarretia, and there may be intermediate taxa between them (see discussion under Gavarretia terminalis).

Conceveiba 109

Key to the Species of Conceveiba 1.

1. 2(1).

2.

3(2). 3.

Stipules deciduous; capsule trigonous, glabrate (minutely stellate); central group of stamens sterile, long-plicate, outer circle shorter and fertile ........................................................................................... C. guianensis Stipules foliaceous, persistent; capsule ellipsoid, densely pubescent; stamens as above or else all fertile ............................................................ 2 Leaves ovate, deeply cordate at base; lateral veins raised, scalariform; pistillate calyx glandular at base; stamens differentiated into a central sterile group and a short, fertile outer group ......................... C. martiana Leaves mostly elliptic, rounded or shallowly cordate at the base; lateral veins not as prominent as above; pistillate calyx not glandular at base; stamens all fertile .................................................................................. 3 Leaves densely tomentose on lower surface ................................ C. ptariana Leaves sparsely stellate-pubescent on lower surface .................... C. latifolia

Conceveiba guianensis Aubl., Hist. Pl. Guiane 924. 1775. —Caniba, Caruana, Cocura, Nicolás, Palo de agua dulce, Palo de mata, Poatoru.

Conceveiba hostmannii Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 332. 1854. Tree 3–15 m tall. Primary or disturbed evergreen forests, 50–1000 m; Delta Amacuro

Fig. 106. Conceveiba guianensis

110

E UPHORBIACEAE

Fig. 107. Conceveiba ptariana

Croton

111

(Serranía de Imataca), Bolívar (Canaima, near Las Claritas, southwest of Maripa, Río Caura, upper Río Cuyuní, Río Nichare), Amazonas (western base of Cerro Duida, Río Casiquiare, Río Pasimoni, San Carlos de Río Negro, Sierra de la Neblina, Sierra Parima, Yavita to Maroa). Monagas. Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 106.

Pflanzenr. IV. 147(Heft 63): 217. 1914. —Palo de mataguaro. Tree 10–25 m tall. Evergreen lowland to lower montane forests, usually on poorly drained white sand, 50–600(–1200) m; Amazonas (Cerro Yapacana, Maroa to Yavita, Río Yaciba, San Carlos de Río Negro, Sierra de la Neblina). Mérida; Colombia (Vaupés), French Guiana, Ecuador, Peru, Brazil (Amazonas).

Conceveiba latifolia Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 332. 1854. —Veconcibea latifolia (Benth.) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 63): 218. 1914. Tree to 15 m tall. Basimontane forests, 100–200 m; Amazonas (Sierra de la Neblina). Brazil.

Conceveiba ptariana (Steyerm.) Jabl., Mem. New York Bot. Gard. 17(1): 134. 1967. —Conceveibastrum ptarianum Steyerm., Fieldiana, Bot. 28: 308. 1952. —Anonti-wa-yek (Arekuna), Warimbay-yek (Arekuna). Tree 8–30 m tall. Montane forests mostly on tepui slopes or summits, 700–1700 m; Bolívar (Aprada-tepui, Cerro Marutaní, Macizo del Chimantá, Ptari-tepui, Sierra de Lema), Amazonas (upper Río Ventuari). Lara, Táchira. ŠFig. 107.

Conceveiba martiana Baill., Adansonia 5: 221. 1865. —Conceveibastrum martianum (Baill.) Pax & K. Hoffm. in Engl.,

18. CROTON L., Sp. Pl. 1004. 1753. Julocroton Mart., Flora 20: 119. 1837. by Paul E. Berry Monoecious or less often dioecious herbs, shrubs, trees, or rarely lianas; indument commonly of stellate to lepidote trichomes; watery or colored latex often present, sometimes aromatic. Leaves alternate or sometimes opposite at congested nodes, simple to deeply lobed, margins entire to variously serrate or dentate, sessile to stalked glands often present at or near the junction of the blade and the petiole, additional disk-shaped glands sometimes present on the lower leaf surface; venation palmate or pinnate. Inflorescences axillary to terminal, bisexual or less often unisexual, spicate or racemose, most often with the pistillate flowers basal and staminate flowers distal, or else pistillate and staminate flowers combined, the staminate flowers often in sessile cymules. Staminate flowers with 4–6 valvate or imbricate calyx lobes; petals usually (0)5(4, 6), receptacle usually pilose; stamens 8–50; filaments inflexed in bud and free; pistillode absent. Pistillate flowers with (4)5–7(–10) imbricate or valvate calyx lobes; petals 0 or 5 and then small; staminodes absent; disk entire or lobed; ovary with (1–)3(4) locules; ovules 1 per locule; styles (2)3, bifid or several times bifid. Fruit capsular, with (2)3 2-valved cocci, columella usually persisting. Seeds oblong, carunculate. Cosmopolitan, mostly tropical and warm-temperate regions; ca. 1200 species, ca. 75 in Venezuela, 45 of these in the flora area. Croton punctatus Jacq. grows commonly along the Venezuelan coast on sandy beaches, and may eventually be found in Delta Amacuro. It is a small, somewhat prostrate shrub with oblong leaves, entire margins, and a rounded apex; it has short-stalked trichomes with a flat rounded top and a fringe of minute thin rays.

112

E UPHORBIACEAE

Key to the Species of Croton 1.

Trees (generally 3–35 m tall with a single main stem); inflorescences mostly elongate (15–30 cm long, but 3–10 cm long in C. schiedeanus) ................................................................................................................ 2 1. Herbs or shrubs (generally 0.2–3.0 m tall and branched close to the base); inflorescences mostly 0.5–15 cm long, but to 30 cm long in some shrubby species .................................................................................... 18 2(1). Indumentum of lepidote (peltate) scales, or sometimes a mixture of lepidote and stellate trichomes ................................................................... 3 2. Indumentum of stellate trichomes .......................................................... 12 3(2). Leaves membranous, eglandular at the base of the blade; pistillate flowers with petals, pedicels 1–2 cm long in fruit; staminate flower buds tiny (1.5–2.5 mm diameter) ................................................. C. schiedeanus 3. Leaves coriaceous or semicoriaceous, with a pair of glands at the base of the blade or apex of the petiole (rarely absent or not easily visible); pistillate flowers without petals, pedicel length various; staminate flower buds larger than above .......................................................................... 4 4(3). Leaves glabrous on upper surface ............................................................. 5 4. Leaves with scattered lepidote scales on upper surface ........................... 7 5(4). Leaves with few, scattered trichomes on lower surface; secondary veins > 1 cm apart and arching towards the apex near the margin ............... ........................................................................................ C. megalodendron 5. Leaves densely and completely covered with indument on lower surface; secondary veins closely parallel (< 1 cm apart) and at close to right angle with midvein ................................................................................ 6 6(5). Leaves silvery-lepidote on lower surface; leaves with a pair of sessile glands at the base of the leaf on the lower surface; lobes of the pistillate calyx not broad or carinate; the staminate buds oblongoid; racemes mostly tightly grouped together at branch tips .................. C. matourensis 6. Leaves brown-lepidote on lower surface; leaves mostly without visible glands at the base of the leaf on the lower surface (although the same plant can have stalked glands on some leaves); pistillate calyx with broad, carinate lobes, persistent in fruit; staminate buds globose; racemes laxly scattered in upper leaf axils or branched stems .... C. sp. D 7(4). Highland tepui plants (1000–2300 m elevation) ....................................... 8 7. Lowland plants (100–300 m elevation) ..................................................... 9 8(7). Leaves clustered at branch apices, margins serrulate; petioles 2–3 cm long; fruits 2–4 cm diameter, densely rusty-yellow lepidote; eastern tepuis > 1500 m .................................................................... C. roraimensis 8. Leaves more spread out on stems, margins entire; petioles 6–10 cm long; fruits unknown; Sierra de la Neblina, ca. 1100 m .................... C. neblinae 9(7). Leaves broadly ovate, palmately or pinnately veined, rounded to usually subcordate at the base; trichomes intermediate between stellate and lepidote ................................................................................................. 10 9. Leaves elliptic, pinnately veined, cuneate to rounded at the base; trichomes entirely lepidote ...................................................................... 11 10(9). Leaves palmately veined at the base, with 4–8 secondary veins per side; leaf margin broadly serrate with saucer-shaped glands in the sinuses; lower surface scattered-appressed pubescent; pedicels 12–20 mm long in fruit, the columella persistent, but the calyx caducous ............. C. sp. B

Croton

10.

11(9). 11. 12(2). 12.

13(12).

13.

14(13).

14. 15(12).

15.

16(15). 16.

17(16).

17.

18(1). 18. 19(18). 19. 20(19). 20.

113

Leaves pinnately veined, with 14–18 secondary veins per side; leaf margin entire, eglandular; lower surface densely pubescent with stalked trichomes; pedicels 4–6 mm long in fruit, the columella and calyx both persistent .........................................................................................C. sp. C Leaves cuneate at the base ........................................................... C. cuneatus Leaves rounded at the base .................................................... C. subcoriaceus Leaves suborbicular or broadly ovate, less than twice as long as wide, palmately veined, entire or markedly 3- or 5-lobed ................................ 13 Leaves elliptic, more than twice as long as wide, pinnately veined (or intermediate between palmately and pinnately veined in C. fragrans), without glands scattered on the lower surface, or if present, these only along the margin .................................................................................. 15 Leaves usually unlobed, if lobed, then the lobes rounded, the leaf apex acute to rounded; lower (and sometimes upper) surface with stalked saucer-shaped glands loosely scattered at vein junctures or along the margin; styles highly divided ........................................... C. palanostigma Leaves lobed, the lobes pointed, lower surface with or without round glands at vein junctures; styles bifid or multifid, but not highly divided .............................................................................................................. 14 Leaves densely scurfy-stellate, especially so when young; basal leaf glands generally hidden by pubescence; lower surface of leaf lacking round glands ..................................................................... C. gossypiifolius Leaves sparsely stellate, basal leaf glands readily visible, lower surface of leaf with scattered round glands .............................................. C. nuntians Leaves with the margin entire, densely pubescent on lower surface, with silky-wooly indument on leaves of young plants or shade leaves (becoming densely lepidote on mature trees or upper leaves), secondary veins closely parallel ............................................................ C. matourensis Leaves with a finely serrulate or serrulate-undulate margin, either nearly glabrous on lower surface or else with dense, stellate trichomes .............................................................................................................. 16 Leaves nearly glabrous, with only scattered trichomes on youngest growth, the base cuneate, without glands along the margin ................. C. icabarui Young leaves with dense, stellate indumentum on lower surface, becoming scattered with age, the base subcordate or rounded, with either saucer-shaped glands or small glandular teeth along the margin .... 17 Leaves finely serrulate with small glandular teeth along the margin; junction of the leaf blade and petiole with several pairs of glands, at least the distalmost pair stalked; stipules laciniate-glandular; calyx lobes carinate (concave), to 10 mm long in young fruit; fruits < 2 cm diameter ........................................................................................ C. fragrans Leaves with saucer-shaped glands along the leaf margin; base of blade on lower surface with a single pair of sessile saucer-shaped glands; stipules minute, not as above; calyx lobes flat, much smaller than above; fruits orangish, ca. 2 cm diameter .............................. C. yavitensis Leaves palmately veined at the base, or else palmately 3- or 5-lobed ... 19 Leaves primarily pinnately veined, unlobed ........................................... 28 Leaves deeply 3- or 5-lobed; herbs ................................................... C. lobatus Leaves not deeply lobed; herbs or shrubs ............................................... 20 Leaves entire-margined ........................................................................... 21 Leaves with toothed and/or lobed margins ............................................. 22

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E UPHORBIACEAE

21(20). Lower surface of leaves stellate-pubescent; blades and petioles mostly larger than below ................................................................... C. vergarenae 21. Lower surface of leaves silvery-lepidote; blades ca. 6 × 4 cm, petiole 2–3 cm long ............................................................................................ C. sp. H 22(20). Leaves ovate, the margin very finely glandular-denticulate, with loosely scattered stellate trichomes on both surfaces, blades 5–8 × 2–4 cm; stipules and calyx of pistillate flowers glandular-laciniate .................... .......................................................................................... C. essequiboensis 22. Leaves narrowly to broadly ovate, the margin coarsely dentate or serrate, with scattered to dense stellate trichomes on both surfaces, blades 3–14 × 2–8 cm; stipules entire, pistillate calyx entire or laciniate (C. argenteus) ............................................................................................. 23 23(22). Leaves broadly ovate, 8–14 × 4–11 cm, rounded to usually broadly cordate at the base, densely soft-pubescent on both surfaces; petioles 2–8 cm long ................................................................................................... C. sp. A 23. Leaves ovate to narrowly ovate or triangular, 3–10 × 2–5 cm, the base cuneate, rounded, oblique, or narrowly cordate, sparsely to densely pubescent on one or both surfaces; petioles 0.5–5 cm long .................... 24 24(23). Base of leaves without glands; inflorescences 1–2.5 cm long; pistillate calyx laciniate .............................................................................. C. argenteus 24. Base of leaves with a pair of sessile or stalked glands; inflorescences 1–8 cm long; pistillate calyx entire ............................................................ 25 25(24). Base of leaves narrowly cuneate, with an evident pair of stalked cupshaped glands at the apex of the petiole; staminate and pistillate flowers separated by a sterile portion of the raceme 2–4 cm long ................ .......................................................................................... C. sipaliwinensis 25. Base of leaves rounded, oblique, or subcordate, with either sessile or stalked glands at the top of the petiole; staminate flowers separated from pistillate flowers by a shorter sterile portion or not at all ........ 26 26(25). Shrubs 1–6 m tall; base of leaves subcordate, with a pair of sessile cupshaped glands; leaf margins irregularly serrate ................. C. nepetifolius 26. Weedy herbs or subshrubs 0.3–1.5 m tall; base of leaves oblique to rounded, with a pair of stalked glands; leaf margins regularly serrate or crenate ............................................................................................. 27 27(26). Leaf blades broadly ovate, margins mostly crenate; stems hispid, with trichomes to 2–3.5 mm long; inflorescences with gland-tipped trichomes ........................................................................................................ C. hirtus 27. Leaf blades triangular in outline, margins regularly serrate; stems shorter-pubescent, not hispid; inflorescences lacking gland-tipped trichomes ....................................................................................... C. trinitatis 28(18). Leaves strongly bicolored, the lower surface covered with a flat, silverywhite, dense covering of lepidote scales or stellate-lepidote trichomes .............................................................................................................. 29 28. Leaves concolorous or bicolored, the lower surface variously pubescent with stellate or lepidote trichomes, but without a flat white-silvery appearance ............................................................................................... 34 29(28). Leaves narrowly elliptic-ovate to lanceolate, the apex long-acuminate, the base without glands on either surface; plants always along rivers ... 30 29. Leaves elliptic to ovate, the apex acute, rounded, or acuminate, the base with a pair of glands on the lower surface; plants in a variety of habitats, but not necessarily restricted to riversides ................................ 31

Croton

115

30(29). Leaf bases smooth, without auriculate projections; stipules linear; indumentum lepidote ....................................................... C. argyrophyllus 30. Leaf bases with irregular, auriculate projections; stipules auriculate; indumentum stellate-lepidote .............................................. C. potaroensis 31(29). Leaves elliptic, stiff-coriaceous, lustrous-waxy on upper surface, the apex rounded ...................................................................................... C. scutatus 31. Leaves ovate, broadly ovate, or narrowly elliptic, membranous, not lustrous-waxy on upper surface, the apex acute to acuminate ............... 32 32(31). Leaf blades 18–28 × 6–12 cm, stems and veins on lower leaf surface with golden-brown porrect trichomes with main arm to ca. 2 mm long; secondary leaf veins ca. 20 per side .................................................... C. sp. G 32. Leaf blades 3–14 × 1.5–9 cm; stems and veins on lower leaf surface without the kind of trichomes described above; secondary leaf veins fewer per side than above .............................................................................. 33 33(32). Leaf blades 3–12 × 1.5–4 cm, subcordate at the base; sepals ca. 5 mm long in fruit ................................................................................... C. orinocensis 33. Leaf blades 7–14 × 3.5–9 cm, rounded at the base; sepals ca. 15 mm long in fruit ................................................................................... C. spruceanus 34(28). Leaves elliptic, stiff-coriaceous, margins entire and revolute, apex rounded to retuse, glabrous and hard-waxy on upper surface; known only from summit of Cerro Guaiquinima ......................................... C. guaiquinimae 34. Leaves variously shaped, but not stiff-coriaceous, apex variously acute to acuminate, occasionally rounded (but not retuse); margins subentire to dentate, crenate, or serrate, not revolute ........................................... 35 35(34). Leaves elliptic-ovate, 14–18 × –8 cm; petiole 5–7 cm long; inflorescence to 30 cm long ...................................................................................... C. pullei 35. Leaves variously shaped, smaller than above, the petiole shorter; inflorescence usually shorter ........................................................................... 36 36(35). Leaf blades lacking sessile or stalked discoid glands at or near the base on the lower surface or on the upper petiole ........................................... 37 36. Leaf blades with a pair of discoid to ellipsoid glands (stalked or sessile) at or near the base on the lower surface or uppermost part of the petiole (this usually requires careful observation with a hand lens or dissecting scope) .............................................................................................. 41 37(36). Leaf margin with glandular crenations; blades narrowly ovate, 8–14 × 3–4 cm, often folded along the margin (conduplicate), sparsely stellate on both surfaces, the lower surface of the leaf easily visible between the trichomes, the base subcordate ............................................. C. guianensis 37. Leaf margin entire to dentate or serrate, sometimes with discoid glands in the sinuses, but never glandular-crenate; blades variously shaped, smaller than above, usually plane, densely soft-stellate with the trichomes covering at least the entire lower surface, the base cuneate, rounded, or subcordate ........................................................................ 38 38(37). Leaves elliptic, subentire to serrulate, 3–5 × 1–2 cm; inflorescence 2– 5 cm long; leaf glands usually present, just hidden by the pubescence ........................................................................................................ C. mollis 38. Leaves ovate, denticulate, larger than above and usually with longer inflorescences; leaf glands entirely lacking ............................................ 39 39(38). Leaf base cuneate to rounded; stipules narrowly triangular or lanceolate, inconspicuous; inflorescences 5–15(–30) cm long, the columella persistent after fruit dehiscence, smoothy 3-parted at the apex (resem-

116

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39. 40(39). 40. 41(36).

41. 42(41).

42.

43(41). 43. 44(43).

44.

45(43).

45.

46(45).

46.

bling a woody, 3-parted stigma) ....................................... C. conduplicatus Leaf base subcordate; stipules auriculate and conspicuous or small and inconspicuous; inflorescences 4–10 cm long; fruits not known .......... 40 Stipules auriculate (broad and wrapping partly around the stem), conspicuous; leaf margin denticulate ....................................... C. bolivarensis Stipules not auriculate, inconspicuous; leaf margin entire ...... C. umbratilis Glands at base of leaf or near apex of the petiole discoid and stipitate (stalked); stalked discoid glands also present in sinuses of serrations along the margin .................................................................................. 42 Glands at or near the base of the leaf on near the petiole apex discoid to ellipsoid and sessile; leaf margin without stalked glands .................. 44 Leaf margins sharply serrate; blade ovate, apex acute; racemes with a single pistillate flower at the base; tepui summits and upper slopes ............................................................................................... C. subserratus Leaf margins serrulate to subentire; blade elliptic, apex rounded to acute; racemes with several pistillate flowers near the base; lowland whitesand shrublands .................................................................. C. spiraeifolius Mature leaves 2–5 × 1–2 cm .................................................................... 44 Mature leaves larger than above ............................................................. 45 Leaves elliptic, margins faintly and shallowly serrulate, basal glands often hidden by pubescence; indumentum never brownish on the branchlets; lowland white-sand areas .......................................... C. mollis Leaves ovate-elliptic, margins sharply serrulate, basal glands readily visible; indumentum usually brownish on the branchlets; upland areas in the Gran Sabana ............................................................... C. subincanus Leaf margins serrulate; lower surface of leaves and branchlets densely covered by stellate-lepidote trichomes, with scattered, darker brown trichomes overlaying them; secondary veins 12–14 per side, closely subparallel; inflorescence 5–12 cm long ..................................... C. sp. F Leaf margins subentire to denticulate; lower surface of leaf and branchlets without darker, brown trichomes, the lower surface leaf only sparsely pubescent; secondary veins fewer than above; inflorescence 8– 20 cm long ............................................................................................ 46 Lower surface of leaves with a pair of flat elliptic glands on the sides of the raised midvein ca. 2–4 mm above the base of the blade; sepals almost entirely connate at anthesis ............................................. C. cajucara Lower surface of leaves with round glands at the base of the blade; sepals narrowly oblong, free for almost the entire length ......................... C. sp. E

Croton argenteus L., Sp. Pl. 1004. 1753. —Julocroton argenteus (L.) Didr., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1857: 134. 1857. Subshrub 0.5–1 m tall; leaves ovate, serrate, the lower surface shortly stellate-pubescent and dull silvery, base cuneate and eglandular, with sharply ascending, palmate venation; leaves grouped in clusters separated by stem sections ca. 10–20 cm long; inflorescence terminal, 2–3 cm long; pistillate sepals laciniate. Open, disturbed, seasonally flooded areas, ca. 50 m; Bolívar (Angosturita

east of Ciudad Bolívar, Hato El Torno near Moitaco). Apure, Cojedes, Zulia; U.S.A. (Texas) to Chile and Argentina. Croton argyrophyllus H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 68. 1817. —Carcanapire rebalsero, Manteco. Croton nervosus Klotzsch, London J. Bot. 2: 50. 1843, nom. illegit. Shrub 1–6 m tall, with watery sap; leaves with > 18 closely subparallel secondary veins per side, eglandular at base, aromatic when crushed; stipules linear. In open rocky situa-

Croton

tions along rivers, often in seasonally flooded areas, 50–300 m; Bolívar (El Tigre, Puerto Ordaz, Río Caura on island 10 km above La Prisión, Río Caura at Raudal La Mura and near Salto Pará, San Félix), Amazonas (Río Orinoco). Apure, Falcón, Lara, Miranda, Nueva Esparta, Sucre, Táchira, Zulia; Colombia, Guyana, Brazil. Liesner & Holst 21598 (MO, NY, PORT), from montane forest openings at 1500–1700 m elevation on Cerro Yutajé, Amazonas state, resembles this species, but has somewhat broader leaves and curious long-stalked, appressed, cup-shaped glands near the base of the leaf on the lower side. Further material may prove it to be an undescribed species. Croton bolivarensis Croizat, J. Arnold Arbor. 21: 89. 1940. Medium to large shrub 1–4 m tall; no noticeable exudate from freshly cut stems and only a mild smell to crushed leaves; stipules conspicuous, auriculate and clasping the stem; branches smooth, brown, and lenticellate. In thickets of seasonally inundated flood plains and river banks, 0–100 m; Delta Amacuro (Santa Catalina), Bolívar (flood plains along middle and lower Río Orinoco, near Ciudad Bolívar). Apure. Croton cajucara Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 376. 1854. —Carurú, Echepeñ (Panare). Shrub or tree 1–6 m tall; leaves narrowly elliptic, apex narrowly acuminate, base cuneate, with sparse stellate trichomes on lower surface and elliptic glands on the raised midvein on the lower surface 2–4 mm above the base, trichomes on upper surface mostly on midvein only; racemes slender, 8– 15 cm long; sepals almost fully connate at anthesis. Riparian forests, deciduous forests on granitic outcrops, 50–400 m; Bolívar (Corozal between Maniapure and Caicara), Amazonas (Isla Ratón, 23 km north of Puerto Ayacucho near Cachama, Río Cataniapo, Río Sipapo, Salto Yureba). Barinas, Zulia; Brazil (Pará, Maranhão, Mato Grosso). Croton conduplicatus H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 80. 1817. —Carcanapire. Well-branched shrub 0.5–2 m tall, with watery sap when cut; leaves narrowly to broadly ovate, noticeably aromatic, apex acuminate, base rounded to cuneate, margin

117

finely denticulate, stellate-pubescent (more densely so and often whiter on lower surface, sometimes pubescence golden-brown), lacking glands at the base near the petiole; inflorescence to 15(–30) cm long; pistillate flowers after fruit dehisces leave a very characteristic persistent columella that is 3-parted at the apex giving the appearance of a 3parted style (but elongate bifid styles are deciduous). Common in secondary scrub, roadsides, shrub savannas, nonflooded areas, near sea level to 400 m; widespread in northernmost Bolívar. Anzoátegui, Aragua, Dependencias Federales, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Sucre, Yaracuy, Zulia; Colombia, Trinidad, Guyana, Brazil, Bolivia. Some collections of this species have previously been identified as Croton rhamnifolius H.B.K., C. fragilis H.B.K., or C. flavens L. The first two names are based on Humboldt and Bonpland collections from coastal Venezuela near Cumaná, as is the type of C. conduplicatus. The microfiche of the Paris collection of this species clearly shows the characteristic ovate, fairly longpetiolate leaves and the persistent columella of the dehisced fruits on long racemes that are typical of the material treated here from the flora area. The leaves sometimes become folded, or conduplicate, during dry periods, and senescing leaves turn a characteristic light orange color. Croton cuneatus Klotzsch in Benth., London J. Bot. 2: 49. 1843. —Arapurima, Cazabe, Mangle de río, Manteco de agua, Reventillo. Croton surinamensis Müll. Arg., Linnaea 34: 82. 1865. Croton monachinoensis Jabl., Mem. New York Bot. Gard. 12: 157. 1965. Shrub or tree 1.5–12 m tall, with reddish sap; leaves oblong to narrowly lanceolate, wedge-shaped at the base, subentire to remotely serrulate, the upper surface with scattered lepidote scales, the lower surface densely to completely covered with lepidote scales, 1 or 2 pairs of round glands at petiole apex; racemes elongate, terminal, of (sub)sessile flowers. Evergreen lowland to lower montane and seasonally flooded riparian forests, 100–700 m; widespread in Bolívar and Amazonas. Apure, Barinas; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 108.

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Croton essequiboensis Klotzsch in Benth., London J. Bot. 2: 49. 1843. —Croton populifolius var. essequiboensis (Klotzsch) Müll. Arg. in A. DC., Prodr. 15(2): 654. 1866. Shrub to 1.5–3 m tall; leaves 4–6 × 2–4 cm, ovate, with scattered stellate trichomes on both surfaces and small basal and marginal glands, margin finely serrulate, venation palmate; petioles 1–4 cm long; stipules and pistillate calyx deeply incised, with prominent stalked glands; racemes 4–6 cm long. Shrub islands in savannas, ca. 300 m; Bolívar (15 km southeast of El Manteco). Guyana, Brazil (Roraima). Croton fragrans H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 81. 1817. Slender tree 6–10 m tall; young leaves densely orange-pubescent, adult leaves broadly ovate, acuminate-caudate, 15–20 × 8– 13 cm, with 5–8 secondary veins per side and several pairs of glands at base, margins finely serrate; petiole 2–4 cm long; sepals carinate, 10 × 7 mm in early fruit, connate for basal third. Secondary vegetation in evergreen lowland and deciduous forests, 100–200 m; Bolívar (Guaniamo, island in Lago Guri 40 km south of dam, Mata Negra south of El Tigre, Reserva Forestal El Caura southeast of Campamento Forestal El Limón), Amazonas (3–4 m west of San Juan de Manapaire along road to Cerro Morrocoy). Scattered in northern Venezuela; Panama, Colombia. The flora area specimens are only doubtfully referred to Croton fragrans; the type of this species comes from central Colombia and was not examined for this treatment. The microfiche of the Paris type collection appears to have leaves with shorter petioles and rounder apices than the material examined from the Venezuelan Guayana. However, this material does agree with C. fragrans in having four sessile glands at the leaf base and in the hispid-pilose stellate pubescence. There is no indication of the habit of the original material of C. fragrans, whose flowers were described by Humboldt and Bonpland as having a citruslike odor when crushed. Croton gossypiifolius Vahl, Symb. Bot. 2: 98. 1791. —Sangre drago, Sangrito, Tarapaurai-yek (Arekuna). Tree 4–15 m tall, with spreading crown and reddish sap; leaves sharply 3- or 5-lobed, with dense (scurfy) stellate pubescence on lower surface, a pair of glands at junction of

petiole and blade but often ± hidden by trichomes, no additional glands evident on either surface of the leaf, young leaves lobed and densely orange-rust-pubescent; columella persistent. Forest edges, thickets, woody islands in savanna, semideciduous forests, 200–800 m; Bolívar (El Palmar, La Vigia 7 km west of El Manteco, Reserva Forestal La Paragua, Río Botanamo, Río Paragua, Santa Elena de Uairén, Serranía de Imataca). Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Trinidad. The sap of Croton gossypiifolius is used to seal skin wounds. Croton guianensis Aubl., Hist. Pl. Guiane 882, t. 339. 1775. Croton tafelbergicus Croizat, Bull. Torrey Bot. Club 75: 401. 1948. Shrub 1–3 m tall with erect racemes of white flowers to 16 cm long; leaves often conduplicate, blades narrowly ovate, margin glandular-crenulate, sparsely stellate on both surfaces, 8–14 × 3–4 cm, base subcordate; stipules dark-linear; petioles 1.5–4 cm long; flowers densely stellate, subtended by bracts with glandular-laciniate margins; calyx with laciniate margins (usually hidden by the pubescence). On exposed diabase dikes in moist, lower montane forests, 400– 600 m; Bolívar (above Piedra de la Virgen in lower section of La Escalera). Guyana, Suriname, French Guiana. Incipient basal leaf glands can be found on leaves of some plants; flora area material is treated under C. guianensis, which has well-developed basal (and sometimes stalked) glands. Croton guaiquinimae Steyerm., Brittonia 32: 19, fig. 2. 1990. Shrub to 2 m tall, with reddish sap; leaves coriaceous, lustrous on upper surface, densely stellate on lower surface; petiole with 2 glands below junction with leaf base; staminate inflorescences axillary, pistillate ones terminal. Meadows of Stegolepis squarrosus with woody patches and edges of gallery forests, 900–1100 m; Bolívar (Cerro Guaiquinima). Endemic. Croton hirtus L’Hér., Stirp. Nov. 17, t. 9. 1785. —Croton glandulosus var. hirtus (L’Hér.) Müll. Arg. in A. DC., Prodr.

Croton

119

Croton lobatus L., Sp. Pl. 1004. 1753. —Piñón montañero. Erect, annual, weedy herb; leaves deeply 3-lobed, with stellate pubescence or else trichomes reduced to one long ray and appearing simple. Disturbed areas, savannas, rocky outcrops, near sea level to 300 m; Delta Amacuro (Tucupita), Bolívar (Altiplanicie de Nuria, Ciudad Bolívar, La Camilera, mouth of Río Parguaza). Widespread in Venezuela north of Río Orinoco; U.S.A. (Southern Florida), Bahamas, Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa.

Croton pakaraimae Jabl., Mem. New York Bot. Gard. 12: 159. 1965. Tree 5–30 m tall, usually with reddish sap; leaves glabrous on upper surface, completely covered by pubescence on lower surface, biglandular at the base on the lower side of the midvein near junction with the petiole, margin entire; pubescence varies from silvery scales (the common condition on mature branches) to golden unbranched trichomes (on younger or lower foliage), even on same plant; secondary veins parallel, ca. 1 cm apart; stipules linear, usually persistent, 9–12 mm long; racemes terminal, elongate, 10–20 cm long. Evergreen lowland to lower montane forests, 100–1200 m; Bolívar (Arabapu, El Manteco, widespread in the Gran Sabana, near Kilómetro 88, 64 km southeast of Los Pijiguaos, Santa Elena de Uairén to Icabarú), Amazonas (Culebra, Platanal, Río Ocamo, upper Río Orinoco, 43 km northeast of Santa Bárbara, Tamatama). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 109. Croton matourensis is particularly variable in its indument. Young plants can be covered with a velvety, golden-brown indumentum of unbranched trichomes, and leaves of most adults have lower surfaces covered by a dense cover or silvery scale-like indumentum. Individual trees have been found with both kinds of trichomes and the full range of intermediate types; in these cases, the lower shade foliage is of the juvenile velvety type, and the more adult, exposed foliage is of the silvery-scale type. Various species have been described based largely on different indumentum types, but they are all just part of the variation present in C. matourensis. Croton pakaraimae was described as lacking a pair of glands at the base of the leaf blade, but examination of the type shows that it does indeed have glands there, just hidden by pubescence.

Croton matourensis Aubl., Hist. Pl. Guiane 879. 1775. —Araburi (Yanomami), Arapurihusi (Yanomami), Canelo, Canelón, Cusapoi-yek (Arekuna), Kusapui (Arekuna), Rabo de zamuro. Croton matourensis var. benthamianus Müll. Arg., Linnaea 34: 95. 1865. Croton kavanayensis Steyerm., Fieldiana, Bot. 28: 313. 1952. Croton lanjouwensis Jabl., Mem. New York Bot. Gard. 12: 158. 1965.

Croton megalodendron Müll. Arg., Flora 55: 4. 1872. —Canelón, Momisi. Croton xanthochloros Croizat, J. Arnold Arbor. 21: 94. 1940. Tree 8–28 m tall, with copious yellowish sap; inflorescences 15–30 cm long, bisexual, nodding. Semideciduous forests, 100–500 m; Bolívar (east of El Palmar, 38 km northeast of Los Rosos near Upata, Río Zariapo in Río Cuchivero basin). Widespread in Venezuelan Coastal Cordillera. ŠFig. 112.

15(2): 684. 1866.—Croton glandulosus subsp. hirtus (L’Hér.) Croizat, Bull. Torrey Bot. Club 75: 401. 1948. Erect annual herb 20–90 cm tall; stems hispid, with trichomes having a central ray 2–3.5 mm long; leaves crenate, unlobed, stellate-pubescent, with stipitate discoid glands at the base; inflorescences terminal, 1–3 cm long, with gland-tipped trichomes. Disturbed areas, savannas, near sea level to 200 m; Delta Amacuro (Caño de Uricoa to San Antonio, road between Tucupita and Los Guires), Bolívar (El Dorado, Maripa to Aripao, Túriba, 8 km southeast of Upata), Amazonas (Cerro San Borja along lower Río Suapure, Isla Ratón, Pintado, Puerto Ayacucho area). Widespread in northern Venezuela; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Argentina, adventive and naturalized in tropical Asia. Croton icabarui Jabl., Mem. New York Bot. Gard. 12: 158. 1965. Medium-sized tree, glabrous except for scattered stellate trichomes on very young leaves; leaves slightly crenate; petiole 3–4 cm long. Upland forests, 400–800 m; Bolívar (Río Icabarú). Endemic.

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Croton mollis Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 375. 1854. Croton crenatus Müll. Arg., Linnaea 34: 129. 1865. Shrub 0.5–1.5 m tall; leaves 3–5 × 1–2 cm, elliptic, acute, serrulate, with dense stellate indument on both surfaces; petiole 4–6 mm long; inflorescence terminal, 3–5 cm long. Soil pockets on edges of granitic outcrops, white-sand river banks, 50–100 m; Amazonas (Río Atabapo, Río Guasacavi, Río Sipapo). Brazil (Amazonas). Croton mollis appears to be closely related to C. spiraeifolius. The description of C. crenatus and the type photographs of the Berlin specimen agree with the treatment of C. mollis here. The type of C. crenatus is cited as coming from the banks of the Orinoco. Croton neblinae Jabl., Mem. New York Bot. Gard. 12: 155. 1965. Tree ca. 12 m tall; leaves 21–25 × 9–12 cm, broadly elliptic, scattered lepidote on both surfaces, biglandular at base, margins entire; petioles 6–10 cm long. Tepui slope forests, 1100–1200 m; Amazonas (Sierra de la Neblina). Endemic. ŠFig. 113. Croton nepetifolius Baill., Adansonia 4: 344. 1863, “nepetaefolius.” —Carcanapire. Shrub 1–6 m tall; leaves 5–9 × 3–5 cm, ovate, irregularly dentate, stellate pubescence denser on lower surface, pair of glands at leaf base on lower surface and sometimes along margin; petioles 1.5–3 cm long; stipules linear, 4–6 mm long; several terminal racemes 4–8 cm long. Disturbed areas along deciduous forests, weedy outskirts of towns, 50–300 m; Bolívar (Caicara, near Guri Dam). Monagas; Brazil (Amazonas, Minas Geraes, Pará). ŠFig. 115. Specimens from the flora area were formerly identified as Croton urticifolius Lam., also from Brazil, but they agree better with specimens identified as C. nepetifolius at MO. Croton nuntians Croizat, Bull. Torrey Bot. Club 75: 402. 1948. —Canelito, Canelo. Tree, sometimes scandent, 4–10 m tall; lower surface of leaves sparsely stellate, with scattered cup-shaped glands near vein junctions; pair of yellowish, readily visible glands at the junction of the blade and petiole; se-

pals persistent, spreading in fruit; seeds finely and regularly verrucose, columella persistent. Secondary vegetation along evergreen lowland forests, 200–400 m, Delta Amacuro (Río Toro, Serranía de Imataca). Guyana. Croton orinocensis Müll. Arg., Fl. Bras. 11(2): 135. 1873. Shrub 1–3 m tall; leaves narrowly 6–12 × 2–4 cm, elliptic-ovate, subcordate at the base, bicolored, densely silvery-white on lower surface with lepidote-stellate trichomes interspersed with scattered brown scales, pinnately veined, with prominent pair of glands at the base on the lower side; petiole 1–2 cm long; racemes 0.5–4(–8) cm long, the flowers usually congested at the tip, brownish-pubescent; pistillate flowers with sepals connate halfway at anthesis, persistent and ca. 5 × 3 mm in fruit; columella persistent. Granitic outcrops (lajas), 50–200 m; Amazonas (Cerro Tigrito and other lajas along road from Puerto Ayacucho to Samariapo, Raudal de Maipures, Tobogán de la Selva south of Coromoto). Adjacent Colombia. Specimens from Guyana have been incorrectly identified as Croton orinocensis (see L. Gillespie, Brittonia 45: 80. 1993). Croton palanostigma Klotzsch, London J. Bot. 2: 48. 1843. —Anontaya-yek (Arekuna), Canelón, Momi (Yekwana), Sangrito. Croton benthamianus Müll. Arg. in Mart., Fl. Bras. 11(2): 105. 1873. Tree 4–15 m tall; leaves large, ovate, the base shallowly cordate, unlobed or slightly to occasionally prominently 3-lobed (the lobes then generally rounded), the lower surface with stellate trichomes and 2 prominent raised circular glands at junction with petiole and numerous smaller glands near vein junctions toward the leaf margin, the latter sometimes also on the upper surface; inflorescence erect to nodding; stigmas highly divided. Secondary lowland to upland forests, riparian vegetation, savannas, 100–1200 m; Bolívar (Kavanayén, Río Kanarakuni, from Santa Elena de Uairén to Icabarú, Urimán, Wonkén), Amazonas (middle slopes of Cerro Marahuaka, Culebra, Río Cunucunuma, Río Mawarinuma, upper Río Orinoco, Río Sipapo, San Carlos de Río Negro, Solano, Yavita). Colombia, French Guiana, Ecuador, Peru, Brazil. ŠFig. 116.

Croton

Croton potaroensis Lanj., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 12: 457. 1934. —Melosa, Tiu-yek (Arekuna). Croton barlettii Lanj., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 12: 460. 1934. Croton cardonei Croizat, Bol. Soc. Venez. Ci. Nat. 11: 79. 1947. Rheophytic shrub 2–6 m tall; leaves narrowly ovate or lanceolate, with long-acuminate apex, the lower surface silvery-gray with appressed stellate-lepidote indumentum, base eglandular with irregular auriculate projections, stipules also auriculate. Among rocks along rivers, along rapids, below waterfalls, river beaches, 50–800 m; Bolívar (Río Apacará near junction with Río Caroní, Río Aparurén, upper Río Caroní at Raidal Kurukuya, Río Caura, Río Chiguao, Río Cuyuní, Río Paragua at Raudal de Guaiquinima, Río Ichún at Salto Ichún, Urimán), Amazonas (Río Cunucunuma near Culebra rapids, Río Siapa at Raudal Gallineta). Guyana. ŠFig. 110. Croton pullei Lanj., Euphorb. Surinam 18, t. 3. 1931. Scrambling shrub; leaves 14–18 × 7–8 cm, elliptic-ovate, apex caudate or long-acuminate, stellate-lepidote, quite scattered on upper surface, denser on lower surface but not covering leaf surface completely, margin serrulate with stalked cup-shaped glands in the sinuses, a pair of glands on the petiole at or 1–3 mm below the junction with the blade; petiole 3–7 cm long; inflorescences to 30 cm long. On gravelly islands, along river edges, 100–200 m; Amazonas (Río Mawarinuma). Brazil (Amazonas: Serra da Neblina), Suriname. Flora area material differs from the type material of Croton pullei in some characters, such as the more serrulate leaves and the petiolar glands, but otherwise shares the distinctive leaves and scrambling (sometimes described as vine-like), habit and riparian habitat. Croton roraimensis Croizat, Bull. Torrey Bot. Club 67: 290. 1940. Croton roraimensis var. subinteger Steyerm., Fieldiana, Bot. 28: 315. 1952. Tree 5–12 m tall; stems ± knobby from prominent leaf bases (leaves clustered at branch apices); leaves 10–15 × 4–7 cm, elliptic, serrate, with scattered lepidote scales on both surfaces, biglandular at the base; petioles stout, 2–3 cm long; stipules persistent,

121

linear-lanceolate; racemes terminal, ridged, stout; fruits large, 2–4 cm diameter, covered by scales. Tepui slopes, gallery forests, 1600– 2300 m; Bolívar (Auyán-tepui, Cerro Jaua, Macizo del Chimantá, Ptari-tepui, Roraimatepui). Guyana. ŠFig. 111. Croton schiedeanus Schlecht., Linnaea 19: 243, fig. 9E. 1847. —Canelo, Rastrojera. Small tree 5–8 m tall; leaves membranous, 10–15 × 4–6.5 cm, elliptic-oblong, pinnately veined, with dense, tiny, appressed silvery scales (reddish brown center and translucent or whitish periphery) on lower surface, scattered on upper surface, eglandular at the base; petioles 1–3 cm long; racemes axillary, 2–6 cm long; staminate flowers tiny, 1–2 mm diameter in bud, pistillate flower with petals 1.5–2 mm long, many style branches, and pedicels 1–2 cm long; fruits covered by peltate scales. Evergreen lowland forests, 100–300 m; Bolívar (between El Dorado and Kilómetro 88, Río Botanamo, San Martín de Turumbán southwest of Tumeremo). Carabobo, Distrito Federal, Falcón, Miranda, Yaracuy; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 114. Croton scutatus P.E. Berry & J.F. Gaskin, Syst. Bot. 23: 171. 1998. Sparsely branched shrub 2–5 m tall; leaves firmly coriaceous, elliptical, apex rounded to retuse, grayish and lustrous on upper surface, completely lepidote and brownish on lower surface; pistillate flower at tip of short inflorescence, sometimes with more basal staminate flowers on the same raceme. Locally gregarious on seasonally flooded lowland white-sand savannas and shrublands on sandstone on low tepuis, 100– 800 m; Amazonas (Caño San Miguel, Caño Cotúa, Cerro Cuao, base of Cerro Yapacana, Río Pasimoni). Colombia (Amazonas, Caquetá). ŠFig. 117. Croton sipaliwinensis Lanj., Recueil Trav. Bot. Néerl. 36: 698, fig. 1. 1939 [1940]. Small shrub 0.7–1 m tall; leaves tripliveined, stellate trichomes on both surfaces but denser on lower surface, irregularly serrate-margined, with a pair of conspicuously stalked glands at the petiole apex. Forest-savanna edges, shrublands on quartzite, 200–500 m; Bolívar (base of Uaipán-tepui),

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Amazonas (slopes of Cerro Mawedi in Río Ocamo basin, between hills north of Laguna Maguiri in southeastern slopes of upper Río Parucito valley). Guyana, Suriname. ŠFig. 119. Croton sipaliwinensis is part of a species complex with C. sclerocalyx (Diedrich) Müll. Arg. and C. lundianus Müll. Arg., both from central Brazil. Croton spiraeifolius Jabl., Mem. New York Bot. Gard. 12: 164. 1965. Shrub 0.5–2 m tall, with slight amount of whitish sap; leaves 4–7 × 1.5–3 cm, elliptic, apex rounded and mucronate to acute; margin entire to serrulate, with round stalked glands in the sinuses; petiole 3–6 mm long; stellate trichomes on both leaf surfaces and fruit; lateral stalked glands at distal end of petiole at leaf base; inflorescences terminal, 3–8 cm long; fruits basal, 4–8 mm diameter. White-sand savannas, 100–200 m; Amazonas (Caño Caname, Caño Cumare, Río Atabapo, Río Autana, Río Guayapo, Santa Cruz, Yapacana savannas). Colombia (Amazonas, Caquetá). Croton spiraeifolius is morphologically most similar to C. mollis. Croton spruceanus Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 375. 1854. Shrub 2–4 m tall; leaves 7–14 × 3.5–8 cm, ovate-elliptic, bicolored, whitish gray on lower surface, biglandular at the base, mostly glabrous on upper surface, densely pubescent on the lower surface with porrect-radiate trichomes (lepidote with separating, diverent arms and central erect branch); racemes 5–15 cm long; pistillate flowers with almost fully connate sepals at anthesis, calyx persistent, lobes splitting to ca. 15 × 5 mm in fruit. Evergreen lowland and premontane forests, 100– 300 m; Amazonas (Caño Negro on southeast side base of Cerro Duida, outskirts of Culebra). Brazil (Amazonas, Pará, Rondônia). Croton subcoriaceus Jabl., Mem. New York Bot. Gard. 12: 156. 1965. Tree 5–7 m tall; leaves with small lepidote scales on upper surface, densely lepidote on lower surface, margin slightly crenate. Riparian forests, 50–200 m; Amazonas (mouth of Cano Yapacana, Río Puruname). Endemic. ŠFig. 121.

Croton subcoriaceus is close to C. cuneatus, possibly even conspecific, but appears to differ in its more rounded leaf base and larger fruits and flowers. Croton subincanus Müll. Arg., Linnaea 34: 139. 1865. Subshrub 0.3–1.5 m tall, often browntinged on branchlets; leaves 2–5 × 1–2 cm, ovate to elliptic, densely covered by whitish, stellate trichomes, with 2 large, lateral round glands at the base of the blade, apex acute, margins serrulate; petiole 4–7 mm long; inflorescences subterminal, (3–)8–11 cm long, several (3–10) pistillate flowers along the lowermost 2–5 cm of the raceme. Upland savannas, rock outcrops along streams, 700–1200 m; Bolívar (Río Karaurín 2–3 km above confluence with Río Yuruaní, Río Kukenán basin between Kun and Uadara-parú, 30 km north-northeast of San Ignacio de Yuruaní, between San Rafael and San Ignacio de Yuruaní). Guyana, Suriname, Brazil (Roraima). ŠFig. 123. Croton subserratus Jabl., Mem. New York Bot. Gard. 12: 164. 1965. Shrub 0.2–1.5 m tall; leaves elliptic-ovate, strongly serrate with marginal stalked glands in teeth sinuses, stellate on both surfaces; petiole 4–6 mm long, with a pair of stalked glands near the apex; racemes terminal, with a single pistillate flower at the base. Tepui scrub, on both sandstone and granite, (1200–)1700–1800 m; Bolívar (Cerro Guanay), Amazonas (Cerro Cuao, Cerro Guanay). Endemic. ŠFig. 122. Croton trinitatis Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 57. 1900. —Conejita, Pata de paloma. Croton miquelensis Ferguson ex Lanj. in Pulle, Fl. Suriname 2: 28. 1932. Herb or subshrub 0.2–1.5 m tall, with stellate trichomes; leaves narrowly triangular with regularly serrate margins, with a pair of stalked glands at the base. Weedy areas, near sea level to 200 m; scattered throughout Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela and the Neotropics. ŠFig. 120. Croton umbratilis H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 82. 1817.

Croton

Shrub(?); leaves ovate-oblong, entire-margined, acuminate at the apex, slightly cordate at the base, with stellate trichomes, scabridous on upper surface, softly canescent-tomentose on lower surface, eglandular. Riparian forests, 50–100 m; Bolívar (along Río Orinoco near Carichana and Uruana). Endemic? Croton umbratilis is apparently only known from its type specimen and needs further collections to help understand its affinities to other species of Croton. Croton vergarenae (Jabl.) L.J. Gillespie, Brittonia 45: 83. 1993. —Julocroton vergarenae Jabl., Mem. New York Bot. Gard. 12: 170. 1965. —Carcanapire. Shrub 2–4 m tall; leaves 8–20 × 5–15 cm, (broadly) ovate, densely stellate on lower surface, less densely pubescent on upper surface, eglandular; petioles 2–12 cm long; stipules lanceolate, semipersistent; inflorescences short and very dense, 2–6 cm long; pistillate calyx laciniate. Semideciduous forests, secondary vegetation, 200–500 m; Bolívar (between La Paragua and El Cristo, La Vergareña, La Vigia 7 km west of El Manteco). Endemic. ŠFig. 125. Croton yavitensis Croizat, J. Arnold Arbor. 26: 189. 1945. Tree 5–8 m tall; leaves narrowly elliptic, base shallowly cordate or rounded, tip acute, margin faintly crenate with cup-shaped glands in sinuses, stellate-pubescent on lower surface and stems (not on upper surface); raceme 15–30 cm long; fruits light orangish brown-pubescent (with dense, stalked-stellate trichomes), 1.2–1.4 cm diameter. Seasonally flooded (black-water) lowland riparian forests, 100–200 m; Amazonas (Caño Ucata above San Juan northeast of San Fernando de Atabapo, Yavita). Endemic. ŠFig. 118. Croton sp. A Shrub 1–2 m tall; leaves broadly ovate, irregularly serrate-margined, stellate-pubescent, with persistent stipules; racemes compact (tightly and densely flowered). Shrub pockets on granitic outcrops, 50–100 m; Amazonas (Caño Ucata southeast of Síquita, Laja Los Oripopos 3–5 km north of Puerto Ayacucho). Endemic. ŠFig. 124.

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Croton sp. B Tree 4–16 m tall; leaves 15–25 × 8–14 cm, broadly ovate, rounded to subcordate at the base, biglandular at the top of the petiole, palmately veined at the base, only 4–6 secondary veins per side, margin entire to serrate with cup-shaped glands in the sinuses, with tiny lepidote-stellate trichomes on both surfaces; petioles 6–10 cm long; inflorescences to 35 cm long, pedicels 12–20 mm long, columella persistent; seeds smooth. Riparian forests, 100–200 m; Amazonas (35 km southeast of La Esmeralda, Río Mavaca, Río Mawarinuma, 44 km southeast of Santa Bárbara). The material included here appears most similar to Croton draconoides Müll. Arg., from Peru and Brazil, but differs from that species in its sparser pubescence, more cordate base, often serrate margin, and glands at the sinuses. Croton sp. C. —Majei (Yanomami). Tree 6–27 m tall; leaves whitish-pubescent on lower surface, biglandular at the base; inflorescences densely scurfy-pubescent; pistillate calyx broad, carinate, persistent in fruit; fruits squatly 3-lobed, densely orange-pubescent with stalked (porrect) trichomes. Riparian forests, 200–400 m; Amazonas (Caño Jayuwapuey in Río Ocamo basin, upper Río Orinoco at Raudal Montserrat [Los Tiestos]). Apparently endemic. This is most likely a new species, known from two collections, Croizat 565 (F, MO, US) and A. Fernández 6780 (MO, PORT). Croizat annotated his collection as “Croton boultonianus Croizat, sp. nov.,” but the name was never published. Croton sp. D. —Momishi (Yekwana). Tree 7–22 m tall, with thick opaque sap, densely brown-lepidote on lower surface of leaves and on stems and inflorescences; leaves usually eglandular; pistillate calyx lobes broad and carinate, persistent in fruit. Granitic outcrops along black-water rivers, streamside thickets on sandstone, 200–400 m; Amazonas (Cerro Mawedi in Río Ocamo basin, Río Cunucunuma at Raudal Wamujatado 4 km northeast of Cerro Huachamacari). Apparently endemic. This appears to be a new species most closely related to Croton matourensis.

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Fig. 108. Croton cuneatus

Croton 125

Fig. 109. Croton matourensis

Fig. 110. Croton potaroensis

Fig. 111. Croton roraimensis

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Fig. 112. Croton megalodendron

Fig. 113. Croton neblinae

Croton 127

Fig. 114. Croton schiedeanus

Fig. 116. Croton palanostigma

Fig. 115. Croton nepetifolius

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Fig. 117. Croton scutatus

Croton 129

Fig. 118. Croton yavitensis

Fig. 119. Croton sipaliwinensis

Fig. 120. Croton trinitatus

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Fig. 121. Croton subcoriaceus

Fig. 122. Croton subserratus

10 mm

Fig. 123. Croton subincanus

Croton 131

Fig. 124. Croton sp. A

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Fig. 125. Croton vergarenae

Croton sp. E Riparian shrub or small tree 2–8 m tall; leaves 5–12 × 2–5 cm, narrowly elliptic or elliptic-ovate, scattered stellate-pubescent on lower surface, margins denticulate with marginal glands on lower surface; stipules linear, dark, ca. 3 mm long; inflorescence 8–20 cm long; sepals linear-lanceolate, almost entirely free from each other. Rocky areas and beaches along rivers and waterfalls, 400–900 m; Bolívar (summit of Cerro Guaiquinima on tributary of Río Carapo, Río Caroní at Salto Erepuchi and above Raudales de Urkará). Endemic. Croton sp. F Shrub or small tree 1.5–6 m tall; leaves 6– 11 × 2.5–5 cm, elliptic-ovate, apex narrowly acute, base obtuse, densely stellate and with a pair of glands at the base of the blade on the lower surface, glabrescent on upper surface, 12–14 subparallel secondary veins per side; inflorescences terminal, 5–12 cm long.

On open granitic outcrops along river, 100– 300 m; Amazonas (Río Siapa 8 km below Raudal Gallineta). Endemic? This is another rheophytic Croton, but one that does not match any of the other ones in the flora area, such as C. argyrophyllus, C. potaroensis, C. pullei, or C. sp. E. Croton sp. G Shrub ca. 2.5 m tall with whitish sap; stems with large, yellow-brown stellate-porrect trichomes (the porrect arm to ca. 2 mm long); leaves elliptic to narrowly ovate, 18–28 × 6–12 cm, apex acute, base narrowly subcordate with a pair of sessile round glands on the lower surface, glabrous on upper surface, densely pubescent lower surface generally whitish with a dense covering of stellatelepidote trichomes scattered with longer yellow-brown trichomes with a long porrect central, erect arm ca. 2 mm long (especially on the veins), ca. 20 subparallel secondary veins per side, margin entire;

Dalechampia 133

stipules linear, pubescent; petioles 3–6 cm long; inflorescences subterminal, ca. 12 cm long, flower clusters spaced about 2 cm apart on the stem, a pedicellate female flower with a cluster of male flowers at the base of the pedicel; stigmas bifid, with whitish trichomes; slender columella persistent after fruit dehiscence, with three separate strands from the base to the coherent tips. River edges and adjacent forest, 200–300 m; Amazonas (upper Río Ocamo near Raudal Arata). Endemic. This species is known from one collection: Angel Fernández 6563 (MO, PORT) and is very likely an undescribed species for which

more material would be desireable. Croton sp. H Erect to clambering shrub 2–3 m tall; stems with sweet, aromatic smell when rubbed; leaves ovate, palmately veined, 6 × 3.5–4 cm, the lower surface densely silverylepidote, the upper surface glabrescent, rounded and eglandular at base; petioles 2–3 cm long; inflorescences 1–2 cm long, only seen in bud. Deciduous forests, 50–100 m; Bolívar (Puerto Ordaz). Better flowering material is needed to determine if this is a new or previously described species.

19. DALECHAMPIA L., Sp. Pl. 1054. 1753. by W. Scott Armbruster Perennial vines or shrubs, monoecious. Stems climbing, spreading, or erect, usually covered with urticating trichomes. Leaves alternate, petiolate, stipulate, with a pair of stipels at base of blade (stipels sometimes obscure or deciduous). Inflorescence pseudanthial, subtended by 2 usually large showy bracts, comprising a cyme of (1)3 pistillate flowers and a 3–5-branched cyme of 4–15 staminate flowers. Staminate cyme subtended by a cup-like involucellar bract, a bilabiate involucel, or 4(5) free bracts; staminate flowers with calyx splitting into 3–6 valvate segments at anthesis; petals absent; stamens 5–90, attached to an elongated column; filaments shorter than anthers; anthers in cluster at column apex, dehiscing longitudinally. Pistillate cyme subtended by 1 bract and 0–2 bractlets; pistillate flowers with 5–12 entire or lobed sepals; petals absent; ovary 3-carpellate, 1 ovule per carpel; styles connate into an elongated column, often dilated at apex. Fruit a 3-seeded capsule, dehiscing explosively by elastic twisting of dry, ± woody cocci. Seeds globose, surface smooth, roughened, or tuberculate, usually gray-brown mottled. Pantropics, primarily Neotropics, absent from Australia; ca. 120 species, 18 in Venezuela, 14 of these in the flora area. Key to the Species of Dalechampia 1.

1.

2(1).

2.

3(2).

Shrubs; leaves pinnately veined; involucral bracts pink (rarely yellowish) at anthesis; inflorescence glands yellow or greenish, the surface papillate, producing spicy fragrance; pistillate sepals 6 .......... D. magnoliifolia Vines; leaves palmately veined; involucral bracts white, green, or pink at anthesis; inflorescence glands, when present, variously colored, producing resin; pistillate sepals 6–12 ....................................................... 2 Leaves compound, 3-foliolate, rarely 4- or 5-foliolate; involucral bracts moderately conspicuous, spathulate, 5–8 mm long, green at anthesis; surface of stigmas papillate .............................................. D. papillistigma Leaves simple; involucral bracts conspicuous, ± ovate, > 1 cm long, white, pink, or green at anthesis, or inconspicuous and stipule-like, lanceolate, < 1 cm long, green at anthesis; surface of stigma smooth ...... 3 Leaves unlobed with 1–3 arcuate secondary veins attached to the midrib in distal half, tertiaries prominent, ladder-like; involucral bracts subequal or dimorphic; involucral bract subtending staminate flowers

134

3.

4(3).

4.

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

6.

7(6).

7.

8(3). 8. 9(8).

9.

E UPHORBIACEAE

inconspicuous, < 5 mm long, stipule-like, green at anthesis; involucral bract subtending pistillate flowers similar to staminate bract, or conspicuous, > 1 cm long, whitish at anthesis; inflorescence gland absent; pistillate sepals 6, unlobed .................................................................... 4 Leaves lobed or unlobed, arcuate secondary veins attached to midrib throughout its length, tertiaries usually less prominent; involucral bracts subequal, conspicuous, > 1 cm in length, petal- or leaf-like, white, pink, or green at anthesis; inflorescence gland present (vestigial and inconspicuous in D. brownsbergensis); pistillate sepals 6–12, lobed or unlobed ............................................................................................... 8 Base of leaves truncate to shallowly cordate; stipules ovate, deciduous prior to full expansion of leaves; fertile arms in staminate pleiochasium 4; staminate flowers 13, style swollen, clavate; pistillate sepals deciduous prior to fruit maturation ................................................. D. micrantha Base of leaves shallowly to deeply cordate (or hastate in immature plants); stipules lanceolate-ovate to lanceolate, deciduous only after leaves have expanded, or persistent; fertile arms in staminate pleiochasium 3 or 4; staminate flowers 10–13, style variously shaped; pistillate sepals persistent into fruit ..................................................... 5 Styles thin, evenly columnar throughout length, stigmatic region not swollen and club-shaped ............................................................. D. liesneri Styles swollen in stigmatic region, club-shaped or widest in middle and tapering toward tip ................................................................................ 6 Sinus at base of leaves usually deltoid, narrow at attachment of blade to petiole; involucral bracts subequal, < 5 mm long, stipule-like, green; style widest near middle, tapering to blunt tip, stigmatic surface green, not contrasting with style base .............................. D. attenuistylus Sinus at base of leaves usually broad and U-shaped; involucral bracts dimorphic or subequal, of various lengths, petal-like or stipule-like, white or green; style widest distal of middle, club-shaped, stigmatic surface white or pale green, contrasting with stylar base ................... 7 Involucral bracts usually dimorphic, the one subtending pistillate flowers > 1 cm long, whitish, the one subtending staminate flower < 5 mm long, stipule-like, green; pistillate sepals densely strigulose; stigmatic surface extending from tip to ca. 2/3 length of stylar column ....................... ........................................................................................... D. heterobractea Involucral bracts usually subequal, < 5 mm long, stipule-like, green (bract subtending the pistillate flowers sometimes 10 mm long, whitish); pistillate sepals glabrescent to minutely puberulent; stigmatic surface extending from tip to ca. 1/4–1/3 length of stylar column .... D. parvibracteata Leaves usually unlobed .............................................................................. 9 Leaves all 3-lobed or both unlobed and 3-lobed present on same plant (except very young plants which may have only unlobed leaves) .......... 12 Staminate pleiochasium with 4 free involucellar bracts; inflorescence gland comprising several concentrically arranged bractlets with laciniate margins, secreting greenish or brownish resin; involucral bracts linear and green or ovate and pink ..................................................... 10 Staminate pleiochasium with involucel of bracts fused for at least basal 1/3 or forming a bilabiate cup; inflorescence gland comprising usually

Dalechampia 135

2 columns of ± laminar bractlets with entire or minutely fimbriate margins, secreting clearish or yellowish resin; involucral bracts broadly ovate, deep green or white ................................................................... 11 10(9). Leaf blades 7–27 cm long with brownish pubescence; involucral bracts ± linear, green at anthesis, 1–2 cm long; inflorescence gland secreting bluish green resin; style ± evenly columnar; pistillate sepals 6, unlobed; fruit 3.5–4 cm diameter ........................................................ D. megacarpa 10. Leaf blades 5–15 cm long, lacking brownish pubescence; involucral bracts ovate, pale pink with reddish veins, 3–5 cm long; inflorescence gland secreting brownish maroon resin; style with distinct peltate stigmatic tip; pistillate sepals 6–12, 2-pinnatifid; fruit < 2 cm diameter .............. ............................................................................................ D. dioscoreifolia 11(9). Involucral bracts white at anthesis, > 3 cm long; inflorescence gland secreting yellowish resin; tips of styles broadly dilated and peltate ......... ....................................................................................................... D. affinis 11. Involucral bracts deep green at anthesis, < 2.5 cm long; inflorescence gland secreting clear to whitish resin; tips of styles only slightly dilated ............................................................................................... D. tenuiramea 12(8). Leaf stipules ovate to lanceolate-ovate, persistent; involucral bracts 3-lobed to about 1/2 their length, < 3.5 cm long at anthesis, with 5 prominent veins; inflorescence gland secreting clear to whitish resin ..... D. scandens 12. Leaf stipules lanceolate-ovate to lanceolate-linear, deciduous; involucral bracts shallowly 3-dentate at apex, > 3 cm long at anthesis, with 7–9 prominent veins; inflorescence gland inconspicuous or secreting yellowish resin ..................................................................................... 13 13(12). Leaf stipules lanceolate-linear; involucral bract whitish cream at anthesis; inflorescence gland prominent, secreting yellowish resin ....... D. tiliifolia 13. Leaf stipules lanceolate-ovate to lanceolate; involucral bracts pale green at anthesis; inflorescence gland inconspicuous and vestigial, not secreting resin ....................................................................... D. brownsbergensis Dalechampia affinis Müll. Arg., Linnaea 34: 223. 1865. Dalechampia dioscoreifolia Pulle, Enum. Vasc. Pl. Surinam 260. 1906, non Poepp. 1841. Twining or sprawling vine. Open woodlands and savannas, near sea level to 500 m; Bolívar (Río Orinoco basin), Amazonas (Río Negro, Río Orinoco). Eastern Apure; Guyana, Suriname, Amazonian Brazil. ŠFig. 126. Dalechampia affinis is very similar and closely related to D. tiliifolia, from which it differs primarily in having unlobed leaves and peltate styles. In Brazil, the copious yellow resin secreted by the inflorescence glands is collected by female euglossine bees (including Euglossa), which pollinate the flowers. Dalechampia attenuistylus Armbr., Brittonia 41: 44. 1989.

Twining vine reaching at least lower canopy. Moist forests, 800–1000 m; Bolívar (near Santa Elena de Uairén). Suriname. A member of sect. Rhopalostylis, Dalechampia attenuistylus is apparently most closely related to D. heterobractea and D. parvibracteata. As is the case for all members of the section, it lacks the prominent inflorescence gland characteristic of most of the rest of genus. Dalechampia brownsbergensis G.L. Webster and Armbr., Syst. Bot. 7: 484. 1982. Twining vine reaching canopy. Wet forests, 200–500 m; southern Delta Amacuro, northeastern Bolívar. Suriname. Phylogenetic analysis has shown that this species has secondarily lost the resiniferous inflorescence gland characteristic of most

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species. Instead, it secretes a reward fragrance from the stigmatic surface of the pistillate flowers, attracting pollinating male euglossine bees which collect the fragrance. Dalechampia dioscoreifolia Poepp., Nov. Gen. Sp. Pl. 3: 20. 1845. Twining vine reaching canopy. Moist or wet forests, 100–1200 m; northern and western Bolívar, northern Amazonas. Venezuelan Coastal Cordillera; Nicaragua, Costa Rica, Panama, Colombia, French Guiana, Peru, probably also in Ecuador and Brazil. ŠFig. 127. In Panama, the maroon resin secreted by the inflorescence glands is collected by female euglossine bees (including Eulaema and Euglossa), which pollinate the flowers. Dalechampia heterobractea Armbr., Syst. Bot. 21: 226, pl. 10–11C–E. 1996. Twining vine reaching canopy. Moist evergreen lowland to lower montane forests, 100–500 m; eastern Bolívar. Guyana, Suriname, French Guiana, Brazil. ŠFig. 128. A member of sect. Rhopalostylis, Dalechampia heterobractea lacks the prominent inflorescence gland. It can be distinguished from the related D. parvibracteata, with which it has been confused, by its dimorphic involucral bracts. Dalechampia liesneri Huft, Ann. Missouri Bot. Gard. 76: 1078. 1989. Twining vine reaching at least lower canopy. Moist forests, 100–500 m; Amazonas (Río Cataniapo, Cerro Yutajé). Brazil. Dalechampia liesneri is a member of sect. Rhopalostylis, but is very distinct from the other member species (including D. attenuistylus, D. heterobractea, D. micrantha, and D. parvibracteata) in having columnar rather than clavate styles. Dalechampia magnoliifolia Müll. Arg., Linnaea 34: 219. 1865. Dalechampia roezliana var. amazonica Ule, Verh. Bot. Vereins Prov. Brandenburg 50: 82. 1909. Monopodial subshrub; leaves slightly serrate-margined. Evergreen lowland forests, 50–300 m; rare in Bolívar and Amazonas. Amazonian Ecuador, Peru, and Brazil.

The inflorescence gland of Dalechampia magnoliifolia secretes fragrance rather than resin. In Peru, the flowers are pollinated by fragrance-collecting male euglossine bees (Eulaema and Euglossa). Dalechampia megacarpa Armbr., Brittonia 41: 47. 1989. Twining vine reaching canopy. Evergreen lowland to montane forests, 100–1000 m; Bolívar (San Ignacio), Amazonas (San Carlos de Río Negro). Amazonian Brazil. The blue-green resin secreted by the inflorescence gland of Dalechampia megacarpa is collected by female Euglossa bees that pollinate the flowers. Dalechampia micrantha Poepp., Nov. Gen. Sp. Pl. 3: 19. 1841. Rhopalostylis buettnerioides Klotzsch ex Baill., Adansonia 5: 317. 1865. Megalostylis poeppigii S. Moore, J. Bot. 54: 208. 1916. Twining vine reaching canopy. Evergreen lowland to lower montane forests, 100–1000 m; eastern Bolívar. Suriname, French Guiana, Peru, Amazonian Brazil. A member of sect. Rhopalostylis, Dalechampia micrantha lacks the resiniferous inflorescence gland and instead secretes a fragrance from the stigmatic surface of the pistillate flowers. It is pollinated in Bolívar state by fragrance-collecting male Euglossa bees. Dalechampia papillistigma Armbr., Brittonia 41: 49. 1989. Twining vine reaching at least lower canopy. Upland forests, 800–1000 m; southeastern Bolívar (between Santa Elena de Uairén and El Paují). Endemic. The inflorescence gland of this species secretes resin as a pollinator reward. Dalechampia parvibracteata Lanj., Recueil Trav. Bot. Néerl. 31: 463. 1934. Twining vine reaching canopy. Evergreen lowland to lower montane forests, 100–500 m; eastern Bolívar. Guyana, Amazonian Brazil. A member of sect. Rhopalostylis, Dalechampia parvibracteata lacks the prominent inflorescence gland. The swollen styles and fragrance secretion by the stigmas suggest pollination by male euglossine bees.

Dalechampia 137

Fig. 126. Dalechampia affinis

Fig. 127. Dalechampia dioscoreifolia

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Fig. 128. Dalechampia heterobractea

Fig. 129. Dalechampia tiliifolia

Fig. 130. Dalechampia scandens

Dendrothrix 139

Dalechampia scandens L., Sp. Pl. 1053. 1753. Dalechampia colorata L. f., Suppl. Pl. 421. 1781 [1782]. Dalechampia latifolia Lam., Encycl. 2: 257. 1786. Dalechampia villosa Lam., Encycl. 2: 257. 1786. Dalechampia mollis Vahl, Eclog. Amer. 3: 44. 1807. Dalechampia ruboides H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 102. 1817. Dalechampia sidaefolia H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 100. 1817. Dalechampia rubiformis Spreng., Syst. Veg. 3: 86. 1826. Dalechampia brevipes Briq., Annuaire Conserv. Jard. Bot. Genève 4: 609. 1900. Twining or sprawling, urticating vine. Open areas, secondary scrub, margins of moist forests, near sea level to 500 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; widely distributed from Mexico and the Caribbean to Argentina. ŠFig. 130. Dalechampia scandens is a variable species with numerous described infraspecific taxa and extensive ecotypic differentiation. The group needs thorough systematic study before any subspecific classification should be attempted. Flowers are pollinated by resin-collecting female euglossine bees (Euglossa, Eulaema), megachilid bees (Hypanthidium), and/or stingless bees (Trigona).

Dalechampia tenuiramea Müll. Arg., Linnaea 34: 222. 1865. Dalechampia cynanchoides S. Moore, Trans. Linn. Soc. London ser. 4, 2: 468. 1895. Twining vine. Openings in seasonally dry forests, 100–200 m; Amazonas (Puerto Ayacucho). Brazil, Bolivia. Dalechampia tenuiramea is apparently related to the D. convoluloides Lam. or D. heteromorpha Pax & K. Hoffm. and D. anomola Pax and K. Hoffm. complexes of Central America and South America. Dalechampia tiliifolia Lam., Encycl. 2: 257. 1786. Dalechampia peruviana Lam., Encycl. 2: 257. 1786. Dalechampia heterophylla Vahl, Eclog. Amer. 3: 44. 1807. Dalechampia pruriens Griseb., Fl. Brit. W. I. 51. 1859. Twining vine, sometimes reaching lower canopy. Open areas, secondary scrub, edges of dry to moist forests, 50–1800 m; widespread in Bolívar, northern Amazonas. Throughout lowland Venezuela; southern Mexico, most of Central America, Colombia, Trinidad, Guyana, French Guiana, Peru, Brazil, Bolivia. ŠFig. 129. The inflorescence glands of Dalechampia tiliifolia secrete yellow resin that is collected by pollinating female euglossine bees (Eulaema, Euglossa).

20. DENDROTHRIX Esser, Novon 3: 245. 1993. by Hans-Joachim Esser Monoecious trees or shrubs, with white latex; trichomes multicellular, dendritic; ultimate ramifications alternate. Leaves alternate, simple, glabrous or pubescent; stipules small and scaly, entire, eglandular, or lacking; petioles distinct, eglandular; blades entire, mostly revolute, eglandular on upper surface, papillate on lower surface, with one pair of marginal glands at the base, without further distinct marginal glands but mostly with small, dispersed pellucid laminar dots; tertiary venation percurrent or reticulate. Inflorescences elongate, terminal, compoundthyrsoid, yellowish, side branches subtended by minute scaly bracts; floral bracts with one or several pairs of cup- to disk-shaped glands, always touching the axis; bracts of cymule with one or several pairs of glands which are cup-shaped or diskshaped when dry, touching the rachis; staminate flowers in 7–20-flowered apical cymules, pistillate flowers one per bract at the base, or sometimes lacking. Staminate flowers erect; bracteoles absent; pedicels short, 0.5–1 mm long at anthesis, articulation just below calyx; perianth 2-lobed, partially fused; stamens (1)2 per

140

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flower; filaments connate, at anthesis as long as or longer than anthers; anthers connate; disk and pistillode absent. Pistillate flowers: bracteoles short or absent; pedicel short; perianth 3-lobed, basally connate; ovary 3-locular, smooth, pubescent, glabrescent; ovules 1 per locule; style column 0.5–1.5 mm long; stigmas 3, undivided; disk and staminodes absent. Fruit a dry septicidal schizocarp, globose, slightly sulcate; exocarp smooth, pubescent, glabrescent; mericarps with glabrous or distinct septa; columella slightly winged. Seeds 3 per fruit, ellipsoid, with caruncule (unknown in D. multiglandulosa); testa dry, smooth, brown. Endemic to the Guyana Shield in southern Venezuela and northern Brazil; 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Dendrothrix 1. 1.

Mature leaves usually glabrous; floral bracts each with several pairs of glands ........................................................................... D. multiglandulosa Mature leaves usually pubescent; floral bracts each with 1 pair of glands ................................................................................................. D. yutajensis

Fig. 131. Dendrothrix yutajensis

Discocarpus 141

Dendrothrix multiglandulosa Esser, Novon 3: 246. 1993. Shrub or tree to 5 m tall. Montane forests and scrub, 1000–1500 m; Amazonas (Cerro Cuao, Cerro Sipapo). Endemic. Dendrothrix yutajensis (Jabl.) Esser, Novon 3: 248. 1993. —Sapium yutajense Jabl., Mem. New York Bot. Gard. 17(1): 184, fig. 24. 1967. —Senefeldera yuta-

jensis (Jabl.) G.L. Webster, Ann. Missouri Bot. Gard. 76: 958. 1989. Shrub or tree to 10 m tall. Montane forests, thickets, and savannas on sandstone, 600–1400 m; Bolívar (Cerro Guaiquinima, Cerro Marutaní, Sierra Pakaraima), Amazonas (Cerro Aracamuni, Cerro Yapacana, Cerro Yutajé, Sierra de la Neblina). Brazil (Amazonas: Serra Aracá). ŠFig. 131.

21. DISCOCARPUS Klotzsch, Arch. Naturgesch. 7(1): 201. 1841. by Grady L. Webster and Paul E. Berry Dioecious trees or shrubs. Leaves alternate, pinnately veined, glabrous or nearly so, coriaceous, entire, eglandular; stipules deciduous. Flowers in axillary clusters; bracts eglandular. Staminate flowers sessile, congested; calyx 4- or 5-lobed, lobes imbricate; disk annular, entire or lobed; petals small and narrow, or absent; stamens (4)5; filaments connate below the level of the disk, free portions 1.5–3 mm long; anthers introrse; pistillode trifid. Pistillate flowers 1–3(–5) per node; pedicels 0–5 mm long; calyx 5-lobed; disk massive, patelliform; petals (0–)5, hyaline, 0.5–3 mm long; ovary 3-carpellate, densely pubescent; styles 3, parted to the base or nearly so, spreading horizontally; stigmas 3, dilated, lobed, horizontal or reflexed; ovules 2 per locule. Fruit capsular, symmetrically 3-lobed or asymmetric due to abortion of 2 carpels, longitudinally dehiscent; pericarp hard, smooth to deeply sculpted, densely pubescent; columella persistent. Seeds 1–3 per fruit, ecarunculate; endosperm scanty or absent. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 3 species, all in the flora area.

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

2.

Ovary and fruit surface smooth; capsules asymmetrical (due to abortion of 2 carpels), 1-seeded; stigmas slender, horizontal ............ D. spruceanus Ovary and fruit surface sculpted; capsules symmetrical, 3-seeded; stigmas dilated, reflexed .............................................................................. 2 Pistillate flowers with pedicels 4–5 mm long; ovary and fruit surface deeply sculpted into long undulate ridges with sharp crests beneath dense indumentum ...................................................................... D. gentryi Pistillate flowers sessile or subsessile; ovary and fruit surface weakly sculpted into shallow depressions or short undulate ridges with rounded crests beneath dense indumentum ................... D. essequeboensis

Discocarpus essequeboensis Klotzsch, London J. Bot. 2: 52. 1843. Discocarpus brasiliensis Klotzsch ex Müll. Arg. in Mart., Fl. Bras. 11(2): 13. 1873. Tree 4–15 m tall. Riparian forests, 50–200 m; Amazonas (Río Orinoco near Cariche). Guyana, Suriname, French Guiana, Brazil.

Discocarpus gentryi S.M. Hayden, Ann. Missouri Bot. Gard. 83: 159. 1996. Tree or shrub 3–8 m tall. Granitic outcrops along rivers, riparian forests, 50–100 m; Amazonas (Río Orinoco at mouth of Río Cataniapo, Santa Bárbara del Orinoco). Peru (Loreto), Brazil (Amazonas). ŠFig. 132.

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Fig. 132. Discocarpus gentryi

Fig. 133. Discocarpus spruceanus

Discocarpus spruceanus Müll. Arg., Linnaea 32: 78. 1863. Tree or shrub 6–12 m tall. Seasonally flooded river plains of black-water rivers,

100–200 m; Amazonas (Caño Yagua near Cerro Yapacana, Río Atacavi basin, lower Río Baría, Río Guainía, Río Yatúa, Río Negro). Colombia (Guainía), Brazil. ŠFig. 133.

22. DITAXIS Vahl ex A. Juss., Euphorb. Gen. 27. 1824. by Grady L. Webster and Paul E. Berry Shrubs or perennial herbs, monoecious or rarely dioecious; stems and foliage with malpighiaceous trichomes. Leaves alternate, entire to serrulate, stipulate. Inflorescences axillary, raceme-like thyrses (sometimes contracted to glomerules), usually bisexual, the pistillate flowers proximal. Sepals normally 5, valvate and coherent in bud; petals 5, free, in staminate flowers basally adherent to the staminal

Drypetes 143

column; disk dissected. Stamens 5–15 (mostly 8–10); filaments connate into a column (sometimes with apical staminodes); anthers generally in 2 whorls, dehiscing longitudinally. Carpels 3; ovules 1 per locule; styles bifid. Fruit a capsule. Seed coat pitted or reticulate, ecarunculate. Southern U.S.A. through Neotropics to Argentina; ca. 40 species, 5 in Venezuela, 1 of these in the flora area. This genus is very close to Argythamnia, but differs in its pollen morphology and larger number of stamens. Ditaxis polygama (Jacq.) L.C. Wheeler, Contr. Gray Herb. 124: 39. 1939. —Croton polygamus Jacq., Enum. Sust. Pl. 32: 1762. —Argythamnia polygama (Jacq.) Kuntze, Revis. Gen. 2: 593. 1891. Shrub 1–2 m tall; flowers white. Deciduous and semideciduous forests interspersed

with Trachypogon savannas, 50–300 m; Bolívar (near Mina de El Manganeso south of El Palmar, 3 km north of Represa Guri). Anzoátegui, Carabobo, Distrito Federal, Falcón, Sucre, Zulia; Lesser Antilles, Colombia. ŠFig. 134.

Fig. 134. Ditaxis polygama

23. DRYPETES Vahl, Eclog. Amer. 3: 49. 1807. by Geoffrey A. Levin Dioecious trees or shrubs, without latex. Leaves alternate, simple, petiolate, base often oblique, margins entire or serrate with glandular or spinose teeth, sometimes dimorphic with the juvenile leaves spinose and the adult leaves entire; stipules large and persistent or more often small and deciduous. Inflorescence axillary, fasciculate. Flowers pedicellate, symmetrical. Staminate flowers with 4 or 5(–7) imbricate sepals; petals absent; stamens 3–12(–50); filaments free; anthers bilocular, dehiscing longitudinally; disk intrastaminal (rarely absent), annular or lobed, the lobes sometimes extending between the bases of the stamens and enclosing them; pistillode small or obsolete. Pistillate flowers with 4 or 5(–7) imbricate, de-

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ciduous sepals; petals absent; disk annular, surrounding base of ovary, or rarely absent; ovary 1–3(–6)-locular; ovules 2 per locule; styles obsolete; stigmas as many as the carpels, elongate, reniform, pulviniform, or subpeltate. Fruit drupaceous; mesocarp leathery or fleshy; endocarp crustaceous. Seeds as many as carpels or solitary, ecarunculate. Pantropics, particularly Paleotropics; ca. 225 species, 4 in Venezuela, 2 of these in the flora area. Recent molecular and biochemical data indicates that Drypetes may not belong in the Euphorbiaceae at all. It may eventually be placed in a family called Putranjivaceae (K. Wurdack, personal communication). G. L. Webster (Madroño 24: 65–68. 1977) reported Drypetes standleyi from Venezuela and the flora area on the basis of two pistillate specimens. Both have the wide stigmas characteristic of D. variabilis, and although neither bears attached mature fruits, both have fruiting pedicels that are longer than those characteristic of D. standleyi and more closely approach the longer pedicels of D. variabilis. A staminate specimen (Cardozo et al. 1990, MO, MY, SD!), which Webster had not seen, shows that D. standleyi does reach Aragua state in northern Venezuela. Three additional species in the flora area, all described from staminate material and known only from their types, do not belong in this genus. Drypetes maguireana Monach. and D. spruceana Müll.Arg. are both synonyms of Chaetocarpus schomburgkianus (Kuntze) Pax & K. Hoffm. The identity of the third species, D. monachinoi Jabl., remains in doubt, but it is probably not a member of the Euphorbiaceae. Key to the Species of Drypetes 1.

1.

Leaf apex definitely acuminate; adult leaves crenate; secondary leaf veins forming loops near the margin; sepals < 2 mm long; stigma clearly reniform, < 1 mm wide, on a short style; staminate flowers 7–12 per fascicle; stamens 4; anthers ca. 0.5 mm long .............................. D. fanshawei Leaf apex acute, obtuse, or rounded, not acuminate; adult leaves entire; secondary leaf veins forming loops well in from the margin; sepals 2.5– 3.5 mm long; stigma subpeltate, > 2 mm wide, essentially sessile and forming a cap on the ovary in flower; staminate flowers 1–6 per fascicle; stamens 4–7; anthers > 1 mm long ................................ D. variabilis

Drypetes fanshawei Sandwith, Bull. Misc. Inform. Kew 7: 258. 1952. —Pan de acure, Querotín. Tree to 15 m tall. Riparian forests, 50–300 m; Delta Amacuro (Río Amacuro near Guyana border, Serranía de Imataca). Adjacent Guyana. Mature fruits of this narrowly distributed species remain unknown.

Drypetes variabilis Uittien, Recueil Trav. Bot. Néerl. 22: 348. 1925. —Kerosén,

Kerosén blanco, Kerosén negro, Querosén. Tree to 45 m tall; bark with prominent lenticels when young, becoming scaly; drupe light yellow when ripe. Evergreen lowland to lower montane forests, 100–600 m; Delta Amacuro (south of Río Orinoco), northeastern Bolívar. Eastern Apure; Guyana, Suriname, French Guiana, Brazil (northeastern Amazonia). ŠFig. 135. Flakes of fresh bark burn easily and produce black smoke with the odor of kerosene or paraffin, hence the common names.

Euphorbia 145

Fig. 135. Drypetes variabilis

24. EUPHORBIA L., Sp. Pl. 450. 1753. by Paul E. Berry Herbs, shrubs, or trees with milky white latex, monoecious. Leaves alternate, opposite, or verticillate, simple, entire or serrate, sometimes lobed, pinnately veined; stipules gland-like, minute or absent. Inflorescences terminal or axillary, composed of 1–many cyathia in fascicles, cymes, or panicles, often subtended by reduced, opposite leaves; cyathium with 5 lobes, (1 or 2)4 or 5 nectar glands alternating with the lobes, the glands with or without petaloid appendages; flowers without perianth (or rudimentary in pistillate flowers of some species). Staminate flowers few to many, consisting of a single stamen; anther with 2 divergent thecae. Pistillate flower terminal, solitary, consisting of a 3-locular ovary, each cell with a single ovule; styles 3, free or connate at the base, usually distally bifid. Fruits capsules or schizocarps, exserted from the cyathium at maturity. Seeds ovoid, smooth or variously sculpted, with or without a caruncle. Cosmopolitan, primarily tropical or subtropical; ca. 1,800 species, ca. 15 in Venezuela, 5 of these in the flora area. The delimitation of Euphorbia is a matter of some dispute. If Chamaesyce is recognized as a distinct genus, then other segregate genera may need to be recognized as well, or else a broader circumscription of Euphorbia should be accepted (M. Christenhusz, personal communication).

146

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Key to the Species of Euphorbia 1.

1. 2(1).

2.

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

Shrubs or trees, sometimes cultivated; leaves broadly ovate to suborbicular, verticillate, round at the base, 3–10 × 2–8 cm; petioles 2–7 cm long ................................................................................................. E. cotinifolia Herbs, often weedy; leaves generally smaller than above ........................ 2 Leaves subtending the inflorescences usually with white or purple spots, leaf blades often pandurate in shape with 2 lateral lobes separated by a rounded sinus; cyathium with a single marginal gland .... E. heterophylla Leaves subtending the inflorescences not brightly colored or with colored spots, leaf blades never pandurate; cyathium with 2–5 marginal glands ................................................................................................................ 3 Petioles usually considerably longer than the leaves ............. E. humayensis Petioles shorter than or as long as the leaves ........................................... 4 Inflorescence with tightly packed, conspicuous bracts ................... E. comosa Inflorescence without tightly packed, conspicuous bracts ........ E. spruceana

Cyathium

Fig. 136. Euphorbia cotinifolia

Gavarretia 147

Euphorbia comosa Vell., Fl. Flumin. 5: 202. 1825 [1829]. Small herb. Mauritia palm swamps, 100– 300 m; Bolívar (near Altamira). Anzoátegui, Aragua, Mérida, Sucre, Zulia; Colombia, Brazil, Bolivia, Argentina. Euphorbia cotinifolia L., Sp. Pl. 454. 1853. Euphorbia cotinoides Miq., Stirp. Surinam. Select. 96. 1850 [1851]. Euphorbia caracasana (Klotzsch & Garcke) Boiss. in A. DC., Prodr. 15(2): 60. 1862. —Alectoroctonum caracasanum Klotzsch & Garcke, Abh. Konigl. Akad. Wiss. Berlin 1860: 40. 1860. Shrub or small tree 2–8 m tall; leaves usually in whorls of 3, 3–10 × 2–8 cm, green or purple, petioles 2–7 cm long. Semideciduous forests, gardens, 100–500 m; Bolívar (Altiplanicie de Nuria), Amazonas (San Carlos de Río Negro [cultivated]). Scattered through northern Venezuela; southern U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 136. There are several variants within this species (acute vs. retuse leaf apices; reddish or green leaves) that can be recognized at a subspecific or varietal level. Some of these will be published in the upcoming Euphorbiaceae treatment for the Flora of the Guianas (M. Christenhusz, personal communication).

Euphorbia heterophylla L., Sp. Pl. 453. 1753. —Poinsettia heterophylla (L.) Klotzsch & Garcke, Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1859: 253. 1859. Erect, weedy herb to 1 m tall, with a central taproot; leaves variable in form on different plants, sometimes panduriform, alternate below but leaves (bracts) opposite with white marks below the inflorescences; leaf blades 2– 11 × 1.5–7 cm; petioles 6–50 mm long. Disturbed areas, savannas, 50–200 m; Bolívar (Ciudad Bolívar, Río Yuruán east of El Dorado), Amazonas (Río Ventuari near mouth of Río Parú). Aragua, Distrito Federal, Guárico, Mérida, Miranda, Portuguesa; native to Mexico, a weed in pantropical and temperate areas. Euphorbia humayensis Brandegee, Zoe 5: 208. 1905. Erect annual herb; leaves ca. 3 cm long, petioles ca. 5 cm long. Trachypogon savannas, 200–400 m; Bolívar (Fila de Chriripón between Upata and El Manteco, La Camilera 40 km west of El Manteco). Guárico, Sucre; Mexico (Baja California). Euphorbia spruceana Boiss. in A. DC., Prodr. 15(2): 53. 1862. Robust perennial herb; leaf blades ca. 3 cm long; petiole ca. 1 cm long. Semideciduous slope forests, 300–500 m; Bolívar (Altiplanicie de Nuria east of Miamo). Colombia, Ecuador, Peru, Bolivia.

25. GAVARRETIA Baill., Adansonia 1: 185, t. 7. 1860. —Conceveiba sect. Gavarretia (Baill.) Müll. Arg. in Mart., Fl. Bras. 11(2): 372. 1874. by Paul E. Berry Dioecious shrubs or trees; indument of small stellate trichomes, without latex. Leaves simple, alternate, margin crenate or dentate with glandular teeth, pinnately veined; stipules lanceolate, 3–14 mm long, deciduous or persistent. Inflorescence a terminal or axillary spike or raceme. Staminate flowers apetalous, in glomerules; stamens numerous, without pistillode; anthers apiculate. Pistillate flowers apetalous, calyx cupular; ovary 2- or 3-locular; stigmas 2 or 3, deeply divided and with long papillae. Fruit capsular, subglobose. Seeds carunculate. Colombia, Venezuela, Guyana, Peru, Brazil; 1 or 2 species. Gavarretia terminalis Baill., Adansonia 1: 185, t. 7. 1860. —Caruguano, Palo de agua dulce, Pico rojo, Temare de arrendajo, Yakui dan (Piaroa).

Shrub or tree (3–)5–25 m tall. Evergreen lowland to lower montane forests, gallery forests, mostly on white sand, 50– 800 m; Bolívar (Icabarú, Río Acanán near

148

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Guadequen), Amazonas (Maroa, Pimichín, 78 km northeast of Puerto Ayacucho, Río Baría, upper Río Cuao, Río Pasimoni, San Carlos de Río Negro, between San Fernando de Atabapo and Santa Bárbara del Orinoco). Other distribution as in genus. ŠFig. 137. Some specimens included here (Gentry 46498, MO, VEN; Williams 14543, VEN; Williams 14556, VEN; Clark 6606, VEN; and

Morillo et al. 3932, VEN) are intermediate with Conceveiba and may represent a distinct taxon (see P. Franco. Las euforbiáceas de la región de Araracuara. Estudios en la Amazonía Colombiana 9: 75. 1995). They have glands on the base of the leaf blade and 3-carpellate ovaries, as opposed to 2-carpellate ovaries and glandless blades in typical Gavarretia terminalis. Franco treated these as a distinct “sp. A.”

Fig. 137. Gavarettia terminalis

Gymnanthes 149

Fig. 138. Glycydendron amazonicum

26. GLYCYDENDRON Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 199. 1922. by Paul E. Berry Dioecious, medium-sized to large trees, with white latex in young shoots. Leaves alternate, tripliveined, elliptic, 9–15 × 4–7 cm, glabrous, nitid on upper surface, apex cuspidate-acuminate, base usually acute-cuneate or rarely rounded, margin entire or undulate, with 2 sunken glands at junction with petiole; petiole 2–6 cm long, with thickened apical pulvinus; stipules small, subulate, caducous. Calyx 4parted, imbricate. Staminate flowers with 25–30 stamens, erect in bud, intrastaminal disk present. Pistillate flower with several staminodia, ovary 2-locular, 1 ovule per locule; stigmas 2, sessile, bifid, contorted. Fruit a spindle-shaped drupe with fleshy mesocarp; endocarp thick-woody, 1-seeded. Seed with crustaceous epidermis and large cotyledons. Colombia(?), Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 of these in the flora area. Glycydendron amazonicum Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 199. 1922. Tree 8–15 m tall; flowers creamy white, fragrant; fruits 2–5 × 1–2 cm. Riparian for-

ests, 50–100 m; Amazonas (mouth of Río Cuao, Río Guayapo, mouth of Río Sipapo). Other distribution as in genus. ŠFig. 138.

27. GYMNANTHES Sw., Prodr. 95. 1788. by Paul E. Berry and Hans-Joachim Esser Monoecious or rarely dioecious shrubs or trees. Leaves alternate, simple, usually coriaceous, pinnately veined; margins glandular or eglandular and entire or crenulate (a few embedded laminar glands in G. hypoleuca); stipules present. Inflorescences axillary, usually bisexual, solitary, spicate, with 1–few proximal pistillate flowers and many distal staminate cymules; bracts mostly biglandular, stiffly ovate-

150

E UPHORBIACEAE

triangular. Staminate flowers lacking perianth or with 1 or 2 rudimentary calyx lobes, disk lacking; stamens mostly (2)3(–6); filaments free or basally connate; anthers extrorse and longitudinally dehiscent. Pistillate flowers lacking perianth or with 3 small sepals; ovary sessile or stipitate, 3-locular, 1 ovule per locule; styles free or basally connate, branches simple. Fruits capsular, the columella persistent. Seeds subglobose, carunculate; testa smooth; endosperm copious. Southern U.S.A., Mexico, Central America, West Indies, northern South America, ca. 20 species, 2 in Venezuela, 1 of these in the flora area. Gymnanthes is sometimes treated to include Actinostemon, but the genera are maintained as distinct in this treatment. Gymnanthes hypoleuca Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 325. 1854. —Ateramnus hypoleucus (Benth.) Rothm., Feddes Repert. Spec. Nov. Regni Veg. 53: 5. 1944. Small tree to 10 m tall; leaves elliptic to lance-oblong, 7–12 × 2.5–6 cm, apex acumi-

Fig. 139. Gymnanthes hypoleuca

nate, margins entire, lower surface glaucous with a few embedded laminar glands. Riparian forests, ca. 100 m; Amazonas (San Carlos de Río Negro). West Indies, Colombia, Trinidad, Guyana, French Guiana, Brazil (Amazonas, Pará). ŠFig. 139.

Haematostemon 151

28. HAEMATOSTEMON (Müll. Arg.) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 68): 31. 1919. —Astrococcus sect. Haematostemon Müll. Arg., Linnaea 34: 158. 1865. by Lynn J. Gillespie Monoecious shrubs or trees. Leaves alternate, simple, stipulate, very shortly petiolate, eglandular; blades acuminate at apex, pinnately veined; margins serrulate to serrate. Inflorescences axillary, spicate, bisexual; lower 2–4 nodes with solitary pistillate flowers or bisexual cymules with a central pistillate flower, upper nodes with condensed staminate cymules, terminal flower pistillate; bracts narrowly triangular, eglandular. Staminate flowers pedicellate; sepals 4, valvate; petals and disk absent; stamens 4; filaments highly dilated and fused at base, sometimes appearing like an intrastaminal pseudodisk; pistillode absent. Pistillate flowers pedicellate; sepals 4–6; petals and disk absent; ovary 3-locular, distinctly 3-lobed with lobes globose and verrucate, ovules 1 per locule; styles connate into a massive, elongated, cup-shaped, hollow-centered column, truncate and papillose at apex. Fruit capsular. Endemic to the Guayana Shield in southern Venezuela and Guyana; 2 species, 1 in Venezuela. The second species in the genus, Haematosetmon guianensis Sandw., is endemic to central Guyana and is distinguished by its larger oblanceolate leaves and larger inflorescences. Haematosetmon is closely related to Astrococcus and Angostyles. Haematostemon coriaceus (Baill.) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 68): 32. 1919. —Astrococcus coriaceus Baill., Adansonia 5: 308. 1864. Shrub to 3 m tall; leaves lanceolate, to 14 × 4 cm, narrowly acute at the apex, coriaceous, finely glandular-crenate; inflorescences 3–8 cm long. Black-water flooded forests, 100–200 m; Amazonas (Río Guainía and tributaries). Endemic. ŠFig. 140. It is very likely that Haematostemon coriaceus will be found on the Colombian side of the Río Guainía.

Pistillate flower Fig. 140. Haematostemon coriaceus

152

E UPHORBIACEAE

29. HEVEA Aubl., Hist. Pl. Guiane 871. 1775. by Paul E. Berry, Grady L. Webster, and Scott V. Heald Monoecious trees; stems with copious milky latex. Leaves alternate, trifoliolate, leaflets acuminate, entire, pinnately veined; petioles long, glandular at apex; stipules small, deciduous. Inflorescences terminal or axillary panicles; bracts eglandular. Flowers pedicellate, gamosepalous, lobes 5, inflexed-valvate, apetalous. Staminate flowers: disk lobed, dissected, or obsolete; stamens 5–10; filaments connate; anthers in 1 or 2 ranks; pistillode at tip of staminal column. Pistillate flowers: disk obsolete or absent; ovary 3-locular, ovules 1 per locule; styles connate at base, stigmatiform. Fruits large capsules, 3-lobed, usually explosively dehiscent. Seeds ecarunculate, mottled. Amazonian Colombia, southern Venezuela, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, and Bolivia; 10 species, 4 in Venezuela, all in the flora area. Hevea rigidifolia (Spruce ex Benth.) Müll. Arg., a species of medium-sized trees to 12 m tall, is known from low, white-sand caatinga forests in the Rio Negro and Rio Uaupés area of northern Brazil and is likely to be found in the flora area. Hevea brasiliensis (Willd. ex A. Juss.) Müll. Arg., a species of larger trees to 50 m tall, is the source of most commercial rubber and is not native to Venezuela. However, it has been planted experimentally in Amazonas state at El Pozo, southeast of San Fernando de Atabapo, as well as at Macuruco and Trapichote. During the latter part of the 19th century and the early part of the 20th century, there were important commerical rubber tapping operations in Venezuela, especially along the Río Casiquiare and parts of the Río Orinoco. San Fernando de Atabapo was the center for these operations, and Hevea benthamiana and H. guianensis were the main species exploited. Key to the Species of Hevea 1.

1.

2(1). 2.

3(2).

Pistillate flower with a conspicuously thickened torus persisting at the base of the ripened capsule; capsule conical-pyramidal, triangular in cross section, apically long-acute, not explosively dehiscent, valves thin; trunks basally enlarged and quickly tapering to a narrow stem with a sparse crown of reclinate leaves; along heavily flooded blackwater river banks ................................................................. H. microphylla Pistillate flower lacking a swelling at base of calyx; capsule subglobose, ovoid or ellipsoid, 3-lobed or round in cross section, dehiscing explosively, valves thick; trunks occasionally basally enlarged, but then without the abruptly tapering stem and reclinate leaves on a small crown as above ....................................................................................... 2 Leaves entirely deciduous before flush of inflorescence and new leaves; anthers 8–10, in two ranks ............................................... H. benthamiana Leaves ± persistent, at least some old leaves retained with the new flush of leaves; anthers either as above or 5 in one rank or 5–7 in two irregular ranks ................................................................................................. 3 Leaflets entirely glabrous, or occasionally lepidote on lower surface; anthers 8–10 in two ranks ......................................................... H. pauciflora

Hevea 153

3.

Leaflets with sparse reddish or brown pubescence on lower surface; anthers 5 in one rank or 5–7 in two irregular ranks ................ H. guianensis

Hevea benthamiana Müll. Arg., Linnaea 34: 204. 1865. —Caucho, Caucho fino, Caucho yapi, Goma fina, Goma seringa, Seringa. Tree 10–30 m tall; leaves from entirely glabrous to reddish- or brown-pubescent on lower surface. Seasonally flooded forests and swamps, 50–300 m; Amazonas (widespread, especially along upper Río Orinoco and Río Casiquiare). Colombia (Vaupés: Río Negro basin), Brazil (Amazonas). ŠFig. 141. Hevea benthamiana was the most widely exploited species of rubber in Amazonian Venezuela earlier in this century. Hevea guianensis Aubl., Hist. Pl. Guiane 871, t. 335, 1775. —Caucho fino, Caucho legítimo, Conuri, Seringa. The most widely distributed species in the genus: Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of H. guianensis 1. Trees 10–25 m tall; anthers 5, in one rank, ca. 1 mm long. ............ var. guianensis 1. Trees 15–30 m tall; anthers 5–7, in two irregular ranks, ca. 0.5 mm long ............ ............................................. var. lutea H. guianensis var. guianensis. —Caucho fino, Goma, Seringa. Evergreen lowland and riparian forests, 50–200 m; Bolívar (El Cácaro between Rio Caura and Rio Paragua), Amazonas (Caño San Miguel, Río Casiquiare, Río Mawarinuma). Other distribution as in the species. H. guianensis var. lutea (Spruce ex Benth.) Ducke & R.E. Schult., Caldasia 3: 249. 1945. —Siphonia lutea Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 370. 1854. —Hevea lutea (Spruce ex Benth.) Müll. Arg., Linnaea 34: 204. 1865. —Caucho, Caucho yapi (Baniba), Goma, Goma concha blanca. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (Cerro Ichún, upper Río Paragua), Amazonas (Río Casiquiare,

upper Río Negro, Yavita). Colombia, Guyana, Suriname, French Guiana, Brazil. ŠFig. 142. Hevea microphylla Ule in Engl., Bot. Jahrb. 35: 669, t. 1, figs j–m. 1905. —Goma brava, Goma rebalsera, Seringa de mono. Tree 5–22 m tall, with relatively sparse, thin white latex for the genus; trunk swollen at the base (where wood is relatively lightweight), then rapidly tapering upward to a particulary small crown with reclinate leaves; bark smooth; fruit elongate, pyramidal-conical, with thin valves, yellow when ripe, not explosively dehiscent; seeds ca. 2.5 cm long. Seasonally flooded banks of blackwater rivers, 50–200 m; Amazonas (Capibara, Río Atabapo basin, Río Guainía). Colombia (Vaupés), Brazil (Amazonas). Hevea microphylla is unsuitable as a commercial source of rubber and was apparently never tapped for that purpose. Hevea pauciflora (Spruce ex Benth.) Müll. Arg., Linnaea 34: 203. 1865. —Siphonia pauciflora Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 370. 1854. Colombia, Venezuela, Guyana, Suriname, Peru, Brazil; 2 varieties, 1 in Venezuela. H. pauciflora var. coriacea Ducke, Arq. Inst. Biol. Veg. 2: 239. 1935. —Caucho banero, Goma, Goma concha blanca. Hevea confusa Hemsl. in Hook., Icon. Pl. 26: t. 2570, 2575. 1898. Hevea minor Hemsl. in Hook., Icon. Pl. 26: t. 2572. 1898. Hevea paludosa Ule in Engl., Bot. Jahrb. 35: 666. 1905. Tree to 22 m tall; leaflets coriaceous, often reclinate; calyx lobes conspicuously calloused; staminate disk lobes short and blunt. Evergreen lowland to lower montane forests, both flooded and nonflooded areas, 50–400 m; Bolívar (upper Río Caura, Río Kanarakuni, upper Río Paragua), Amazonas (Capibara, Maroa, Macuruco, Río Atabapo and tributaries, Río Sipapo, San Carlos de Río Negro, Sierra de la Neblina, Yavita). Colombia, Guyana, Suriname, Peru, Brazil. ŠFig. 143.

154

E UPHORBIACEAE

Pistillate flower

Staminate flower Fig. 141. Hevea benthamiana

Hevea 155

Pistillate flower

Staminate flower

Fig. 142. Hevea guianensis var. lutea

156

E UPHORBIACEAE

Staminate flower

Fig. 143. Hevea pauciflora var. coriacea

Pistillate flower

Hura 157

Fig. 144. Hura crepitans

158

E UPHORBIACEAE

30. HURA L., Sp. Pl. 1008. 1753. by Paul E. Berry Monoecious, small to large trees; bark with hard, sharp-tipped conical spines; sap clear. Leaves alternate, simple, long-petiolate with 2 rounded glands at petiole apex, stipulate; blades broadly ovate to subcordate; margins serrate, pinnately veined. Inflorescences unisexual; staminate flowers in narrow, cone-like, long-pedunculate spikes; each flower enclosed by a thin bract that ruptures at anthesis, calyx united to form a membranous denticulate cup, petals and disk absent; stamens numerous, united; anthers sessile, verticillate. Pistillate flowers solitary in upper leaf axils, on thick pedicels; calyx cup-shaped; ovary 5–20-locular, each with a single ovule; styles connate into a long, thick column, style branches as many as the locules. Fruit a large, woody, depressed, explosively dehiscent capsule. Seeds large, suborbicular, laterally compressed, ecarunculate. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela. Hura crepitans L., Sp. Pl. 1008. 1753. —Jabillo. Tree to 40 m tall; trunk with conical spines. Riparian and semideciduous forests, ca. 100 m; Bolívar (Temblador on lower Río Caura). Widespread elsewhere in Venezuela;

Nicaragua, Costa Rica, Panama, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, widely cultivated throughout the tropics. ŠFig. 144.

31. HYERONIMA Allemão, Pl. Novas Brasil 1848. by Grady L. Webster Dioecious trees or shrubs; indument lepidote. Leaves alternate, pinnately veined, entire; stipules deciduous. Inflorescences axillary, ± compound spiciform or racemiform thyrses; bracts eglandular; flowers apetalous. Staminate flowers subsessile; calyx basally connate, 4- or 5-lobed; disk pulviniform, extrastaminal; stamens 3–6; filaments free; anthers with enlarged connective and pendent anther sacs; pistillode present. Pistillate flowers pedicellate; calyx as in staminate flowers; disk cupular; ovary glabrous or lepidote, 2-locular; stigmas sessile; ovules 2 per locule. Fruit drupaceous; endocarp ornamented. Seed 1 per fruit, ecarunculate. Mexico, Central America, Cuba, South America to Bolivia and southern Brazil; ca. 15 species, 5 in Venezuela, 2 of these in the flora area. Key to the Species of Hyeronima 1.

1.

Stamens 4; leaves mostly 10–30 cm long, sparsely to densely lepidote on lower surface; petioles 4–22 cm long; stipules 5–20 mm long; endocarp < 3.5 mm long .................................................................... H. alchorneoides Stamens 5; leaves usually smaller than above, subglabrous on lower surface; petioles 1–4 cm long; stipules obsolete or absent; endocarp > 4 mm long ............................................................................................. H. oblonga

Hyeronima alchorneoides Allemão, Pl. Novas Brasil (icon.) 1848. Stilaginella laxiflora Tul., Ann. Sci. Nat. Bot. sér. 3, 15: 244. 1851. —Hyeronima

laxiflora (Tul.) Müll. Arg., Linnaea 34: 67. 1865. Tree to 30 m tall; stipules foliaceous. Evergreen lowland forests, near sea level to 500

Hyeronima 159

m; Delta Amacuro (Caño Jobure, Río Amacuro, Serranía de Imataca), northern Bolívar, Amazonas (Río Siapa, San Carlos de Río Negro, base of Sierra de la Neblina). Barinas, Lara, Mérida, Táchira, Yaracuy, Zulia; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 146. Hyeronima oblonga (Tul.) Müll. Arg., Linnaea 34: 66. 1865. —Stilaginella oblonga Tul., Ann. Sci. Nat. Bot. sér. 3, 15: 248. 1851.

Tree 5–18 m tall. Evergreen lower montane to upper montane forests, 300–2300 m; Bolívar (between Betania and Santa Elena de Uairén, Cerro Jaua, widespread on tepuis of the Gran Sabana, 45 km east of Los Pijiguaos, upper Río Paragua), Amazonas (Cerro Aracamuni, Río Coro Coro, Río Matacuni, Sierra de la Neblina, Sierra Parima). Distrito Federal, Falcón, Mérida, Táchira; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 145.

Fig. 145. Hyeronima oblonga

160

E UPHORBIACEAE

Fig. 146. Hyeronima alchorneoides

Jablonskia 161

32. JABLONSKIA G.L. Webster, Syst. Bot. 9: 229. 1984. by Grady L. Webster Monoecious shrubs or small trees, glabrous. Leaves alternate, evergreen, pinnately veined, entire, with small basal laminar glands, short-petiolate; stipules caducous. Inflorescences axillary, bracteate glomerules of sessile to subsessile flowers. Sepals 5, imbricate (pistillate sepals persistent in fruit), entire; petals absent; disk present (staminate segmented, pistillate entire). Stamens 5, free; anthers versatile, dehiscing longitudinally, introrse; pistillode present. Carpels 3; ovules 2 per locule; styles free, bifid. Fruit a thin-walled, irregularly dehiscing capsule. Seed coat fleshy. Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil; 1 species. Jablonskia congesta (Benth. ex Müll. Arg.) G.L. Webster, Syst. Bot. 9: 232. 1984. —Phyllanthus congestus Benth. ex Müll. Arg., Linnaea 32: 25. 1863. Shrub or small tree 1–8 m tall; leaves 7– 14 cm long, blade with minute pigment

Fig. 147. Jablonskia congesta

streaks, decurrent on petiole. Evergreen lowland forests, seasonally flooded riparian forests, 50–200 m; Amazonas (Río Casiquiare between Guachupita and El Porvenir, Río Siapa, San Carlos de Río Negro). Other distribution as in genus. ŠFig. 147.

162

E UPHORBIACEAE

33. JATROPHA L., Sp. Pl. 1006. 1753. Curcas Adanson, Fam. Pl. 2: 356. 1763. by Grady L. Webster Monoecious or rarely dioecious trees, shrubs, or perennial herbs, sometimes tuberous or succulent; stems exuding clear or reddish latex; trichomes simple or glandular. Leaves alternate, simple, petioles without glands, blades very variable in form, often with palmate lobes, margins entire to dentate; stipules simple or glandular and branched. Inflorescences axillary or terminal pedunculate compound dichasia. Staminate flowers with 5 imbricate sepals; petals 5, free or connate to form a short tube, greenish, whitish, or reddish; disk entire or 5-segmented; stamens usually 8(6–14) in 2 whorls, anthers longitudinally dehiscent; pistillode small or absent. Pistillate flowers with 5 imbricate sepals, petals as in staminate flowers; disk annular or dissected, staminodes absent; ovary with 3(1, 2) locules, ovules 1 per locule, styles free or united, simple or bifid distally. Capsules dehisent into 3(1, 2), 2-valved cocci or ± fleshy and tardily dehsicent. Seeds carunculate; testa crustaceous, smooth; endosperm copious. Pantropics, subtropics, and southwestern U.S.A.; ca. 175 species, 6 in Venezuela (some cultivated only), 2 of these in the flora area.

Fig. 148. Jatropha gossypiifolia

Fig. 149. Jatropha curcas

Mabea 163

Key to the Species of Jatropha 1.

1.

Stipules glandular-dissected; leaf blades deeply 3- or 5-lobed, with marginal glandular teeth; petals dark reddish, glabrous; stamens mostly 8; fruits dry, seeds mottled and < 1 cm long .......................J. gossypiifolia Stipules entire, rudimentary; leaf blades unlobed or coarsely and shallowly 3–7-lobed; petals greenish to whitish, hirsute on inner surface; stamens mostly 10; fruits fleshy and tardily dehiscent, seeds blackish and > 1 cm long .............................................................................. J. curcas

Jatropha curcas L., Sp. Pl. 1006. 1753. Shrub or small tree 2–5 m tall. Dry, lowland, disturbed areas, 50–300 m; Bolívar (Ciudad Bolívar, northwest of El Manteco). Dry lowland areas of northern Venezuela; probably native to Mexico and Guatemala. ŠFig. 149. Jatropha curcas is widely cultivated for its medicinal fruits and as living fence posts in many areas of the Neotropics.

Jatropha gossypiifolia L., Sp. Pl. 1006. 1753. Large herb or shrub 1–3 m tall. Granitic outcrops, deciduous forests, savannas, disturbed areas near towns, near sea level to 200 m; Delta Amacuro (Tucupita), dry areas of northern Bolívar, Amazonas (Puerto Ayacucho). Widespread in dry areas of northern Venezuela; widespread weed throughout the Neotropics. ŠFig. 148.

34. MABEA Aubl., Hist. Pl. Guiane 867, t. 334. 1775. by Hans-Joachim Esser Monoecious trees or shrubs, sometimes climbing, or lianas, with white latex; trichomes multicellular, dendroid or rarely uniseriate, reddish to brownish or pale; lower branches whorled, terminal ones often alternate. Leaves alternate, simple, glabrous or pubescent; blades entire or serrate with glandular, mostly caducous teeth, sometimes revolute, tertiary venation reticulate, upper surface eglandular, lower surface papillose, glaucous, or smooth, with 0–40 marginal or submarginal glands on each side, basal ones sometimes enlarged; stipules basally serrate and glandular, apically entire, caducous; petioles 4–15 mm long. Inflorescences elongate, terminal and axillary, simple or compound thyrsoid, yellowish, brownish, or deep red, side branches subtended by foliaceous bracts; floral bracts with (0)1 pair of elliptic-cylindrical glands in variable position to the axis, touching it or remote or decurrent; staminate flowers in (1–)3–5(–7)-flowered apical cymules; pistillate flowers single per bract at the base sometimes with additional staminate flowers. Staminate flowers erect; bracteoles absent; pedicel distinct, 3–120 mm long, hardly elongating when flowering; perianth 5- or 6-lobed, partly connate; stamens ca. 3–80 per flower; filaments absent to longer than anthers, free; anthers free; disk and pistillode absent. Pistillate flowers: bracteoles absent or irregular; pedicel distinct; perianth 6lobed (very rarely 3-lobed), connate at base; disk and staminodes absent; ovary 3locular, smooth or with 3 pairs of protuberances, with dense, persistent, short, pale indument and sometimes additionally with brown, lanate trichomes; style column 1–21(–33) mm long; stigmas 3, undivided, recurved. Fruit a dry, septicidal schizocarp, globose to transversely ellipsoid, sulcate or not; pericarp thick, woody; exocarp smooth or rarely with 3 pairs of protuberances, densely and persistently pubescent; mericarps with distinct septa; columella slightly winged. Seeds 3 per fruit, globose to ellipsoid, with or without caruncule; testa dry, smooth, brown or sometimes variegated.

164

E UPHORBIACEAE

Mexico, Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 38 species, 16 in Venezuela, 15 of these in the flora area. The name Mabea caudata Pax & K. Hoffm. has sometimes been applied to plants from the flora area. It is, however, a synonym of M. speciosa Müll. Arg., a species not present in Venezuela. A single collection from Bolívar (Steyermark 59105, NY, Gran Sabana, Río Kukenán valley near Kun, 1065–1220 m), a tree ca. 12 m tall with leaves distinctly whitish-farinose and pale-pubescent on the whole lower surface and with numerous marginal glands, fruits 17–20 mm long, and unknown flowers might represent another undescribed species, but is insufficiently known and not included below. Key to the Species of Mabea 1.

1.

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

3.

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

5.

6(5).

6.

Leaf blades sharply serrate, distance between teeth 0.5–1(–1.5) mm; petioles and midvein pubescent but lower surface of leaf blades otherwise nearly glabrous ...................................................................... M. subsessilis Leaf blades entire or shallowly serrate, distance between teeth (2–)3–5 mm apart; petioles and lower surface of leaf blades either glabrous or both with similar pubescence ........................................................................ 2 Lower surface of leaf blades completely and densely tomentose, the surface not visible ................................................................................ M. sp. A Lower surface of leaf blades glabrous or moderately pubescent, the surface visible .............................................................................................. 3 Liana to 25 m tall; leaves with 40–55 minute marginal glands (ca. 0.15 mm diameter) per side; staminate part of thyrse (6–)8–12 cm diameter; staminate pedicels regularly fused for at least half of their length; fruits excluding style ca. 20 mm long ................................ M. pulcherrima Shrub or tree 0.2–15(–25) m tall; leaves with 0–10(–30) minute to larger marginal glands per side (0.15–0.4 mm diameter); staminate part of thyrse 0.4–4.5 cm diameter; staminate pedicels free or only slightly and irregularly fused at base; fruits excluding style 8–18 mm long .... 4 At least young leaves distinctly pubescent on lower surface ................... 5 Leaves glabrous (very rarely shortly pilose-papillate on lower surface when young) ........................................................................................... 8 Leaf blades (13–)17–25 × 5.5–10 cm; stipules 20–25 mm long; staminate part of inflorescences 13–16 × 3.5–5 cm; floral bracts 5–8 mm long; pistillate sepals 4–6 mm long; style column 15–25 mm long ...................... ......................................................................................... M. longibracteata Leaf blades 4.5–11(–17) × 2–5(–6) cm; stipules 3–5 mm long; staminate part of inflorescences 3–11 × 2–3 cm; floral bracts 2–4 mm long; pistillate sepals 2–4 mm long; style column 2–9(–15) mm long ................... 6 Leaf blades narrowly acuminate (acumen more than twice as long as wide); staminate pedicels free; fruits excluding style 13–18 mm long, with rusty-brown pubescence ........................................................ M. piriri Leaf blades broadly acuminate (acumen < twice as long as wide); staminate pedicels free or fused at base; fruits excluding style 10–13 mm long, without rusty-brown pubescence ................................................. 7

Mabea 165

7(6).

7.

8(4).

8.

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

11.

12(11). 12. 13(12).

13.

14(13).

14.

15(13).

Leaf blades cordate at base, with complete venation distinct on lower surface in dried leaves (blade whitish only between veinlets); glands of floral bracts never decurrent along axis, fruits sometimes with dorsal protuberances (muricate) ................................................................. M. taquari Leaf blades rounded-obtuse, hardly cordate at base, with only larger veins distinct on lower surface in dried leaves (blade nearly completely whitish); glands of floral bracts often decurrent along axis; fruits never with dorsal protuberances (not muricate) .............................. M. montana Inflorescences and infructescences repeatedly compound with numerous branches; floral bracts eglandular; stamens with distinct filament when flowering; fruits with rusty-brown pubescence, often with three pairs of dorsal protuberances (muricate) ..................................... M. nitida Inflorescences and infructescences simple or slightly compound with few branches; floral bracts glandular; stamens without visible filament when flowering; fruits with or without rusty-brown pubescence, without dorsal protuberances (not muricate) ............................................... 9 Leaf blades mucronate to retuse (rarely acute) at apex; fruits transversely ellipsoid (wider than long) ................................................................... 10 Leaf blades acuminate at apex; fruits globose to ellipsoid (not wider than long) ...................................................................................................... 11 Leaf blades 2–7 mm wide, staminate cymules all 1-flowered .................... ............................................................................................... M. linearifolia Leaf blades (3–)10–48 mm wide, at least some staminate cymules 3-flowered .......................................................................................... M. frutescens Leaf blades ovate-elliptic with cordate base; staminate part of thyrse 4–8 mm diameter, completely glabrous; pistillate calyx 3-lobed ........... .................................................................................................. M. anomala Leaf blades oblong-elliptic, with acute to rounded (rarely subcordate) base; staminate part of thyrse 12–30 mm diameter, at least partly papillate to pubescent; pistillate calyx 6-lobed (rarely 3-lobed) ......... 12 Bract glands removed from the inflorescence axis by a peduncle, at least in the staminate part ........................................................................... 13 Bract glands always touching the inflorescence axis ............................. 16 Leaf blades 5–11 × 2.5–4 cm; sepals of pistillate flowers and fruits sometimes glandular; fruits excluding style 11–13(–15) mm long, with pale but without reddish brown trichomes ................................................. 14 Leaf blades 10–19 × 3–8 cm; sepals of pistillate flowers and fruits eglandular; fruits excluding style 14–18 mm long, with reddish brown trichomes .............................................................................................. 15 Leaf blades densely whitish-farinose on most of the lower surface with only larger veins of different color and distinct, with 5–15 marginal glands per side; sepals of pistillate flowers and fruits eglandular .................................................................................................. M. montana Leaf blades slightly whitish-farinose on lower surface but with complete venation of different color and distinct, with 0–3 marginal glands per side; sepals of pistillate flowers and fruits often glandular ...... M. trianae Leaf blades subentire, densely whitish-farinose on nearly the whole lower surface including venation, with 5–20 marginal glands per side; stami-

166

E UPHORBIACEAE

nate part of thyrses 15–20 mm diameter; staminate cymules with 3–7 flowers each ....................................................................... M. arenicola 15. Leaf blades serrate, slightly whitish-farinose on lower surface but complete venation of different color and distinct, with 0–4 marginal glands per side; staminate part of thyrse 25–35 mm diameter; staminate cymules consistently with 3 flowers each ..................................... M. piriri 16(12). Leaf blades partly whitish-farinose on lower surface but complete venation of different color (when dried); axis of inflorescence with scattered trichomes; sepals of pistillate flowers similar, 2.5–4.5 mm long and usually longer than the ovary; style column 12–20 mm long; pedicels of staminate flowers free ......................................................... M. occidentalis 16. Leaf blades whitish-farinose on lower surface or not, but usually only largest veins of different color (when dried); axis of inflorescence with nearly continuous papillae or trichomes; sepals of pistillate flowers similar, 2–3 mm long, or 3 larger ones and 3 smaller ones, all usually shorter than the ovary; style column 3–7 mm long; pedicels of staminate flowers free or slightly fused at base .......................................... 17 17(16). Leaf blades 5–10 × 2–4 cm, with 5–15 marginal glands per side, the lower surface whitish-farinose .......................................................... M. montana 17. Leaf blades 20–25 × 6–8 cm, with 20–30 marginal glands per side, the lower surface not whitish-farinose ................................................. M. sp. B Mabea anomala Müll. Arg. in Mart., Fl. Bras. 11(2): 526. 1874. —Rebentillo, Reventilla. Shrub or small tree to 8 m tall, sometimes climbing. River banks, seasonally flooded forests and secondary vegetation on varied, especially sandy, soils, 100–200 m; Amazonas (Río Casiquiare, mouth of Río Cuao, Río Guainía, Río Negro, San Fernando de Atabapo). Colombia, northern Brazil. This species was long overlooked and often confused with Mabea subsessilis, from which it differs by the mostly ovate shape of the leaves, their widely spaced and rounded teeth, the totally missing vegetative indument, the smaller thyrses, and the apparently 3-parted calyx of pistillate flowers and fruits. Mabea arenicola Esser, Novon 3: 341. 1993. —Rebentillo, Rebentillo amarillo. Tree to 10 m tall, sometimes climbing. Roadsides, thickets, and nonflooded forests on white sand, 100–200 m; Amazonas (San Carlos de Río Negro, San Fernando de Atabapo to Santa Bárbara, Yavita to Maroa road). Colombian and Brazilian Amazonia. ŠFig. 150. Mabea frutescens Jabl., Mem. New York Bot. Gard. 17(1): 175. 1967.

Mabea orbiculata Jabl., Mem. New York Bot. Gard. 17(1): 176. 1967. Xylopodious shrub or small tree 0.2–4 m tall. Savannas, scrubland, river banks, on white sand, locally very common, 50–200 m; Amazonas (Río Atabapo, Río Guasacavi, Río Orinoco between Río Atabapo and Río Yagua). Colombia (Guainía). ŠFig. 153. Mabea frutescens is vegetatively a very variable species, the plants with broader and membranaceous leaves growing in shade, the ones growing in full sun sclerophyllous; flowers and fruits are homogeneous. Mabea linearifolia Jabl., Mem. New York Bot. Gard. 17(1): 176. 1967. Wiry shrub 20–50 cm tall. White-sand savannas, 100–200 m; Amazonas (base of Cerro Yapacana). Endemic. ŠFig. 154. Because of the vegetative variability of Mabea frutescens, M. linearifolia is best distinguished from that species only by the strictly 1-flowered staminate cymules. Mabea longibracteata Esser, Novon 3: 343. 1993. Tree to 10 m tall. Evergreen primary forests along rivers, swamps, 100–200 m; Amazonas (base of Sierra de la Neblina). Brazil (Amazonas: base of Serra da Neblina.

Mabea 167

Mabea montana Müll. Arg. in A. DC., Prodr. 15(2): 1151. 1866. Mabea lucida Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 16. 1912. —Mabea montana subsp. lucida (Pax & K. Hoffm.) Hollander, Proc. Kon. Ned. Akad. Wetensch., C. 89: 150. 1986. Mabea verrucosa Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 37. 1912. ?Mabea atroviridis Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 68): 55. 1919. Mabea longepedicellata Pittier, J. Wash. Acad. Sci. 19: 353. 1929. Mabea floribunda Jabl., Mem. New York Bot. Gard. 17(1): 174. 1967. Mabea parguazae Jabl., Mem. New York Bot. Gard. 17(1): 168. 1967. Shrub or tree 0.2–7 m tall. Rocky river banks, dry forests, savannas on sandstone and granitic soils, 50–800 m; Bolívar (Cerro Bolívar, Cerro Guaiquinima, Río Orinoco, Río Paragua, Río Parguaza), Amazonas (Puerto Ayacucho to Samariapo). Venezuelan Coastal Cordillera, Venezuelan Andes; Costa Rica, Panama, Colombia, Trinidad, Guyana. ŠFig. 152. The wood of Mabea montana is quite hard. Although clearly distinguishable from vicariant M. taquari in most of the flora area, there is a transition zone of some characters, e.g., indument and leaf shape, in some areas such as the northern Gran Sabana. In this area, muricate fruits are not known, and some specimens cannot be determined unambiguously as one of the two species. Instead, they may be better classified as hybrids. Mabea nitida Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 367. 1854. —Cachimbo, Manteca rebalsera, Reventillo. Mabea nitida var. albiflora Müll. Arg. in A. DC., Prodr. 15(2): 1152. 1866. Mabea muricata Jabl., Mem. New York Bot. Gard. 17(1): 169. 1967. Tree to 12(–20) m tall. Seasonally flooded river banks and forests of different water and soil types, 100–200 m; Bolívar (lower Río Caura, Río Parguaza, Río Suapure), Amazonas (Ocamo, Río Baría, Río Casiquiare and its tributaries, Río Cuao, Río Manapiare, Río Orinoco at least up to Río Ocamo, Río

Sipapo). Anzoátegui, Apure, Guárico; very common and widespread in the western and central Amazon Basin and in the Colombian and Venezuelan Llanos, absent in the eastern Amazon Basin. ŠFig. 151. As in Mabea taquari, the muricate fruits are a common, though inconsistant character. Mabea occidentalis Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 364. 1854. Mabea pallida Müll. Arg. in A. DC., Prodr. 15(2): 1150. 1866. Tree to 8 m tall. Humid forests, secondary forests, 100–200 m; Amazonas (Raudal Ceguera on Río Autana, Río Cataniapo, Río Cuao, lower Río Sipapo). Widespread in northern Venezuela from Zulia to Sucre; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru. Populations in the flora area differ from the typical, understory species found north of the Llanos, by having larger leaves (16–20 × 5–7 cm) with more secondary veins (16–20 pairs below acumen) and larger thyrses (4.5 cm diameter). Mabea piriri Aubl., Hist. Pl. Guiane 867, t. 334, fig.1. 1775. —Molenillo, Pata de gallina, Pata de grulla, Pata de paují, Usibe. Mabea maynensis Spruce ex Müll. Arg. in A. DC., Prodr. 15(2): 1150. 1866. Tree to 15(–25) m tall. Evergreen lowland to lower montane forests, less often riversides or secondary forests, near sea level to 900 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Gran Sabana, lower Río Caroní, upper Río Caura, Río Icutu, Serranía de Imataca), Amazonas (Caño Culebra, near Puerto Ayacucho, Río Cunucunuma, Río Ocamo, Sierra Parima). Monagas; Guyana, Amazon Basin, Brazilian Atlantic coastal forest. ŠFig. 155. Mabea piriri is recognizable by the always brown-haired ovaries and fruits, together with mostly glabrous leaves. The wood is used for construction. Mabea pulcherrima Müll. Arg., Flora 55: 44. 1872. Mabea eximia Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 107. 1925

168

E UPHORBIACEAE

Fig. 150. Mabea arenicola

Fig. 151. Mabea nitida

Fig. 152. Mabea montana

Mabea 169

Fig. 153. Mabea frutescens

Fig. 154. Mabea linearifolia

Fig. 155. Mabea piriri

170

E UPHORBIACEAE

Fig. 156. Mabea subsessilis

Fig. 157. Mabea taquari

Manihot 171

Liana to 25 m tall. Nonflooded evergreen lowland to lower montane forests, occasionally in riparian forests, 300–800 m; Bolívar (Gran Sabana, upper Río Paragua). Amazon Basin, not yet known from Colombia. Mabea subsessilis Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 282. 1912. —Cachimpapudek (Pemón), Kashimujuru (Yekwana), Pandara-yek (Arekuna), Shakuru (Yekwana). Mabea argutissima Croizat, Bull. Torrey Bot. Club 67: 288. 1940. Shrub or tree to 15 m tall. Evergreen lowland to montane forests, secondary growth, 100–1500 m; Bolívar (Gran Sabana, upper and middle Río Paragua, Roraima-tepui), Amazonas (Río Casiquiare, Río Cunucunuma). Amazonian Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 156. Mabea taquari Aubl., Hist. Pl. Guiane 870, t. 334, fig. 2. 1775. Mabea schomburgkii Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 365. 1854. Mabea taquari var. angustifolia Müll. Arg. in A. DC., Prodr. 15(2): 1150. 1866. Shrub or tree to 8 m tall, sometimes climbing. Riverbanks, sandy beaches, forests, scrubland and granitic outcrops near rivers, 100–900 m; Delta Amacuro (Serranía de Imataca), Bolívar (Río Caura near Salto

Pará, Río Kukenán, Serranía de Imataca, near Urimán), Amazonas (Río Yagua). Monagas; Trinidad, Guyana, eastern Amazon Basin. ŠFig. 157. Mabea trianae Pax, Bot. Jahrb. Syst. 26: 506. 1899. —Boborutepe (Warekena), Duara blanca, Guachimaca, Reventillo. Mabea parvifolia Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 282. 1912. Shrub or tree to 9 m tall. Wooded river banks, gallery forests, granitic outcrops, 50– 300 m; Bolívar (Cerro Guaiquinima, Río Caura, 36 km east of Río Cuchivero, Río Orinoco, Río Paragua, Río Parguaza), Amazonas (Río Orinoco, Río Parucito, Río Ventuari). Llanos of Venezuela and Colombia. Glands at the basal margins of the inner sepals of pistillate flowers are much more common than in all other species in the flora area but are not a constant character. Mabea sp. A Small shrub to 2.5 m tall. Scrub savannas on quartzitic substrate, along rivulets, 1100– 1200 m; Amazonas (Cerro Parú). Endemic. Mabea sp. B Shrub to 3 m tall. Monate forests, ca. 1000 m; Bolívar (expected). Guyana (known from base of Mount Roraima).

35. MANIHOT Mill., Gard. Dict. Abr. ed. 4. 1754. by Grady L. Webster and Paul E. Berry Subshrubs, shrubs, trees, or clambering vines, monoecious or less often dioecious; stems usually with white latex; roots often with tubers. Leaves alternate, simple, though often deeply lobed, petiolate to subsessile, blades with stipels at the base; margins entire to serrate or lobed, glabrous to pubescent, sometimes the lower surface waxy-glaucous; stipules small, deciduous. Inflorescences terminal or pseudoaxillary, solitary or several, racemose or paniculate, usually bisexual, flowers apetalous, usually pedicellate. Staminate flowers with petaloid calyx usually connate in lower part to form a tube, lobes 5, imbricate, disk large and intrastaminal with 5 bifid lobes; stamens 10 in 2 unequal series; anthers introrse; pistillode absent or minute. Pistillate flowers with petaloid calyx, usually 5 sepals united near the base, disk thick and fleshy, subtending the ovary, staminodes absent; ovary 3-locular, ovules 1 per locule; styles 3, short, united at the base, style branches broadly dilated and multilobed. Fruits capsular, smooth or with longitudinal wings, breaking into three 2-valved cocci, columella usually persistent. Seeds smooth, with thincrustaceous testa, carunculate, endosperm copious.

172

E UPHORBIACEAE

Southwestern U.S.A. (Arizona), Mexico, Central America, West Indies, South America except Chile; ca. 100 species, 8 in Venezuela, 6 of these in the flora area. Despite a 1973 monograph of the genus (D. J. Rogers & S. G. Appan. Flora Neotropica Monograph 13: 1–272. 1973), the circumscription of many species is unclear, and more taxonomic work will be required to better resolve the species. Key to the Species of Manihot 1. 1. 2(1). 2. 3(2). 3. 4(1).

4. 5(4). 5.

Leaves 3- or 5-lobed, the area between lobes < 5 mm across at the base ................................................................................................................ 2 Leaves 3- or 5-lobed or simple, when lobed, the area between lobes > 5 mm across at the base ................................................................................... 4 Vines or scandent shrubs; leaves always 3-lobed, not peltate .................... ............................................................................................... M. brachyloba Shrubs, usually not climbing; leaves 3- or 5-lobed, usually narrowly peltate at the base .................................................................................. 3 Leaf lobes linear-lanceolate, < 1 cm wide ............................. M. surinamensis Leaf lobes elliptic to oblong, usually > 1 cm wide ............................ M. tristis Stems distinctly enlarged at the nodes; staminate calyx pubescent within; fruits with distinct wings; widely cultivated, occasionally locally escaped ......................................................................................... M. esculenta Stems not distinctly enlarged at the nodes; staminate calyx glabrous within; fruits without distinct wings; wild plants ................................ 5 Leaves 3-lobed, the lobes entire, never pandurate; flowers in racemes; staminate pedicels < 1 cm long ............................................... M. anomala Leaves (3)5-lobed, the lobes often pandurate; flowers in panicles; staminate pedicels 1–2 cm long ............................................... M. carthaginensis

Manihot anomala Pohl, Pl. Bras. Icon. Descr. 1: 27, t. 21. 1827. Shrubs with white latex. Venezuela, Peru, Brazil, Bolivia, Paraguay, northern Argentina; 5 subspecies, 1 in Venezuela.

Araguao), Amazonas (southeast of Puerto Ayacucho, Río Casiquiare). Táchira, Zulia; Costa Rica, Hispaniola, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

M. anomala subsp. pubescens D.J. Rogers & Appan, Fl. Neotrop. Monogr. 13: 167, fig. 74D, 75A. Shrub ca. 2 m tall. Dry forests along stream on granitic outcrops, 100–300 m; Bolívar (km 103 on road between Caicara and Puerto Ayacucho, Serranía Cerbatana). Brazil (Goias, Minas Gerais, Pará).

Manihot carthaginensis (Jacq.) Müll. Arg. in A. DC., Prodr. 15(2): 1073. 1866. —Jatropha carthaginensis Jacq., Select. Stirp. Amer. Hist. 256, t. 162, fig. 1. 1763. Shrub or small tree. Semideciduous forests, 100–200 m; Bolívar (Ciudad Piar). Widespread in dry areas of northern Venezuela. Bonaire, Colombia, Trinidad. Brazil.

Manihot brachyloba Müll. Arg. in Mart., Fl. Bras. 11: 451. 1874. —Jo-to-rocoanejóro (Warao). Manihot rusbyi Britton, Bull. Torrey Bot. Club 28: 302. 1901. Vine or scandent shrub to 5 m tall. Gallery forests, evergreen lowland forests, near sea level to 100 m; Delta Amacuro (Caño

Manihot esculenta Crantz, Inst. Rei Herb. 1: 167. 1766. —Yuca, Yuca amarga, Yuca dulce. Jatropha manihot L., Sp. Pl. 1007. 1753. Manihot diffusa Pohl, Pl. Bras. Icon. Descr. 1: 55. 1827. Manihot utilissima Pohl, Pl. Bras. Icon. Descr. 1: 32. 1827.

Manihot 173

Large herb or shrub 1–4 m tall; leaves deeply lobed. Mainly cultivated in fields and clearings, near sea level to 1000 m; widespread throughout the flora area. Probably originated in Brazil, now widely cultivated in the rest of lowland Venezuela and throughout the tropics. Manihot esculenta is the main staple of many of the Amerindians of southern Venezuela. There are many varieties and indigenous names that are not listed here. Manihot surinamensis D.J. Rogers & Appan, Fl. Neotrop. Monogr. 13: 80. 1973. Shrub ca. 1 m tall. Shrubby savannas, 100–500 m; Bolívar (trail from Río Ambutuir to Urimán, Río Trueno north of Cerro Guaiquinima), Amazonas (lower Río Ventuari). Guyana, Suriname, French Guiana.

Manihot tristis Müll. Arg. in Mart., Fl. Bras. 11(2): 449. 1874. Shrubs. Venezuela, Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas); 3 subspecies, 2 in Venezuela, both in the flora area. Key to the Subspecies of M. tristis 1. Base of leaf lobes very narrow, < 3 mm wide; petiole attachment basal ............... ....................................... subsp. tristis 1. Base of leaf lobes > 3 mm wide; petiole attachment peltate ....... subsp. saxicola M. tristis subsp. saxicola D.J. Rogers & Appan, Fl. Neotrop. Monogr. 13: 84, figs. 31C, 32A–C. 1973. Manihot maguireana D.J. Rogers & Appan, Fl. Neotrop. Monogr. 13: 161; 24: 169, figs. 71D, 72 A–B. 1973.

Fig. 158. Manihot tristis subsp. saxicola

174

E UPHORBIACEAE

Shrub 1–2 m tall. Savannas and forest edges, 300–500 m; Bolívar (Canaima, La Vergareña, upper Río Paragua, Urimán). Apure. ŠFig. 158. M. tristis subsp. tristis Manihot orinocensis Croizat, J. Arnold Ar-

bor. 24: 169. 1943. Shrub 1–4 m tall; perianth magenta and yellow. Semideciduous forests, shrub patches on granitic outcrops, savannas, 50–300(– 800) m; northern Bolívar, Amazonas (southwest of La Reforma, Puerto Ayacucho area, Río Cuao). Apure; adjacent Colombia.

36. MAPROUNEA Aubl., Hist. Pl. Guiane 895. 1775. by Paul E. Berry and Hans-Joachim Esser Monoecious or less often dioecious shrubs or trees, glabrous throughout, with scanty white latex. Leaves alternate, simple, petiolate, blades entire, pinnately veined, glandular on the lower surface (rarely eglandular); stipules small, persistent. Inflorescences terminal, usually bisexual, of 1–4 solitary, pedicellate, pistillate flowers at basal nodes, the staminate flowers densely aggregated in a strobiform mass at the end of the rachis, separated from the pistillate part by an elongated internode; bracts biglandular. Staminate flowers mostly 3 per bract; pedicel very short; calyx 2(3)-lobed, the lobes imbricate; petals and disk absent; stamens usually 2; filaments completely connate into a slender tube exserted from the calyx at anthesis; anthers apiculate, extrose and longitudinally dehiscent. Pistillate flowers solitary to each bract; calyx 3-parted, imbricate; petals and disk absent; carpels 3, each with 1 ovule; styles usually connate into a column (but free in M. amazonica), the

Fig. 159. Maprounea guianensis

Margaritaria 175

tips unbranched. Fruits capsular; columella weakly persistent. Seeds with large caruncle partly covering top of seed, the surface deeply crateriform-foveolate (in the Venezuelan taxa). Tropical America and Africa; 5 species, 2 in Venezuela, both in the flora area. Key to the Species of Maprounea 1.

1.

Leaves apically obtuse to rounded or emarginate, abaxially eglandular at the base; 1 or 0 pistillate flowers per inflorescence; staminate inflorescences at least as wide as long; fruits 8–10 mm long .......... M. amazonica Leaves apically acute to acuminate, abaxially with 0, 1, or 2 pairs of basal glands; pistillate flowers (1) 2 or 3 per inflorescence; staminate inflorescence oblong-elliptic in outline, longer than wide; fruits 4–6(–8) mm long ........................................................................................ M. guianensis

Maprounea amazonica Esser, Novon 9: 32. 1999. Maprounea guianensis var. obtusata (Müll. Arg.) Müll. Arg. in Mart., Fl. Bras. 11(2): 543. 1874. —Maprounea guianensis var. guianensis forma obtusata Müll. Arg., Linnaea 32: 115. 1863. Shrub or tree to 10 m tall; leaves ovate to orbicular. Seasonally flooded black-water riverbanks, white-sand caatinga, 100–200 m; Amazonas (Isla Ratón, Río Sipapo, Río Temi, near Samariapo). Colombia (Vichada), Brazil (Amazonas).

Maprounea guianensis Aubl., Hist. Pl. Guiane 895, t. 342. 1775. Tree 3–10 m tall; leaves elliptic to ovate. Evergreen lowland to lower montane forests, often riparian or disturbed habitats, near sea level to 900 m; Delta Amacuro (Jotajana), Bolívar (scattered), Amazonas (widespread). Widespread in northern Venezuela; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 159.

37. MARGARITARIA L. f., Suppl. Pl. 66. 1781 [1782]. by Grady L. Webster and Paul E. Berry Dioecious shrubs or trees; stems without latex. Leaves alternate, entire, deciduous, stipulate, pinnately veined, eglandular, petiolate. Flowers apetalous, axillary or on short shoots; staminate flowers in clusters; pistillate flowers 1–5 per node. Staminate sepals 4, 2-seriate; disk annular; stamens 4; filaments free; pistillode absent. Pistillate calyx and disk as in staminate flowers; ovary 2–6-locular; styles free or nearly so, bifid; ovules 2 per locule. Fruit irregularly dehiscent; endocarp thin and papery. Seeds 2 per locule, outer seed coat fleshy and bluish, inner coat thick and bony. Pantropics except for the Pacific islands, 14 species, 1 in Venezuela. Margaritaria nobilis L. f., Suppl. Pl. 66. 1781 [1782]. —Phyllanthus nobilis (L. f.) Müll. Arg. in A. DC., Prodr. 15(2): 414. 1866. —Ojo de grulla. Semideciduous tree 5–12 m tall; branches lenticellate; seeds covered by a thin, bluish, fleshy layer. Evergreen or semideciduous lowland to lower montane forests, riparian forests, near sea level to 900 m; Delta Amacuro (northeast of El Palmar, between

Macareo and Río Grande), Bolívar (Represa Guri, Río Icutu, Río Orinoco near Angosturita, middle Río Paragua), Amazonas (Samariapo, 43 km northeast of Santa Bárbara, Sierra Parima). Widespread in northern Venezuela; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 160.

176

E UPHORBIACEAE

Fig. 160. Margaritaria nobilis

38. MICRANDRA Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 371. 1854. Cunuria Baill., Adansonia 4: 287. 1864. by Paul E. Berry Monoecious trees with white latex, often with buttress roots. Leaves alternate, simple, entire, pinnately veined, often with a pair of prominent basal glands on the upper surface; stipules usually conspicuous, enveloping the terminal bud, caducous, leaving a prominent scar. Inflorescences in axillary or subterminal panicles, the pistillate flowers usually terminal; flowers apetalous. Staminate flowers with 5 imbricate or valvate sepals, with or without a disk; stamens 5–10; anthers dorsifixed; 3lobed pistillode present. Pistillate flowers with 5 sepals, calyx deeply cup-shaped or more open and shallowly bowl-shaped, disk cup-shaped or annular, lobed or unlobed; ovary 3-locular, 1 ovule per locule; stigmas sessile, 3, each shortly bifid. Fruit a capsule; endocarp woody. Seeds large, ellipsoid, with or without a caruncle.

Micrandra 177

Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; ca. 15 species, 7 in Venezuela, all in the flora area. Key to the Species of Micrandra 1.

1.

2(1).

2.

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

5(4).

5.

6(5). 6.

Upper surface of leaves with faint or no basal glands; young inflorescences and ovaries usually sericeous; inflorescences generally shorter than the leaves; stamens 5 .................................................................... 2 Upper surface of leaves with a prominent pair of slightly raised round glands at the base; young inflorescences and ovaries glabrous to sericeous; inflorescences usually longer than the leaves; stamens 7–10 ................................................................................................................ 4 Leaves generally obovate, 4–10 × 2–4 cm, petioles 1–2.5 cm long, rounded to less often acute at the apex; mature fruit 1–2 × 1–1.5 cm, pedicels 1–2 cm long; trees 5–12 m tall with broad, shallowly fluted trunks, re stricted to seasonally flooded black-water river banks ................. M. sp. A Leaves elliptic or ovate to obovate, generally larger than above, at least in length or width, apex shortly acuminate or rounded; fruits larger than above, pedicels > 3 cm long; trees larger than above, neither fluted nor restricted to black-water river banks .................................................... 3 Leaves narrowly or obovate-oblong, (5–)8–12 × (3–)4–6 cm, apex rounded to shortly acuminate; petioles 2–4 cm long .................................. M. minor Leaves elliptic, (9–)14–22 × 5–10 cm, apex generally short-acuminate; petioles 4–5 cm long ........................................................... M. siphonioides Leaves rounded to subcordate at the base; petioles very thick, 3–4 mm diameter; stamens 10; mature fruits ca. 6 × 3 cm ......................... M. glabra Leaves subcuneate to rounded at the base; petioles more slender than above; stamens 7–10; mature fruits generally smaller or else more globose ......................................................................................................... 5 Leaves firmly coriaceous and strongly revolute-margined, drying glaucous on lower surface, the apex rounded; petioles ca. 1/5 the length of blade; stamens 7 ......................................................................... M. sprucei Leaves chartaceous, the margin flat or only slightly revolute, the apex acute to acuminate; petioles 1/5–1/2 the length of the blade; stamens 8 or 10 ............................................................................................................ 6 Axils of leaf veins glabrous; stamens 10; petioles 1/3–1/2 as long as the blade ................................................................................................ M. spruceana Axils of leaf veins with tufts of hairs; stamens 8; petioles 1/5–1/4 as long as the blade .................................................................................... M. rossiana

Micrandra glabra (R.E. Schult.) R.E. Schult., Bot. Mus. Leafl. 15: 203. 1952. —Cunuria glabra R.E. Schult. ex Baldwin & R.E. Schult., Bot. Mus. Leafl. 12: 339, t. 44. 1947. —Surúba, Suru-wai-yek (Arekuna). Tree to 35 m tall, with buttress roots; leaves coriaceous, large (10–21 × 8–13 cm), petioles short, thick and rigid, veins promi-

nent; fruits large, to 6 × 3 cm. Montane gallery forests, 1100–1400 m; Bolívar (Gran Sabana, Sierra de Lema). Guyana, Suriname. ŠFig. 161. The seeds of Micrandra glabra are edible after cooking. Micrandra minor Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 372. 1854. —Micran-

178

E UPHORBIACEAE

dra siphonioides var. minor (Benth.) Mull. Arg. in A. DC., Prodr. 15(2): 709. 1866. —Kunudi, Kunuri (Yekwana). Tree to 35 m tall; evergreen lowland and riparian forests, lower montane forests on granitic hills, 50–500 m; Bolívar (Río Caura basin 55 km south of Jabillal, Río Nichare basin, Río Yudi), Amazonas (base of Piedra Cocuy, Río Matacuni). Amazonian Colombia and Brazil. ŠFig. 164. Micrandra rossiana R.E. Schult., Bot. Mus. Leafl. 15: 211. 1952. —Aramasa (Curripaco), Cunuri, Cunuri blanco, Katajai (Yekwana). Tree 10–35 m tall, without buttress roots; leaves coriaceous, with 2 prominent glands at base, the base cuneate to occasionally rounded; ovary and young growth sericeous; capsule ellipsoid, ca. 4 × 2 cm, turning yellow at maturity; seeds carunculate. Evergreen lowland to montane forests (usually nonflooded ground), 100–1000 m; Bolívar (Cerro Camarón near Cerro Guaiquinima, Cerro Ichún, Macizo del Chimantá [slopes of Abacapá-tepui and Toronó-tepui], Río Icabarú, mouth of Río Nichare), Amazonas (78 km northeast of Puerto Ayacucho, Río Matacuni, upper Río Siapa, San Carlos de Río Negro, base of Sierra de la Neblina). Apure, Zulia; Colombia, Guyana, Ecuador, Brazil. ŠFig. 162. The seeds of Micrandra rossiana are roasted and eaten by various Amerindian groups. Micrandra siphonioides Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 371. 1854. —Arbol de la vaca, Caucho, Caucho tomoro (Arekuna). Tree 10–30 m tall. Evergreen lowland and flooded riparian to lower montane forests, 50– 500 m; Bolívar (Río Caroní, upper Río Caura, Río Ichún, Río Nichare, Río Paragua to Cerro Guaiquinima, Río Torono, Salto Pará), Amazonas (base of Piedra Cocuy, Río Cuao, Río Metacuni in Río Casiquiare basin). Apure; French Guiana, Peru, Brazil. ŠFig. 165. Micrandra spruceana (Baill.) R.E. Schult., Bot. Mus. Leafl. 15: 217. 1952. —Cunuria spruceana Baill., Adansonia 4: 288. 1864. —Conuri de tierra firme, Conuri propio, Momo (Yanomami).

Micrandra cunuri Baill. in A. DC., Prodr. 15(2): 2. 1866. Tree to 35 m tall, with conspicuous buttress roots; fruits to 6 cm long. Evergreen lowland forests on ultisols, 100–200 m; Amazonas (Maroa to Yavita, Piedra Cocuy, Río Casiquiare, Río Guainía, upper Río Ocamo). Amazonian Colombia, Peru, and Brazil. The seeds of Micrandra spruceana are sometimes eaten by local inhabitants (after boiling and soaking in water to detoxify them). Micrandra sprucei (Müll. Arg.) R.E. Schult., Bot. Mus. Leafl. 15: 218. 1952. —Clusiophyllum sprucei Müll. Arg., Flora 57: 518. 1864. —Conori, Conuri banero, Cunuri, Temare montañero. Cunuria crassipes Müll. Arg. in Mart., Fl. Bras. 11(2): 510. 1874. Tree to 25 m tall, with buttress or more commonly stilt roots; leaves stiffly coriaceous, drying bicolored (with the upper surface drying a glaucous bluish gray), secondary veins 8–11, fine, tertiary veins parallel, margins often revolute, with 2 prominent glands at the base; stamens 7, with a prominent reddish lobed disk; pistillate calyx strongly cupular; pedicels 3–4 cm long; fruit to 4 cm diameter; seeds ecarunculate. Caatinga forests, forests on white-sand, podzolic soils, 50–200 m; Amazonas (Maroa to Yavita, base of Piedra Cocuy, Río Yatúa, Río Pasimoni, San Carlos de Río Negro, base of Sierra de la Neblina). Amazonian Colombia and Brazil. ŠFig. 163. Micrandra sp. A. —Palo de boya chivudo. Tree 5–12 m tall, the base usually much enlarged, either shallowly fluted or with aerenchymous, short, bristly roots lining the lower trunk; bark rugose, light brown or tan; wood very lightweight; leaves 4–10 × 2–4 cm, obovate-elliptic, mostly rounded to occasionally acute at the tip, eglandular or only faintly glandular at the base of the blade; fruits green, 1–2 × 1–1.5 cm. Along banks of black-water river banks in areas flooded for most of the year, 50–200 m; Amazonas (Caño San Miguel, Caño Yagua, Río Atabapo and Río Temi basins, Río Baría). ŠFig. 166.

Micrandra 179

Fig. 161. Micrandra glabra

Fig. 162. Micrandra rossiana

180

E UPHORBIACEAE

Fig. 163. Micrandra sprucei

Fig. 164. Micrandra minor

Fig. 165. Micrandra siphonioides

Micrandra 181

Pistillate flower

Seed

Staminate flower

Fig. 166. Micrandra sp. A

182

E UPHORBIACEAE

39. MICROSTACHYS A. Juss., Euphorb. Gen. 48, pl. 15, fig. 50. 1824. —Sebastiana sect. Microstachys (A. Juss.) Müll. Arg., DC., Prodr. 15(2): 1166. 1866. Tragiopsis H. Karst. in Wochenschr. Gärtnerei Pflanzenk. 2: 5. 1859. by Hans-Joachim Esser and Paul E. Berry Monoecious herbs or small shrubs; indument consisting of multicellular trichomes. Leaves alternate; margins densely serrate (teeth < 0.5 mm apart) with sharp or peculiarly enlarged, persistent teeth, sometimes entire but then margin distinctly prominent (by fusion of the teeth?). Staminate inflorescences spiciform, leaf-opposed racemes; pistillate flowers supra-axillary. Staminate flowers with 3parted calyx, subtended by small serrate bracts, corolla and disk lacking; stamens 3; anthers globose, extrorse; pistillode absent. Pistillate flowers with reflexed, deeply 3-parted style; locules 3, 1 ovule per locule; ovaries and fruits with longitudinal rows of multiple (> 3 pairs) excrescences, rarely smooth. Fruit a 3-parted capsule breaking into 2-valved cocci; columella persistent. Seeds oblongoid, with a large caruncule. Pantropics; at least 13 or 14 species, 1 in Venezuela. Microstachys chamaelea (L.) Müll. Arg. is found in Asia, Africa, and Australia, two or three additional local endemics are found in Africa, and at least ten additional species in South America, most of them local endemics in Brazil. The genus is much in need of revision.

Fig. 167. Microstachys corniculata

Omphalea

183

Microstachys has often been included in Sebastiania Spreng., but that genus consists of usually glabrous shrubs or trees with the following differences: leaf margins usually serrate with shallow, inconspicuous, and caducous glandular teeth (> 1 mm apart); both sexes in same inflorescence, these regularly terminal and axillary; ovaries and fruits smooth; and seeds ellipsoid, with a small caruncle that separates from the seed. Microstachys corniculata (Vahl) Griseb., Fl. Brit. W. I. 49. 1854. —Tragia corniculata Vahl, Eclog. Amer. 2: 55, pl. 19. 1798. —Sebastiania corniculata (Vahl) Müll. Arg. in A. DC., Prodr. 15(2): 1168. 1866. —Palo de sardina, Yuquilla. Tragiopsis fruticulosa H. Karst in Wochenschr. Gärtnerei Pflanzenk. 2: 5. 1859. Weedy herb; leaves lanceolate to lanceovate, rounded at the base, hirsutulous; staminate racemes filiform, nearly as long as

the opposite petiole, pistillate flowers borne on the stems between the nodes, shortly pedicellate; fruits with triangular spur-like projections. Edges of Mauritia palm swamps, dry savannas, deciduous forests, granitic outcrops, disturbed areas, open river banks, 50–400 m; widespread in northern Bolívar and Amazonas. Widespread in the Llanos and scattered in the rest of Venezuela; Costa Rica, Panama, West Indies, Guyana, Suriname, French Guiana, Brazil, Paraguay. ŠFig. 167.

40. OMPHALEA L., Syst. Nat. ed. 10, 2: 1264. 1759. by Lynn J. Gillespie Monoecious trees, shrubs, or lianas, with red or orange-red latex (sometimes scant). Leaves alternate, simple, stipulate, petiolate; blades unlobed or lobed, pinnately or palmately veined; margins entire, biglandular at petiole apex or blade base; laminar glands usually present on abaxial blade surface. Inflorescences terminal, thyrsoid but often appearing spicate, racemose, or paniculate, bisexual; cymules highly condensed to lax, bisexual with central flower(s) pistillate or distally unisexual and staminate; bracts subtending cymules large, foliose, green or colored, often biglandular; bracteoles small, triangular. Staminate flowers subsessile or pedicellate: sepals 4 or 5, imbricate; petals absent; disk extrastaminal, annular, rarely 5segmented or obsolete; androecium mushroom-shaped with massive hemispheroidal cap formed from highly expanded connate anther connectives, or rarely anthers free and connectives not expanded; filaments entirely connate into a slender staminal column, or rarely anthers subsessile; anthers 2 or 3; pistillode absent. Pistillate flowers subsessile or pedicellate: sepals 4 or 5, imbricate; petals and disk absent; ovary 3locular, locules uniovulate; styles entirely connate, often massive. Fruit a large, 3seeded, schizocarpous capsule or berry, 3-lobed or subglobose, sometimes with conical beak; pericarp woody or fleshy. Seeds large, ellipsoid, ovoid, or globose, ecarunculate. Neotropics, Australasia, Madagascar, Tanzania; 17 species, 1 in Venezuela. Omphalea diandra L., Syst. Nat. ed. 10, 2: 1264. 1759. Omphalea megacarpa Hemsl. in Hook., Icon. Pl. 4: pl. 2537. 1897. Omphalea elaeophoroides Steyerm. ex Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 152. 1940. Liana climbing to 30 m or more in forest canopy; leaves to 27 × 16 cm; inflorescence paniculate, 15–70 cm long; fruit subglobose,

to 12 cm diameter. Moist to wet lowland forests, along rivers, near sea level to 100 m; Delta Amacuro (Caño Araguaito above Los Rastrojos, Caño Araguao, Río San José). Monagas; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 168. Young individuals are vine-like, climbing by means of terminal tendrillate shoots. The seeds are reported to be edible, but may be mildly toxic.

184

E UPHORBIACEAE

Fig. 168. Omphalea diandra

Pausandra 185

41. PAUSANDRA Radlk., Flora 53: 92. 1870. by Ricardo de S. Secco Dioecious trees, rarely shrubs. Leaves simple, alternate, stipulate, chartaceous, subcoriaceous to coriaceous, slightly pilose on the lower surface or hirsute on both surfaces; margin serrulate, apex acuminate; petiole thickened at apex, with a pair of glands at the petiole-leaf blade junction. Inflorescences spicate thyrses, axillary, subaxillary, or terminal, glomerules few-flowered, rarely densely flowered; flowers subsessile to short-pedicellate, the pistillate ones solitary. Staminate flowers small to medium-sized; calyx lobes 2–5, unequal, imbricate, pilose, rarely glabrous; corolla 5-lobed, the lobes slightly emarginate or revolute, frequently connate to above the middle or only at the base, inner surface pilose, rarely glabrous; stamens 6 or 7, epipetalous. Pistillate flowers larger; calyx 2–5-lobed, pilose; petals 5, free or connate to above the middle, inner surface pilose or glabrous; disk flat or undulate, glabrous; ovary 3-locular, pilose, 1 ovule per locule; styles 3, bilobed. Fruit a tricoccous capsule. Seeds oblong to obovate, the surface marbled, blackish or brownish, carunculate. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonia and southeastern areas), Bolivia; 6 species, 1 in Venezuela.

Fig. 169. Pausandra martinii

186

E UPHORBIACEAE

Pausandra martinii Baill., Adansonia 11: 92. 1873. —Mangle, Manglillo, Redetu koru (Piaroa). Pausandra flagellorachis Lanj., Euphorb. Surinam 30. 1931. Shrub or tree 2–20 m tall; corolla creamcolored. Evergreen lowland to lower montane forests, river margins, roadsides, rocky formations, 100–700 m; Delta Amacuro (Río Cuyubini, Serranía de Imataca), Bolívar

(Altiplanicie de Nuria, 85 km south-southeast of Entrerios, Río Cuyuní, Serranía de Imataca), Amazonas (upper Río Cuao, Río Putaco 20 km above junction with Río Ocamo, San Carlos de Río Negro, Serranía Batata 55 km southeast of Puerto Ayacucho, base of Sierra de la Neblina, Sierra Parima, Yavita to Maroa). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amapá, Amazonas, Pará). ŠFig. 169.

42. PEDILANTHUS Neck., Elem. Bot. 2: 354. 1790. by Paul E. Berry Monoecious shrubs with woody or succulent greenish stems, usually with few distal lateral branches, with white latex. Leaves alternate, distichous, simple, sessile or petiolate; blades fleshy; margins entire; stipules small. Inflorescences axillary or terminal cymes of few to many cyathia, these bilaterally symmetrical, slipperlike, partly or entirely reddish. Staminate flowers numerous, each consisting of a single pedicellate stamen; anthers 2-lobed. Pistillate flower consisting of a single pedicellate 3-locular ovary, ovules single in each locule; styles 3, connate for most of their length, distally bifid. Fruits capsules or indehiscent, 3-lobed. Seeds ovoid or rounded-angular, smooth or tuberculate, ecarunculate. Southern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil; 15 species, 1 in Venezuela. Pedilanthus tithymaloides (L.) Poit., Ann. Mus. Natl. Hist. Nat. 19: 390. 1812. —Euphorbia tithymaloides L., Sp. Pl. 453. 1753. Semisucculent shrub 0.5–1.5 m tall with milky latex; stems zigzag, often leafless; leaves elliptic-ovate, 3–13 × 1–7 cm; cyathia red. Southern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil; 8 subspecies, 1 in Venezuela. P. tithymaloides subsp. tithymaloides. —Ipecacuana, Ipecacuana de monte. Semideciduous forests, dry slopes, 50–400 m; Bolívar (Altiplanicie de Nuria, Cerro Paja 25 km east of Upata, lower Río Caura). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Nueva Esparta, Sucre, Zulia; Mexico, Central America, Curaçao, Colombia, Trinidad, Guyana, Suriname, Ecuador. ŠFig. 170. The plant is used as a purgative and to induce vomiting and is occasionally cultivated as an ornamental (e.g. Ciudad Bolívar).

Fig. 170. Pedilanthus tithymaloides subsp. tithymaloides

Pera 187

43. PERA Mutis, Bih. Kongl. Svenska Vetensk.-Acad. Handl. 5: 299, t. 8. 1784. by Lynn J. Gillespie Dioecious shrubs or trees. Trichomes lepidote (peltate, scale-like) or stellate, rarely simple. Leaves alternate or rarely opposite, simple, exstipulate, petiolate, eglandular; blades pinnately veined; margins entire. Inflorescence axillary, pseudanthial, unisexual, comprising 3 or 4 flowers subtended by a showy involucral bract and (1)2 bracteoles; pseudanthia pedunculate, fasciculate in axils of older leaves or on short shoots below leaves, often fragrant; involucral bract subglobose, ovoid, or ellipsoid, consisting of 2 connate bracts, completely enclosing flowers in bud and opening along one side at anthesis; staminate involucre sometimes with pistillode(s) representing reduced pistillate flower(s), pistillate involucre sometimes with rudimentary calyx or calyces representing reduced staminate flower(s). Staminate flowers sessile; calyx irregularly lobed or laciniate, often reduced or absent; petals and disk absent; stamens (2)3 or 4; filaments free to partly connate. Pistillate flowers sessile or shortly pedicellate; calyx, corolla, and disk absent; ovary 3-locular, globose or ellipsoid, locules uniovulate; styles absent or very short; stigmas 3, often sessile, connate into a thick-lobed peltate structure. Fruit a 3-seeded schizocarpous capsule, globose or ellipsoid; pericarp woody. Seeds ovoid or ellipsoid, somewhat flattened, smooth, shiny, black or dark brown, carunculate, the caruncle large and usually red or orange. Central America, Bahamas, Greater Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 30 species, 6 in Venezuela, all in the flora area. Key to the Species of Pera 1.

1.

2(1). 2.

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

Leaf blade glabrescent, sometimes with scattered simple trichomes near base on lower surface; involucral bract opening wide at anthesis, subtended by a single bracteole; filaments mostly fused, staminal column 1.5–2.5 mm long ................................................................ P. distichophylla Leaf blade with lepidote or stellate trichomes on lower surface; involucral bract opening by a slit along one side and remaining subglobose at anthesis, subtended by 2 bracteoles; filaments free or fused near base only, staminal column < 1 mm long ...................................................... 2 Lower surface of leaf blade very densely covered with trichomes, the surface not visible between trichomes; filaments slender, > 2 mm long ...... 3 Lower surface of leaf blade sparsely to moderately densely covered with trichomes, the surface clearly visible between trichomes; filaments stout, < 2 mm long ................................................................................. 4 Trichomes lepidote; lower surface of leaf blade and capsule densely covered with flat, scale-like trichomes .......................................... P. decipiens Trichomes stellate; lower surface of leaf blade and capsule appearing densely tomentose .................................................................... P. tomentosa Capsule glabrous, ovary glabrous or covered with flat lepidote trichomes; lower surface of leaf blade and petiole sparsely lepidote, trichomes distinctly scale-like often with an uneven or irregularly fringed margin .................................................................................................... P. glabrata

188

4.

5(4).

5.

E UPHORBIACEAE

Capsule and ovary very densely covered with stellate trichomes and appearing tomentose; lower surface of leaf blade and petiole sparsely to moderately densely stellate-lepidote, trichomes with long radiating arms ........................................................................................................ 5 Leaf blade subcoriaceous, apex rounded or obtuse, sometimes very shortly acuminate, lower surface sparsely to moderately densely stellate-lepidote, midrib lacking tufted stellate trichomes; capsule 1.4–1.8 cm diameter ........................................................................................... P. bicolor Leaf blade membraneous to chartaceous, apex acute or acuminate, lower surface very sparsely to sparsely stellate-lepidote, midrib often with longer tufted stellate trichomes; capsule 1–1.2 cm diameter ........... .................................................................................................. P. citriodora

Pera bicolor (Klotzsch) Müll. Arg. in A. DC., Prodr. 15(2): 1028. 1866. —Peridium bicolor Klotzsch, London J. Bot. 2: 44. 1843. —Cururú luiro, Laurel de sapo. Pera schomburgkiana Müll. Arg. in A. DC., Prodr. 15(2): 1027. 1866. Shrub or tree to 33 m tall; leaf blade obovate or elliptic, 5–11 × 3–6 cm, the upper surface shiny and drying blackish, the lower surface dull and distinctly paler; involucral bract yellow, moderately densely stellate. Savannas and forests on white sand or sandy soils, 100–600 m; Bolívar (base of Cerro Guaiquinima, north edge of Gran Sabana), Amazonas (base of Cerro Yapacana and vicinity, Río Negro area). Guyana, Suriname, French Guiana, Brazil (Amazonas). ŠFig. 172. Pera citriodora Baill., Adansonia 5: 222. 1865. —Majagua verde, Perro de agua banero, Tabaquillo. Tree 2.5–30 m tall; leaf blade elliptic, 6– 9.5 × 2.5–4 cm, the lower surface dull but not distinctly paler; involucral bract yellow, sparsely stellate. Riparian forests, scrubby forests on white sand, 100–200 m; Amazonas (Caño Yagua, near San Carlos de Río Negro). Brazil (Amazonas, Pará). ŠFig. 174. Pera decipiens (Müll. Arg.) Müll. Arg. in A. DC., Prodr. 15(2): 1029. 1866. —Peridium decipiens Müll. Arg., Linnaea 34: 201. 1865. —Peridium bicolor var. nitidum Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 323. 1854. —Pera nitida (Benth.) Jabl., Mem. New York Bot. Gard. 17(1): 148. 1967. —Congrillo blanco, Congrio, Palo candela.

Shrub or tree to 30 m tall; leaf blade elliptic or narrowly elliptic, 6–14 × 3–6 cm, obtuse or acute at apex, subcoriaceous, the lower surface paler; involucral bract creamcolored or pale yellow, often turning pinkish. Savannas, gallery forests, evergreen lowland to montane forests, often on sandy soils, 50– 1200 m; Bolívar (El Cácaro, Gran Sabana, 70 km southeast of La Vergareña, Río Caura, Río Nichare basin, middle Río Paragua), Amazonas (Caño Chimoni in Río Casiquiare basin, between Río Atabapo and La Esmeralda on the upper Río Orinoco, Río Atacavi, near San Carlos de Río Negro). Amazonian Colombia, Guyana, Suriname, Amazonian Peru and Brazil. ŠFig. 175. Pera distichophylla (Mart.) Baill., Étude Euphorb. 434. 1858. —Spixia distichophylla Mart., Flora 24(2) Beibl.: 30. 1841. Shrub or tree to 8 m tall; leaf blade narrowly elliptic, narrowly oblong-elliptic, or lanceolate, 9–19 × 3–7 cm, acuminate or acute at apex, chartaceous; involucral bract white, cream-colored, or yellow; ovary and capsule tomentose. Evergreen lowland and flooded forests, often on sandy soils, 50– 200(–500) m; eastern Bolívar (mouth of Río Cuao, Río Orinoco near La Urbana), eastern Amazonas (Caño San Miguel, Río Atabapo, Río Casiquiare, Río Guainía, Río Negro, Río Orinoco, Yavita to Maroa). Amazonian Colombia and Brazil. ŠFig. 173. Pera glabrata (Schott) Baill., Étude Euphorb. 434. 1858. —Peridium glabratum Schott in Spreng., Syst. Veg. 4(cur. post.): 410. 1827. —Uanden (Arekuna).

Pera 189

Fig. 171. Pera glabrata

Fig. 173. Pera distichophylla

Fig. 172. Pera bicolor

190

E UPHORBIACEAE

Fig. 174. Pera citriodora

Fig. 176. Pera tomentosa

Fig. 175. Pera decipiens

Phyllanthus 191

Peridium ferrugineum Schott in Spreng., Syst. Veg. 4(cur. post.): 410. 1827. —Pera ferruginea (Schott) Müll. Arg. in A. DC., Prodr. 15(2): 1031. 1866. Shrub or tree to 35 m tall; leaf blade elliptic, 5–12 × 2.5–5.5 cm, obtuse to broadly acuminate at apex; chartaceous to subcoriaceous; involucral bract pale yellow or creamcolored, lepidote. Evergreen lowland forests, gallery forests, forest-savanna edges, shrub savannas, often on white sand, 100–1300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Gran Sabana, near La Paragua, 6 km from Maniapure toward Caicara, Serranía de Imataca), Amazonas (base of Cerro Duida, Cerro Mawedi in Río Ocamo basin, Cerro Parú, Macabana–Río Ventuari region, Río Cunucunuma, Santa Bárbara del Orinoco).

Aragua, Táchira, Trujillo, Yaracuy; Trinidad, Guyana, Suriname, French Guiana, Amazonian Peru, Amazonian and eastern Brazil. ŠFig. 171. Pera tomentosa (Benth.) Müll. Arg. in A. DC., Prodr. 15(2): 1028. 1866. —Peridium bicolor var tomentosum Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 323. 1854. Shrub or tree to 20 m tall; leaf blade elliptic or oblong-elliptic, 7–16 × 3.5–7 cm, shortly acuminate at apex, subcoriaceous; involucral bract yellow turning reddish. Evergreen lowland forests on sandy soils, 100– 400 m; Amazonas (Río Cunucunuma basin near base of Cerro Duida and Cerro Huachamacari, Río Putaco). Amazonian Peru, Brazil, and Bolivia. ŠFig. 176.

44. PHYLLANTHUS L., Sp. Pl. 981. 1753. Conami Aubl., Hist. Pl. Guiane 926. 1775. Asterandra Klotzsch, Arch. Naturgesch. 7(1): 200. 1841. by Grady L. Webster Monoecious or less often dioecious trees, shrubs, or herbs, branchlets in some species deciduous and floriferous, glabrous or with indument of unbranched trichomes. Leaves alternate, evergreen or deciduous, entire, pinnately veined, shortpetiolate, stipulate, sometimes with a single conspicuous large gland in the middle or towards the apex of the blade. Flowers mostly in axillary clusters. Sepals mostly 5 or 6, entire, imbricate; petals absent; disk usually present, entire or segmented. Stamens mostly 3–6, free or the filaments connate; anthers dehiscing vertically or horizontally. Carpels 3 (rarely 2 or 4 or more); ovules 2 per locule; styles mostly bifid (sometimes dilated or unlobed). Fruit usually capsular (rarely baccate or drupaceous), usually with 6 ecarunculate seeds. Seeds with dry seed coat; endosperm copious. Pantropics, some warm-temperate regions; ca. 700 species, 46 in Venezuela, 43 of these in the flora area. The majority of species, both herbaceous and woody, have distinctive branching with flowers produced only on leafy deciduous branchlets. Two species, Phyllanthus bolivarensis Steyerm. and P. orinocensis Steyerm., are of uncertain status because of the lack of flowers, and are not included in the key. Phyllanthus acidus (L.) Skeels, whose common name in some areas is Grosella, is cultivated for its fleshy fruits. Key to the Species of Phyllanthus 1. 1.

Herbs, mostly annual or short-lived; stamens 2, 3, or 5; flowers on main axes or only on deciduous branchlets .................................................... 2 Shrubs or trees; stamens 3–6; flowers only on deciduous branchlets .... 13

192

2(1). 2.

E UPHORBIACEAE

Floating aquatic; leaves inflated; seeds smooth .............................. P. fluitans Terrestrial, or, if aquatic, rooted and leaves not inflated; seeds smooth or ornamented ............................................................................................ 3 3(2). Leaves on main axis distichous, not reduced; flowering axes not deciduous ................................................................................................................ 4 3. Leaves on main axis spirally arranged, on upper (distal) portions of axes reduced to stipule-like scales; flowers produced only on lateral, leafy, ± deciduous branchlets .......................................................................... 5 4(3). Stems terete, not winged; staminate and pistillate flowers at the same axils; seeds verruculose ...................................................... P. caroliniensis 4. Stems compressed, narrowly winged; staminate and pistillate flowers at different axils; seeds smooth ......................................... P. hyssopifolioides 5(3). Stamens 5, free; fruiting pedicels capillary, 3–8 mm long; seeds papillose; proximal cymules on branchlet with both staminate and pistillate flowers ................................................................................................. P. tenellus 5. Stamens 2 or 3, free or filaments connate; fruiting pedicels not capillary; seeds verruculose, ribbed, or striate; proximal cymules on branchlet (in monoecious species) staminate .............................................................. 6 6(5). Ovary verruculose; seeds transversely ribbed on back; pistillate flowers sessile, at proximal axils of branchlet, sepals 6; lower surface of leaves hispidulous marginally and submarginally .............................. P. urinaria 6. Ovary smooth; seeds verruculose or longitudinally ribbed or striate on back; pistillate flowers pedicellate, at distal axils of branchlet, sepals 5 or 6; leaves glabrous ........................................................................... 7 7(6). Leaves suborbicular, mostly as wide as or wider than long; pistillate flowers each accompanied by 1 or 2 staminate flowers; sepals 6; seeds 1.2–1.3 mm long, verruculose ................................................ P. orbiculatus 7. Leaves oblong or elliptic to lanceolate, distinctly longer than wide; pistillate flowers each accompanied by 0 or 1 staminate flower; sepals 5 (very rarely 6); seeds ribbed or striate ............................................... 8 8(7). Pistillate flowers each accompanied by 1 staminate flower; fruiting sepals distinctly acute at tip, scarcely over 1 mm long; seeds 0.9–1 mm long, ribbed............................................................................................ P. amarus 8. Pistillate flowers at solitary distal nodes of branchlet, staminate flowers 1–3 at proximal nodes; fruiting sepals obtuse or rounded at tip; seeds 0.8–1.3 mm long, ribbed or striate ........................................................ 9 9(8). Plants dioecious or subdioecious (occasional plants with 1 or 2 flowers of the other sex); anthers dehiscing vertically or nearly so, filaments completely connate; seeds striate, banded; leaves mostly acute .............. 10 9. Plants monoecious, nearly all branchlets with both staminate and pistillate flowers; anthers dehiscing horizontally to slightly obliquely, filaments completely or partially connate; leaves mostly obtuse ............ 11 10(9). Staminate sepals 1.5–2.3 mm long; anther column 0.5 mm high or more; staminate pedicels 0.5–1.5 mm long; seeds 1.8–2 mm long ... P. lindbergii 10. Staminate sepals 0.7–1 mm long; anther column 0.3–0.5 mm high; staminate pedicels 0.2–0.6 mm long; seeds 1.1–1.3 mm long .... P. microphyllus 11(9). Stamens 2; leaves mostly oblong, narrowed to tip, < 3 mm wide; plants 10–20 cm tall ........................................................................... P. minutulus

Phyllanthus 193

11.

12(11).

12.

13(1). 13. 14(13). 14. 15(13). 15.

16(15).

16. 17(16).

17.

18(17). 18. 19(17). 19. 20(19). 20. 21(19). 21.

Stamens 3 (sometimes 1 reduced, rarely 2); leaves mostly obtuse to rounded (sometimes apiculate) at tip, mostly > 3 mm wide; plants often exceeding 20 cm tall ............................................................................. 12 Stems often with conspicuous aerenchyma at base; leaf margins often reddish; pistillate disk 5-angled or -lobed; filaments completely connate ........................................................................................... P. stipulatus Stems without basal aerenchyma; leaf margins never reddish; pistillate disk asymmetrically 3-lobed; filaments connate in lower half ............... ................................................................................................... P. caribaeus Branchlets bipinnatiform (branchlet axis with 1–several lateral branches) .............................................................................................................. 14 Branchlets pinnatiform (branchlet axis lacking lateral branches) ........ 15 Branchlets with 1–4 lateral axes, glabrous; leaves obtuse to acute, glabrous; ovary hirtellous ...........................................................P. brasiliensis Branchlets with 10–20 lateral axes, hirtellous; branchlets and leaves usually copiously hirtellous; ovary glabrous ............................... P. piscatorum Dioecious trees, cauliflorous or ramiflorous; sepals 4; stamens 4, filaments free; disk absent; fruits indehiscent .................................... P. elsiae Monoecious shrubs or small trees (dioecious shrubs in P. attenuatus), not cauliflorous; sepals and stamens 2–6, filaments free or connate; disk present; fruits dehiscent (except in P. attenuatus) ............................. 16 Leaves 1–5(–6) mm wide, without a gland at the apex or on lower surface near (< 0.5 mm from) the tip (glands, if present, very small, not over 0.2 mm across); anthers generally muticous, often emarginate; styles free or nearly so, bifid or dilated at the tip ......................................... 17 Leaves > 5 mm wide, laminar gland on lower surface present or absent; anthers muticous or apiculate; styles bifid or unlobed ...................... 23 Leaves distinctly falcate, with subapical laminar glands on lower surface 0.1–0.2 mm across; staminate disk of 3 segments; filaments free, anthers apiculate; styles connate, unlobed ............................................. 18 Leaves not falcate, laminar glands obscure or absent; staminate disk of 3 or 6 segments; filaments free or connate; styles free or connate, bifid or unlobed ............................................................................................. 19 Leaves mostly 1–2 mm wide, over 40 per branchlet; seeds 2.3–2.7 mm long ........................................................................................... P. obfalcatus Leaves well over 2 mm wide, 20–35 per branchlet; seeds ca. 3 mm long ............................................................................................. P. pycnophyllus Styles unlobed, ± connate; anthers apiculate; ovary sometimes stipitate .............................................................................................................. 20 Styles bifid, free; anthers emarginate; ovary not stipitate ..................... 21 Leaf blades obcuneate, emarginate at apex; branchlets with 5–15 leaves; staminate pedicel ≤ 1 mm long; styles stigmatiform ................. P. carrenoi Leaf blades oblong to linear, obtuse at apex; branchlets with 20–50 leaves; staminate pedicel 1.5–4 mm long; styles 1–3 mm long ........ P. subapicalis Filaments connate; staminate disk segments 3; fruiting pedicel ca. 2 mm long; leaves 4–5 mm long ..................................................... P. minutifolius Filaments free; staminate disk segments 6; fruiting pedicel 6–10 mm long .............................................................................................................. 22

194

E UPHORBIACEAE

22(21). Leaf blades linear-lanceolate, 15–25 mm long; staminate pedicels 4–5 mm long; pistillate pedicels 8–10 mm long ......................................P. maguirei 22. Leaf blades narrowly elliptic, 3–6 mm long; staminate pedicels 1–2 mm long; pistillate pedicels 5–6 mm long ........................................ P. tepuicola 23(16). Leaves without laminar glands on lower surface; anthers muticous, sometimes distinctly emarginate ................................................................. 24 23. Leaves with prominent laminar glands on lower surface (mostly at least 0.3 mm across); anthers mostly apiculate ........................................... 31 24(23). Styles connate below, tips dilated (not clearly bifid); sepals 5; staminate disk massive, 5-angled, pitted; stamens 5–7, filaments connate, anthers horizontal; branchlets 50–100 cm long; seeds 4.5–5.5 mm long ........................................................................................... P. juglandifolius 24. Styles free or connate, bifid, tips slender; sepals usually 6; staminate disk tenuous; stamens 3, filaments free or connate; branchlets < 50 cm long; seeds < 4 mm long ................................................................................ 25 25(24). Plants dioecious; leaves cuspidate-acuminate, chartaceous; staminate disk not dissected; filaments connate below; fruits slightly fleshy, indehiscent ..................................................................................... P. attenuatus 25. Plants monoecious; leaves rounded to obtuse or emarginate, membranous to coriaceous; staminate disk entire to dissected; filaments distinct or connate; fruits dehiscent ..................................................................... 26 26(25). Leaves coriaceous or subcoriaceous, 4–8 cm long; branchlets 1–1.5 mm thick, pilose, with 3–7 leaves; filaments distinct; pistillate pedicels pilose ................................................................................... P. atabapoensis 26. Leaves chartaceous, mostly 2–4 cm long; branchlets < 1 mm thick, glabrous or pilose, with 3–15 leaves; pistillate pedicels glabrous ........... 27 27(26). Branchlets pilose (at least when young), with mostly 10–12 leaves; fruiting pedicel < 20 mm long; pistillate calyx pilose; filaments connate ................................................................................................... P. paezensis 27. Branchlets glabrous, with 3–15 leaves; fruiting pedicel 3–30 mm long; filaments distinct ................................................................................. 28 28(27). Leaves membranous or chartaceous; branchlets slender (ca. 0.5 mm thick or less), terete, smooth ......................................................................... 29 28. Leaves coriaceous or subcoriaceous; branchlets mostly 1 mm thick or more, often compressed, angled, or ridged ......................................... 30 29(28). Branchlets with only 3–5 leaves; leaf blades mostly obovate, obtuse to rounded at tips; staminate flowers in few-flowered clusters; fruiting pedicel capillary, 25–30 mm long .............................................. P. rupestris 29. Branchlets with 10–15 leaves; leaf blades suborbicular, rounded to retuse at tip; staminate flowers in dense many-flowered clusters; fruiting pedicel < 20 mm long ............................................................... P. borjaensis 30(28). Leaves obovate, cuneate at base; fruiting pedicel 3–7 mm long ................ ........................................................................................... P. jablonskianus 30. Leaves ± orbicular, rounded at base; fruiting pedicel 14–20 mm long .................................................................................................... P. neblinae 31(23). Styles free except at base; leaves narrowly elliptic, rigid, prominently revolute, laminar gland subapical; seeds 3.5 mm long; branchlets with 4–8 leaves ................................................................................. P. lediformis

Phyllanthus 195

31.

32(31). 32. 33(32). 33. 34(32). 34. 35(34).

35.

36(34).

36.

37(36). 37.

38(37). 38. 39(38). 39. 40(39). 40.

Styles connate into a column; leaves obovate to orbicular, margins plane or revolute, laminar gland usually 1 mm or more from the tip; branchlets with 5–20 leaves ................................................................ 32 Filaments connate at least halfway; staminate pedicels 6–8 mm long; styles 1.5–2.5 mm long ........................................................................ 33 Filaments free; staminate pedicels mostly < 6 mm long; styles 0.5 mm long or more ......................................................................................... 34 Ovary sessile; styles completely connate; leaves rounded at tip ................ ................................................................................................. P. longistylus Ovary stipitate; styles free at tips; leaves ± emarginate at tip .... P. chimantae Pistillate pedicel < 1 cm long; styles unlobed .......................................... 35 Pistillate pedicel > 1 cm long; styles bifid or unlobed ............................. 36 Stylar column 1–1.3 mm long; leaves ≤ 1 cm wide, chartaceous; branchlets terete, mostly with > 10 leaves; fruiting pedicels 3–9 mm long ............................................................................................ P. vacciniifolius Stylar column < 1 mm long; leaves often > 1 cm wide, coriaceous or subcoriaceous; branchlets angled, mostly with ≤ 10 leaves; fruiting pedicels 1–2.5 mm long ................................................................... P. major Leaves obovate, bright green with whitish midrib; fruiting pedicel slender, 20–25 mm long; styles reduced to stigmas capping the ovary; staminate disk of 6 segments ............................................... P. strobilaceus Leaves obovate to broadly obovate or orbicular, not bright green with whitish midrib; fruiting pedicel mostly shorter than above; styles present; staminate disk of 3 segments ................................................ 37 Leaves obovate, shiny on the upper surface, cuneate at base; fruiting pedicels 18–20 mm long; styles bifid, free.............................. P. myrsinites Leaves broadly obovate to orbicular (often wider than long), not shiny on the upper surface, rounded to cordate at base; fruiting pedicels of various lengths; styles unlobed, connate or stigmatiform ........................ 38 Leaves deeply clasping-cordate; fruiting pedicel < 10 mm long ...... P. duidae Leaves shallowly cordate to truncate; fruiting pedicel > 10 mm long ... 39 Leaves opposite on branchlet, emarginate at base; styles ca. 2 mm long ............................................................................................. P. paraqueensis Leaves alternate on branchlet; styles < 1 mm long ................................ 40 Leaves emarginate at apex, obtuse or rounded at base; styles 0.3–0.5 mm long; filaments > 2 mm long ..................................................... P. jauaensis Leaves rounded at apex, cordate at base; styles stigma-like; filaments < 0.5 mm long ............................................................................ P. ventuarii

Phyllanthus amarus Schumach., Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Afh. 4: 195. 1829. Herb; branchlets with 15–30 leaves. Disturbed areas, near sea level to 500 m; Delta Amacuro (Pedernales), Bolívar (Paramichi), Amazonas (near Puerto Ayacucho). Widespread in lowland Venezuela and New World and Old World tropics.

Phyllanthus atabapoensis Jabl., Mem. New York Bot. Gard. 17(1): 101. 1967. Shrub 1–3 m tall; leaves coriaceous, mostly 4–8 cm long. Along streams in savannas or seasonally flooded forests, 50–200 m; Amazonas (Caño Cotúa, Caño Momoni in Río Casiquiare basin, base of Cerro Yapacana, Río Atabapo, Río Pasimoni, Río Sipapo); Colombia (Guainía), Brazil (Amazonas). ŠFig. 179.

196

E UPHORBIACEAE

Phyllanthus attenuatus Miq., Linnaea 21: 479. 1848. Dioecious shrub or tree 3–12 m tall; leaves acuminate, mostly 5–8 cm long. Riparian or flooded forests, 50–500 m; Amazonas (Caño Yureba, La Esmeralda, near Raudal de Maypures, San Juan de Manapiare), Bolívar (Canaima). Barinas, Mérida, Táchira; Colombia, Guyana, Suriname, Peru, Brazil. ŠFig. 182.

Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil.

Phyllanthus bolivarensis Steyerm., Fieldiana, Bot. 28: 317. 1952. Shrub 1–2 m tall. Semideciduous forests, 50–300 m; Bolívar (1–10 km northwest of Upata on road to San Félix). Endemic. The affinities of Phyllanthus bolivarensis are uncertain, as flowers are unknown; it may prove not even to be a species of Phyllanthus.

Key to the Subspecies of P. caroliniensis 1. Stipules 0.8–1.2 mm long; leaves mostly rounded at tip .... subsp. caroliniensis 1. Stipules 1.5–2 mm long; leaves mostly obtusely pointed at tip ................................. ............................... subsp. guianensis

Phyllanthus borjaensis Jabl., Mem. New York Bot. Gard. 17(1): 108. 1967. Shrub 1–3 m tall; branchlets slender, with 8–15 rounded chartaceous leaves. Semideciduous forests, 100–200 m; Bolívar (Cerro San Borja on middle Río Orinoco), Amazonas (Puerto Ayacucho). Apure; Colombia (Guainía).

P.

Phyllanthus brasiliensis (Aubl.) Poir. in Lam., Encycl. 5: 296. 1804. —Conami brasiliensis Aubl., Hist. Pl. Guiane 927. 1775. —Phyllanthus conami Sw., Prodr. 28. 1788. —Barbasco, Caicareño. Shrub to 3 m tall; branchlets ca. 20 cm long, with 2–4 lateral axes; leaves ovate, obtuse or apiculate, glabrous, 2–3.5 cm long. Semideciduous forests, 100–200 m; Bolívar (El Tigre). Guyana, Suriname, French Guiana, northern Brazil. This species has been erroneously reported from elsewhere in South America (usually based on confusion with Phyllanthus piscatorum H.B.K.). Phyllanthus caribaeus Urb., Symb. Antill. 5: 382. 1908. Herb; branchlets with 15–30 leaves. Evergreen lowland forests, near sea level to 400 m; Delta Amacuro (Río Toro), Bolívar (Serranía de Imataca). Portuguesa, Zulia; West

Phyllanthus caroliniensis Walter, Fl. Carol. 228. 1788. Herb 10–50 cm tall; leaves distichous, flowers on all axes. U.S.A., Central America, West Indies, South America (except Chile); 4 subspecies, 3 in Venezuela, 2 of these in the flora area.

P. caroliniensis subsp. caroliniensis 0–100 m; Delta Amacuro (Caño Macareo). Aragua, Guárico, Portuguesa, Zulia; other distribution as for the species. caroliniensis subsp. guianensis (Klotzsch) G.L. Webster, Contr. Gray Herb. 176: 46. 1955. —Phyllanthus guianensis Klotzsch, London J. Bot. 2: 51. 1943. Phyllanthus schomburgkianus Müll. Arg. in A. DC., Prodr. 15(2): 387. 1866. 0–300 m; Delta Amacuro (scattered), northern Bolívar. Central America, West Indies, Guyana, Suriname, French Guiana, Brazil. ŠFig. 180. Phyllanthus carrenoi Steyerm., Bol. Soc. Venez. Ci. Nat. 32: 343. 1976. Shrub 2–3 m tall; branchlets with 5–15 rigid obcuneate leaves < 1 cm long, lamina gland 0.1–0.2 mm wide. Tepui meadows and scrub, 1900–2300 m; Bolívar (Cerro Jaua). Endemic. Phyllanthus chimantae Jabl., Mem. New York Bot. Gard. 17(1): 100. 1967. Shrub 0.5–3 m tall; branchlets 2–8 cm long, with 3–10 suborbicular leaves, laminar glands 0.5–1 mm long. Tepui meadows and shrublands, 2000–2500 m; Bolívar (Macizo del Chimantá). Endemic.

Phyllanthus 197

Phyllanthus duidae Gleason, Bull. Torrey Bot. Club 58: 382. 1931. Shrub 1–3 m tall; branchlets 5–15 cm long; leaves rigid, orbicular with cordate bases clasping stem. Tepui meadows and scrub, 1000–2300 m; Amazonas (Cerro Duida). Endemic. ŠFig. 188. Phyllanthus elsiae Urb., Repert. Spec. Nov. Regni Veg. 15: 405. 1919. Tree 3–10 m tall, cauliflorous or ramiflorous; branchlets 10–30 cm long, with 10–20 chartaceous, broadly elliptic, bluntly pointed leaves 3–6 cm long; fruits indehiscent. Woods along shores of lagoons and rivers, near sea level to 100 m; Delta Amacuro (scattered), Bolívar (along Río Orinoco). Anzoátegui, Apure, Barinas, Falcón, Miranda, Monagas, Zulia; Mexico, Central America, Lesser Antilles, Colombia. ŠFig. 181. Phyllanthus elsiae is often confused with the cultivated P. acidus L. Phyllanthus fluitans Benth. ex Müll. Arg., Linnaea 32: 36. 1863. Floating aquatic; leaves distichous, suborbicular, 1–1.5 cm across. Streams and swamps, near sea level to 100 m; Delta Amacuro (Caño Cocuina, La Florida). Apure, Barinas; Ecuador, Peru, Brazil, Bolivia, Paraguay. Phyllanthus hyssopifolioides H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 108. 1817. Phyllanthus monocladus Urb., Repert. Spec. Nov. Regni Veg. 15: 404. 1919. Herb, stem often unbranched; leaves distichous, 0.5–1 cm long. Swamps, savannas, 50–200 m; Bolívar (Río Caroní), Amazonas (Puerto Ayacucho area, Río Samariapo, San Juan de Manipiare). Guárico, Monagas; Central America, Hispaniola, Colombia, Suriname, French Guiana, Brazil, Bolivia, Paraguay. ŠFig. 189. Phyllanthus jablonskianus Steyerm. & Luteyn, Ann. Missouri Bot. Gard. 71: 317. 1984. Subshrub or shrub 0.1–1.5 m tall; branchlets 4–12 cm long, with 10–20 leaves; leaf blades rigid, obovate, 1–2 cm long, the upper surface shiny. Montane scrub and meadows, 1700–2300 m; Amazonas (Sierra

de la Neblina). Adjacent Brazil (Amazonas: Serra da Neblina). Phyllanthus jauaensis Jabl., Mem. New York Bot. Gard. 23: 865. 1972. Shrub 0.5–1 m tall; branchlets only 1–2 cm long, with 3–5 suborbicular rigid leaves 1–1.5 cm wide. Montane scrub, 1900–2200 m; Bolívar (Cerro Jaua). Endemic. Phyllanthus juglandifolius Willd., Enum. Pl. suppl. 64. 1813. Guatemala, Belize, Honduras, West Indies, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil; 2 subspecies, 1 in Venezuela. P. juglandifolius subsp. cornifolius (H.B.K) G.L. Webster, J. Arnold Arbor. 39: 151. 1958. —Phyllanthus cornifolius H.B.K., Nov. Gen. Sp. (quarto ed.) Pl. 2: 115. 1817. Shrub or tree 2–10 m tall; branchlets 40– 100 cm long, with 25–45 leaves; leaf blades acuminate, 10–20 cm long. Semideciduous forests, 100–300 m; northern Bolívar (El Manteco, La Paragua). Zulia; Colombia, Ecuador, Peru, Brazil (Acre and Pará to Pernambuco). ŠFig. 183. Phyllanthus lediformis Jabl., Mem. New York Bot. Gard. 17(1): 103. 1967. Shrub 0.3–1 m tall; branchlets 3–5 cm long, with 4–10 leaves; leaf blades narrowly elliptic or oblong, 2–3 cm long, margins revolute, laminar gland 3–4 mm from tip. Montane scrub, 1200–1500 m; Amazonas (Cerro Yutajé). Endemic. Phyllanthus lindbergii Müll. Arg. in Mart., Fl. Bras. 11(2): 35. 1873. Herb, often with basal aerenchyma, dioecious or nearly so; branchlets mostly 2–5 cm long, with 10–15 leaves. Swamps, wet savannas, 50–200 m; Amazonas (Puerto Ayacucho area). Colombia, Brazil, Paraguay. Phyllanthus longistylus Jabl., Mem. New York Bot. Gard. 17(1): 100. 1967. Shrub 1–3 m tall; branchlets with 10–20 leaves; leaf blades obovate, 1–1.5 cm long, with gland near apex. Montane scrub, 1100– 2200 m; Bolívar (Auyán-tepui, Macizo del

198

E UPHORBIACEAE

Chimantá [Chimantá-tepui]), Amazonas (Cerro Parú, Cerro Sipapo). Endemic. Phyllanthus longistylus is only questionably distinct from P. chimantae. Phyllanthus maguirei Jabl., Mem. New York Bot. Gard. 17(1): 105. 1967. Shrub 0.5–2 m tall; branchlets 8–10 cm long, with 20–25 leaves; leaf blades linear, 1.5–2.2 cm long and only 1–2 mm wide. Montane forests, along streams, 1000–2000 m; Amazonas (Cerro Duida, Sierra de la Neblina). Endemic. Phyllanthus major Steyerm., Fieldiana, Bot. 28: 318. 1952. Shrub 1–2 m tall; branchlets 2–7 cm long, with 5–15 leaves; leaf blades obovate, rigid, 1–2 cm long, laminar gland 0.5–1 mm long. Montane scrub, savannas, 1000–2300 m; Bolívar (Gran Sabana, northern tepuis), Amazonas (Duida-Marahuaka-Huachamacari massif). Guyana. Phyllanthus microphyllus H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 109. 1817. Herb, often with basal aerenchyma, dioecious or nearly so; branchlets 2–5 cm long, with 10–20 leaves. Swamps, wet savannas, 50–300 m; Bolívar (El Manteco, along Río Orinoco between Caicara and Las Ventanas), Amazonas (Culebra, Puerto Ayacucho area). Apure, Guárico; Brazil. Phyllanthus minutifolius Jabl., Mem. New York Bot. Gard. 17(1): 115. 1967. Subshrub 10–30 cm tall; branchlets 4–5 cm long, with 60–70 leaves; leaf blades linear-lanceolate, 4–5 mm long. Tepui meadows, ca. 1800 m; Amazonas (Cerro Sipapo). Endemic. ŠFig. 178. Phyllanthus minutulus Müll. Arg. in Mart., Fl. Bras. 11(2): 54. 1873. Herb; branchlets 1–4 cm long, with 8–15 leaves. Wet savannas, along streams, 50–800 m; Bolívar (Arekuna, San Ignacio), Amazonas (Puerto Ayacucho area). Guárico, Sucre; Colombia, Guyana, Brazil. ŠFig. 185. Phyllanthus myrsinites H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 111. 1817. Phyllanthus adenophyllus Müll. Arg., Linnaea 32.24. 1863. Phyllanthus dinizii Huber, Bull. Soc. Bot. Genève sér. 2, 6: 182. 1915.

Phyllanthus gallinetae Jabl., Mem. New York Bot. Gard. 17(1): 111. 1967. Phyllanthus glaucoviridis Jabl., Mem. New York Bot. Gard. 17(1): 101. 1967. Shrub 1–3 m tall; branchlets terete, slender, 10–40 cm long, mostly with 15–30 leaves; leaf blades obovate, rigid, glandular near tip. White-sand savannas, granitic outcrops, riparian forests, 50–600 m. Eastern Colombia, southern Venezuela, northern Brazil, disjunct in Peru; 2 varieties, both in the flora area. Phyllanthus myrsinites is a variable species; additional subspecific taxa can possibly be recognized. Key to the Subspecies of P. myrsinites 1. Branchlets ± terete, < 1 mm diameter; leaves mostly obovate, broadest above the middle, laminar gland 0.5–2 mm diameter, mostly 1–2 mm from tip ............ ................................ subsp. myrsinites 1. Branchlets flattened, > 1 mm diameter; leaves obovate to oblong, often broadest at or below the middle, laminar gland ca. 0.5 mm diameter, mostly < 1 mm from tip ........... subsp. francavillanus P. myrsinites subsp. myrsinites Shrub 1–3 m tall. White-sand savannas, lajas, riparian forests, 50–600 m; Amazonas (Caño Caname, Cerro Marahuaka, base of Cerro Yapacana, Patacame, Río Siapa, Río Sipapo, Samariapo, Sierra de la Neblina). Eastern Colombia (Vaupés, Amazonas), northeastern Peru (San Martín), northern Brazil (Amapá, Amazonas, Mato Grosso, Pará). P. myrsinites subsp. francavillanus (Müll. Arg.) G.L. Webster, comb. & stat. nov. —Phyllanthus francavillanus Müll. Arg., Linnaea 32: 20. 1863. Phyllanthus pimichinianus Jabl., Mem. New York Bot. Gard. 17(1): 111. 1967. Shrub 1–3 m tall. White-sand savannas, seasonally flooded banks of black-water rivers, 50–200 m; Amazonas (Caño Caname, Caño San Miguel, Pimichín, Río Temi). Brazil (Pará). ŠFig. 190. Phyllanthus neblinae Jabl., Mem. New York Bot. Gard. 17(1): 107. 1967. Shrub 0.1–2 m tall; branchlets 5–8 cm long, with 8–10 suborbicular leaves. Mon-

Phyllanthus 199

tane scrub, 1000–2300 m; Amazonas (Sierra de la Neblina). Adjacent Brazil (Amazonas: Serra da Neblina). Phyllanthus obfalcatus Lasser & Maguire, Brittonia 7: 79. 1950. Shrub 0.5–4 m tall; branchlets 5–10 cm long, with 40–75 leaves; leaves falcate, 0.7–1 cm long. Montane scrub, 1300–2200 m; Amazonas (Cerro Camani, Cerro Ualipano, Cerro Yaví, Cerro Yutajé, Serranía Uasadi, Sierra de Maigualida). Endemic. ŠFig. 192. Phyllanthus orbiculatus Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. Herb; branchlets mostly 3–6 cm long, with 10–15 leaves. Disturbed areas, forests, savannas, 100–700 m; widespread in northern Bolívar, Amazonas (Cerro Yapacana, San Carlos de Río Negro). Scattered in northern Venezuela; Colombia, Trinidad, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. ŠFig. 184. Phyllanthus orinocensis Steyerm., Fieldiana, Bot. 28: 321. 1952. Shrub ca. 1.5 m tall; leaves ovate-elliptic, acuminate, 2–4 cm long. Premontane forests, 200–300 m; Amazonas (base of Cerro Duida). Endemic. The affinities of Phyllanthus orinocensis are uncertain, as flowers are unknown. Phyllanthus paezensis Jabl., Mem. New York Bot. Gard. 17(1): 113. 1967. Shrub to 3 m tall; branchlets 10–12 cm long, with 10–12 leaves; leaf blade ovate-elliptic, obtuse, 2–4 cm long, pilose on both faces. Riparian forests, 50–100 m; Bolívar (between Puerto Paez and Orupe, island in Río Orinoco). Colombia (Vichada). Phyllanthus paraqueensis Jabl., Mem. New York Bot. Gard. 17(1): 104. 1967. Low shrub; branchlets 10–12 cm long, with 10–20 opposite leaves; leaf blade suborbicular, rigid, 1.5–2 cm wide. Montane scrub, 1500–1800 m; Amazonas (Cerro Sipapo). Endemic. Phyllanthus piscatorum H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 113. 1817. Phyllanthus pseudoconami Müll. Arg. in Mart., Fl. Bras. 11(2): 42. 1873. Phyllanthus pseudoconami var. pubescens

Müll. Arg. in Mart., Fl. Bras. 11(2): 42. 1873. Phyllanthus ichthyomethius Rusby, Mem. New York Bot. Gard. 7: 282. 1927. Shrub 1–3 m tall; branchlets with ca. 15 lateral axes, each with 10–15 elliptic acute leaves; leaf blades 1.5–3 cm long, usually hirsutulous, at least abaxially. Riparian forests, 50–400 m; Bolívar (El Tigre, Imataca, Santa María de Erebato), Amazonas (Culebra, Mavaca, Raudal de Atures, Río Manapiare, Tamatama). Anzoátegui; Ecuador, Peru, Brazil, Bolivia. ŠFig. 177. Most, if not all, of the Phyllanthus used as fish-poison plants belong to this variable species. It is closely related to P. brasiliensis Aubl., which differs in less ramified branchlets with blunter leaves. Phyllanthus pycnophyllus Müll. Arg. in A. DC., Prodr. 15(2): 322. 1866. Shrub 1–4 m tall; branchlets 5–10 cm long, with 15–20 leaves; leaf blade falcately oblanceolate, rigid, 1.5–2 cm long. Montane scrub, 1900–2400 m; Bolívar (Cerro El Sol, Ilú-tepui, Roraima-tepui). Guyana. Phyllanthus rupestris H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 110. 1817. Phyllanthus brachycladus Müll. Arg., Linnaea 32: 35. 1863. Phyllanthus delicatissimus Jabl., Mem. New York Bot. Gard. 17(1): 107. 1967. Shrub 1–3 m tall; branchlets slender, 2–4 cm long, with 3–5 leaves; leaf blade membranaceous, elliptic or obovate, blunt, 2–3 cm long. Riparian thickets or forests, 50–300 m; Amazonas (Puerto Ayacucho area, Río Negro, Río Orinoco between Raudal de Maipures and Río Atabapo, Salto Yureba). Apure; Colombia, Brazil (Amazonas). ŠFig. 187. Phyllanthus stipulatus (Raf.) G.L. Webster, Contr. Gray Herb. 176: 53. 1955. —Moeroris stipulata Raf., Sylva Tellur. 91. 1838. Phyllanthus diffusus Klotzsch in Seem., Bot. Voy. Herald 105. 1853. Herb, base of stem often with aerenchyma; branchlets 2–6 cm long, with 15–30 leaves. Swamps, moist forests, near sea level to 900 m; Delta Amacuro (lower Río Orinoco), Bolívar (scattered), Amazonas (scattered). Anzoátegui, Cojedes, Guárico; widespread in Central America and South America. ŠFig. 186.

200

E UPHORBIACEAE

Phyllanthus strobilaceus Jabl., Mem. New York Bot. Gard. 17(1): 96. 1967. Shrub 1–2 m tall; branchlets 10–25 cm long, with 20–35 leaves; leaf blade obovate, rounded at tip, ca. 1.5 cm long. Montane scrub, 1000–1800 m; Bolívar (Ilú-tepui, Macizo del Chimantá), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Endemic.

Phyllanthus tenellus Roxb., Fl. Ind. ed. 2, 3: 668. 1832. Phyllanthus corcovadensis Müll. Arg. in Mart., Fl. Bras. 11(2): 30. 1873. Annual herb 1–30 cm tall; branchlets 5– 15 cm long, with 10–25 leaves. Weedy areas, 100–200 m; Bolívar (Entrerios). Widespread weed in tropical to warm-temperate regions.

Phyllanthus subapicalis Jabl., Mem. New York Bot. Gard. 17(1): 101. 1967. Shrub 2–3 m tall; branchlets 3–8 cm long, with 30–60 leaves; leaf blade linear-oblong, rigid, 0.5–1 cm long; laminar gland apical, small and often indistinct or obsolete. Venezuela, Brazil; 3 subspecies, all in the flora area.

Phyllanthus tepuicola Steyerm., Acta Bot. Venez. 10: 236. 1975. Shrub; branchlets 5–9 cm long, with 25– 45 leaves; leaf blade elliptic-oblong, ca. 0.5 cm long. Montane scrub, ca. 1000 m; Amazonas (Cerro Duida). Endemic.

Key to the Subspecies of P. subapicalis 1. Leaves not closely imbricate; styles shortly bifid (0.1–0.2 mm) at tip; anthers ca. 1 mm long ........................ subsp. A 1. Leaves closely imbricate; styles entire; anthers < 1 mm long or > 1.5 mm long .......................................................... 2 2. Stylar column 1 mm long; anthers > 1.5 mm long; seeds 3.2 mm long ................... ............................ subsp. sequoiifolius 2. Stylar column (1.5–) 2–4 mm long; anthers < 1 mm long; seeds 2.5–2.8 mm long ....................... subsp. subapicalis P. subapicalis subsp. sequoiifolius (Jabl.) G.L. Webster, comb. & stat. nov. —Phyllanthus sequoiifolius Jabl., Mem. New York Bot. Gard. 17(1): 93. 1967. Forests, 1700–1900 m; Bolívar (Macizo del Chimantá [Sarvén-tepui]). Endemic. P. subapicalis subsp. subapicalis Phyllanthus anadenus Jabl., Mem. New York Bot. Gard. 17(1): 93. 1967. Tepui meadows and scrub, 1000–1800 m; Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Autana, Cerro Guanay, Cerro Jaua, Cerro Sipapo, Cerro Yapacana). Brazil (Roraima: Serra Parima). ŠFig. 193. P. subapicalis subsp. A Riparian scrub on sandstone, 1200–1300 m; Amazonas (Río Coro Coro west of Serranía Yutajé. Endemic.

Phyllanthus urinaria L., Sp. Pl. 982. 1753. Phyllanthus croizatii Steyerm., Fieldiana, Bot. 28: 317. 1952. Herb; branchlets wing-angled, 5–10 cm long, with 20–35 leaves; leaf blades oblong, submarginally hispidulous on abaxial face. Disturbed areas, wet or moist forests, near sea level to 100 m; Delta Amacuro (scattered). Several states of northern Venezuela; native of tropical Asia, widespread weed in the Neotropics. Phyllanthus vacciniifolius (Müll. Arg.) Müll. Arg. in A. DC., Prodr. 15(2): 322. 1866. —Glochidion vacciniifolium Müll. Arg., Linnaea 32: 63. 1863. Phyllanthus vacciniifolius subsp. vinillaensis Steyerm., Ann. Missouri Bot. Gard. 71: 317. 1984. Shrub 0.5–2.5 m tall; branchlets mostly 5–10 cm long, with 15–25 leaves; leaf blade obovate, rounded, 1–1.5 cm long. Savannas, river banks, 400–2400 m; Bolívar (widespread on tepuis, Gran Sabana), Amazonas (Cerro Aracamuni, Cerro Vinilla, Sierra de la Neblina). Guyana, Brazil. ŠFig. 191. Phyllanthus ventuarii Jabl., Mem. New York Bot. Gard. 17(1): 104. 1976. Shrub 0.1–1.5 m tall; branchlets 10–15 cm long, with 6–15 leaves; leaf blade 3–4 cm long, suborbicular to reniform, usually wider than long. Montane thickets, ca. 2000 m; Amazonas (Cerro Parú). Endemic. Phyllanthus ventuarii is closely related to P. duidae.

Phyllanthus

Fig. 177. Phyllanthus piscatorum

Fig. 179. Phyllanthus atabapoensis

201

Fig. 178. Phyllanthus minutifolius

Fig. 180. Phyllanthus caroliniensis subsp. guianensis

202

E UPHORBIACEAE

Fig. 181. Phyllanthus elsiae Fig. 182. Phyllanthus attenuatus

Phyllanthus 203

Fig. 183. Phyllanthus juglandifolius subsp. cornifolius

Fig. 184. Phyllanthus orbiculatus

Fig. 185. Phyllanthus minutulus

204

E UPHORBIACEAE

Fig. 186. Phyllanthus stipulatus

Fig. 188. Phyllanthus duidae

Fig. 187. Phyllanthus rupestris

Fig. 189. Phyllanthus hyssopifolioides

Phyllanthus 205

Fig. 190. Phyllanthus myrsinites subsp. francavillanus

Fig. 192. Phyllanthus obfalcatus

Fig. 191. Phyllanthus vacciniifolius

Fig. 193. Phyllanthus subapicalis subsp. subapicalis

206

E UPHORBIACEAE

45. PIRANHEA Baill., Adansonia 6: 235. 1866. Celaenodendron Standl., Contr. Dudley Herb. 1: 76. 1927. by Grady L. Webster and Paul E. Berry Dioecious shrubs or trees. Leaves alternate, trifoliolate, long-petiolate; stipules caducous. Inflorescences axillary, spicate. Flowers apetalous; staminate sepals 4–6, imbricate; stamens 3–15; filaments free, disk central with interstaminal lobes. Pistillate sepals 6, 2-seriate, imbricate; disk 6-lobed; ovary 3-locular, 2 ovules per locule; styles unlobed. Fruit capsular, valves woody, each locule with one ecarunculate seed. Mexico, Colombia, Venezuela, Guyana, Brazil; 4 species, 2 in Venezuela, both in the flora area. Key to the Species of Piranhea 1. 1.

Fruit smooth, depressed-globose, pedicel 5–10 cm long ........................ ..................................................................................... P. longepedunculata Fruit verrucose, angular, apiculate, sessile or short-pedicellate .......... .................................................................................................... P. trifoliata

Piranhea longepedunculata Jabl., Mem. New York Bot. Gard. 17(1): 122. 1967. —Caramacate, Pata de gallina. Tree 3–30 m tall. Evergreen lowland and gallery forests, 100–500 m; Delta Amacuro

(Río Toro), Bolívar (20–35 km southwest of El Manteco on road to San Pedro de Las Dos Bocas, Serranía de Imataca). Mérida, Táchira, Trujillo. ŠFig. 194.

Fig. 194. Piranhea longepedunculata

Plukenetia 207

Fig. 195. Piranhea trifoliata

Piranhea trifoliata Baill., Adansonia 6: 236, t. 6. 1866. Tree 5–25 m tall. Seasonally flooded riparian forests, granitic outcrops, evergreen lowland forests, 50–200 m; Bolívar (Río Orinoco at Cerro Carichana), Amazonas

(Caño Asisa near junction with Río Parú, savannas at base of Cerro Yapacana, Isla Ratón, lower Río Sipapo). Anzoátegui, Apure, Guárico; Guyana, Brazil (Amazonas, Mato Grosso, Pará). ŠFig. 195.

46. PLUKENETIA L., Sp. Pl. 1192. 1753. Apodandra Pax & K. Hoffm.in Engl., Pflanzenr. IV. 147(Heft 68): 20. 1919. Elaeophora Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 112. 1925. by Lynn J. Gillespie Monoecious or rarely dioecious lianas or twining vines. Leaves alternate, simple, stipulate, petiolate; blades pinnately or palmately veined; margins subentire to serrulate; 1–several pairs of flat glands present near base on upper surface of blade; flat glands scattered or absent on lower surface. Inflorescences axillary or terminal on short shoot, racemose, bisexual, with pistillate flower(s) at base and staminate flowers above in condensed cymules, or rarely unisexual (P. polyadenia); bracts small, triangular, eglandular. Staminate flowers pedicellate, green or yellowish green; sepals 4 or 5, valvate; petals absent; disk interstaminal, segmented or annular, often reduced or absent; stamens 15–40, free, on convex or subglobose receptacle; filaments short to elongate or anthers sessile; pistillode absent. Pistillate flowers pedicellate; sepals 4; petals and disk absent; ovary 4-locular, 4-angled to deeply

208

E UPHORBIACEAE

4-lobed, with lobes tuberculate, carinate, or horned, locules uniovulate; styles partly to entirely connate, column cylindrical to globose or obovoid. Fruit a 4-seeded schizocarpous capsule or berry, deeply 4-lobed to subglobose, each carpel carinate or with central tubercle or horn. Seeds lenticular and laterally compressed, subglobose, or ovoid, ecarunculate. Neotropics, southeast Asia, Africa, Madagascar; ca. 13 species, 5 in Venezuela, all in the flora area. Key to the Species of Plukenetia 1.

1.

2(1).

2.

3(1).

3.

4(3).

4.

Leaf blades palmately veined or 3-veined at base, broadly obtuse to cordate at base; styles partly connate (sometimes free only at tip), 4–30 mm long; all stamens with distinct filaments; fruit fleshy, indehiscent or tardily dehiscent, 2.5–11 cm diameter .................................................. 2 Leaf blades pinnately veined, acute to rounded at base; styles entirely connate, 1–4 mm long; all anthers sessile or stamens dimorphic with inner anthers sessile and outer with short filaments; fruit capsular, 1–2 cm diameter .................................................................................... 3 Leaf blade base rounded or obtuse; stylar column 3–7 mm long; fruit 6–11 cm diameter; staminate bud narrowly oblong-ellipsoid; filaments slender, 2–3 mm long ............................................................. P. polyadenia Leaf blade base cordate or truncate; stylar column 15–30 mm long; fruit 2.5–6 cm diameter; staminate bud subglobose; filaments conical, ca. 0.5 mm long ................................................................................ P. volubilis Ovary and capsule lobes distinctly horned; stylar column slender-cylindrical, 2–4 mm long; androecium consisting of 15–25 anthers sessile on globose receptacle ..................................................................... P. loretensis Ovary and capsule lobes with rounded tubercle (fruit unknown in P. multiglandulosa); stylar column stout-cylindrical, 1–2(–3) mm long; androecium consisting of outer whorl of 4(5) stamens with filaments and inner cluster of 6–12 sessile anthers on globose receptacle .......... 4 Young shoots and petioles densely hirsute, lower surface of blade hirsute, trichomes persistent; basilaminar glands in 3–5 pairs ...................... ....................................................................................... P. multiglandulosa Young shoots and petioles puberulent, lower surface of blade sparsely puberulent, becoming glabrous; basilaminar glands in 1 or 2(3) pairs ...... ............................................................................................... P. penninervia

Plukenetia loretensis Ule, Verh. Bot. Vereins. Prov. Brandenburg 81. 1908. —Apodandra loretensis (Ule) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 68): 21. 1919. —Bejuco de morrocoy. Twining vine; leaf blade elliptic, 6–19 × 3–9 cm, acute to obtuse and shortly decurrent at base, chartaceous; inflorescence 1–6 cm long. Evergreen lowland and lower montane forests, often on white sand, 50–800 m; Bolívar (Cerro Camarón southwest of Cerro Guaiquinima, Icabarú area, Santa María de Ere-

bato), Amazonas (widespread). Guyana, Amazonian Peru, Brazil (Amazonas, Mato Grosso, Rondônia), Amazonian Bolivia. ŠFig. 196. Plukenetia multiglandulosa Jabl., Mem. New York Bot. Gard. 17(1): 143, fig. 23. 1967. Twining vine; leaf blade narrowly elliptic, 6–9 × 1.8–3 cm, cuneate at base, chartaceous; inflorescence ca. 2 cm long. Talus forests at escarpment bases, ca. 1800 m; Amazonas (Cerro Parú). Endemic

Plukenetia 209

Plukenetia penninervia Müll. Arg., Linnaea 34: 158. 1865. Twining vine; leaf blade elliptic, 6–19 × 2.5–8.5 cm, obtuse to rounded at base, subcoriaceous; inflorescence 1–3 cm long. Disturbed areas in evergreen lowland forests or semideciduous forests, 100–300 m; Delta Amacuro (Serranía de Imataca), Bolívar (near Guri, Serranía de Imataca). Distrito Federal, Falcón, Miranda, Portuguesa, Zulia; southern Mexico, Central America, Colombia. Plukenetia polyadenia Müll. Arg. in Mart., Fl. Bras. 11(2): 334. 1874. —Elaeophora polyadenia (Müll. Arg.) Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 146. 1930. Elaeophora abutifolia Ducke, Arch. Bot. Rio de Janeiro 4: 112. 1925. —Plukenetia abutifolia (Ducke) Pax & K. Hoffm. in Engl. & Prantl, Pflanzenfam. ed. 2, 19c: 141. 1931. Liana climbing to 20 m or more in forest

Fig. 196. Plukenetia loretensis

canopy; leaf blade elliptic or ovate-elliptic, 7– 14 × 4–10 cm; staminate and bisexual inflorescences 5–25 cm long, pistillate inflorescences 1.5–7 cm long; seeds ovoid, ca. 5 cm long. Evergreen lowland to lower montane forests, 50–900 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), southern Bolívar (scattered), Amazonas (headwaters of Río Orinoco). Guyana, Suriname, French Guiana, Amazonian Ecuador, northern Peru, Amazonian Brazil. ŠFig. 197. Plukenetia volubilis L., Sp. Pl. 1192. 1753. Plukenetia macrostyla Ule, Verh. Bot. Vereins. Prov. Brandenburg 80. 1908. Twining vine or slender liana; leaf blade triangular-ovate, 7–13 × 4–10 cm; inflorescence 5–18 cm long; seeds lenticular, 1.5–2 × 0.7–0.8 cm. River banks in evergreen lowland forests, 100–200 m; Amazonas (base of Sierra de la Neblina). Distrito Federal, Miranda; Lesser Antilles, Colombia, Suriname, Ecuador, Peru, Brazil (western Amazonas, Pará), Amazonian Bolivia.

210

E UPHORBIACEAE

Fig. 197. Plukenetia polyadenia

Podocalyx 211

Pistillate branch

Staminate branch

Fig. 198. Podocalyx loranthoides

212

E UPHORBIACEAE

47. PODOCALYX Klotzsch, Arch. Naturgesch. 7: 202. 1841. by Grady L. Webster and Paul E. Berry Dioecious trees, without latex. Leaves simple, alternate, entire, eglandular, pinnately veined; stipules small; petioles thickened at both ends. Inflorescences raceme-like thyrses, 1–3(–8) per node. Flowers 1–3 per axil (pistillate) or several in tomentose capitula (staminate), apetalous. Sepals 5, imbricate. Staminate flowers with massive, lobed interstaminal disk; stamens 4–7; filaments free. Pistillate flowers 3-carpellate, 2 ovules per locule; styles 3; stigmas sessile, dilated, undivided. Fruit capsular, 3-lobed, pubescent; pedicel thickened and generally with evident lenticels. Seed smooth, ovoid, ecarunculate. Colombia, Venezuela, Peru, Brazil; 1 species. Podocalyx loranthoides Klotzsch, Arch. Naturgesch. 7: 202. 1841. —Richeria loranthoides (Klotzsch) Müll. Arg. in DC., Pordr. 15: 49. 1866. —Palo de agua dulce, Reventillo. Cunuria ? casiquiarensis Croizat, J. Arnold Arbor. 26: 191. 1945. Small tree 5–10(–20) m tall; leaves with strong subparallel secondary veins (7–10 per side) and mostly inconspicuous lower order

venation; fruits split open characteristically into 3 segments both septicidally and loculicidally with a persistent columella. Evergreen lowland and riparian forests, riparian scrub on white sand, 50–400 m; Bolívar (Río Caroní below Urimán), Amazonas (Caño Negro off Río Cunucunuma, Caño San Miguel, Río Atabapo basin, Río Orinoco, Río Pasimoni). Colombia, Peru, Brazil (Amazonas, Pará). ŠFig. 198.

48. POGONOPHORA Miers ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 372. 1854. by Ricardo de S. Secco Dioecious trees or shrubs. Leaves simple, alternate, subcoriaceous to coriaceous; stipules very short; margins entire. Inflorescences axillary, paniculate, with dense silky trichomes, the pistillate ones larger than the staminate. Staminate flowers subsessile; sepals 5, free, imbricate, the 2 external ones smaller than the 3 internal ones; petals 5, imbricate, internally bearded; stamens 5, attached to the perimeter of the staminal disk; rudimentary gynoecium present. Pistillate flowers shortpedicellate; calyx as in the staminate flowers, persistent; petals 5 free, imbricate, internally bearded; ovary 3-locular, 1 ovule per locule; style trifid. Fruit a tricoccous capsule, the cocci flattened. Seeds ovoid; testa crustaceous, carunculate. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, western Africa; 2 species, 1 in Venezuela. Pogonophora schomburgkiana Miers ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 373. 1854. —Flor de mayo. Pogonophora schomburgkiana var. longifolia Miers ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 373. 1854. Pogonophora schomburgkiana f. elliptica Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 47): 109. 1911. Poraresia anomala Gleason, Bull. Torrey

Bot. Club 58: 385. 1931 [1932]. Shrub or tree 1.5–14 m tall. Nonflooded and seasonally flooded forests, granitic outcrops, roadsides, 100–200 m; Bolívar (Serranía de Imataca), Amazonas (Caño Yapacana, Río Casiquiare basin, Río Cunucunuma, Río Guainía, Río Guasacavi, Río Negro, Río Orinoco 35 km southeast of La Esmeralda). Colombia, Guyana, Suriname, French Guiana, Brazil (widespread). ŠFig. 199.

Richeria 213

Pistillate flower

Dehiscent fruit

Fig. 199. Pogonophora schomburgkiana

49. RICHERIA Vahl, Eclog. Amer. 1: 30. 1796 [1797]. by Ricardo de S. Secco Dioecious trees or shrubs. Leaves simple, alternate, chartaceous, subcoriaceous, at times having a pair of glands at the base of the blade; stipules deciduous; margins entire or distantly crenulate. Inflorescence axillary, a raceme or spike, 1 or more per axil. Staminate flowers apetalous, sessile; calyx lobed, imbricate; stamens 3–6, free, alternating with the disk glands; pistillode present. Pistillate flowers apetalous, pedicellate; calyx 3–5-lobed, imbricate; annular disk slightly lobed; ovary 3-locular, glabrous or pubescent, 2 ovules per locule; styles 3, entire or bipartite. Fruit a fleshy tricoccous capsule, columella 3-winged, the wings papyraceous. Seeds 1 per locule, elliptic, with endosperm, ecarunculate. Costa Rica, Panama, Antilles, Colombia, Venezuela, Guyana, French Guiana, Peru, Brazil; 3 species, 1 in Venezuela. Richeria grandis Vahl, Eclog. Amer. 1: 30. 1796 [1797]. Guarania laurifolia Baill., Adansonia 5: 348. 1865. —Richeria laurifolia (Baill.) Baill., Adansonia 6: 16. 1865. Richeria grandis var. divaricata Müll. Arg. in A. DC., Prodr. 15(2): 467. 1866.

Richeria grandis var. genuina Müll. Arg. in A. DC., Prodr. 15(2): 468. 1866. Richeria grandis var. obovata Müll. Arg. in A. DC., Prodr. 15(2): 468. 1866. —Richeria obovata (Müll. Arg.) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 81): 29. 1922.

214

E UPHORBIACEAE

Staminate branch Pistillate branch

Fruiting branch

Fig. 200. Richeria grandis

Richeria grandis var. racemosa Müll. Arg. in A. DC., Prodr. 15(2): 467. 1866. —Richeria racemosa (Müll. Arg.) Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 81): 29. 1922. Tree or shrub 2–15 m tall; fruits often mottled or tuberculate when dried. Ever-

green lowland flooded and nonflooded forests, secondary and riparian forests, montane forests, 100–2000 m; widely scattered in Bolívar and Amazonas. Apure, Mérida, Trujillo, Yaracuy; Lesser Antilles, Colombia (Vaupés), Guyana, French Guiana, Ecuador, Brazil (widespread). ŠFig. 200.

50. RICINUS L., Sp. Pl. 1007. 1753. by Grady L. Webster Monoecious shrubs or small trees, appearing herbaceous when young. Leaves alternate, palmately veined and lobed, peltate; stipules connate, deciduous; petiole long, glandular at apex; margins serrate. Flowers apetalous, in terminal panicles, staminate flowers below, pistillate above. Staminate calyx splitting into 3–5 segments; disk absent; stamens many; filaments connate into irregular fascicles; pistillode absent. Pistillate flowers pedicellate; calyx splitting into 3–5 segments;

Sagotia 215

Fig. 201. Ricinus communis

disk absent; ovary 3-locular, muricate; styles free, bifid, branches papillate; ovules 1 in each cell. Fruit capsular. Seeds carunculate; endosperm copious. Native to east Africa or India, widely cultivated and naturalized throughout the tropics and subtropics; 1 species. Ricinus communis L., Sp. Pl. 1007. 1753. —Tártago. Shrub or small tree 2–5 m tall. Waste areas, near sea level to 300 m; Delta Amacuro (Sacupana) and also probably scattered in

populated areas of Delta Amacuro, northern Bolívar, and Amazonas. Widespread in northern half of Venezuela; widely naturalized in Neotropics and subtropics, native to east tropical Africa or India. ŠFig. 201.

51. SAGOTIA Baill., Adansonia 1: 53. 1860, nom. cons., non Duchass. & Walp., Linnaea 23: 737. 1850. by Ricardo de S. Secco Monoecious trees or shrubs. Leaves simple, alternate, chartaceous to subcoriaceous; stipules lacking; margins entire. Inflorescences terminal racemes, or simple or corymbose panicles, the staminate flowers generally at the apex, the pistillate ones toward the base. Staminate flowers with very slender pedicels; calyx 5- or 6parted, imbricate, glabrous or pilose; petals 5, free, imbricate; stamens numerous, subsessile. Pistillate flowers with thick pedicels; calyx accrescent; petals lacking; ovary 3-locular, 1 ovule per locule, densely pilose; styles 3, bipartite, wrinkled to papillose. Fruit a tricoccous capsule, the cocci 2-valved. Seeds ovoid, smooth, marbled, carunculate. Amazonian Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 species, both in Venezuela.

216

E UPHORBIACEAE

Key to the Species of Sagotia 1.

1.

Inflorescence rachis and pedicels pilose; calyx of pistillate flowers with sepals oblong-elliptic to obovate; style 3–11 mm long; stamens inserted on a pilose receptacle ......................................................... S. brachysepala Inflorescence rachis and pedicels glabrous; calyx of pistillate flowers with sepals linear-lanceolate to elliptic-lanceolate; style 1.5–3.5 mm long; stamens inserted on a glabrous receptacle .............................. S. racemosa

Sagotia brachysepala (Müll. Arg.) Secco, An. 1° Simp. Trop. Umido 2: 59. 1986. —Sagotia racemosa var. brachysepala Müll. Arg., Flora 33: 516. 1864. —Caicareño, Tunari. Tree 5–35 m tall. Nonflooded forests, river margins, roadsides, 300–500 m; Bolívar

(base of Abacapá-tepui, Río Caura at La Angostura, Río Karún, Río Uaiparú). Zulia; Colombia (Río Vaupés), Guyana, Suriname, French Guiana, Brazil (Amazonas, Mato Grosso, Rondônia, Amapá, Pará). The wood is light and used for firewood and crates.

Fig. 202. Sagotia racemosa

Sandwithia 217

Sagotia racemosa Baill., Adansonia 1: 54. 1860. Sagotia racemosa var. genuina Müll. Arg., Flora 33: 516. 1864. Sagotia racemosa var. ligularis Müll. Arg., Flora 33: 516. 1864. Sagotia racemosa var. microsepala Müll. Arg., Flora 33: 517. 1864. Sagotia tafelbergii Croizat, Bull. Torrey Bot. Club 75: 404. 1948.

Shrub or tree 2–9 m tall. Riverbanks, lowland forests, 100–200 m; Bolívar (south of Aprada-tepui along Río Chirca, between La Lira and Kilómetro 88), Amazonas (La Esmeralda, Río Casiquiare near Solano, Río Sipapo, near San Carlos de Río Negro, base of Sierra de la Neblina). Zulia; Amazonian Colombia, Guyana, Suriname, French Guiana, Brazil (Amazonas, Amapá, Pará, Maranhão). ŠFig. 202.

52. SANDWITHIA Lanj., Bull. Misc. Inform. Kew 1932: 184. 1932. by Ricardo de S. Secco Monoecious trees. Leaves simple, alternate, chartaceous; stipules lacking; margins entire; petiole pulvinate at the apex. Staminate inflorescences in terminal panicles or fascicles, the pistillate ones simple racemes, usually terminal, sometimes axillary. Staminate flowers with a 2-laciniate calyx, valvate in bud; petals 3 or 4, imbricate, oblong to suborbiculate, margins ciliate; stamens numerous; filaments

Pistillate flower

Fig. 203. Sandwithia guianensis

218

E UPHORBIACEAE

short or long. Pistillate flowers with a 3-laciniate calyx, rarely tubular, undulate or with 5 free sepals; petals 5, minute, lanceolate or obovate, pilose; ovary 3-locular, densely pilose, 1 ovule per locule, style trifid. Fruit tricoccous, subglobose, conical. Seeds ovoid, carunculate. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 species, both in the flora area. Key to the Species of Sandwithia 1.

1.

Staminate inflorescence paniculate; stamens 5–10 mm long, filaments 3.5–8 mm long, anthers suborbicular, thecae voluminous, connectives truncated at the apices; pistillate flowers with connate calyx; style branches slender, the branch ends entire ............................. S. guianensis Staminate inflorescence fasciculate, sometimes borne on leafless nodes; stamens ca. 1.5 mm long, filaments ca. 0.5 mm long, anthers ellipticlanceolate, thecae linear, discrete, connectives acuminate at the apices; pistillate flower with 5 free sepals, style branches thick, the branch ends bifid ................................................................................ S. heterocalyx

Sandwithia guianensis Lanj., Bull. Misc. Inform. Kew 1932: 184. 1932. —Caspadillo, Gaspadillo. Tree 2–24 m tall. Evergreen lowland and lower montane forests, 200–500 m; border between Delta Amacuro and Bolívar (Serranía de Imataca, upper Río Caura above Araguaña), Amazonas (bases of Cerro Huachamacari and Sierra de la Neblina). Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas). ŠFig. 203.

Sandwithia heterocalyx Secco, Bol. Mus. Paraense Emilio Goeldi, Sér. Bot. 4: 179. 1988. Tree 2–18 m tall. Evergreen lowland and lower montane forests, 100–600 m; Bolívar (Río Caura), Amazonas (Río Cataniapo, upper Río Cunucunuma, Serranía Batata 55 km southeast of Puerto Ayacucho). Colombia, Brazil (Amazonas: upper Río Negro).

53. SAPIUM Jacq., Enum. Syst. Pl. 9, 31. 1760, nom. cons. Sapium subg. Eusapium Pax & K. Hoffm. sect. Americana Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 200. 1912, nom. inval. by Hans-Joachim Esser Monoecious shrubs or trees, with white latex; glabrous; ultimate ramification alternate. Leaves alternate, simple, glabrous; stipules scaly, nearly entire, eglandular, persisting and often seemingly 4 because of similar axillary bud scales; petioles short but distinct, apically usually biglandular, rarely eglandular; blades entire to serrate or ciliate, hardly revolute in the flora area, not tripliveined, tertiary venation reticulate, the upper surface eglandular, the lower surface smooth to papillate, often with few to several strictly marginal glands that may even be stalked. Inflorescences elongate, terminal, simple, yellowish green; floral bracts with a pair of large, undivided, disk- to cup-shaped glands often decurrent along the axis; staminate flowers in apical, 3–18-flowered cymules, pistillate flowers single per bract at base, sometimes with additional staminate flowers. Staminate flowers erect; bracteoles divided into cilia-like lobes; pedicel short (ca. 1 mm) at anthesis, absent in bud; perianth 2-lobed, partly connate; stamens 2 per flower; filaments distinct, free;

Sapium 219

anthers free; disk and pistillode absent. Pistillate flowers: bracteoles not visible; pedicel short to nearly absent; perianth 2- or 3-lobed, partially connate; disk and staminodes absent; ovary smooth, 1–3-locular; style column short; stigmas 1–3, undivided. Fruit a dry to slightly fleshy schizocarp, opening septicidally and loculicidally (in variable sequence), globose, hardly sulcate; pericarp thin, woody to nearly membranous or somewhat fleshy; exocarp smooth; mericarps with thin and caducous septa; columella with caducous wings. Seeds 1–3 per fruit; irregularly globose-elliptic, ecarunculate; testa brownish to blackish, completely covered by a red fleshy aril. Southeastern U.S.A., Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; ca. 25 species, 6 in Venezuela, 4 of these in the flora area. Key to the Species of Sapium 1.

1. 2(1).

2.

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

Leaves with distinctly stalked petiolar glands, stalks at least 1 mm long and twice as long as wide (often inflexed below and not visible from above) ..................................................................................................... 2 Leaves with shortly stalked to sessile petiolar glands, stalks < 1 mm long and < 2 times as long as wide (rarely inflexed below) .......................... 3 Leaves (except the largest) usually distinctly whitish-papillate on the lower surface, with 7–13 pairs of side veins below the apex, margin serrate with teeth 1–1.5 mm apart, marginal glands not to slightly (to 0.3 mm) stalked ................................................................. S. paucinervium Leaves not or indistinctly whitish-papillate on the lower surface, with (11–)16–26 pairs of side veins below the apex, margin entire or serrate with teeth 1.5–5 mm apart, marginal glands distinctly (0.4–1 mm) stalked ................................................................................ S. glandulosum Leaves subentire; ovary and fruit 1-locular, with one oblique stigma (rarely a second rudimentary stigma present) ........................ S. jenmanii Leaves subentire to serrate; ovary and fruit 2- or 3-locular, with 2 or 3 distinct stigmas ........................................................................................... 4 Ovary and fruit 2-locular, with 2 stigmas .................................. S. contortum Ovary and fruit 3-locular, with 3 stigmas ............................. S. glandulosum

Sapium contortum Croizat, J. Arnold Arbor. 26: 193. 1945. Tree to 6 m tall. Forests along streams, roadsides, 100–300 m, Bolívar (north of Puerto Ayacucho), Amazonas (Puerto Ayacucho, lower Río Ventuari). Apure, Táchira; probably also in Colombia. This species is intermediate between Sapium glandulosum and S. jenmanii in carpel number and size of the petiolar glands (the latter being smaller than in typical S. glandulosum and slightly larger than in S. jenmanii, although some overlap exists). It was omitted by Jablonski (Mem. New York Bot. Gard. 17(1): 181-186. 1967). Kruijt in

his recent Sapium monograph (Biblioth. Bot. 146. 1996) proposed it as a doubtful synonym of S. glandulosum, but determined most of the relevant collections as S. jenmanii, while at the same time restricting the latter species to strictly 1-locular fruits. Sapium glandulosum (L.) Morong, Ann. New York Acad. Sci. 7: 227. 1893. —Hippomane glandulosa L., Sp. Pl. 1191. 1753. —Sapium aucuparium Jacq., Enum. Syst. Pl. 31. 1760, nom. illeg. —Hippomane biglandulosa L., Sp. Pl. ed. 2, 2: 1431. 1763, nom. illeg. —Sapium hippomane G. Mey., Prim. Fl. Esseq. 275, fig.

220

E UPHORBIACEAE

Fig. 204. Sapium glandulosum

Savia 221

21. 1818, nom. illeg. —Sapium biglandulosum (L.) Müll. Arg., Linnaea 32: 116. 1863, nom. illeg. —Sapium biglandulosum var. aubletianum Müll. Arg., Linnaea 32: 117. 1863, nom. illeg. —Sapium aubletianum (Müll. Arg.) Huber, Bull. Herb. Boissier sér. 2, 6: 362. 1906, nom. illeg. —Caucho, Lechero, Lechero blanco, Lechero morado, Mukonyo (Panare). Sapium prunifolium Klotzsch, London J. Bot. 2: 45. 1843. Sapium moritzianum Klotzsch in Seem., Bot. Voy. Herald 100. 1852. Sapium biglandulosum var. klotzschianum Müll. Arg., Linnaea 32: 117. 1863. —Sapium klotzschianum (Müll. Arg.) Huber, Bull. Herb. Boiss., sér. 2, 6: 438, fig. 30. 1906. Sapium biglandulosum var. lanceolatum Müll. Arg., Linnaea 32: 118. 1863. —Sapium lanceolatum (Müll. Arg.) Huber, Bull. Herb. Boiss., sér. 2, 6: 441. 1906. Sapium obtusilobum Müll. Arg., Linnaea 32: 116. 1863. Sapium hemsleyanum Huber, Bull. Herb. Boissier sér. 2, 6: 362. 1906. Sapium albomarginatum Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 85): 203. 1924. Sapium guaricense Pittier, J. Wash. Acad. Sci. 19: 355. 1929. Sapium naiguatense Pittier, J. Wash. Acad. Sci. 19: 355. 1929. Sapium paucistamineum Pittier, J. Wash. Acad. Sci. 19: 356. 1929. Shrub or tree to 20 m tall. Riparian or swampy forests, often on seasonally flooded ground, secondary forests, forest remnants in pastures, swampy savannas, evergreen and semideciduous forests on granitic slopes

or sandstone, sand, or clay, near sea level to 600 m; Delta Amacuro (west of Caño JotaSuburu, Pedernales), Bolívar (Altiplanicie del Nuria, eastern Gran Sabana, basins of lower Río Caroní, lower Río Caura, Río Cuyuní, and Río Suapure). Widespread elsewhere in Venezuela; Mexico, much of the West Indies, South America (except Chile), sometimes cultivated for latex production in other parts of the tropics, e.g., Malesia. ŠFig. 204. The fruits of this species are edible, and the latex yields rubber of high quality. Although it has been experimentally grown in different parts of the tropics, its harvest is more difficult than in Hevea, and it has never become commercially important. Sapium jenmanii Hemsl. in Hook., Icon. Pl. 27: pl. 2649. 1900. —Caucho, Caucho blanco, Lechero. Tree to 40 m tall. Lower montane forests, 300–700 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Santa Elena de Uairén, Serranía de Imataca). Trinidad (introduced from Guyana), Guyana, Brazil (eastern Amazon basin). The latex, which is used for gum balls, is of similar quality to the preceding species, and likewise not important commercially. Sapium paucinervium Hemsl. in Hook., Icon. Pl. 27: pl. 2648. 1900. —Caucho blanco. Sapium microdentatum Lanj., Euphorb. Surinam 46, fig. 12. 1931. Tree to 20 m tall. Lower montane forests, 200–600 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil (eastern Amazon basin).

54. SAVIA Willd., Sp. Pl. 4: 771. 1806. by Grady L. Webster Dioecious shrubs or small trees, without latex. Leaves simple, alternate, entire, eglandular, pinnately veined; stipules caducous. Inflorescence axillary, flowers in glomerules (pistillate flowers fewer in number or solitary), calyx and corolla 5merous, lobes imbricate, disk annular. Staminate flowers subsessile; stamens usually 5; filaments free; anthers introrse. Pistillate flowers pedicellate, 3-carpellate, ovules 2 per locule; styles free, bifid. Fruit a capsule, columella persistent. Seeds usually 1 per locule, smooth, ecarunculate. Mexico, West Indies, Venezuela, Brazil, east Africa, Madagascar; 20–25 species, 1 in Venezuela.

222

E UPHORBIACEAE

Fig. 205. Savia sessiliflora

Savia sessiliflora (Sw.) Willd., Sp. Pl. 4: 771. 1806. —Croton sessiliflorum Sw., Prodr. 100. 1788. Tree 5–18 m tall; leaves chartaceous, deciduous, generally long-acuminate; petiole very short. Semideciduous forests, locally

dominant and gregarious, ca. 400 m; Bolívar (Cerro Panamo east of Lago Guri). Aragua, Carabobo, Distrito Federal, Falcón, Yaracuy; Mexico, Cuba, Puerto Rico, Hispaniola, Virgin Islands. ŠFig. 205.

55. SENEFELDERA Mart., Flora 24(2) Beibl.: 29. 1841. by Hans-Joachim Esser Monoecious trees, rarely shrubs, with white latex; trichomes multicellular, uniseriate; ramification in whorls. Leaves in whorls of 4(8), simple, glabrous or pubescent; stipules longer than wide, entire, caducous; petioles distinct, eglandular; blades entire, not revolute, upper surface with or without a single, sometimes divided gland at the base, lower surface smooth (neither papillose nor glaucous), with 2–10 laminar glands on each side, never with marginal glands; tertiary venation percurrent to reticulate. Inflorescences elongate, axillary, compound thyrsoid, yel-

Senefeldera 223

Staminate inflorescence

Fig. 206. Senefeldera inclinata

224

E UPHORBIACEAE

lowish, side branches subtended by larger, scaly, sometimes hairy bracts; floral bracts eglandular or with one pair of oblong-cylindrical glands; staminate flowers in 1–3(–7)-flowered apical cymules, pistillate flowers single per bract at the base; bracts of cymules eglandular or with one pair of oblong-cylindrical glands touching the rachis or decurrent. Staminate flowers erect to inclined; bracteoles undivided, similar to bracts, eglandular; pedicel short or distinct; perianth totally connate, without free lobes; stamens (4–)6–17; filaments very short to longer than anthers, free or basally connate; anthers free; disk and pistillode absent. Pistillate flowers: bracteoles often absent; pedicel short to nearly absent; perianth 3-lobed, basally connate; ovary 3-locular, smooth, glabrous; ovules 1 per locule; style column 0–1.5 mm long; stigmas 3, undivided; disk and staminodes absent. Fruit a dry schizocarp, primarily septicidal, globoid to slightly ellipsoid, sulcate; pericarp thick, woody; exocarp smooth, glabrous; mericarps with narrow septa or these absent; columella distinctly winged. Seeds 3 per fruit, globoid to ellipsoid, with or without a caruncule; testa dry, smooth or brownish or bicolored and partly sculptured. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 4 species, 2 in Venezuela, 1 of these in the flora area. Under its current circumscription, Senefeldera is heterogeneous and presumably polyphyletic, and the atypical S. inclinata should probably be excluded and placed in a new genus. Senefeldera inclinata Müll. Arg. in Mart., Fl. Bras. 11(2): 530, fig. 27, O–V. 1874. —Palo de cuca, Palo de cuchara, Palo de perro de agua. Senefeldera multiflora var. acutifolia Müll. Arg. in A. DC., Prodr. 15(2): 1154. 1866, pro parte. Senefeldera karsteniana Pax & K. Hoffm. in Engl., Pflanzenr. IV. 147(Heft 52): 25. 1912. Tree 4–15(–30) m tall; leaves eglandular on upper surface; floral bracts always

eglandular; staminate flowers inclinate, distinctly pedicellate, with partly fused stamens; fruits 15–17 mm long; seeds globoid, often dorsally sometimes apiculate, caruncule missing to rudimentary; testa smooth, brown. Evergreen lowland forests, 100–200 m; Amazonas (between Maroa and Yavita, Río Casiquiare, Río Negro, Río Temi, between San Carlos de Río Negro and Solano). Zulia; Amazonian Colombia, Ecuador, Peru, northwestern Brazil, Bolivia. ŠFig. 206.

56. SENEFELDEROPSIS Steyerm., Bot. Mus. Leafl. 15: 45. 1951. by Hans-Joachim Esser Monoecious trees or shrubs with white latex; trichomes multicellular, uniseriate; bark papery; branching in whorls. Leaves alternate, simple, glabrous or pubescent; stipules ca. 1 mm long, shortly triangular, eglandular, caducous; petioles distinct, eglandular; blades entire, revolute, the upper surface usually with one pair of basal glands, lower surface papillose or glaucous, with (0–)2–5 submarginal glands on each side; tertiary venation percurrent. Inflorescences elongate, terminal, compound thyrsoid, yellowish, side branches subtended by leafy or else minute scaly bracts; floral bracts with one pair of elliptic-cylindric glands, touching the axis and often decurrent; staminate flowers in 6–12-flowered apical cymules; pistillate flowers single per bract at base, often lacking. Staminate flowers erect; bracteoles similar to bract, undivided, eglandular; pedicel 0–2 mm long; perianth 3-lobed, partially connate; stamens 3–5 per flower; filaments free, as long as anthers; anthers free. Pistillate flowers: bracteole short to absent; pedicel short; perianth 3(6)-lobed, basally connate; ovary 3-locular, smooth, pubescent; ovules 1 per locule; style column

Senefeldera 225

0.5–2 mm long; stigmas 3, undivided. Fruit a partly fleshy, septicidal, oblongoid, sulcate schizocarp; pericarp very thick, woody but leathery-fleshy outside; exocarp smooth, sparsely pubescent, glabrescent; mericarps with distinct septa; columella slightly winged. Seeds 3 per fruit, oblongoid-ellipsoid, with or without a caruncule; testa dry, smooth, brown. Endemic to the Guayana Shield in eastern Colombia, southern Venezuela, and Guyana; 2 species, both in the flora area. Key to the Species of Senefelderopsis 1. 1.

Leaves elliptic, apically acuminate, usually chartaceous; indument yellowish ............................................................................. S. chiribiquetensis Leaves obovate, apically mucronate to emarginate, usually coriaceous; indument reddish ....................................................................... S. croizatii

Senefelderopsis chiribiquetensis (R. Schult. & Croizat) Steyerm., Bot. Mus. Leafl. 15: 47. 1951. —Senefeldera chiribiquetensis R. Schult. & Croizat, Caldasia 3: 122. 1944. Shrub or small tree to 8 m tall; bark greyish, stems subsucculent; leaf blades papillose on lower surface; seeds carunculate. Forested slopes, xerophytic shrublands or savannas on sandstone or granitic soils, granitic outcrops, gallery forests, 100–1400 m; Amazonas (Cerro Baraco 15 km west of La Esmeralda, Cerro Mawedi in Río Ocamo basin, slopes of Cerro Moriche, north of Cerro Vinilla, Cerro Yapacana, southern slopes of Cuao-Sipapo massif, 5 km south of Pendare on Río Sipapo, Río Coro Coro at base of Cerro Yutajé, Río Cunucunuma). Amazonian Colombia.

Fig. 207. Senefelderopsis croizatii

Senefelderopsis croizatii Steyerm., Bot. Mus. Leafl. 15: 46. 1951. Senefelderopsis sipapoensis Jabl., Mem. New York Bot. Gard. 12(3): 175. 1965. Senefelderopsis venamoensis Jabl., Mem. New York Bot. Gard. 12(3): 176. 1965. Tree to 30 m tall; lower surface of leaf blades glaucous-farinose; seeds ecarunculate. Montane and gallery forests, 800–1600 m; Bolívar (base of Carrao-tepui, 3 km south of El Paují, base of Ilú-tepui, La Escalera), Amazonas (Cerro Aracamuni, Cerro Sipapo). Guyana. ŠFig. 207.

226

E UPHORBIACEAE

Pistillate plant

Staminate plant

Fig. 208. Tetrorchidium rubrivenium

57. TETRORCHIDIUM Poepp., Nov. Gen. Sp. Pl. 3: 23. 1842. by Paul E. Berry Dioecious trees or shrubs; stems with simple or T-shaped trichomes. Leaves alternate, rarely opposite; stipules glandular; petioles with prominent paired glands near the apex; blades entire or dentate; venation pinnate. Staminate inflorescences axillary, spiciform or racemiform, occasionally branched, flowers usually 3–7 in

Tragia 227

sessile glomerules; flowers small, subsessile; calyx lobes 3, imbricate, ribbed on the inner face, petals and disk absent; stamens 3, free, opposite the calyx lobes; filaments short; anthers peltate, rounded, 4-thecate. Pistillate inflorescences axillary, short racemes or panicles, solitary; flowers subsessile or pedicellate; calyx lobes 3; petals absent, disk cup-shaped or 3-lobed; ovary 2- or 3-locular, 1 ovule per locule; styles short, free, bifid, often with broad stigma-like style branches. Fruits capsular, thin-walled, 2- or 3-lobed. Seeds rounded, ecarunculate; outer seed coat fleshy, inner seed coat foveolate. Neotropics, Africa; ca. 20 species, 2 in Venezuela, 1 of these in the flora area. Tetrorchidium rubrivenium Poepp., Nov. Gen. Sp. Pl. 3: 23, t. 227. 1842. Shrub or tree 3–20 m tall. Montane forests, 600–1300 m; Bolívar (Sierra de Maigualida, Serranía Marutaní), Amazonas (Ca-

ño Piedra near Cerro Cuao). Widespread in montane areas of northern Venezuela; Central America, West Indies, Colombia, Ecuador, Peru, Brazil. ŠFig. 208.

58. TRAGIA L., Sp. Pl. 980. 1753. by Lynn J. Gillespie Monoecious or rarely dioecious herbs, twining vines, or subshrubs. Stems and foliage usually covered with stinging trichomes. Leaves alternate, simple, petiolate, eglandular; stipules usually small, deciduous; blades pinnately or palmately veined; margin entire, lobed, or serrate. Inflorescences axillary or terminal, racemose, sometimes with a single branch near base, bisexual with pistillate flowers at basal node(s) or unisexual; bracts small, eglandular. Staminate flowers pedicellate; sepals 3–6, valvate; petals absent; disk interstaminal and segmented or intrastaminal, often absent; stamens (1)2–50; filaments often connate; pistillode absent. Pistillate flowers pedicellate; sepals 3–6; petals and disk absent; ovary 3-locular, locules uniovulate, densely covered with stinging hairs; styles slender, partly connate. Fruit a 3-seeded, 3-lobed schizocarpous capsule, covered with stinging trichomes. Seeds globose to ellipsoid, ecarunculate. Tropical and warm-temperate regions of the New World, Africa, southern and southwestern Asia, and northeastern Australia; ca. 130 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Tragia 1.

1.

Filaments free, stamens 7–10; staminate disk segments 6 or 7; staminate sepals 3(4); inflorescence bisexual, appearing dichotomous, consisting of a racemose staminate main axis and a pistillate branch ....... T. fendleri Filaments connate into a central elongate column bearing a cluster of ca. 5 sessile anthers; staminate disk absent; staminate sepals 5; inflorescence unisexual, racemose ................................................... T. guayanensis

Tragia fendleri Müll. Arg., Linnaea 34: 179. 1865. Tragia japurensis Müll. Arg. in Mart., Fl. Bras. 11(2): 404. 1874. Twining vine; leaf blade oblong-ovate or ovate, 7–15 × 4–8 cm, base cordate; styles 3–

3.5 mm long, papillose on inner surface. Savannas, disturbed areas in forests, 50–300 m; Delta Amacuro (near Tucupita), Bolívar (near El Manteco), Amazonas (base of Sierra de la Neblina). Zulia; Guyana, Brazil (Amazonas). ŠFig. 209.

228

E UPHORBIACEAE

Pistillate branch and flower

Staminate flower

Fig. 209. Tragia fendleri

Tragia guayanensis L.J. Gillespie, Novon 4: 331. 1994. Climbing subshrub; young stems and petioles hirsute; leaf blade elliptic or ovateelliptic, 12–17 × 6–12 cm, base narrowly cor-

date; styles 6–10 mm long, smooth. Evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, Río Matacuni, mouth of Río Ventuari). Endemic.

EUPHRONIACEAE by Julian A. Steyermark and Luis Marcano-Berti Trees or shrubs. Young branches pubescent with simple trichomes. Stipules small, deciduous. Leaves alternate, simple; blades epunctate, pinnately veined; margins entire. Inflorescence terminal or axillary, generally racemose; bracts deciduous. Flowers bisexual, perigynous. Calyx with 5 subequal, imbricate sepals or lobes inserted on a campanulate or turbinate-campanulate hypanthium or calyx tube, lacking a spur or pouch; petals 3, free, contorted in bud, without a spur. Androecium monadelphous, 6–10-merous, with 4 fertile stamens in 2 opposite pairs, separated on one side by a long staminode with a sterile anther and on the other side by 1–5 short, denticulate staminodes opposite the long staminode; anthers bithecate, longitudinally dehiscent. Ovary superior, 3-locular; ovules 2 per locule, superposed, the lower one pendent, the upper one erect and shorter; style 1, simple. Fruit a 3-locular, septicidal capsule, the calyx and androecium persistent. Seed 1 per locule, winged at the base. Endemic to the Guayana Shield in Colombia, Venezuela, Guyana, and Brazil; 1 genus and 3 species, all in the flora area.

Euphronia 229

Euphronia has previously been placed in the Trigoniaceae or the Vochysiaceae, but was designated as a distinct family by Marcano-Berti (Pittieria 18: 15. 1989). 1. EUPHRONIA Mart., Nov. Gen. Sp. Pl. 1: 121, t. 73. 1824 [1826]. Characters and distribution as in the family description. Key to the Species of Euphronia 1.

1.

2(1).

2.

Sepals or calyx lobes caudate-acuminate, the exterior portion covered by a dense, uniform, appressed, shortly woolly tomentum; inflorescence subfasciculate, the flowers 1–6, crowded near the upper portion of the rachis; pedicels 7–8 mm long, nearly equaling the calyx, covered by an appressed, shortly woolly tomentum ........................... E. acuminatissima Sepals or calyx lobes acute, subobtuse, or subrounded at the apex, the exterior portion with loose, ascending to spreading trichomes throughout or at least on midrib; inflorescence racemose, the flowers 3–23, the pedicels arising along an elongated rachis; pedicels 1.5–5 mm long, shorter than the calyx, with loosely ascending to spreading pubescence .................... 2 Floriferous portion of rachis usually conspicuously elongated beyond the leafy branch, mainly (7–)15–23-flowered, 4–10 cm long; leaf blades 4.5–7.5 cm long, mainly lance- to oblong-elliptic, generally acute to subacute at the apex, 2–2.5 times longer than wide; sepals or calyx lobes ± uniformly gray-green when dry; exterior of sepals or calyx lobes with loose sericeous or ascending hairs along midportion, elsewhere with shortly appressed tomentum ................................................ E. hirtelloides Floriferous portion of rachis only shortly elongated, mainly 3–8-flowered, 1–4 cm long; leaf blades (0.6–)1.5–3(–4.8) cm long, mainly ovate to ovate-elliptic or obovate, generally rounded to obtuse at the apex, 1.7–2 times longer than wide; inner sepals or calyx lobes with conspicuous broad white borders when dry; exterior of sepals or calyx lobes with dense, loosely ascending to spreading trichomes over the entire surface ................................................................................................ E. guianensis

Euphronia acuminatissima Steyerm., Ann. Missouri Bot. Gard. 74: 91. 1987. Subshrub or tree 0.2–5 m tall; leaves oblong, elliptic or obovate. White-sand savannas and shrublands, 100–200(–2000) m; Bolívar (Uéi-tepui near La Ciudadela), central and western Amazonas. Endemic. ŠFig. 212. Euphronia guianensis (R.H. Schomb.) Hallier f., Meded. Rijks-Herb. 35(in obs.): 13. 1918. —Lightia guianensis R.H. Schomb., Linnaea 20: 757. 1847. —Curataquilla, Sacaraiyek (Arekuna). Shrub or tree (0.2–)2–10 m tall. Rocky open sandstone outcrops and savannas, 300– 1400(–1600) m; Bolívar (Gran Sabana west

to Cerro Guaiquinima). Guyana. ŠFig. 210. Euphronia hirtelloides Mart., Nov. Gen. Sp. Pl. 1: 121, t. 73. 1825 [1826]. Lightia licanioides Spruce ex Warm. in Mart., Fl. Bras. 13(2): 121. 1875. —Euphronia licanioides (Spruce ex Warm.) Hallier f., Meded. Rijks-Herb. 35: 13. 1918. Shrub or tree 2–10 m tall. Shrub savannas, riparian forests, 100–200 m; Amazonas (Caño San Miguel, base of Cerro Yapacana, Río Atacavi, Río Baría, Río Casiquiare, Río Guayapo, Río Pasiba, Río Pasimoni, San Carlos de Río Negro). Colombia (Caquetá, Guainía), Brazil (Rio Negro basin). ŠFig. 211.

230

E UPHRONIACEAE

Fig. 210. Euphronia guianensis

Fig. 211. Euphronia hirtelloides

Fig. 212. Euphronia acuminatissima

F ABACEAE 231

FABACEAE by Gerardo A. Aymard C., Nidia L. Cuello A., Paul E. Berry, Velva E. Rudd, Richard S. Cowan, Paul R. Fantz, Richard H. Maxwell, Charles H. Stirton, Hans-Helmut Poppendieck, Haroldo Cavalcante de Lima, Renée H. Fortunato, Basil Stergios, Nereida Xena de Enrich, David A. Neill, R. Toby Pennington, and Celia Gil Herbs, sometimes twining, shrubs, trees, or woody vines, rarely cauliflorous; roots commonly bearing root nodules with nitrogen-fixing bacteria. Leaves alternate, rarely opposite, pinnately or less often palmately compound or trifoliolate, sometimes unifoliolate or simple, the petiole and individual leaflets commonly with a swollen basal pulvinus; stipules present, sometimes modified into spines or prickles. Inflorescences mostly racemes, spikes, panicles, or heads. Flowers bisexual, usually strongly irregular, mostly papilionaceous. Sepals mostly 5, generally connate into a lobed, often bilabiate tube; petals usually 5, dissimilar [rarely 0–4 or 6, rarely 5 and similar (Etaballia)], the uppermost (adaxial; in resupinate flowers the lowermost, abaxial) petal the standard (sometimes called the vexillum and other parts associated with it are referred to by the adjective “vexillar”), borne externally to the other petals, usually the largest and ± enclosing the others in bud, 2 lateral petals, the wings, usually similar to each other and distinct, sometimes connivent basally, 2 lower petals innermost, similar to each other, and mostly connate to form a keel enclosing the androecium and gynoecium. Stamens mostly 10, seldom 5–9 or 11, rarely numerous, 9 of the filaments usually connate into an open sheath around the pistil, the 10th, adaxial one, usually partly separated from the rest, or filaments rarely free; anthers 4-sporangiate, 2-thecal, mostly opening longitudinally. Gynoecium a single carpel or rarely 2 or several separate carpels, each with a terminal style and stigma; ovules 1–many on a marginal placenta. Fruit commonly dry and dehiscent along both sutures, but sometimes a follicle, or indehiscent and then sometimes winged or breaking transversely into 1-seeded segments (loment), sometimes drupaceous, samaroid, or nut-like. Seeds with a short funiculus and usually a hard seed coat; embryo with thickened cotyledons. Cosmopolitan, ca. 400 genera and 10,000 species, 67 genera and 368 species in the flora area. This treatment follows the Cronquist system (A. Cronquist. An Integrated System of Classification of Flowering Plants. Columbia University Press. 1991), which recognizes three different families among the legumes.

Key to the Genera of Fabaceae by Paul E. Berry and Gerardo A. Aymard C. 1. 1. 2(1). 2.

Herbs, vines, small shrubs, or lianas ........................................................ 2 Large shrubs or trees ............................................................................... 38 Leaves digitately 2- or 4-foliolate; flowers enclosed by 2 bracts ... 67. Zornia Leaves 1-foliolate, 3(4 or 5)-foliolate, or pinnately compound; flowers not enclosed by 2 bracts, sometimes surrounded by several bracts ........... 3

232

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

F ABACEAE

Inflorescences spicate ................................................................................ 4 Inflorescences racemes, pseudoracemes, or solitary flowers .................... 5 Flowers surrounded by a series of floral bracts; stamens monadelphous; fruit a loment ..................................................................... 58. Stylosanthes 4. Flowers not surrounded by a series of floral bracts; stamens diadelphous; fruit not a loment .................................................................. 34. Indigofera 5(3). Leaves pinnately compound ...................................................................... 6 5. Leaves 1-foliolate, 3(4 or 5)-foliolate ....................................................... 18 6(5). Leaves even-pinnate .................................................................................. 7 6. Leaves odd-pinnate .................................................................................... 8 7(6). Vines; standard obovate; stamens monadelphous; seeds bicolored, orange and black ......................................................................................... 1. Abrus 7. Herbs to small shrubs or trees; standard nearly round; stamens diadelphous; seeds brown ................................................................. 55. Sesbania 8(6). Mostly prostrate herbs; standard scarious, persistent; stamens in 2 lateral bundles ..................................................................... 56. Soemmeringia 8. Vines, herbs, or small shrubs; standard usually deciduous; stamens not in 2 lateral bundles .................................................................................... 9 9(8). Herbs to small shrubs .............................................................................. 10 9. Woody or herbaceous vines ...................................................................... 13 10(9). Leaflets alternate or subopposite; fruit a loment ................ 3. Aeschynomene 10. Leaflets opposite; fruit not a loment ....................................................... 11 11(10). Pubescence of minutely hooked (view at 20–30×) trichomes; flowers resupinate; ovary stipitate ................................................................ 17. Clitoria 11. Pubescence absent, or if present, never with minutely hooked trichomes; flowers not resupinate; ovary sessile .................................................. 12 12(11). Pubescence of malpighiaceous (T-shaped) trichomes; anthers with an apical projection ......................................................................... 34. Indigofera 12. Pubescence not of malpighiaceous trichomes; anthers without an apical projection ................................................................................ 61. Tephrosia 13(9). Woody vines .............................................................................................. 14 13. Herbaceous vines ..................................................................................... 16 14(13). Pubescence of minutely hooked trichomes .................................... 17. Clitoria 14. Plants glabrous or pubescent, but never with minutely hooked trichomes .............................................................................................................. 15 15(14). Fruits with distinct submarginal wings; pseudoracemes with 6–12-flowered lateral short shoots .............................................................. 22. Derris 15. Fruits diverse, but never with distinct submarginal wings; pseudoracemes either 6–12-flowered on lateral short shoots or with paired flowers on a short common peduncle or flowers arising directly from the rachis and appearing verticillate, paired, or single ......................... 36. Lonchocarpus 16(13). Flowers red; calyx long-tubular; style densely barbate, 1–1.5 cm long ................................................................................................... 8. Barbieria 16. Flowers yellow, blue, white, or purple; calyx cupular; style to 8 mm long .............................................................................................................. 17 17(16). Stipules striate; calyx without tubercular-based trichomes; standard orbicular with a dorsal spur; stamens pseudodiadelphous; fruit mostly linear, 2-valved, dehiscent .................................................. 14. Centrosema

F ABACEAE 233

17.

Stipules not striate; calyx with tubercular-based trichomes; standard obovate to suborbicular without a dorsal spur; stamens monadelphous; fruit a loment ......................................................................15. Chaetocalyx 18(5). Erect or prostrate herbs to small shrubs ................................................ 19 18. Vines or lianas .......................................................................................... 25 19(18). Fruit a loment .......................................................................................... 20 19. Fruit not a loment .................................................................................... 22 20(19). Leaves 4-foliolate; calyx glandular-punctate; stamens monadelphous ................................................................................................... 52. Poiretia 20. Leaves 1- or 3-foliolate; calyx not glandular-punctate; stamens diadelphous ...................................................................................................... 21 21(20). Stipules scarious; calyx deeply divided, lobes unequal, the upper 2 glumaceous; fruit a loment, terete to compressed, submoniliform, puberulent ............................................................................................... 6. Alysicarpus 21. Stipules not scarious; calyx subentire to bifid, the lower lobe 3-toothed with the central tooth longer than the laterals, or the calyx almost equally lobed; fruit a loment, not submoniliform, glabrous or with hooked trichomes ................................................................ 23. Desmodium 22(19). Stamens monadelphous; anthers dimorphic; fruit inflated, not septate ............................................................................................... 19. Crotalaria 22. Stamens diadelphous; anthers uniform to subuniform; fruit not inflated, septate .................................................................................................. 23 23(22). Lower surface of leaflets not gland-dotted; stipules setaceous; petioles not canaliculate; flowers reddish purple or pink; anthers with an apical projection ............................................................................... 34. Indigofera 23. Lower surface of leaflets gland-dotted; stipules not setaceous; petioles canaliculate; flowers yellow; anthers without an apical projection....... 24 24(23). Shrubs; corolla yellow, lined with red; calyx not gland-dotted; standard ovate; seeds 4–6 ........................................................................ 10. Cajanus 24. Perennial herbs or subshrubs; corolla yellow, sometimes with the standard veins reddish purple; calyx glandular; standard obovate; seeds 2 ............................................................................................... 27. Eriosema 25(18). Pubescence of minutely hooked trichomes (view at 20–30×) ....... 17. Clitoria 25. Plants glabrous or pubescent, if pubescent, never with minutely hooked trichomes .............................................................................................. 26 26(25). Inflorescences nodose, 2–8 flowers at each node ..................................... 27 26. Inflorescences not nodose ........................................................................ 29 27(26). Calyx appearing bilabiate; fruits frequently with extra longitudinal ribs ............................................................................................... 12. Canavalia 27. Calyx 4- or 5-lobed; fruits without extra ribs .......................................... 28 28(27). Flowers reddish, the standard straight; stamens diadelphous; fruits with long, down-curved beaks .................................................... 20. Cymbosema 28. Flowers shades of purple, occasionally white or lilac, the standards mostly reflexed; stamens pseudomonadelphous; fruits short-beaked or beak absent ................................................................................ 24. Dioclea 29(26). Leaflets and calyx gland-dotted ............................................. 54. Rhynchosia 29. Leaflets and calyx not gland-dotted ........................................................ 30 30(29). Calyx with irritating trichomes; anthers barbate; glandular disk present;

234

F ABACEAE

30. 31(30). 31. 32(31).

32.

33(31). 33. 34(33).

34.

35(33). 35. 36(35). 36. 37(36).

37.

38(1). 38. 39(38). 39. 40(39). 40. 41(39). 41. 42(41).

42.

43(42). 43.

fruit densely covered with stiff irritating trichomes ............... 40. Mucuna Calyx without irritating trichomes; anthers glabrous; glandular disk absent; fruit without stiff irritating trichomes ....................................... 31 Stamens monadelphous or pseudomonadelphous .................................. 32 Stamens diadelphous ............................................................................... 33 Twining herbs; inflorescences not nodose, bracteoles subtending the calyx; calyx 5-lobed; stamens pseudomonadelphous; fruit with a hooked tip ........................................................................................... 62. Teramnus Climbing or erect subshrubs; inflorescences nodose; bracteoles present but not subtending the flowers; calyx 4-lobed; stamens monadelphous; fruit without hooked tip ........................................................... 31. Galactia Style glabrous ........................................................................................... 34 Style pubescent ........................................................................................ 35 Leaflets somewhat rhombic; calyx 5-lobed; standard with a pair of inflexed lateral auricles at base; fruit transversely grooved; anthers 8 or 9 ................................................................................ 11. Calopogonium Leaflets linear or lanceolate; calyx 4-lobed; standard without a pair of inflexed lateral auricles at base; fruit not transversely grooved; anthers 10 ................................................................................ 31. Galactia Roots tuberous; fruit articulate ............................................. 46. Pachyrhizus Roots not tuberous; fruit not articulate .................................................. 36 Pubescence of uncinate trichomes; style coiled through 1.5–2.5 revolutions; bracteoles persistent in fruit ....................................... 47. Phaseolus Uncinate trichomes absent; style not coiled, or, if coiled, then through 3–5 revolutions; bracteoles caducous .................................................. 37 Inflorescence nodes not glandular; upper calyx teeth free; corollas purplish upon drying, with elongated wing petals; thickened part of style bent sharply ..................................................................... 38. Macroptilium Inflorescence nodes glandular; upper calyx 2-lipped; corollas yellowish or white, sometimes with some purple or violet upon drying, with wing petals equal to the others and with erect, sigmoid-shaped, or coiled keel petals; thickened part of style gradually bent or coiled .............. 65. Vigna Leaves simple or unifoliolate ................................................................... 39 Leaves compound ..................................................................................... 45 Leaf margin serrate; fruits indehiscent .................................................. 40 Leaf margin entire; fruits dehiscent or indehiscent ............................... 41 Fruit globose to ovoid or broadly turbinate; seeds 1 or 2 .......... 35. Lecointea Fruit ellipsoid, laterally compressed, stipitate, apiculate; mesocarp fleshy; seeds 1–few ............................................................................. 66. Zollernia Inflorescences axillary spikes; petals linear .............................. 29. Etaballia Inflorescences racemes or panicles; petals not linear ............................. 42 Calyx entire and completely enclosing the flower in bud, opening irregularly or in distinct lobes; flowers not papilionaceous, petals 0–6; stamens numerous (> 15) ......................................................................... 43 Calyx lobes apparent in bud (entire and enclosing the bud in Alexa); flowers papilionaceous or not, petals generally 5; stamens generally < 15 ....................................................................................................... 44 Petals 3–6; fruit a single-seeded drupe ............................................ 4. Aldina Petals 0 or 1; fruit a 1- to multiseeded legume .......................... 59. Swartzia

F ABACEAE 235

44(42). 44. 45(38). 45. 46(45). 46. 47(45). 47. 48(47).

48.

49(48). 49. 50(48).

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

55.

56(55). 56. 57(56). 57. 58(56).

Anthers not versatile; fruits 1-seeded ....................................... 21. Dalbergia Anthers versatile; fruits 2- or 3-seeded .................................. 51. Poecilanthe Leaves opposite ........................................................................................ 46 Leaves alternate ....................................................................................... 47 Leaf rachis terete; inflorescence axillary; calyx tubular, not pellucid-punctate; petals yellow ............................................................ 49. Platymiscium Leaf rachis usually winged; inflorescence terminal; calyx pellucid-punctate, bilabiate; petals violet ....................................................... 60. Taralea Stamens free or nearly so ........................................................................ 48 Stamens connate to a considerable extent .............................................. 60 Calyx entire and completely enclosing the flower in bud, opening irregularly or in distinct lobes; flowers not papilionaceous, petals 0–6; stamens numerous (> 15) ......................................................................... 49 Calyx lobes apparent in bud (but entire and enclosing the bud in Alexa); flowers papilionaceous or not, petals generally 5; stamens generally fewer ..................................................................................................... 50 Petals 3–6; fruit a single-seeded drupe, less often 2–4-seeded ....... 4. Aldina Petals 0 or 1; fruit a multi-seeded legume ................................. 59. Swartzia Flowers ± regular, the adaxial petal a little wider than the others; fruit compressed, indehiscent, round to oblong, transversely rugulose ................................................................................................. 2. Acosmium Flowers zygomorphic or subactinomorphic (Myrocarpus); fruits various, but often either winged or dehiscent .................................................. 51 Fruit 1- or 2-seeded; leaves pellucid-punctate ........................................ 52 Fruit 1- or more-seeded; leaves mostly not pellucid-punctate ............... 54 Fruit narrowly elliptic or fusiform, the seed(s) flat and situated in the middle of the winged fruit .................................................. 42. Myrocarpus Fruit with a bulbous apical seed chamber and a large basal wing ........ 53 Leaflets 14–25; anthers shorter than the filaments; plants native in the flora area ......................................................................... 43. Myrospermum Leaflets 5–15; anthers longer than the filaments; plants mostly cultivated in the flora area .................................................................... 44. Myroxylon Calyx large, enclosing the bud; ovary long-stipitate; inflorescence a terminal, woody raceme .......................................................................... 5. Alexa Calyx various, not completely enclosing the bud; ovary sessile or shortstipitate; inflorescence axillary or terminal, generally not woody .... 55 Calyx 2-lobed, the upper lobe enlarged and covering the petals; petals pink; fruit flattened, 1-seeded, tapering at both ends, elastically dehiscent ...................................................................................... 39. Monopteryx Calyx without a large lobe covering the petals in bud; petals mostly blue or purple, less often white with darker nectar spot; fruit not as above .............................................................................................................. 56 Fruits indehiscent .................................................................................... 57 Fruits dehiscent ....................................................................................... 58 Fruits linear-oblong to elliptic, membranous, upper suture shortly winged ................................................................................................. 9. Bowdichia Fruits oblong, tapering at both ends, apex acute, not winged ................... .............................................................................................. 57. Spirotropis Stigma lateral; seeds reddish, hard, shiny .................................. 45. Ormosia

236

F ABACEAE

58. Stigma terminal; seeds not as above ....................................................... 59 59(58). Standard without lateral appendages; fruit with 1 or 2 seeds ................... .............................................................................................. 25. Diplotropis 59. Standard with 2 lateral appendages; fruit with more than 2 seeds .......... .......................................................................................... 16. Clathrotropis 60(47). Leaves 3-foliolate ..................................................................................... 61 60. Leaves pinnately compound with more than 3 leaflets .......................... 62 61(60). Flowers large, > 15 mm long; fruit dehiscent; seeds hard, red .................. ................................................................................................ 28. Erythrina 61. Flowers small, < 15 mm long; fruit indehiscent, drupaceous; seeds soft, not red ........................................................................................... 7. Andira 62(60). Lower surface of leaflets gland-dotted; flowers yellow ................ 10. Cajanus 62. Lower surface of leaflets not gland-dotted; flowers red, orange, rose, violet, or white .......................................................................................... 63 63(62). Fruits drupaceous .................................................................................... 64 63. Fruits nondrupaceous, dry at maturity, dehiscent or indehiscent ......... 65 64(63). Leaf rachis terete; calyx not pellucid-punctate, truncate or with short obsolete teeth or shortly 5-dentate; petals pink, violet, or white ..... ....................................................................................................... 7. Andira 64. Leaf rachis flattened or winged; calyx pellucid-punctate, bilabiate, upper lip with 2 large lobes and lower lip very short, entire, or 3-dentate; petals violet to whitish mauve ...................................................... 26. Dipteryx 65(63). Leaflets opposite on the rachis ................................................................ 66 65. Leaflets alternate or subopposite on the rachis ...................................... 73 66(65). Fruit with basal seed and apical wing, the basal part spinose with a stiff lateral spur; leaves with numerous reddish orange resinous dots; petals yellowish ..................................................................... 13. Centrolobium 66. Fruit without a basal spinose portion and lateral spur; leaves without resin dots; petal color various .............................................................. 67 67(66). Fruit flat, ovate-elliptic, with broad lateral wings; calyx spathaceous; petals yellow to orange; anthers opening by 2 apical pores ...... 30. Fissicalyx 67. Fruit not as above; calyx not spathaceous; petal color various; anthers longitudinally dehiscent ...................................................................... 68 68(67). Fruit longitudinally 4-winged ....................................................... 48. Piscidia 68. Fruit not 4-winged ................................................................................... 69 69(68). Fruits long and narrow, either cylindrical and constricted between the seeds or somewhat flattened and dehiscent ....................................... 70 69. Fruits thin and flattened, rounded to oblong, mostly indehiscent ......... 71 70(69). Fruit cylindrical, indehiscent, constricted between the seeds ... 41. Muellera 70. Fruit dehiscent, flattened, often wider toward the apex, very little constricted between the seeds, the 2 valves hard and often coiling in dehiscence ....................................................................................... 32. Gliricidia 71(69). Fruit 1-seeded, elastically dehiscent, 2-valved, flat; calyx pellucid-punctate, bilabiate ............................................................................. 60. Taralea 71. Fruit 1- or more-seeded, not elastically dehiscent; calyx not pellucid-punctate ....................................................................................................... 72 72(71). Fruit with 2 subparallel prominent veins near the margin; seeds 1(2); inflorescence a terminal panicle; vexillar stamen connate to staminal tube ................................................................................ 33. Hymenolobium

Abrus 237

72.

73(65). 73.

74(73). 74. 75(74).

75.

76(75). 76. 77(76). 77. 78(77). 78. 79(77). 79.

Fruit without subparallel marginal veins; seeds 1–12; inflorescence a lateral or terminal raceme or panicle; vexillar stamen free at base ......................................................................................... 36. Lonchocarpus Fruits long and narrow, many-seeded, without appendages; leaves evenpinnate .................................................................................... 18. Coursetia Fruits rounded to oblong, but not long and narrow, 1–few-seeded, sometimes with appendages; leaves odd-pinnate or less often even-pinnate .............................................................................................................. 74 Fruit with a central seed and surrounding wing (see also Dalbergia) ............................................................................................. 53. Pterocarpus Fruit not as above .................................................................................... 75 Leaves even-pinnate, the rachis often flattened and ± winged; calyx pellucid-punctate, bilabiate; fruit with dehiscent twisting woody valves .................................................................................................... 60. Taralea Leaves odd-pinnate, the rachis terete; calyx not pellucid-punctate or bilabiate; fruit generally indehiscent, often 1-winged with an apical or basal seed ............................................................................................. 76 Fruit with an apical seed and basal wing ............................ 50. Platypodium Fruit various, but not as above ................................................................ 77 Flowers in pyramidal, usually terminal panicles; fruits with a basal seed and distal wing or turgid and spongy-fibrous ..................................... 78 Flowers in subcorymbose, usually axillary panicles; fruits winged by flattened valves, seeds either basal or central ......................................... 79 Fruit with 2 small, basal, lateral wings ................................ 64. Vataireopsis Fruit without basal wings, apically winged or wingless with spongy-fibrous mesocarp ......................................................................... 63. Vatairea Seeds basally positioned in the fruit; anthers longitudinally dehiscent; stipules sometimes spinescent ......................................... 37. Machaerium Seed(s) medially positioned in the fruit; anthers with short, transverse slits; stipules never spinescent .............................................. 21. Dalbergia

1. ABRUS L., Syst. Nat. ed. 12, 2: 472. 1767. by Nidia L. Cuello A. Mostly twining vines, seldom shrubs. Leaves alternate, even-pinnate; rachis grooved; leaflets 11–27, oblong to ovate, mucronate, sparingly to densely pubescent with ascending trichomes, opposite along the rachis; stipels minute, appressed to the rachis; stipules subulate, persistent. Inflorescences axillary or lateral pseudoracemes, terminating leafy or leafless branches, with 1–several sessile flowers at a node; bracts and bracteoles often caducous. Flowers white, reddish brown, lavender, or pinkish. Calyx campanulate, lobed or short-toothed, sparingly shortpubescent; standard obovate, apically notched and with a short claw; wing petals upcurved, shorter than keel petals. Stamens 9, not exserted, monadelphous, the staminal tube adnate to the standard. Style curved, glabrate. Fruit oblong to linear, somewhat oblique, hardly stipitate, much-compressed or not, the valves softly pubescent, often becoming indurated, septate, the short, slender beak downturned. Seeds subglobose or slightly compressed-ovoid, arillate, sometimes with an annulus around the hilum, the hilum depressed.

238

F ABACEAE

Fig. 213. Abrus precatorius subsp. africanus

Fig. 214. Abrus pulchellus subsp. tenuiflorus

Acosmium

239

Paleotropics, now widely introduced and circumtropical; 17 species, 2 in Venezuela, both in the flora area. The brightly colored orange-red and black seeds are highly toxic, but are sometimes used as an adornment or in artwork by Panare Amerindians. Key to the Species of Abrus 1.

1.

Leaflets oblong-ovate, 3–8 mm wide; flowers in pedunculate racemes; fruits usually thick and woody, the surface muriculate, densely appressed-pubescent, ca. 2 times as long as broad; seeds subglobose, shiny scarlet with black ........................................................ A. precatorius Leaflets oblong-obovate, 7–13 mm wide; flowers sessile in small clusters; fruits mostly not woody, the surface smooth and sparsely pubescent, ca. 3 times as long as broad; seeds compressed, shiny brown .... A. pulchellus

Abrus precatorius L., Syst. Nat. ed. 12, 2: 472. 1767. Paleotropics, introduced to the Neotropics; 2 subspecies, 1 in Venezuela. A. precatorius subsp. africanus Verdc., Kew Bull. 24: 241. 1970. —Peonía. Abrus minor Desv., Ann. Sci. Nat. (Paris) 9: 418. 1826. Woody vine over shrubs. Disturbed or cultivated areas, near sea level to 50 m; Delta Amacuro (Santa Catalina), Bolívar (Caicara, Ciudad Bolívar). Anzoátegui, Aragua, Barinas, Falcón, Guárico, Miranda, Monagas, Nueva Esparta, Trujillo; U.S.A. (southern Florida), Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama, West Indies, Colombia, Guyana, Peru, Brazil (Bahia), Africa,

Asia, Oceania. ŠFig. 213. Abrus pulchellus Wall. ex Thwaites, Enum. Pl. Zeyl. 91. 1864 [1859]. Paleotropics, introduced to Neotropics; 5 subspecies, 1 in Venezuela. A. pulchellus subsp. tenuiflorus (Benth.) Verdc., Kew Bull. 24: 250. 1970. —Abrus tenuiflorus Benth. in Mart., Fl. Bras. 15(1): 216. 1859. —Arepillo, Bejuco arepillo. Vine over shrubs. Forest borders, 100– 1000 m; Bolívar (near Santa Elena de Uairén), Amazonas (near confluence of Río Negro and Brazo Casiquiare). Táchira; Brazil (Amazonas, Goiás, Mato Grosso), Africa, Sri Lanka. ŠFig. 214.

2. ACOSMIUM Schott. in Spreng., Syst. Veg. 4(cur. post.): 406. 1827. by Charles H. Stirton and Gerardo A. Aymard C. Large shrubs or trees. Stipules small, caducous. Leaves 3–31-foliolate, odd-pinnate, alternate. Leaflets membranaceous or coriaceous, often discolorous, opposite on the rachis. Inflorescences paniculate, terminal, rarely axillary, racemes short to very elongate. Flowers white, cream, or yellow, scarcely zygomorphic, pedicellate, bracteate and bracteolate, with axillary clusters of stalked glands. Calyx turbinatecampanulate, upper pair of teeth much larger and broader than lateral and keel teeth; petals free, erect-ascending, uppermost petal outermost, differently shaped than rest. Stamens free, 10 (rarely 5), subequal, usually longer than petals; anthers uniform, ovate. Ovary sessile or shortly stipitate, 2–4-ovulate, glabrous or shaggy; style filiform; stigma small or truncate, terminal. Legume indehiscent, compressed and flat or turgid, 1–4-seeded. Seeds ovate or orbicular, compressed. Central America, Colombia, Venezuela, Guyana, Suriname, Brazil, Bolivia, Paraguay, Argentina, Madagascar; 17 species, 3 in Venezuela, 2 of these in the flora area.

240

F ABACEAE

Key to the Species of Acosmium 1.

1.

Leaves 5–9-foliolate; leaflets ovate to broadly lanceolate, subcoriaceous, not prominently reticulate on the upper surface; fruit sessile or subsessile, turgid, woody; ovary glabrous ..................................... A. nitens Leaves 5- or 7-foliolate; leaflets ovate to ovate-elliptic, chartaceous to subcoriaceous, reticulate on the upper surface; fruit distinctly stipitate, plano-compressed, papery, winged; ovary pubescent ..................... A. sp. A

Acosmium nitens (Vogel) Yakovlev, Notes Roy. Bot. Gard. Edinburgh 29: 353. 1969. —Leptolobium nitens Vogel, Linnaea 11: 394. 1837. —Sweetia nitens (Vogel) Benth., J. Linn. Soc., Bot. 8: 262. 1865. —Chimaco, Congrio, Congrio rebalsero. Small tree to 10 m tall; leaves 5–9foliolate; flowers cream. Semideciduous to evergreen lowland forests along rivers and adjacent slopes, savannas, riparian shrubland along black-water rivers, 50–200 m; northern Bolívar, widespread in Amazonas. Apure, Guárico; Colombia, Guyana, Suriname, Brazil (Amazonas, Para, Mato Grosso, Rondônia), Bolivia. ŠFig. 215.

Acosmium sp. A. —Congrio, Inchapemen (Panare), Karamate yo. Slender, spreading tree to 20 m tall, branched from low down on the trunk; leaves 5- or 7-foliolate; flowers white; fruit winged. Savannas, savanna/forest ecotone, semideciduous forests, granitic outcrops, 50–300 m; Bolívar (Ciudad Piar, Corozal, east of Miamo, Morichal El Caballo, San Félix to Upata road, San Pedro de los Bocas, islands in Lago Guri), Amazonas (10 km south of Puerto Rico). Guyana, Brazil (Roraima).

Fig. 215. Acosmium nitens

Aeschynomene 241

3. AESCHYNOMENE L., Sp. Pl. 713. 1753. by Velva E. Rudd Herbaceous annuals, shrubs, or small trees. Leaves alternate, pinnate, 5– many-foliolate; stipules usually persistent, peltate, appendiculate below the point of attachment or attached at the base and not appendiculate; stipels lacking; leaflets alternate or subopposite, usually small, entire to serrate-denticulate. Inflorescences axillary or terminal, simple or compound racemes, sometimes fasciculate; bracts mostly similar to and intergrading with the stipules; bracteoles paired at the base of the calyx. Flowers ca. 5–25 mm long. Calyx bilabiate or campanulate with 5 subequal lobes; petals 5, white, yellow, orange, or purplish, glabrous or pubescent. Stamens 10, monadelphous or diadelphous (5 + 5); anthers uniform, elliptic, dorsifixed, or, sometimes nearly basifixed. Ovary (1)2–18-ovulate, sessile or shortstipitate; style glabrous with base usually persisting as a cuspidate apex to the fruit; stigma terminal, minutely capitate or penicillate. Fruit a loment, (1)2–18-articulate, commonly with the upper (adaxial) margin essentially entire, the lower margin crenate, sometimes both margins crenate or both subentire. Seeds reniform, smooth, lustrous, the hilum orbicular. Mainly tropical or subtropical America, Africa, Asia, and Australia, with a few species in warm-temperate areas; 150 species, 15 in Venezuela, 10 of these in the flora area. Key to the Species of Aeschynomene 1. 1. 2(1).

2.

3(2).

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

Stipules attached at the base, not peltate; calyx campanulate with 5 subequal lobes; standard pubescent on the outer face ............................... 2 Stipules peltate, appendiculate below the point of attachment; calyx bilabiate with the vexillar lip bifid, the keel lobe trifid; petals glabrous ... 4 Stems suffrutescent, erect; inflorescence terminal, paniculate, or some axillary and racemose; fruit moniliform, 4–6-articulate; stipe 4–5 mm long, glabrous ......................................................................... A. paniculata Stems herbaceous, prostrate or decumbent; inflorescences axillary, racemose; fruit with upper margin straight, the lower margin crenate with constriction; stipe < 4 mm long, usually hispid .................................... 3 Leaves (5–)8–15(–20)-foliolate; leaflets obovate to obovate-oblong, (2–) 3–8 mm wide; fruit 3(4)-articulate; articles 2.5–5 × 2–4 mm; stipe 3–4 mm long ............................................................................ A. brasiliana Leaves (11–)15–30-foliolate; leaflets oblong-elliptic, 1–2 mm wide; fruit 2-articulate; articles 2–2.5 × 2–2.5 mm; stipe 1–2 mm long ....... A. histrix Leaflets asymmetrical, appearing to be 2–several-costate ....... A. americana Leaflets asymmetrical or not, 1-costate .................................................... 5 Costa of leaflet excentric; fruit hispid and tuberculate ........... A. fluminensis Costa of leaflets essentially central; fruit glabrous or pubescent, the surface variously raised but not tuberculate ............................................. 6 Plants in general, especially in fruits, blackening on drying; calyx lips entire or subentire ..................................................................................... 7 Plants, including fruits, turning brownish to straw-colored or remaining green on drying; calyx lips clearly lobed or toothed ............................. 8

242

F ABACEAE

7(6). 7.

8(6).

8. 9(8). 9.

Fruit submoniliform at maturity, stipe and basal article separated by a suture; stipe 8–15 mm long; flowers 7–12 mm long ................ A. pratensis Fruit with one margin entire, the other crenate, stipe and basal article continuous, not separated by a suture; stipe 4–8 mm long; flowers 4–9 mm long ..................................................................................... A. sensitiva Flowers (8–)10–15 mm long, standard reflexed; fruit 4–6 mm wide; stipe 3–6(–10) mm long; leaflets entire or with a slight tendency toward marginal trichomes ............................................................................... A. rudis Flowers (4–)5–12 mm long; standard essentially erect; fruit 2.5–4 mm wide; stipe 3–15 mm long; leaflets serrulate-ciliate to entire .............. 9 Stipe of fruit 3–4(–6) mm long; flowers (4–)5–7(–9) mm long; calyx 4–5 mm long ................................................................................................ A. evenia Stipe of fruit (5–)10–15 mm long; flowers 8–12 mm long; calyx 6–9 mm long ................................................................................................ A. scabra

Aeschynomene americana L., Sp. Pl. 715. 1753. —Cujicillo. Mexico, Central America, West Indies, South America, introduced in Africa and Asia; 2 varieties, 1 in Venezuela. A. americana var. americana Herb, somewhat weedy, usually erect, to ca. 2.5 m tall. Wet or moist roadsides, ditches, fields, forest edges, near sea level to 500 m; Delta Amacuro (Caño Manamo near Tucupita), Bolívar (Canaima). Northern and western Venezuela; Mexico, Central America, West Indies, widespread in South America, introduced in Africa and Asia. Aeschynomene brasiliana (Poir.) DC., Prodr. 2: 322. 1825. —Hedysarum brasilianum Poir. in Lam., Encycl. 6: 448. 1804 [1805]. Aeschynomene guarica Pittier, Bol. Técn. Minist. Agric. 5: 41. 1844. Prostrate or decumbent herb. Wet roadsides, savannas, scrub. Mexico, Central America, Cuba, Colombia, Venezuela, Trinidad, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 varieties, all in Venezuela, 2 of these in the flora area.

elsewhere in Venezuela; other distribution as in species. A. brasiliana var. carichanica Rudd, J. Wash. Acad. Sci. 49: 48. 1959. 200–400 m; Bolívar (between Caicara and Maniapure, Cerro Carichana along Río Orinoco, Cerro El Médano 23 km southwest of Caicara). Endemic. ŠFig. 220. Aeschynomene evenia C. Wright in Sauvalle, Anales Acad. Ci. Méd. Habana 5: 334. 1868. West Indies, western Colombia, Venezuela, Trinidad, eastern Brazil; 2 varieties, both in Venezuela, 1 of these in the flora area. A. evenia var. serrulata Rudd, Contr. U.S. Natl. Herb. 32: 61. 1955. —Dormidera, Gusana. Erect herb, often suffrutescent, to ca. 50 cm tall; leaflets serrulate-ciliate. Moist or wet places, near sea level to 100 m; Delta Amacuro (Capure), Bolívar (Ciudad Bolívar), Amazonas (Maroa, San Carlos de Río Negro). Aragua, Barinas, Guárico, Monagas; West Indies, western Colombia, Trinidad, eastern Brazil. ŠFig. 216.

Key to the Varieties of A. brasiliana 1. Flowers (4–)5–8 mm long; segments of fruit 2.5–3 × 2–3 mm ......... var. brasiliana 1. Flowers ca. 10 mm long; segments of fruit 4–5 × 3–4 mm .......... var. carichanica

Aeschynomene fluminensis Vell., Fl. Flumin. 310. 1825 [1829]. Cuba, Hispaniola, Dominican Republic, Colombia, Venezuela, Brazil, Bolivia, Paraguay; 2 varieties, 1 in Venezuela.

A. brasiliana var. brasiliana 50–200 m; northeastern Bolívar, Amazonas (Culebra, Isla Ratón). Widespread

A. fluminensis var. fluminensis Suffrutescent, shrubby herb; stems erect to ca. 4 m tall. In water, wet savannas, river

Aeschynomene 243

banks, beaches, 50–300 m; eastern Bolívar (between Caicara and Río Cuchivero, between Tumeremo and El Dorado). Cuba, Dominican Republic, Brazil, Bolivia, Paraguay. Aeschynomene histrix Poir. in Lam., Encycl. suppl. 4: 77. 1816. Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; 5 varieties, 3 in Venezuela, 1 of these in the flora area. A. histrix var. histrix Herb, commonly prostrate, sometimes erect; stems to ca. 1 m long. Disturbed areas, savannas, forest edges, 50–300 m; Delta Amacuro (between Piacoa and Castillos de Guayana), northern Bolívar, Amazonas (around and north of Puerto Ayacucho). Anzoátegui, Guárico; Guatemala, Honduras, El Salvador, Nicaragua, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Paraguay. ŠFig. 217. Some specimens trend toward var. incana (Vogel) Benth., and some have been previously cited as var. incana, but in the strict sense, var. incana occurs in southern South America, particularly in Argentina and Uruguay.

Aeschynomene paniculata Willd. ex Vogel, Linnaea 12: 95. 1838. Suffrutescent, erect herb to ca. 2.5 m tall. Rocky slopes, savannas, open woods, roadsides, 100–500 m; northern Bolívar, Amazonas (Puerto Ayacucho). Anzoátegui, Aragua, Trujillo; widespread in Mexico, Central America, Colombia, Guyana, Suriname, Brazil, Bolivia. Aeschynomene pratensis Small, Bull. New York Bot. Gard. 3: 423. 1905. Nicaragua, Costa Rica, Panama, Cuba, Hispaniola, Colombia, northern Brazil, Bolivia; 2 varieties, 1 in Venezuela. A. pratensis var. caribaea Rudd, Contr. U.S. Natl. Herb. 32: 47. 1955. Erect herb, sometimes suffrutescent; flowers 7–8 mm long; calyx 4–5 mm long; fruit with articles 5–6 × 4–5 mm; stipe 8–10 mm long. Marshy areas, river banks, 100– 800 m; Bolívar (La Paragua, Santa Elena de Uairén). Anzoátegui, Cojedes, Trujillo; Nicaragua, Costa Rica, Panama, Cuba, Hispaniola, Colombia, Guyana, northern Brazil, Bolivia. ŠFig. 219. Aeschynomene rudis Benth., Pl. Hartw. 116. 1839 [1843]. Erect herb to ca. 2 m tall. Moist or wet places, river banks or in water, ca. 50 m;

Fig. 216. Aeschynomene evenia var. serrulata

Fig. 217. Aeschynomene histrix var. histrix

244

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Fig. 218. Aeschynomene scabra

Fig. 219. Aeschynomene pratensis var. caribaea

Fig. 220. Aeschynomene brasiliana var. carichanica

Aldina 245

Bolívar (Ciudad Bolívar). Guárico; widely distributed from southern U.S.A., Mexico, Central America, Cuba, Colombia, Brazil, Bolivia, to Argentina. Aeschynomene scabra G. Don, Gen. Hist. 2: 284. 1832. Herb, sometimes suffrutescent, to ca. 3 m tall. Swamps, humid savannas, 50–300 m; Bolívar (Ciudad Bolívar, Río Orinoco between Río Horeda and Cerro Carichana). Mexico, Central America, Ecuador, Peru, Brazil, Bolivia, Paraguay. ŠFig. 218. The two collections from the flora area (Wurdack & Monachino 39868, F, NY, US; Wurdack & Monachino 39941, NY, US, VEN) were previously referred to Aeschynomene rostrata Benth., a species known only from Bahia, Brazil. The leaflets on specimens from the flora area have more serrulate-ciliate mar-

gins than most collections, and thus A. scabra seems a better determination. Aeschynomene sensitiva Sw., Prodr. 107. 1788. Southern Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 3 varieties, 1 in Venezuela. A. sensitiva var. sensitiva Erect, suffrutescent herb 1–2(–4) m tall. In water or on wet river banks and mud flats, near sea level to 100 m; Delta Amacuro (Caño Araguao, Caño Güiniquina, Sacupana, between Tucupita and Los Güires), Bolívar (Ciudad Bolívar, south of Tumeremo). Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Táchira; other distribution as in species.

4. ALDINA Endl., Gen. Pl. 1322. 1840, nom. cons. Allania Benth., J. Bot. (Hooker) 2: 91. 1840. by Basil Stergios and Richard S. Cowan Trees. Leaves odd-pinnate or the upper sometimes 1-foliolate, exstipulate; leaflets opposite to subopposite, blades mostly coriaceous to rigid with rounded to cordate base. Inflorescences terminal or axillary to leaves nearest end of the branch, racemose or paniculate-racemose; bracts and bracteoles minute; pedicels short, 2bracteolate apically. Calyx entire in bud, opening about halfway in 2–5 recurved or reflexed segments, tube turbinate; petals (3)4–6, elliptic to oblanceolate, obovate or oblong, usually concave to cucullate, white to yellowish, glabrous. Stamens numerous, glabrous; filaments borne on margin of short hypanthial cup adnate to calyx base; anthers large, uniform, narrowly oblong to linear, opening lengthwise by slits. Gynophore arising from the bottom of hypanthium cup, swollen-articulate at junction with the incurved, somewhat oblong ovary; ovary usually 1, but potentially 2 or 3; style subulate, straight or usually apically curved; stigma punctiform. Fruit to 10 cm long, dehiscent, thick, woody, ovoid, and 1-seeded or else oblong-fusiform and 2– 4-seeded. Colombia, Venezuela, Guyana, northwestern Brazil; 22 species, 18 in Venezuela, all in the flora area. The genus is currently under revision by the senior author. The treatment here of the published taxa is largely an extract from an earlier, preliminary treatment. Key to the Species of Aldina 1. 1. 2(1).

Leaflets completely glabrous on all surfaces; racemes simple or multiple ................................................................................................................ 2 Leaflets pubescent on lower surface, sometimes minutely so; inflorescence an elongated, open, multiple raceme or terminal panicle of racemes ..... 8 Flowers showy, calyx tube 2.5–3 cm, petals 3–4 × 2 cm; ovaries 2 or 3 on the stipe; fruits elongated, sausage-shaped to thickly fusiform and no-

246

FABACEAE

ticeably constricted between the 2–4 seeds; leaflets thickly coriaceous, almost cartilaginous, noticeably cordate at the base ..................... A. sp. A 2. Flowers half again as large or smaller than above; ovary solitary on stipe; fruits ovoid to oblong, 1- or 2-seeded; leaflets chartaceous to ± coriaceous, but not cordate at the base ......................................................... 3 3(2). Leaflets pallid and waxy on lower surface, broadly lanceolate to acuminate, 10–11 × 4–4.5 cm; calyx and unopened buds very sparsely whiteappressed-puberulent near the base or usually glabrous; calyx tube truncate, indentedly barrel-shaped at the base; pedicels 11–12 mm long; bracteoles 3–3.5 mm below base of calyx tube ....................... A. sp. B 3. Leaflets shiny on lower surface, without a waxy coating, oblong to ovateelliptic, obtuse-mucronate, (8–)9–14 × 4.5–6 cm; calyx and unopened buds uniformly and usually densely pubescent; calyx tube truncate, claviform at the base; pedicels 1–3 mm long; bracteoles 0–1 mm below base of calyx tube ................................................................................... 4 4(3). Leaflets noticeably tessellate with dark red lines on lower surface; mature buds 15–17 × 8 mm; racemes simple and axillary or occasionally paired .......................................................................................... A. latifolia 4. Leaflets not tessellate, smooth and homogeneous on lower surface; mature buds 9–11 × 5–7 mm; racemes multiple-branched or in small panicles ................................................................................................... 5 5(4). Leaflets glaucous or with an ash-gray-green bloom on lower surface; ovary mostly glabrous .................................................................... A. berryi 5. Leaflets entirely shiny on both surfaces, not at all discolorous; ovary pubescent .................................................................................................... 6 6(5). Leaves 1- or 3-foliolate ............................................................ A. heterophylla 6. Leaves 5- or 7-foliolate ............................................................................... 7 7(6). Leaflets coriaceous, 7–8 × 5–6 cm, elliptic, mucronate with no extended acumen; inflorescence and flower buds ferruginous-velutinous, with a golden brown lustre ......................................................................... A. sp. C 7. Leaflets chartaceous, 11–13 × 4.5–5.5 cm, lanceolate to obtuse, acuminate, the acumen 1–1.5 cm long; inflorescence and flower buds tanbrown, appressed-pubescent .......................................................... A. sp. G 8(1). Ovary glabrous or nearly so ....................................................................... 9 8. Ovary either densely and uniformly pubescent, or densely golden-brownpuberulent on lower, dorsal portion only, and either ventral suture or both sutures glabrous .......................................................................... 10 9(8). Pistil as long as or longer than the stamen filaments at anthesis; leaflets pruinulose-lepidote on lower surface, with semierect or reclining trichomes; petals 5, 18–20 × 6–6.5 mm .............................................. A. sp. D 9. Pistil noticeably shorter than the stamen filaments at anthesis; leaflets appressed-puberulent, subglaucous with a uniform bloom on lower surface; petals 4, 20–25 × ca. 10 mm ....................................... A. macrophylla 10(8). Leaflets rich erect-brown-villous on lower surface; racemes and flower buds notably dark, golden-brown-velutinous; calyx 2-lobed at anthesis ........................................................................................... A. kunhardtiana 10. Leaflets appressed-puberulent, either pale lilac or with a pruinose, waxy bloom on lower surface; racemes and flower buds tan-brown, appressed-puberulent; calyx 3–5-lobed at anthesis ................................ 11 11(10). Stipe ≥ 10 mm long, 2 or more times as long as the ovary at anthesis .. 12

Aldina 247

11. Stipe 2–2.5 mm long, never longer than the ovary at anthesis .............. 13 12(11). Leaflets 10–12 × 6–7.5 cm, obtuse-mucronulate to rounded, pale lilac on lower surface with a pruinulose, waxy bloom; calyx tube cupular, 7–8 × 9 mm wide at anthesis; petals 15 mm long and strongly cucullate .......................................................................................................... A. sp. E 12. Leaflets 16–17 × 7–8 cm, obtuse-acuminate, monocolorous and nearly shiny, waxy bloom not prominent on lower surface; calyx tube funnelshaped, 10 × 8–9 mm at anthesis; petals 20–25 mm long and somewhat flattened ...................................................................................... A. latifolia 13(11). Petals 3 or 4 .............................................................................................. 14 13. Petals 5 or 6 .............................................................................................. 15 14(13). Leaflet venation stongly elevated, conspicuous on both surfaces; petals 2–2.5 × 1–1.8 cm; leaflets oblong to oblong-ovate, 7.5–18 × 4.5–9 cm, scantily appressed-puberulent but shiny and the same color on the lower surface ............................................................................ A. reticulata 14. Leaflet venation inconspicuous, usually obscure on upper surface; petals 1–2 cm long; leaflets elliptic, obtuse to rounded at the apex, 8–9 × 5–6 cm, dark wine-colored on the lower surface .......................................... A. sp. F 15(13). Leaflets acute to acuminate apically, upper and lower surfaces markedly discolorous ............................................................................................ 16 15. Leaflets obtuse to rounded-obtuse, upper and lower surfaces not markedly discolorous .................................................................................... 17 16(15). Leaflets tessellate and appressed-puberulent on lower surface; petals 5, 2–2.3 cm long .......................................................................... A. discolor 16. Leaflets not tessellate, golden-pubescent on lower surface; petals 6, ca. 1.8 cm long ...................................................................................... A. aurea 17(15). Leaf axis and petiole appressed-puberulent, petiolules 5–7 mm long; petals 6 ............................................................................................. A. elliptica 17. Leaf axis and petiole glabrous when mature, petiolules 10–25 mm long; petals 5 ................................................................................................. 18 18(17). Leaflets ovate to orbicular, 7–15 × 6–14 cm, lateral veins conspicuous, 7–13 per side; racemes paired, 7–19-flowered ....................... A. paulberryi 18. Leaflets elliptical, 8.5–10 × 3.5–4.5 cm, lateral veins inconspicuous, 22–25 per side; racemes paniculate, many-flowered, elongate ....... A. petiolulata Aldina aurea R.S. Cowan, Mem. New York Bot. Gard. 10(1): 145. 1958. Tree 3–4 m tall; leaflets discolorous, pallid on lower surface, apically acuteacuminate; ovary densely pubescent; petals 6, ca. 1.8 cm long. Montane forests, 500–1300 m; Amazonas (Cerro Yutajé). Endemic. Aldina berryi R.S. Cowan & Steyerm., Ann. Missouri Bot. Gard. 71: 312. 1984. Tree ca. 20 m tall; leaflets completely glabrous, shiny on both surfaces, ± coriaceous; petals 5–7, white; ovary glabrous. Montane and gallery forests, 800–1000 m; Bolívar (near Cerro Guaiquinima, Río Chiguao). Endemic.

Aldina discolor Spruce ex Benth. in Mart., Fl. Bras. 15(2): 12. 1870. —Cojón de verraco. Tree 18–25 m tall; leaflets completely glabrous, discolorous, tessellate, appressed-puberulent on lower surface; inflorescence in multiple, axillary racemes; petals 5, 2–2.3 cm long, yellowish. Nonflooded evergreen lowland forests, 100–200 m; Amazonas (Yavita). Brazil (Amazonas). Aldina elliptica R.S. Cowan, Mem. New York Bot. Gard. 10(1): 146. 1958. Tree 10–25 m tall; leaf stalk including petiole appressed-puberulent into maturity, petiolules 5–7 mm; leaflets appressed-pubes-

248

FABACEAE

cent on lower surface, obtuse to rounded-obtuse, not discolorous; petals 6, ca. 1.5 cm long; ovary uniformly pubescent, not exserted at anthesis. Tepui slope forests, ca. 1800 m; Amazonas (Cerro Yutajé). Endemic. Aldina heterophylla Spruce ex Benth. in Mart., Fl. Bras. 15(2): 13. 1870. Small tree (in the flora area) ca. 2 m tall; leaves 5-foliolate; leaflets glabrous and shiny, ovate-elliptic, obtuse to fairly mucronate, 8–10 × 4–5 cm; ovary golden-sericeous; fruit 1–4 cm long, irregularly shaped. White-sand shrub savannas, ca. 500 m; Amazonas (12 km west-southwest of Cerro Autana). Brazil. Aldina kunhardtiana R.S. Cowan, Mem. New York Bot. Gard. 8: 106. 1958. Small (10 m tall) to giant (35 m tall) tree; leaflets “lauraceous” with pronounced obtuse tip, the lower surface with distinctive rich, brown-villous pubescence; inflorescence an elongate, open, terminal panicle of racemes, dark, golden-brown-velutinous; fruits globose, to 8 cm diameter. Evergreen lowland forests, forest-savanna ecotones, 100–200 m; Amazonas (Río Casiquiare, from Río Sipapo south to San Fernando de Atabapo, upper Río Temi, San Carlos de Río Negro to Solano, Yavita to Maroa road). Brazil. ŠFig. 222. Aldina latifolia Spruce ex Benth. in Mart., Fl. Bras. 15(2): 12. 1870. Tree 5–17 m tall. Southern Venezuela, adjacent Brazil; 3 varieties, all in the flora area. Key to the Varieties of A. latifolia 1. Leaflets appressed-puberulent on the lower surface ............... var. pubescens 1. Leaflets glabrous on both surfaces ........ 2 2. Leaves 1- or 3-foliolate .............................. ........................... var. auyantepuiensis 2. Leaves mostly 3- or 5-foliolate .................. ........................................ var. latifolia A. latifolia var. auyantepuiensis Pittier ex H.S. Irwin, Acta Bot. Venez. 2: fig. 13. 1967. —Automoyek (Arekuna). Tree 5–8 m tall. Dwarf montane forests, 700–1100 m; Bolívar (Guayaraca, east-northeast of Icabarú, Kamarata, Río Carapo along Cerro Guaiquinima). Endemic. ŠFig. 221.

This is perhaps the most weakly defined variety, for the leaves at the apex of the branchlets often have a reduced number of leaflets. A. latifolia var. latifolia. —Cojón de verraco. Tree 5–15 m tall. Lowland, usually riparian forests, white-sand savannas, 100–200 m; Amazonas (Maroa, Pimichín, Río Autana, Río Guayapo). Endemic. A. latifolia var. pubescens R.S. Cowan, Mem. New York Bot. Gard. 8: 107. 1953. Bushy tree ca. 8 m tall. White-sand savannas at edges of riparian forests, 100–200 m; Amazonas (Caño Cupueni near San Fernando de Atabapo, near Yavita). Brazil (Amazonas). Aldina macrophylla Spruce ex Benth. in Mart., Fl. Bras. 15(2): 13. 1870. Tree to 17 m tall; leaflets appressed-pubescent on lower surface; ovary glabrous or nearly so. Venezuela; 2 varieties, both in the flora area. Endemic. Key to the Varieties of A. macrophylla 1. Stipe 6–10 mm with ovary noticeably exserted in mature flower; calyx tube cupulate, 6–8 × (6–)7–9 mm; petals 18–25 mm long; bracteoles inserted 1.5–2 mm below base of calyx tube; stipe completely and densely appressed-puberulent .......................... var. macrophylla 1. Stipe ≤ 2 mm with ovary not exserted in mature flower; calyx tube 3–4 × 5–6 mm; petals 8–10 × 4–5 mm; bracteoles inserted at base of calyx tube; stipe pubescence restricted to a central circular band ....................... var. yapacanensis A. macrophylla var. macrophylla Tree to 17 m tall. Riparian forests, whitesand savannas, 100–200 m; Amazonas (Caño Chimoni in Río Siapa basin, Caño San Miguel, Río Yatúa). Endemic. ŠFig. 224. A. macrophylla var. yapacanensis (R.S. Cowan) Stergios, comb. & stat. nov. —Aldina yapacanensis R.S. Cowan, Mem. New York Bot. Gard. 8: 108. 1953. Tree 8–10 m tall; flowers brownish to white; ovary generally not exserted in ma-

Aldina 249

ture flowers; calyx funnel-shaped, 3–4 × 5–6 mm; bracteoles inserted at base of calyx tube; pubescence on stipe restricted to a band in the middle. Black-water riparian and swamp forests, 100–200 m; Amazonas (Caño Yapacana, Río Emoni in lower Río Siapa basin). Endemic. This variety was reduced to synonomy by Cowan (Mem. New York Bot. Gard. 10(1): 146–147. 1958), because of relatively high variability in leaflet size and shape, and in variation in petal and stipe size as flowers opened and matured. However, with further scrutiny and an additional, copiously flowered collection, the ovary was found to consistently not be exserted. Also, as shown by the type collection, the bracteoles are inserted 1.5–2 mm below the base of the calyx tube in var. macrophylla, but in var. yapacanensis, they are at the base. In var. yapacanensis, the pubescence on the stipe is restricted to a band in the middle. Although these characters may seem weak, they furnish more reliable criteria than those distinguishing some varieties of Aldina latifolia. Aldina paulberryi Aymard, Novon 8: 330. 1998. Tree ca. 15 m tall; leaves 1- or 3-foliolate; leaflets ovate-orbicular, the apex rounded-obtuse, 7– 15 × 6–14 cm, not discolorous, lateral veins 7–13 per side; racemes 2–many, 7–19-flowered. Evergreen lowland forests, 100–200 m; Amazonas (near Macuruco on upper Río Orinoco). Endemic. ŠFig. 225. This species can be distinguished from its nearest apparent relative, Aldina petiolulata, by orbicular, not elliptical leaflets, more prominent but less numerous lateral veins, and paniculate, fewer flowered, shorter racemes. Aldina petiolulata R.S. Cowan, Mem. New York Bot. Gard. 10(4): 70, fig. 59. 1961. Tree to 40 m tall; leaflets appressed-puberulent on lower surface, elliptical, with 22–25 lateral veins; inflorescence a manyflowered panicle of elongate racemes. Montane forests, 700–800 m; Amazonas (Sierra de la Neblina). Endemic. Aldina reticulata R.S. Cowan, Mem. New York Bot. Gard. 8: 107. 1953. Small tree; lower surface of leaflets and ovary appressed-puberulent; venation

strongly elevated and conspicuous on both surfaces; stipe ≤ ovary length; petals 4. Lower montane forests, lowland swampy forests, 100–800 m; Amazonas (headwaters of Caño Grande in Cerro Sipapo, Maroa area, Río Temi). Endemic. ŠFig. 223. Aldina sp. A Shrubby tree 2–6(–8) m tall; leaflets glabrous, cartilaginous-coriaceous, cordate at the base, 7–8 × 4–5 cm; flowers showy, petals 3–4 × 2 cm; ovaries 2 or 3 on a single stipe; fruits thickly fusiform, 2–4-seeded. Open, low-rocky, shrubby slopes, 400–1000 m; Bolívar (Cerro Kampe, middle Río Paragua basin, Serranía Senkopirén). Endemic. This spectacular species is probably the most distinct one in the genus, easily recognized by very large flowers with a long-exserted, thick stipe bearing 2 or 3 ovaries. Aldina sp. B Tree 8–10 m tall; leaflets 10–11 × 4–4.5 cm; calyx and unopened buds usually glabrous, but sometimes sparsely white-appressed-puberulent near the base; pedicels 11–12 mm long; bracteoles borne 3–3.5 mm below the base of the calyx tube. Humid, lower montane forests, ca. 500 m; Bolívar (Quebrada El Trueno 90 km south of La Paragua and at the base of Cerro Guaiquinima). Endemic. This species is readily distinguished by the glabrous, broadly lanceolate-acuminate leaflets that have an off-white, pallid-waxy coating on the lower surface. It is probably closest to Aldina latifolia, which also has simple, axillary racemes and densely appressed-tomentose stipe and ovary. Mature flowers of both species have a noticeably exserted pistil. Aldina sp. C Shrubby tree 2–5 m tall, unique by the combination of glabrous leaflets and branched, multiple racemes; leaflets ovate-elliptic, mucronate, coriaceous; inflorescence appressedferruginous-tomentose; ovary densely puberulent-tomentose. White-sand shrub savannas, forest-savanna ecotones, 100–1300 m; Amazonas (Cerro Autana, Cerro Parú). Aldina sp. C is closely related to A. latifolia, A. berryi, and A. heterophylla, all of which have shiny, glabrous leaflets without the waxy coating on the lower surface. It can be distinguished from A. latifolia by the non-

250

FABACEAE

tessellate leaflets and multiple-branched racemes; from A. berryi by the entirely shiny leaflets and pubescent ovary; and from A. heterophylla by the 5–7 leaflets and lustrous, ferruginous pubescence. Aldina sp. D Tree ca. 10 m tall; leaflets glabrous with a glaucous, somewhat ashy bloom on the lower surface, ovate-elliptic, obtuse-rounded at the apex, 9–11 × 6–6.5 cm; ovary glabrous. Semiblack-water, riparian swamp forests, 100– 200 m; Amazonas (Río Emoni in the lower Río Siapa basin). Endemic. This species most resembles Aldina berryi in its 4 petals and glabrous ovary, but differs in the more notably pallid-discolorous, obtuse to rounded leaflets, in the longer, exserted pistil, and in the longer pedicellate but smaller flowers. Aldina sp. E Tree 4–6 m tall, occasionally 10–30 m tall. Riparian or swamp forests, 100–200 m; Amazonas (Caño Momoni in the Río Casiquiare basin, Caño Yagua, Macuruco, Río Negro). Endemic. This species is similar to Aldina macrophylla var. pubescens in the 3–5-lobed calyx and the long stipe, but differs in having smaller, obtuse-mucronate to rounded, discolorous leaflets.

Fig. 221. Aldina latifolia var. auyantepuiensis

Aldina sp. F Tree 3–6 m tall. Swamp forests, flooded riparian alluvial plains, 50–200 m; Amazonas (Laguna Pasiba, Río Negro). Endemic. Aldina sp. F and and A. reticulata are the only two species that have 3 or 4 petals instead of the usual 5 or 6. Aldina sp. F differs from A. reticulata in the obscure venation on the upper leaflet surface, more obtuse, ovateelliptic leaflets, smaller flowers, and a dark, wine-colored lower leaflet surface. Aldina sp. G Medium-sized tree; leaves 5- or 7foliolate; leaflets completely glabrous on both surfaces and shiny without any waxy coating on lower surfaces and concolorous, chartaceous; petiolules 4–6 mm long; inflorescence and flower buds tan-brown, appressed-puberulent. Savannas, 800–1000 m; Amazonas (Cerro Sipapo). Colombia (Vaupés), Brazil (Amazonas: Manaus) . This species is most closely related to Aldina sp. C, both with 5- or 7-foliolate leaves, entirely shiny, discolorous leaflets, and pubescent ovary. It can be distinguished from Aldina sp. C by its chartaceous rather than coriaceous leaflets, which are obtuse and acuminate rather than elliptic and mucronate, the inflorescence and flower buds are brown-tan appressed-puberulent rather than loosely ferruginous-velutinous.

Aldina 251

Fig. 222. Aldina kunhardtiana

252

FABACEAE

Fig. 223. Aldina reticulata

Fig. 224. Aldina macrophylla var. macrophylla

Aldina 253

Fig. 225. Aldina paulberryi

254

F ABACEAE

5. ALEXA Moq. in A. DC., Prodr. 13(2): 168. 1849. Alexandra R.H. Schomb., London J. Bot. 4: 12. 1845, non Alexandra Bunge 1843. by Charles H. Stirton and Gerardo A. Aymard C. Trees 10–36 m tall, rarely cauliflorous. Stipules minute, rapidly caducous. Leaves 5–17-foliolate, odd-pinnate, alternate. Leaflets membranaceous or coriaceous. Inflorescences terminal, racemes short to very elongate, becoming quite woody with age. Flowers white, green, cream, yellow, orange, or crimson, strongly zygomorphic, pedicellate, bracteate (the bracts sometimes swollen and glandular); bracteoles minute and caducous; pearl glands absent. Calyx turbinate-campanulate to campanulate, incurved or erect, densely pubescent, teeth 3–5, minute. Petals free, erect or ascending; standard outermost, elliptic to obovate, base truncate, apex emarginate, densely pubescent or glabrous; wing and keel petals scarcely differentiated, mostly falcate. Stamens 10–12, equal, free, staminodes sometimes present; anthers dorsifixed, versatile, longitudinally dehiscent, elongate. Ovary stipitate, 4– 9-ovulate, glabrous or pubescent; style filiform; stigma inconspicuous, terminal. Legume dehiscent, woody, apex acute, calyx woody and persistent in fruit, densely reddish brown, glabrescent, 1–7-seeded. Seeds swollen, ovate or elliptic, glabrous. Venezuela, Guyana, Suriname, French Guiana, Brazil; 10 species, 6 in Venezuela, all in the flora area. Key to the Species of Alexa 1.

1.

2(1). 2.

3(1). 3. 4(3) 4. 5(3). 5.

Leaves membranaceous, with well-developed drip tips; flowers tubular, but petals reflexing very quickly; flower bracts swollen and glandular; corolla cream to greenish white ............................................................ 2 Leaflets subcoriaceous, with drip tips absent or scarcely developed; flowers typically papilionoid without petals reflexing; flower bracts not swollen and glandular; corolla pale yellow to crimson and orange ..... 3 Inflorescence dense; stipe of ovary glabrous; calyx as wide as long; style finely pubescent, to 2 cm long, uncinate at the apex ................. A. cowanii Inflorescence lax; stipe of ovary pubescent; calyx 2–3 times longer than broad; style glabrous, 6–8 cm long, curved at the middle ...................... ........................................................................................ A. canaracunensis Leaflets densely yellow- to brown-puberulous and opaque on the upper surface .................................................................................................... 4 Leaflets glabrous or glabrescent and shiny on the upper surface ............ 5 Leaflets 7 or 9, densely pubescent on the lower surface only; pedicels 2–3.5 cm long ..................................................................... A. bauhiniiflora Leaflets 11 or 13(15, 17), pubescent on both surfaces; pedicels 0.4–1.5 cm long .............................................................................................. A. confusa Inflorescences in terminal racemes 10–21 cm long; corolla densely pubescent, pale yellow; petals membranous ................................ A. herminiana Inflorescences borne on trunk, to 7 cm long; corolla glabrous, crimson to orange; petals subcoriaceous ............................................... A. imperatricis

Alexa 255

Fig. 226. Alexa cowanii

256

F ABACEAE

Fig. 227. Alexa herminiana

Alexa 257

Fig. 228. Alexa imperatricis

258

F ABACEAE

Alexa bauhiniiflora Ducke, Bull. Mus. Hist. Nat. (Paris), sér. 2: 732. 1932. Tree 5–10 m tall. Along rivers, 100–500 m; Amazonas (Río Mawarinuma, Río Paduari, headwaters of Río Siapa, Sierra de la Neblina). Brazil (Amazonas, Pará).

guaza, Río Tabaro, Río Tonoro, Santa María de Erebato, Serranía Pia-Zoi), Amazonas (Ocamo, upper Río Cunucunuma, Río Padamo). Guyana, Brazil (Amazonas, Roraima). Some branches become hollowed out and are occupied by stinging ants.

Alexa canaracunensis Pittier, Bol. Soc. Venez. Ci. Nat. 7: 308. 1942. —Caicareño, Kadaka (Arekuna), Kayapa, Tinajito, Tunadi (Yekwana). Small tree 3–12 m tall, rarely taller; leaves firmly membranaceous; corolla greenish cream. Evergreen lowland to montane forests, 100–1500 m; Bolívar (Amaruaytepui, Cerro Paují, El Dorado, Las Claritas, Macizo del Chimantá [Chimantá-tepui], Río Canaracuni, Río Caura, Río Cuyuní, Río Icabarú, Río Padauri, upper Río Paragua, Río Tírica), Amazonas (Sierra Parima). Brazil (Roraima). Young stems subtending the inflorescences become hollowed out and occupied by ants, which feed on nectar provided by swollen pear-shaped flower bracts.

Alexa cowanii Yakovlev, Bot. Zurn. (Moscow & Leningrad) 62(3): 436. 1977. —Tinajito. Tree to 20 m tall; leaves membranaceous, glabrous; corolla white to pale greenish white. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (near El Dorado, along the Guyana border, Kilómetro 88, Río Uiri-yuk near La Escalera). Guyana (upper reaches of the Kamarang River watershed). ŠFig. 226. The branches often become hollowed out and are occupied by ants.

Alexa confusa Pittier, Bol. Soc. Venez. Ci. Nat. 8: 262. 1943. —Caicareño montañero, Dequira-arepillo (Piaroa), Guamo peludo, Kure-yek (Arekuna), Leche de cochino, Pitopitosiji (Yanomami), Tuneene (Yekwana). Alexa superba R.S. Cowan, Mem. New York Bot. Gard. 8: 117. 1953. Alexa imperatricis auct. non R.H. Schomb. 1845, nec Baill. 1870. Tree 10–30 m tall; leaves subcoriaceous, the upper surface dark green, the lower surface yellowish green and densely pubescent; corolla cream-colored on inner faces, buff brown outside. Evergreen lowland to lower montane forests, occasionally along savanna borders, 50–1300 m; Bolívar (Icabarú, Río Caura, Río Karún, Río Pacairao, Río Par-

Alexa herminiana N. Ramírez, Ernstia 37: 41. 1986. Tree 10–20 m tall; leaves coriaceous, the upper surface rich green, the lower surface paler; corolla pale yellow. Evergreen lowland to lower montane forests, 50–500 m; Amazonas (Caño Piedra on Cerro Sipapo massif, Raudal Rabipelado near Samariapo, Río Cataniapo). Endemic. ŠFig. 227. Alexa imperatricis (R.H. Schomb.) Baill., Hist. Pl. 2: 362. 1870. —Alexandra imperatricis R.H. Schomb., London J. Bot. 4: 12. 1845, non Baker 1888, nec Pittier 1942. —Coreco, Leche de cochino. Tree 30–40 m tall, cauliflorous; leaves coriaceous, slightly glossy, the upper surface darker green; corolla orange, one petal and stamens crimson to orange. Evergreen lowland forests, 50–300 m; Delta Amacuro (eastnortheast of El Palmar, Río Toro, Serranía de Imataca), Bolívar (El Dorado, Kilómetro 88, Río Cuyuní, Río Toro). Adjacent Guyana. ŠFig. 228.

6. ALYSICARPUS Desv., J. Bot. Agric. 1: 120. 1813. by Nidia L. Cuello A. Herbs, suberect to prostrate; stems ascending to procumbent, terete, striate. Leaves alternate, 1(–3)-foliolate; leaflets linear-lanceolate to ovate, reticulatevenose; stipules scarious, acuminate, free or connate; bistipellate. Inflorescence terminal or rarely axillary, racemose; bracts scarious, deciduous; pedicels paired, short. Flowers small. Calyx deeply 5-lobed, the lobes unequal, the upper 2 connate nearly to the apex, glumaceous; corolla reddish purple, rarely orange; standard obovate to

Andira 259

orbicular, clawed; wing petals obliquely oblong; keel petals slightly incurved, obtuse. Stamens 10, diadelphous, the upper connate only basally; anthers uniform. Ovary sessile or shortly stipitate; ovules numerous; style filiform, incurved at the apex; stigma broadly capitate, terminal. Fruit a loment, terete to compressed, submoniliform, indehiscent. Seeds suborbicular or globose, smooth, lustrous, estrophiolate. Old World tropics; 25 species, 1 introduced into tropical America as forage, 1 in Venezuela. Alysicarpus vaginalis (L.) DC., Prodr. 2: 353. 1825. —Hedysarum vaginale L., Sp. Pl. 746. 1753. Annual herb, suberect to prostrate. In Trachypogon savannas on low hills and disturbed areas, 200–500 m; Delta Amacuro (Tucupita), north-central and eastern Bolívar (Altiplanicie de Nuria, 16 km north of El Manteco, El Pao, Tumeremo). Aragua, Barinas, Guárico, Miranda, Portuguesa; U.S.A. (Florida), Mexico, Honduras, Panama, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, India, Asia, Oceania, Madagascar. ŠFig. 229.

Fig. 229. Alysicarpus vaginalis

7. ANDIRA Juss., Gen. Pl. 363. 1789, nom. cons. by Gerardo A. Aymard C., Nidia L. Cuello A., and R. Toby Pennington Trees or shrubs. Leaves alternate, odd-pinnate, rarely trifoliolate or unifoliolate; leaflets opposite or rarely alternate; stipels setaceous, persistent or absent. Inflorescences terminal and axillary panicles, often crowded and subsessile. Flowers rose, violet, or white; bracts and bracteoles small, caducous. Calyx truncate or shortly 5-dentate; petals long-clawed; standard suborbicular, exappendiculate; wing petals straight, oblong, free; keel petals similar, overlapping at the back. Stamens 10, diadelphous; vexillar stamen free; anthers ovate, small, versatile; ovary stipitate, ovules (1)2–8; style short, incurved; stigma small, terminal. Fruit drupaceous, ovoid, obovoid, or globose, indehiscent, shortly stipitate with fleshy mesocarp and hard, woody endocarp. Seeds 1(–3), large, filling the entire cavity, exalbuminate. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa (1 species); ca. 30 species, 5 in Venezuela, all in the flora area.

260

F ABACEAE

Key to the Species of Andira 1. 1. 2(1). 2. 3(2). 3. 4(2). 4.

Leaves all trifoliolate ................................................................. A. trifoliolata Leaves with 5–15 leaflets (occasionally some leaves trifoliolate) ............ 2 Leaves with 5–7 leaflets (occasionally trifoliolate) ................................... 3 Leaves with 9–15 leaflets (never trifoliolate) ............................................ 4 Leaflets minutely pubescent on lower surface, the apex rounded to emarginate ................................................................................... A. tervequinata Leaflets glabrous on lower surface, the apex cuspidate ..................... A. sp. A Branches and branchlets glabrous; leaflets glabrous, the apex acuminate; stipules 5–10 mm long ................................................................ A. inermis Branches and branchlets brown-tomentose; leaflets minutely pubescent on lower surface, the apex obtuse or retuse; stipules ca. 3 mm long ............................................................................................ A. surinamensis

Andira inermis (W. Wright) DC., Prod. 2: 475. 1825. —Geoffroea inermis W. Wright, London Med. J. 8: 256. 1787. —Pilón. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina; 2 subspecies, 1 in Venezuela.

white spot. Deciduous to evergreen lowland forests, 50–100 m; Delta Amacuro (Río Cuyubini), Bolívar (southwest of Caicara, Maniapure). Barinas, Distrito Federal, Falcón, Miranda, Portuguesa, Yaracuy, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, cultivated elsewhere in the tropics. ŠFig. 230.

A. inermis subsp. inermis Tree to 35 m tall; stipules linear; petals pink or purplish, the standard with a central

Fig. 230. Andira inermis subsp. inermis

Andira 261

Fig. 231. Andira surinamensis

Fig. 232. Andira trifoliolata

262

F ABACEAE

Andira surinamensis (Bondt) Splitg. ex Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 60. 1939. —Geoffroea surinamensis Bondt, Cort. Geoffr. Surinam. 13, figs. 1–8. 1788. —Arisoru (Warao), Canelito negro, Palo blanco, Pilón, Pilón rebalsero, Sobó (Yekwana), Winka, Wonka (Panare). Geoffroea retusa Poir. in Lam., Encycl. 8: 182. 1808. —Andira retusa (Poir.) DC., Prod. 2: 475. 1825. Tree to 40 m tall; petals pink to violet, the standard with a central white spot. Deciduous to evergreen and riparian forests, sometimes on granitic outcrops, 50–400 m; Delta Amacuro (Caño Arature, east-northeast of El Palmar), northern Bolívar, Amazonas (Caño Yutajé, Capibara, La Esmeralda, Puerto Ayacucho, Río Casiquiare, Río Cataniapo, Samariapo). Anzoátegui, Apure, Barinas, Táchira; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 231. The fruits are used by the Warao Amerindians to treat skin diseases. Andira tervequinata R.T. Penn., Aymard & Cuello, Novon 7: 72. 1997. Tree to 20 m tall. Lower montane forests

with sandy savannas, 300–500 m; Bolívar (Río Aparamán at base of Amaruay-tepui, banks of Río Trueno at base of Cerro Guaiquinima). Endemic. Andira trifoliolata Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938. —Congrio. Tree to 20(–30) m tall; petals white or dull flesh-colored. Evergreen lowland forests, edges of white-sand savannas, riparian forests, 100–200 m; Amazonas (Caño San Miguel, lower Río Baría, Río Casiquiare, San Carlos de Río Negro, Santa Bárbara, Solano). Brazil (Amazonas). ŠFig. 232. A collection currently included here, Gentry and Tillett 10889 (FHO, MO), may represent a new species. It is a shrub, while all other records of Andira trifoliolata are trees. Moreover, the leaflets have sparse appressed trichomes on their lower surfaces, whereas all other specimens of A. trifoliolata are glabrous. Andira sp. A. —Congrio. Small tree ca. 5 m tall; petals white. Shrubby white-sand savannas, 100–200 m; Amazonas (Caño San Miguel, Caño Ucata southeast of Síquita, lower Río Ventuari). Brazil (Amazonas, Rondônia).

8. BARBIERIA DC., Prodr. 2: 234. 1825. by Paul R. Fantz Woody (not in flora area) or herbaceous vine, scandent or trailing; branches striate-terete, densely hirsute, trichomes reddish brown becoming whitish with age; pubescence of minutely hooked trichomes common (view at 20–30×). Leaves alternate, odd-pinnate, petiolate, stipulate, stipellate; leaflets (9–)13–21(–25), opposite, oblong to elliptic-oblong, entire, broadly acute to obtuse, mucronate, appressed-pilose on lower surface; stipules persistent, striate, lanceolate, subulate-acuminate; stipels persistent, 1-veined, linear-subulate; petiolules quadrate. Inflorescence axillary or terminal, pseudoracemose, few-flowered, long-pedunculate; pedicels paired at nodes; bracts striate, lance-subulate; bracteoles paired at calyx base, lanceolate, subulate-acuminate. Flowers resupinate, showy, papilionaceous, red. Calyx narrowtubular, persistent in fruit, 5-lobed, the lobes deltoid to rapidly subulate above; standard complicate, long-clawed, spurless, oblong-oblanceolate; wing petals oblong, long-clawed, much shorter than the long-clawed, elliptic-oblong keel petals which nearly equals length of standard. Stamens diadelphous, often persistent in fruit. Pistil enclosed within staminal sheath; ovary sessile; style linear, densely barbate. Legume sessile, linear, straight, without costa, enclosed at base by persistent calyx, valves puberulent-hirsute, strongly transversly impressed between the seeds, spirally twisting upon dehiscence 0.5–1.5 turns. Seeds dark brown to black, smooth, transverse-oblong.

Bowdichia 263

Fig. 233. Barbieria pinnata

Widespread in the Neotropics, but apparently rare locally since few collections exist in herbaria as compared to its range; 1 species. Barbieria pinnata (Pers.) Baill., Hist. Pl. 2: 263. 1870. —Galactia pinnata Pers., Syn. Pl. 2: 302. 1807. —Clitoria pinnata (Pers.) R.H. Sm. & G.P. Lewis, Kew Bull. 46: 320. 1991. Clitoria polyphylla Poir. in Lam., Encycl. suppl. 2: 300. 1811. —Barbieria poly-

phylla (Poir.) DC., Mém. Légum. 242, t. 39. 1825. Moist soils in open areas of forests and forest edges, 500–1000 m; Bolívar (Río Icabarú, Santa Elena de Uairén). Barinas, Sucre; Southern Mexico, Central America, West Indies, Colombia, Peru, Brazil, Bolivia. ŠFig. 233.

9. BOWDICHIA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 376. 1823. by Charles H. Stirton Large shrubs or small trees to 10 m tall. Leaves 9–21-foliolate, odd-pinnate, alternate, deciduous; stipules 3–7 mm long; leaflets chartaceous or coriaceous, alternate to opposite, stipellate. Inflorescences axillary or terminal racemes. Flowers blue, bisexual, zygomorphic, bracts and bracteoles small, triangular, fringed with pearl glands. Calyx turbinate with subequal lobes, sinus of vexillar lobes broad, teeth fused for half their length; petals 5, unequal, free, glabrous, clawed; standard blue with yellow or rose nectar patch, very broadly ovate, emarginate; wing petals 2–3 times larger than the keel petals, asymmetrically obovate; keel petals narrowly cultrate (knife-blade-like). Stamens 10, free, filaments subequal with thickened bases; anthers subequal, often staminodal, oblong, dorsifixed. Ovary stipitate, 5–10ovulate, densely or sparsely pubescent; style curved, shorter than ovary; stigma large, terminal, capitate. Fruits stipitate, compressed, indehiscent, linear-oblong to

264

F ABACEAE

Fig. 234. Bowdichia virgilioides

elliptic, membranous, upper suture shortly winged. Seeds oblong, compressed, asymmetric, with subterminal hilum. Colombia, Venezuela, Suriname, Guyana, Brazil, Bolivia; 2 species, 1 in Venezuela. Bowdichia virgilioides H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 376. 1823. —Alcornoque, Alcornoque sabanero, Cornoco. Tree 6–10 m tall; corolla bluish violet. Trachypogon-Curatella-Byrsonima savannadry forest ecotone, granitic outcrops, 50–500 m; Bolívar (Caicara, Ciudad Bolívar, Puerto Ordaz, Represa Guri, Río Cuchivero), Ama-

zonas (Cacuri, Puerto Ayacucho, lower Río Ocamo, Río Ventuari). Anzoátegui, Apure, Aragua, Barinas, Cojedes, Guárico, Mérida, Monagas, Portuguesa, Sucre, Trujillo, Zulia; Colombia, Guyana, Brazil, Bolivia. ŠFig. 234. This species is quite variable throughout its range.

10. CAJANUS DC., Cat. Pl. Horti Monsp. 85. 1813, nom. cons. —Cajan Adans., Fam. Pl. 2: 326. 1763. by Nidia L. Cuello A. Shrubs or subshrubs. Leaves alternate, pinnately or sometimes digitately 3foliolate; leaflets with vesicular glands below, membranous, puberulent, oblanceolate to lanceolate, basally cuneate, apically acute; stipules triangular-lanceolate, caducous; stipels present or absent. Inflorescence an axillary or terminal raceme, pedunculate or almost sessile; flowers 6–12 per rachis; bracts small, caducous. Calyx campanulate, 5-lobed, the upper lobe longer, lobes acute or acuminate; corolla yellow

Cajanus 265

or lined with red or standard dorsally reddish, to 3 cm long; standard ovate, auriculate; wing petals obliquely-obovate; keel petals auriculate, rounded-oblique, obtuse. Stamens 10, diadelphous, the vexillar stamen free; anthers uniform. Ovary sessile; ovules 4–6; style slender, glabrous; stigma capitate. Fruit linear-oblong, 2-valved with oblique constrictions between the seeds, yellowish green with brown mottling, puberulent. Seeds reniform to suborbicular, the hilum oblong, strophiolate. Paleotropics (2 species sometimes escaping from cultivation in Neotropics); 32 species, 1 in Venezuela. Cajanus cajan (L.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 53. 1900. —Cytisus cajan L., Sp. Pl. 739. 1753. —Quinchoncho. Shrub 1–3 m tall; widely cultivated for food and forage, sometimes escaped on disturbed ground, 50–1000 m; Delta Amacuro (Santa Catalina), Bolívar (Gran Sabana, Isla Anacoco, Serranía de Imataca). Aragua, Cojedes, Distrito Federal, Guárico, Mérida, Miranda, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Goiás, Mato Grosso, Minas Gerais, São Paulo), Bolivia, widespread in Africa and Asia. ŠFig. 235.

Fig. 235. Cajanus cajan

266

F ABACEAE

11. CALOPOGONIUM Desv., Ann. Sci. Nat. (Paris) 9: 423. 1826. Stenolobium Benth., Ann. Wiener Mus. 2: 125. 1837. by Richard H. Maxwell Small twining to large climbing vines. Leaves alternate, pinnately 3-foliolate; stipules acute or lanceolate; leaflets mostly ovate, entire. Inflorescence axillary, erect and pseudoracemose or fasciculate; bracts and bracteoles mostly linear, ciliate, mostly caducous. Flowers medium-sized to small, numerous. Calyx tube 5-lobed, the upper lobes united, the apex entire or emarginate; corolla blue or violet; standard reflexed, basally biauriculate; wing and keel petals adherent. Vexillar stamen free; anthers mostly 8 or 9. Ovary sessile, many-ovulate; style filiform or subulate, usually incurved; stigma terminal, capitate, glabrous. Fruit a legume, linear to linearoblong, several- to many-seeded, compressed, septate, dehiscent; exocarp transversely furrowed between seeds. Seeds reniform to cuboid, the hilum small, oval. Neotropics; 6–8 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Calopogonium 1.

1.

Calyx and fruit with appressed or spreading trichomes to ca. 1 mm long; calyx lobes acute, shorter than or about equal to the tube length; fruit ca. 8 mm wide; terminal leaflets broadly ovate to rhomboidal, occasionally lobed ................................................................................. C. coeruleum Calyx and fruit with erect or spreading trichomes to ca. 2 mm long or longer; calyx lobes linear, greatly exceeding the tube length; fruit ca. 4 mm wide; terminal leaflets ovate to elliptic or lanceolate ......... C. mucunoides

Fig. 236. Calopogonium coeruleum

Fig. 237. Calopogonium mucunoides

Canavalia 267

Calopogonium coeruleum (Benth.) C. Wright in Sauvalle, Anales Acad. Ci. Méd. Habana 5: 337. 1869. —Stenolobium coeruleum Benth., Ann. Wiener Mus. Naturgesch. 2: 125. 1838. —Caraotillo. Vine, may become long and high-climbing. Roadsides, fields, disturbed areas, 50–400 m; Delta Amacuro (Río Toro), northern Bolívar, Amazonas (Río Ocamo). Anzoátegui, Apure, Barinas, Carabobo, Guárico, Miranda, Monagas, Portuguesa, Sucre, Zulia; Mexico, Central America, West Indies, Guyana, Suriname, French Guiana, throughout Brazil and Colombia, extending south through Ecuador and Peru into Bolivia and tropical Argentina. ŠFig. 236. Calopogonium mucunoides Desv., Ann. Sci. Nat. (Paris) 9: 423. 1826. —Frijol de

lagartilla, Peine del diablo. Vine; the long trichomes, especially on the fruit, may be yellow or a darker rusty color. Disturbed, semi-open, humid sites, near sea level to 300 m; Delta Amacuro (Caño Angosturito, Caño Güiniquina), Bolívar (middle Río Orinoco), Amazonas (San Carlos de Río Negro). Guárico, Monagas, Zulia, and probably throughout rest of Venezuela; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amapá, the drier northeastern states south to Bahía, Pará, Amazonas, Rondônia). ŠFig. 237. This has been widely used as a cover crop and apparently escapes as a weed throughout central Brazil and in the Old World tropics.

12. CANAVALIA DC., Prodr. 2: 403. 1825, nom. cons. Canavali Adans., Fam. Pl. 2: 325, 531. 1763. Wenderothia Schltdl., Linnaea 12: 330. 1838. by Gerardo A. Aymard C. and Richard H. Maxwell Climbing, twining, or trailing vines. Leaves alternate, 3-foliolate; stipules small, lanceolate or oblong, deciduous or caducous; leaflets mostly ovate to suborbicular, entire, unlobed. Inflorescence axillary, pseudoracemose, mostly pendent, tuberculate; each tubercle 2–6-flowered, bracteate. Flowers mostly resupinate, showy, bracteolate. Calyx 5-lobed, appearing bilabiate, the upper lobes largest, entire or emarginate, the lower and lateral lobes crowded, minute; corolla whitish, pink, rose, or shades of violet or purple; standard reflexed; wing petals free; keel petals wider than wings, upcurved, frequently beaked, united distally. Stamens 10, monadelphous or pseudomonadelphous. Pistil mostly curved; ovary pubescent, sessile or short-stipitate; stigma capitate. Fruit a legume, oblong, compressed, dehiscent, the upper sutures with 2 parallel ribs, frequently each valve with 1 or 2 extra ribs, several- to many-seeded. Seeds ellipsoid, mostly compressed, the hilum oblong or linear. Pantropics; ca. 60 species, 11 in Venezuela, 5 of these in the flora area. Key to the Species of Canavalia 1.

1.

2(1). 2.

Common trailer on seacoast beaches or twining vines on backbeach vegetation; leaflets somewhat orbicular, the apices frequently emarginate; seed hilum ca. 6 mm long ............................................................... C. rosea Inland species; leaflets ovate, elliptic, lanceolate, or combinations, not orbicular, the apices rounded, acute, to acuminate; seed hilum 6–22 mm long ......................................................................................................... 2 Fruits without an extra rib; pubescence of leaflets long-silky, very dense on lower surface of blade ..................................................... C. sericophylla Fruits with an extra rib; leaflets glabrous or pubescence sparse to moderately dense on lower surface of blade .................................................... 3

268

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

4.

F ABACEAE

Upper calyx lobe gradually concave behind the apex; flowers to ca. 4 cm long; seeds ca. 3 mm thick .................................................... C. grandiflora Upper calyx lobe depressed behind the apex; flowers 2–3 cm long; seeds 9–12 mm thick ....................................................................................... 4 Leaflets broadly ovate, chartaceous; standard ca. 2.5 cm long; seed hilum 8–12 mm long, shorter than the seed; fruit to ca. 2.8 cm wide .............. ............................................................................................... C. brasiliensis Leaflets lanceolate-elliptic, coriaceous; standard to ca. 3 cm long; seed hilum 15–18 mm long, nearly as long as the seed; fruit to ca. 3.5 cm wide ..................................................................................................... C. dictyota

Fig. 238. Canavalia brasiliensis

Canavalia brasiliensis Mart. ex Benth., Ann. Wiener Mus. Naturgesch. 2: 135. 1838. Canavalia fendleri Piper, Contr. U.S. Natl. Herb. 20: 570. 1925, pro parte major, fide Sauer, Brittonia 16: 144. 1964. Trailing vine in the open, or twining over shrubs. Mud flats, lakeside lowlands, into upper savannas, disturbed areas, 50–200 m; Bolívar (near Caicara, Ciudad Bolívar). Widespread elsewhere in Venezuela; U.S.A. (Florida), Mexico, Central America, West Indies, widespread in South America although apparently rare in Amazon basin. ŠFig. 238. Canavalia brasiliensis is planted occasionally as a cover crop and is sometimes a weedy invader, usually in lowlands.

Centrolobium 269

Canavalia dictyota Piper, Contr. U.S. Natl. Herb. 20: 574. 1925. Vine trailing, twining and climbing on undershrubs, or occasionally high-climbing. Open ground, thickets, rocky outcrops, near sea level to 100 m; Delta Amacuro (Caño Angosturita, Caño Güiniquina, Tucupita), Bolívar (lower Río Caura, Río Pargueni), Amazonas (Puerto Ayacucho). Distrito Federal, Lara, Nueva Esparta, Sucre, Zulia; Panama, West Indies, Colombia (Magdalena), Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Bahia). Canavalia grandiflora Benth., Ann. Wiener Mus. Naturgesch. 2: 135. 1837. Liana or trailing or climbing vine. Humid forests, but extending from forest edges into open areas, including rocky outcrops, 50–100 m; Bolívar (Anacoco, Caicara, Represa Guri, lower Río Caroní, Río Caura, Río Cuyuní, Santa María de Erebato, Tumeremo), Amazonas (Puerto Ayacucho). French Guiana, Peru, Brazil (Goiás, Mato Grosso, Minas Gerais, Pará, Rondônia).

Canavalia rosea (Sw.) DC., Prodr. 2: 404. 1825. —Dolichos roseus Sw., Prodr. 105. 1788. Dolichos maritimus Aubl., Hist. Pl. Guiane 765. 1775. Canavalia maritima Thouars, J. Bot. Agric. 1: 80. 1813. Woody prostrate vine on beach, or twining, climbing over vegetation on the back beach, from the drift line to ca. 100 m; Delta Amacuro (Caño Güiniquina). Anzoátegui, Aragua, Distrito Federal, Falcón, Sucre, Zulia; Pantropics. The name Canavalia maritima (Aubl.) Thouars has frequently been used, but Thouars’s epithet is here considered to be a new name rather than a combination from Aublet’s epithet. Thus, C. rosea is the correct name. Canavalia sericophylla Ducke, Arq. Inst. Biol. Veg. 4: 23. 1938. Vine. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma at base of Sierra de la Neblina). Amazonian Peru, Brazil (Amazonas).

13. CENTROLOBIUM Mart. ex Benth., Comm. Legum. Gen. 31. 1837. by Velva E. Rudd Trees to ca. 30 m tall, commonly buttressed at the base; bark grayish, smooth or fissured; sap red. Leaves deciduous, alternate, odd-pinnate, 7–21-foliolate; leaflets oblong, ovate, or elliptic, the lower surface with numerous reddish orange resinous dots; stipules deltoid to broadly orbicular or rhomboid, often caducous; stipels lacking. Bracts and bracteoles stipule-like but smaller. Flowers ca. 1–2 cm long in terminal panicles. Calyx turbinate-campanulate with 4 subequal lobes, the vexillar lobe emarginate or bipartite; petals yellowish, sometimes with red or violet markings. Stamens 10, monadelphous; anthers uniform, dorsifixed. Ovaries sessile or short-stalked, 1–3(4)-ovulate; style filiform, usually persistent as a stout spine; stigma small, terminal. Fruit samaroid, indehiscent with a large 1–3(4)-seeded basal body and a large terminal, sterile wing, the body echinate with spines to 4 cm long, the wings spatulate or cultriform, the persistent stylar spine terminal. Seeds oblong-subreniform. Panama, Colombia, Venezuela, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia; ca. 6 species, 1 in Venezuela. Centrolobium paraense Tul., Arch. Mus. Hist. Nat. 4: 87. 1844. —Cartán. Tree to 30 m tall. Evergreen lowland forests, savannas. Panama, Colombia, Venezuela, Trinidad (cultivated?), Guyana, Brazil; 2

varieties, both in the flora area. The wood of Centrolobium paraense is fine-grained and has beautiful shades of reddish orange. It is used for furniture and cabinet work as well as for general construction.

270

F ABACEAE

Fig. 239. Centrolobium paraense var. orinocense

Centrosema 271

Key to the Varieties of C. paraense 1. Leaflets rounded to subcordate at the base, subglabrous on the upper surface or often with some pubescence along the midvein, glabrous on lower surface at maturity; flowers 12–15 mm long .................................... var. orinocense 1. Leaflets subcordate at the base, sometimes revolute, tomentose on upper surface, subglabrous on lower surface except for tomentum on major veins; flowers 15–17 mm long ....... var. paraense C. paraense var. orinocense Benth. in Mart., Fl. Bras. 15(1): 266. 1862, “orenocense.” —Centrolobium orinocense

(Benth.) Pittier, Bol. Mus. Comerc. Venez. 1: 19. 1927 [1926]. —Balaústre, Cartán. Centrolobium patinense Pittier, J. Wash. Acad. Sci. 5: 570. 1915. 100–300 m; Bolívar (Ciudad Bolívar, Represa Guri area, La Prisión, middle Río Caura, San Mateo, Serranía de Guayapo in lower Caura basin). Anzoátegui, Aragua, Guárico, Zulia; Panama, Colombia. ŠFig. 239. C. paraense var. paraense. —Cartán. 100–300 m; Bolívar (El Manteco, La Paragua, Represa Guri, Río Asa, mouth of Río Paragua, Upata). Anzoátegui, Guárico, Sucre, Zulia; Trinidad (cultivated?), Guyana, Brazil (Roraima).

14. CENTROSEMA (DC.) Benth., Comm. Legum. Gen. 53. 1837, nom. cons. —Clitoria sect. Centrosema DC., Prodr. 2: 234. 1825. Bradburya Raf., Fl. Ludonear 104. 1817. by Paul R. Fantz Scandent herbs, trailing to high-climbing, suffrutescent at base from perennial xylopodium; pubescence of minutely hooked trichomes common (view at 20–30×); aerial stems wiry to lianaceous, twining, angular-terete. Leaves alternate, odd-pinnate or occasionally digitate, 3-, 5-, or 7-foliolate, entire, stipulate, stipellate, petiolate; stipules ovate to deltoid, striate, persistent; stipels linear to linear-deltoid, striate, persistent; petiolules subquadrangular. Inflorescences axillary, usually solitary, pseudoracemose, few to several-flowered, crowded apically, often with one flower open at a time; peduncle elongate, rachis straight to weakly flexuose, nodose; pedicels paired at nodes; bracts crowded, in 3 series, striated, smaller than bracteoles, outer pair largest; bracteoles paired at calyx base, striated, enfolding buds, appressed to calyx and often obscuring calyx tube and upper and lateral teeth, deciduous in fruit. Flowers resupinate, papilionaceous, showy, lilac to purple or white with purplish veins. Calyx short-campanulate, usually persisting in early fruiting stages before deteriorating, upper 2 lobes connate, lower ventral tooth elongated, often conspicuous, extending from between bracteoles; standard complicate, orbicular, emarginate, spurred or gibbous above claw, pubescent on outer surface; wing petals falcate-obovate, auriculate above claw; keel petals incurved, clawed, subequal to scarcely shorter than wings, ventral margin broad U-shaped. Stamens pseudodiadelphous, vexillar stamen fused basally, filaments dilated apically. Pistil enclosed in staminal sheath; ovary sessile, linear, appressed-pubescent; style strongly incurved, broadly U-shaped, ± dilated apically, persistent as a beak in fruit; stigma marginal at style apex, barbellate basally. Legume sessile, linear to occasionally broadened, compressed and flat, beaked; valves costate, longitudinal veins near each margin, occasionally winged, sutures slightly thickened, subseptate within and between the seeds; dehiscence by spiral twisting of valves. Seeds transverse-oblong, thick, numerous to occasionally few per pod.

272

F ABACEAE

New World tropics and subtropics, a single species extending into temperate southern U.S.A., a few species introduced into Old World tropics; 34 species, 12 in Venezuela, 10 of these in the flora area. Key to the Species of Centrosema 1. 1. 2(1). 2.

3(2).

3.

4(2). 4. 5(4).

5.

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

7.

8(6).

8.

Leaves digitate, rachis absent; leaflets 3, 5, or 7 ......................... C. venosum Leaves pinnate, rachis 0.2–4 cm long; leaflets 3 ....................................... 2 Fruit 8–50 mm wide; flowers 4–6 cm long; petiolules 5–8 mm long; stipels 5–8 mm long; leaflets 6–12 cm wide; inflorescences 6–15 cm long ...... 3 Fruit 4–7 mm wide; flowers 1.5–4 cm long; petiolules 1–5 mm long; stipels 2–5 mm long; leaflets 0.5–6(–7) cm wide; inflorescences 1–8(–11) cm long ......................................................................................................... 4 Fruit 8–13 mm wide, beak 10–15 mm long; calyx tube 4–5 mm long, teeth obscure or ventral tooth to 2 mm long; bracteoles 8–16 × 6–9 mm; pedicels 5–10 mm long; stipules 4–9 mm long .......................... C. plumieri Fruit 35–50 mm wide, beak 25–40 mm long; calyx tube 6–9 mm long, teeth 2–3 mm long with ventral tooth 4–5 mm long; bracteoles 15–20 × 9–13 mm; pedicels 10–20 mm long; stipules 10–16 mm long ................. ................................................................................................ C. triquetrum Bracteoles 14–23 × 6–10 mm; bracts 6–10 mm wide; legume beak 18–27 mm long ......................................................................................................... 5 Bracteoles 4–15 × 2–7 mm; bracts 1.5–5 mm wide; legume beak 3–16 mm long ......................................................................................................... 6 Calyx tube 5–6 mm wide, ventral tooth 2.5–4 mm long, upper and lateral teeth 1–2 mm long; fruit 55–90 × 4–4.5 mm, beak 16–27 mm long; pedicels 5–11 mm long ........................................................ C. brasilianum Calyx tube 6–10 mm wide, ventral tooth 5–8 mm long, upper and lateral teeth 2–3 mm long; fruit 100–180 × 7–8 mm, beak 25–52 mm long; pedicels 10–15 mm long ................................................. C. tetragonolobum Calyx tube 2–3 mm long, 3–5 mm wide; fruit 3–4 mm wide, 5–10 mm long; leaflets linear, 0.3–1 cm wide ................................................................ 7 Calyx tube 3–6 mm long, 5–10 mm wide; fruit 4–7 mm wide, (8–)10–20 mm long; leaflets ovate to elliptic to rhombic to oblong, 1–7 cm wide ........ 8 Flowers 1–1.5 cm long; bracteoles 4–7 × 2–3 mm; calyx teeth subequal, upper and lateral teeth 3–4 mm long, ventral tooth 4–6 mm long; fruits 5–8 cm long, beak 3–7 mm long; leaflets with main lateral veins 14–19; rachis 0.5–1.2 cm long; stipules 5–6 mm long ........... C. pascuorum Flowers 2–4 cm long; bracteoles 9–13 × 3–5 mm; calyx teeth unequal in length, upper and lateral teeth 0.4–1 mm long, ventral tooth 5–8 mm long; fruits 7–10 mm long, beak 10–15 mm long; leaflets with main lateral veins 20–40; rachis nearly lacking, 0.1–0.3 mm long; stipules 3–4 mm long .................................................................................... C. angustifolium Bracteoles 10–15 mm long; bracts 8–10 mm long; calyx tube 8–10 mm wide, 4–6 mm long, ventral tooth 9–15 mm long; leaf rachis 1.5–3.5 cm long, leaflets commonly large, 6–15 cm long; robust vine or lianaceous ........................................................................................... C. macrocarpum Bracteoles 6–10 mm long; bracts 3–7 mm long; calyx tube 5–7 mm wide,

Centrosema 273

9(8).

9.

3–4 mm long, ventral tooth 5–10 mm long; leaf rachis 0.4–1.5(–2) cm long, leaflets commonly smaller, 2–7(–10) cm long; slender vine ........ 9 Upper and lateral calyx teeth subequal to calyx tube, 2–4 mm long, ventral tooth 5–8 mm long; fruit 6–7 mm wide, beak 8–10 mm long, valve dehiscence 1–1.5 turns; bracts 6–7 mm long; pedicels 6–8 mm long in flower, 8–13 mm long in fruit; petioles 3–4 mm long .................... C. molle Upper and lateral calyx teeth longer than calyx tube, 6–8 mm long, ventral tooth 8–10 mm long; fruit 4–5 mm wide, beak 10–15 mm long, valve dehiscence 1.5–5 turns; bracts 3–4 mm long; pedicels 2–5 mm in flower, 4–8 mm long in fruit; petioles 2–3 mm long .......... C. virginianum

Centrosema angustifolium (H.B.K.) Benth., Comm. Legum. Gen. 54. 1837, non (H.B.K.) Benth. in Mart., Fl. Bras. 15(1): 129. 1859. —Clitoria angustifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 417. 1823 [1824]. —Bradburya angustifolium (H.B.K.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Perennial herbaceous vine with wiry stem and linear leaves. Savannas, edges of semideciduous forests, 200–800 m; Bolívar (Altiplanicie de Nuria east of Miamo, northeast of El Manteco, Represa Guri, Serra do Sol on Brazilian border), Amazonas (Caño Picure in Río Ventuari basin). Anzoátegui, Apure, Guárico, Monagas; Central America, Colombia, Guyana, Suriname, Brazil, cultivated in Peru. Centrosema brasilianum (L.) Benth., Comm. Legum. Gen. 54. 1837. —Clitoria brasiliensis L., Sp. Pl. 753. 1753. —Bradburya brasilianum (L.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Clitoria angustifolia auct. non H.B.K. 1823 [1824]: sensu Centrosema angustifolium (H.B.K.) Benth., Comm. Legum. Gen. 54. 1837; Centrosema angustifolium (H.B.K.) Benth. in Mart., Fl. Bras. 15(1): 129. 1859; Centrosema brasilianum var. angustifolium (H.B.K.) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 62. 1939. Perennial herbaceous vine with narrow to broad leaflets. Savannas, roadsides, forest edges, 50–500 m; Delta Amacuro (near Los Castillos de Guayana, Sacupana), Bolívar (widespread across northern part of state, Río Hacha), Amazonas (Puerto Ayacucho to Samariapo). Apure, Distrito Federal, Monagas, Nueva Esparta, Sucre, Zulia; Colombia,

Guyana, Suriname, French Guiana, Brazil, Bolivia. ŠFig. 245. Centrosema macrocarpum Benth., Ann. Mag. Nat. Hist. ser. 1, 3: 436. 1839. —Bradburya macrocarpa (Benth.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. —Centrosema macrocarpon auct., orthographic error. Liana or robust vine. Open areas and secondary regrowth of forests, sandy soils, 50– 300 m; Bolívar (Altiplanicie de Nuria, Río Cuchivero, Río Orinoco), Amazonas (Isla Ratón, Río Cataniapo). Anzoátegui, Aragua, Cojedes, Zulia; Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 240. Centrosema molle Mart. ex Benth., Comm. Leg. Gen. 55. 1837; Ann. Wein. Mus. Naturgesch. 2: 119. 1839. Centrosema pubescens auct. non Benth. 1837: sensu Benth. in Mart., Fl. Bras. 15: 131. 1839; Barbosa-Fevereio, Rodriguésia 42: 184. 1977. Perennial herbaceous vine. Open areas of evergreen lowland forests, river banks, savannas, sandy areas, near sea level to 300 m; Delta Amacuro (Río Orinoco), Bolívar (Ciudad Bolívar, La Paragua, Río Cuyuní, Río Pargueni off middle Orinoco), Amazonas (Puerto Ayacucho to Samariapo, Santa Bárbara del Orinoco). Widespread elsewhere in Venezuela north of the Orinoco; Mexico, Central America, West Indies, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, introduced into tropical Africa and Asia. This is the most commonly collected species of Centrosema and has often been confused with and misidentified as C. pubescens (see Fantz, Sida 17: 321–332. 1996).

274

F ABACEAE

Fig. 240. Centrosema macrocarpum

Fig. 241. Centrosema pascuorum

Fig. 242. Centrosema plumieri

Centrosema 275

Fig. 243. Centrosema triquetrum

Fig. 244. Centrosema venosum

Fig. 245. Centrosema brasilianum

276

F ABACEAE

Centrosema pascuorum Mart. ex Benth., Comm. Legum. Gen. 561. 1837. —Bradburya pascuora (Mart. ex Benth.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Suffrutescent herb with narrow leaves, trailing to weakly climbing. Rocky area and roadsides, ca. 800 m; Bolívar (near Santa Elena de Uairén). Anzoátegui, Cojedes, Guárico, Miranda, Monagas, Zulia; Mexico, Central America, Ecuador, Brazil. ŠFig. 241. Centrosema plumieri (Turpin ex Pers.) Benth., Comm. Legum. Gen. 54. 1837. —Clitoria plumieri Turpin ex Pers., Syn. Pl. 2: 303. 1807. —Bradburya plumieri (Turpin ex Pers.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Liana or robust vine. Evergreen lowland forests, disturbed forests, 50–200 m; Delta Amacuro (Sacupana), Bolívar (La Unión in middle Río Caura), Amazonas (Capibara, Mavaca, Río Mawarinuma). Apure, Guárico, Monagas, Portuguesa; Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, introduced into tropical Africa. ŠFig. 242. Centrosema tetragonolobum SchultzeKraft & R.J. Williams, Caldasia 16: 133. 1990. Perennial herbaceous vine with narrow to broad leaflets. Borders of gallery forests in savanna zone between 4°–6° latitude on both sides of the Río Orinoco, 50–200 m; Bolívar (18 km southeast of Río Parguaza on the road to Puerto Ayacucho). Amazonas (Samariapo to north of Puerto Ayacucho). Colombia (Vichada). Centrosema triquetrum (Hoffmanns. ex Benth.) Benth. in Benth. & Hook. f.,

Gen. Pl. 1(2): 528. 1865. —Platysema triquetra Hoffmanns. ex Benth., Ann. Wiener Mus. Naturgesch. 2: 122. 1839. —Bradburya triquetra (Hoffmanns. ex Benth.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Liana or robust vine. Evergreen lowland forests, riparian bamboo thickets, 50–200 m; Delta Amacuro (Río Cuyubini), Amazonas (Río Mawarinuma). Táchira; Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas, Goiás, Maranhão, Pará). ŠFig. 243. Centrosema venosum Mart. ex Benth. in Mart., Fl. Bras. 15(1): 133. 1859. —Bradburya venosum (Mart. ex Benth.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Perennial herb, stems creeping or trailing, weakly twining. Open grassy savannas, 50–100 m; Bolívar (between Caicara and Puerto Ayacucho, Río Parguaza), Amazonas (Canaripó, from north of Puerto Ayacucho to Samariapo, Rincones, Río Ventuari). Anzoátegui; Colombia, Brazil (Bahia, Goias, Minas Gerais, Pará). ŠFig. 244. Centrosema virginianum (L.) Benth., Ann. Wiener Mus. Naturgesch. 2:120. 1837. —Clitoria virginiana L., Sp. Pl. 753. 1753. —Bradburya virginianum (L.) Kuntze, Revis. Gen. Pl. 1: 164. 1891. Perennial herbaceous vine. Semideciduous woods and savannas, 100–500 m; Bolívar (Altiplanicie de Nuria, Upata to Guasipati), Amazonas (near Puerto Ayacucho). Aragua, Distrito Federal, Miranda; U.S.A., Mexico, Central America, West Indies, South America to Argentina, introduced into India, Reunión.

15. CHAETOCALYX DC., Prodr. 2: 243. 1825. by Velva E. Rudd Perennial vines, herbaceous or suffrutescent. Stems slender, terete, longitudinally striate. Leaves alternate, odd-pinnate, 5–17-foliolate; leaflets oblong, elliptic, ovate, obovate, or suborbicular; stipules deltoid or deltoid-ovate to lanceolate, attached at the base; stipels absent. Bracts intergrading with the stipules, often caducous; bracteoles lacking. Flowers ca. 1.2–3 cm long, solitary or in axillary, racemose, paniculate, or fasciculate inflorescences. Calyx campanulate with 5 subequal lobes or teeth, base symmetrical or asymmetrically gibbous, often setose with glandular trichomes; petals cream-white to yellow, sometimes marked with red or violet, pubescent or glabrous. Stamens 10, monadelphous; anthers uniform, elliptic,

Chaetocalyx 277

Fig. 246. Chaetocalyx scandens var. pubescens

dorsifixed. Ovaries sessile or stipitate, 6–16-ovulate, glabrous or pubescent; stigma terminal, minute, capitate. Fruit a loment, linear to subelliptic, laterally compressed or subterete, longitudinally striate or reticulate-striate, to 16-articulate, the segments approximately isometric to several times longer than wide. Seeds reniform-rod-shaped, sublustrous, brownish, the hilum elliptic, subterminal. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; 14 or 15 species, 1 in Venezuela. Chaetocalyx scandens (L.) Urb., Symb. Antill. 2: 292. 1900. —Coronilla scandens L., Sp. Pl. 743. 1753. Herbaceous vine; leaflets densely pubescent to subglabrous on one or both surfaces; calyx and fruit puberulent. Stream banks, open woods, savannas, rocky hillsides. Southern Mexico, Dominican Republic, Lesser Antilles, northern Colombia, Venezuela, eastern Brazil; 2 varieties, both in Venezuela, 1 of these in the flora area. The other variety, Chaetocalyx scandens var. scandens, is glabrous or subglabrous and is found in Venezuela along the Coastal Cordillera in Aragua, Carabobo, and Distrito Federal. C. scandens var. pubescens (DC.) Rudd, Contr. U.S. Natl. Herb. 32: 236. 1958.

—Chaetocalyx pubescens DC., Prodr. 2: 243. 1825. Chaetocalyx vestita Standl., Field Mus. Nat. Hist., Bot. Ser. 8: 14. 1930. Chaetocalyx fissa Pittier, Bol. Soc. Venez. Ci. Nat. 6: 189. 1940. Chaetocalyx magniflora Pittier, Bol. Soc. Venez. Ci. Nat. 6: 186. 1940. Chaetocalyx nigrescens Pittier, Bol. Soc. Venez. Ci. Nat. 6: 188. 1940. Chaetocalyx paucifolia Pittier, Bol. Soc. Venez. Ci. Nat. 6: 185. 1940. Chaetocalyx perglandulosa Pittier, Bol. Soc. Venez. Ci. Nat. 6: 187. 1940. Woody, rocky slopes, savannas, 300–700 m; Bolívar (Altiplanicie de Nuria). Venezuelan Coastal Cordillera; southern Mexico, Dominican Republic, Lesser Antilles, northern Colombia, eastern Brazil. ŠFig. 246.

278

F ABACEAE

16. CLATHROTROPIS Harms in Dalla Torre & Harms, Gen. Siphon. 221. 1901. by Charles H. Stirton and Gerardo A. Aymard C. Tall trees. Leaves 5- or 7(9)-foliolate, odd-pinnate, alternate; leaflets chartaceous or coriaceous, opposite, the upper surface dark green, the lower surface paler; stipules small, caducous. Inflorescences terminal, racemose-paniculate. Flowers white with purple nectar patch, fragrant, bisexual, zygomorphic, pedicellate, bracteate and bracteolate. Calyx unequally 5-toothed, upper pair fused for 1/4–1/2 their length; petals clawed, ± auriculate; standard orbicular to obovate, glabrous; wing petals ± same length as keel petals; sculpturing present, lamellate. Stamens 10, free or nearly so; filaments subequal; anthers uniform, oblong, versatile. Ovary subsessile or subsessile, 4- or 5-ovulate, hairy; style filiform; stigma small, terminal. Fruit dehiscent, compressed to flattened, sutures often thickened and dilated, valves woody or coriaceous, glabrous or densely golden-pubescent, elliptic, oblong, trapezoid or semireniform. Seeds few, large, compressed; testa fragile. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 6 species, 5 in Venezuela, all in the flora area. Key to the Species of Clathrotropis 1. 1. 2(1). 2. 3(1).

3.

4(3). 4.

Lower surface of leaves minutely pubescent; fruits densely velutinous, with thickened and dilated sutures ...................................................... 2 Lower surface of leaves glabrous; fruits glabrous, without prominent sutures ....................................................................................................... 3 Lower surface of leaves glaucous; fruits crispate, copper-colored, shortly ridged along upper sutures ................................................ C. glaucophylla Lower surface of leaves green; fruits densely velutinous, rusty brown, strongly ridged along upper sutures ................................... C. macrocarpa Leaflets ovate-oblong to sublanceolate, apex attenuate, the secondary and tertiary venation scarcely visible on both surfaces; flowers borne in dense short panicles; fruits < 3.5 cm long ..................................... C. nitida Leaflets broadly elliptic to oblong-elliptic, apex rounded to short-acuminate, the secondary and tertiary venation prominently reticulate on both surfaces; flowers borne in large lax panicles; fruits > 3.5 cm long ................................................................................................................ 4 Young branches, inflorescences, and buds finely silvery to yellowish white-pubescent ................................................................. C. brachypetala Young branches, inflorescences, and buds densely chocolate brown pubescent ............................................................................................. C. brunnea

Clathrotropis brachypetala (Tul.) Klein., Recueil Trav. Bot. Néerl. 22: 398. 1925. —Diplotropis brachypetala Tul., Arch. Mus. Hist. Nat. 4: 111. 1844. —Caicareño, Mahomo montañero, Montanallari (Warao). Tree to 30 m tall; leaves 5- or 7-foliolate, broadly elliptic to oblong-elliptic, glabrous, nitid, to ca. 25 cm long; fruits dark, glabrous, flattened; seeds to 5 cm long and 1.5 cm

thick. Evergreen lowland and riparian forests, near sea level to 300 m; Delta Amacuro (Caño Joba-Suburu west of Caño Guayo and east of Caño Sacupana, Río Cuyubini), Bolívar (Jabillal on Río Caura, Río Ariza, Río Cuyuní between Isla Anacoco and Acarabisi), Amazonas (23 km northeast of Puerto Ayacucho near Cachama). St. Vincent, Trinidad, Guyana, Suriname. ŠFig. 247.

Clathrotropis 279

Fig. 247. Clathrotropis brachypetala

280

F ABACEAE

Clathrotropis brunnea Amshoff, Acta Bot. Neerl. 17: 103. 1968. —Kajadi (Yekwana), Ma’nour’uruwe’dasu (Piaroa). Clathrotropis colombiana Yakovlev, Nauch. Dokl. Near Sch. Biol. Nauki 1: 58. 1971. Tree to 40 m tall; leaves (3)5(7)-foliolate, oblong to obovate-oblong, pubescent on venation, otherwise glabrescent, coriaceous or chartaceous; inflorescences and buds densely chocolate brownish-pubescent; fruits dark blue. Evergreen lowland forests, 100–300 m; Bolívar (Río Tabaro off Río Nichare), Amazonas (Caño Iguana, Caño Mosquito off Caño Marieta). Colombia, Guyana, Suriname. White phloem tissue of Clathrotropis brunnea is rubbed on insect stings to relieve pain. Clathrotropis glaucophylla R.S. Cowan, Bol. Soc. Venez. Ci. Nat. 15: 99. 1954. —Cabari, Guapocoji (Yanomami), Kajadi (Yekwana). Tree 30–40 m tall; leaves 3- or 5-foliolate, elliptic, obovate to oblong-obovate, the lower surface minutely puberulent and glaucous; fruits crispate, light copper-colored, shortly ridged along upper sutures. Evergreen lowland to lower montane forests, 100–600 m; Bolívar (Amaruay-tepui, Aparamán-tepui,

Auyán-tepui, Río Cácaro, upper Río Caura, upper Río Erebato, Río Icabarú), Amazonas (Cerro Huachamacari, Maroa, Río Coromoto, Río Cuao, Río Ocamo, Río Padamo, near San Carlos de Río Negro, Yavita). Guyana. Clathrotropis macrocarpa Ducke, Trop. Woods 31: 16. 1932. —Barbasco, Cabarí. Tree to 40 m tall; leaves 5- or 7-foliolate, obovate, coriaceous; fruits velutinous, strongly ridged along upper sutures. Evergreen lowland to montane, nonflooded forests, 100–900 m; Bolívar (widespread), Amazonas (slopes of Cerro Huachamacari, Río Cataniapo, Río Cuao, Río Mawarinuma, San Carlos de Río Negro, Sierra de la Neblina, Solano road). Colombia, Guyana, Brazil (Amazonas, Roraima). Clathrotropis nitida (Benth.) Harms, Bot. Jahrb. Syst 33: 27. 1903. —Diplotropis nitida Benth., J. Bot. (Hooker) 2: 71. 1841. —Chimako, Guacapú. Tree to 20 m tall; leaves 5- or 7-foliolate, ovate-oblong to sublanceolate, glabrous, coriaceous, nitid; fruits small, oval-oblong, glabrous. Riparian forests, 50–400 m; Amazonas (Río Baría, Río Casiquiare between Culimacare and junction with Río Negro, Río Guianía, Río Pacimoni, Río Siapa, Río Venamo, Río Yatúa). Colombia, Brazil (Amazonas).

17. CLITORIA L., Sp. Pl. 753. 1753. Ternatea Tourn. ex Mill., Gard. Dict. Abr. ed. 4, 3: Ternatea. 1754. Neurocarpum Desv., J. Bot. Agric. 1: 119. 1813. by Paul R. Fantz Trees, shrubs, lianas, or subshrubs to perennial herbs or vines; pubescence of minutely hooked trichomes common (view at 20–30×). Leaves alternate, odd-pinnate, 3-foliolate, petiolate to subsessile or 1-foliolate, sessile; stipules and stipels persistent, striate, rarely caducous; petioles and rachis longitudinally striated, sometimes canaliculate; petiolules subquadrate, rugose. Inflorescences axillary, terminal or cauliferous, bearing chasmogamous or infrequently cleistogamous flowers, 1–many-flowered; peduncles usually solitary or several-fascicled; pedicels paired at nodes; bracts striate, in 3 series below pedicels; bracteoles paired at calyx base, persistent, striate, usually appressed to calyx, rarely borne below on pedicels. Chasmogamous flowers resupinate, showy, papilionaceous, pink, blue to violet, or white fading to pale yellow. Calyx funnel-shaped, persistent in fruit, 5-lobed, upper 2 lobes subconnate, lowermost narrower, often longer. Standard complicate, emarginate, short-clawed, spurless, veins dark-colored, converging in throat, abaxial surface pubescent. Wings long-clawed, extending beyond the falcate-incurved, shortclawed keel. Stamens 10, diadelphous, often persistent in fruit. Pistil enclosed

Clitoria 281

within staminal sheath; ovary stipitate; style geniculate, bearded longitudinally apically. Cleistogamous flowers uncommon, inconspicuous unless with fruit; petals lacking or remnants hidden in calyx; calyx funnel-shaped, small, persistent in fruit, bracteolate, 5-lobed; pistil similar to that of chasmogamous flowers, but style bent abruptly back toward base, in contact with anthers. Legume stipitate, linear, straight to subfalcate, thickened at sutures, valves flat or convex, weak to strongly depressed between seeds, ecostate or costate, beaked, spirally twisting upon dehiscence. Seeds dark brown to black, smooth or viscid. Tropical to subtropical areas, mostly Neotropics, a few species in temperate areas; 60 species, 15 in Venezuela, 12 of these in the flora area. The other three species in Venezuela are Clitoria fairchildiana Howard, a cultivated Brazilian endemic that has been collected in Miranda state and in Ciudad Bolívar; C. glaberrima Pittier, from Panama, rare elsewhere in Central America, in Carabobo and Miranda states in Venezuela; and C. ternatea L., a cultivated and naturalized African introduction found in Carabobo, Cojedes, Mérida, Sucre, and Zulia states. Key to the Species of Clitoria 1.

1.

2(1). 2.

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

4.

5(1).

5.

6(5).

6.

Calyx 10-veined; fruits turgid, costate with medial longitudinal vein, 3.5– 6(–7) × 0.6–1.1 cm; stipe 7–12(–14) cm long, enclosed within calyx; cleistogamy present (calyx 1–1.2 cm long); petiolules 2–5 mm long; seeds viscid; subshrubs to perennial herbs, rarely vines ..................... 2 Calyx many-veined or striate; fruits flat, ecostate, (6–)8–24 × 1–2 cm; stipe 15–37 mm long, exerted beyond calyx; cleistogamy absent; petiolules 4–10 mm long; seeds smooth; trees, shrubs, or lianas ............... 5 Leaves petiolate, petiole 2–8 cm long; inflorescence 5–20 cm long; stem pubescence rufous; flowers white fading dull yellow; vines ....... C. falcata Leaves subsessile to sessile, petiole to 1.5 cm long; inflorescence 0.5–7 cm long; stem pubescence whitish; flowers pale bluish, lilac, or lavender; subshrubs or perennial herbs, aerial stems erect ................................. 3 Leaves sessile, 1-foliolate; leaflets 3–7 mm wide ................... C. simplicifolia Leaves subsessile, 3-foliolate; leaflets 1–3 mm wide ................................ 4 Calyx tube 16–22 mm long, 9–13 mm broad at throat, lobes 9–15 mm long; bracteoles 8–14 mm long; flowers 5.5–7.5 cm long; petioles to 10 mm long; lower surface of leaflets glabrate ................................. C. guianensis Calyx tube 11–15 mm long, 6–9 mm broad at throat, lobes 6–9 mm long; bracteoles 6–9 mm long; flowers 4–5.5 cm long; petioles 2–4 mm long; lower surface of leaflets pubescent ......................................... C. laurifolia Flowers 2.5–4 cm long, purple or white with purple marginally; calyx tube 4–7 mm wide at throat, lobes 2–4 mm long; valves of fruit depressed between seeds (fruit unknown in C. steyermarkii); trees or shrubs .... 6 Flowers 4–8 cm long, lilac-purple to pinkish mauve; calyx tube 8–13 mm wide at throat, lobes 4–13 mm long; valves of fruit not depressed between seeds; lianas or virgate shrubs ................................................... 8 Inflorescence to 0.5 cm long; flowers white with purple marginally; bracteoles linear, 3–5 × 0.3–0.5 mm; calyx pubescence uncinate; petiolules 4–6 mm long; leaflets 5–11 cm long ................................... C. steyermarkii Inflorescence 1–3.5 cm long; flowers dull purple; bracteoles ovate, 1.5–3 ×

282

F ABACEAE

1–2.5 mm; calyx pubescence densely strigose; petiolules 8–10 mm long; leaflets 10–20(–30) cm long ................................................................... 7 7(6). Calyx tube 7–11 mm long, lobes 2–3 mm long; bracteoles 1.5–3 mm wide; leaflets broadly ovate to orbicular, 8–18 cm wide; trees .......... C. dendrina 7. Calyx tube 13–15 mm long, lobes 3–4 mm long; bracteoles 1–1.5 mm wide; leaflets lanceolate-elliptic, 6–8 cm wide; shrubs .................... C. canescens 8(5). Lower surface of leaflets waxy, margins revolute; petiolules 2–4 mm long; fruits 9–15 mm wide; fruit dehiscence twisting 1–1.3 turns; virgate shrubs ......................................................................................... C. coriacea 8. Lower surface of leaflets not waxy, margins not revolute; petiolules 4–8 mm long; fruits 15–26 mm wide; fruit dehiscence twisting 0.2–0.5 turns; lianas .............................................................................. 9 9(8). Inflorescence 5–22 cm long; flowers 4–6 cm long, bluish violet; bracteoles 10–15 mm long; stipe 14–20 mm long; lower surface of leaflets rufoustomentose .............................................................................. C. arborescens 9. Inflorescence to 5 cm long; flowers 6–8 cm long, pink-rose to pinkish mauve; bracteoles 2–10 mm long; stipe 24–37 mm long; lower surface of leaflets glabrate ................................................................................... 10 10(9). Calyx tube 11–16 mm long, pubescence predominately uncinate; bracteoles 7–10 mm long; inflorescence to 0.5 cm long; staminal tube 3–4 cm long; vexillar claw 6–9 mm long; stipels 1–3 mm long ................ C. sagotii 10. Calyx tube 16–24 mm long, pubescence strigose; bracteoles 2–7 mm long; inflorescense 0.5–5 cm long; staminal tube 4–5 cm long; vexillar claw 14–19 mm long; stipels 3–7 mm long .................................................. 11 11(10). Style 16–25 mm long, subequal to ovary length; fruit pubescence rufostrigose; leaflets with 9–13(–15) pairs of lateral veins; inflorescences 1–5 cm long .............................................................................. C. javitensis 11. Style 24–32 mm long, ca. twice the ovary length; fruit pubescence uncinate; leaflets with 7–9 pairs of lateral veins; inflorescences 0.5–1 cm long ........................................................................................ C. cavalcantei Clitoria arborescens W.T. Aiton, Hortus Kew ed. 2, 4: 302. 1814. —Ternatea arborescens (W.T. Aiton) Kuntze, Revis. Gen. Pl. 1: 210. 1891. Clitoria poitaei DC., Prodr. 2: 234. 1825. Clitoria amoena Miq., Stirp. Surinam. Select. 24. 1850 [1851]. Liana, occasionally scandent shrub. Open forest areas, along forest margins of river banks, 50–500 m; Delta Amacuro (Río Amacuro, Río Cuyubini, Río Orinoco, Serranía de Imataca), Bolívar (widespread in northern part of state, base of Auyán-tepui). Miranda, Monagas, Sucre, Zulia; Colombia, Guyana, Suriname, French Guiana. Clitoria arborescens is not in Central America as has been reported in the literature. These erroneous reports are based upon misidentified specimens, usually Clitoria javitensis or Clitoria glaberrima.

This species has been cultivated for its climbing habit and showy floral display. The seeds are used as a fish poison in Guyana. Clitoria canescens Pittier ex Fantz, Sida 8: 305. 1980. Shrub. Sandy places along rivers, ca. 100 m; Amazonas (Río Guainía at Caño San Miguel). Endemic. Clitoria cavalcantei Fantz, Sida 9: 201. 1982. Small shrub with a climbing apex or liana. Riparian forests and clearings, 500–600 m; Bolívar (Río Curutu in upper Río Paragua). Brazil (Amazonas, Pará). Clitoria coriacea Schery, Fieldiana, Bot. 28: 260. 1952. Clitoria cerifera R.S. Cowan, Mem. New York Bot. Gard. 9: 349. 1957.

Clitoria

Virgate shrub 1.5–2.5 m tall. White-sand savannas and shrublands, 50–200 m; scattered in western Amazonas. Endemic. Clitoria coriacea will likely be found in adjacent Colombia (Guianía). Clitoria dendrina Pittier, Contr. U.S. Natl. Herb. 20: 126. 1918. —Generala, Oreja de tigre, Peonío de cerro. Tree 2–11(–25) m tall. Open areas of semideciduous forests, along forest borders, savannas, 50–400 m; Bolívar (Caño Asisa off Río Parú, La Prisión, middle Río Orinoco, Río Parguaza, east of Túriba), Amazonas (Caño Asisa, Puerto Ayacucho). Aragua, Barinas, Carabobo, Cojedes, Lara, Mérida, Portuguesa; Colombia. ŠFig. 248. Clitoria falcata Lam., Encycl. 2: 51. 1786. Clitoria rubiginosa Juss. ex Pers., Syn. Pl. 2: 303. 1807. Clitoria glycinoides DC., Prodr. 2: 234. 1825. Herbaceous vine. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, introduced and naturalized in Africa and Asia; 4 varieties, 1 in Venezuela. Variety aurantiaca (Benth.) Fantz and var. latifolia (Rizzini) Fantz are endemic to Brazil; var. glabrescens (Verdc.) Fantz occurs in Africa and is introduced into Guadeloupe and Tobago. C. falcata var. falcata Savannas, forest borders, 50–1000 m; Bolívar (Gran Sabana), Amazonas (between Caicara and Puerto Ayacucho, near Yavita). Widespread throughout most of Venezuela north of the Orinoco; other distribution neotropically as in species, and also introduced into Africa (Gold Coast, Liberia, Nigeria) and Asia (Java, Taiwan). ŠFig. 253. Clitoria guianensis (Aubl.) Benth., J. Proc. Linn. Soc., Bot. 2: 40. 1858. —Crotalaria guianensis Aubl., Hist. Pl. Guiane 761, t. 305. 1775. —Clitoria guyanensis Benth. in Mart., Fl. Bras. 15(1): 121. 1862. Subshrub to perennial, suffrutescent herb 10–60 cm tall. Southern Mexico, Central America, western Cuba, Colombia, Venezuela, Brazil, Bolivia; 3 varieties, 1 in Venezuela.

283

Clitoria guianensis is not in eastern Cuba and elsewhere in the West Indies as has been reported in the literature. These reports are usually misidentified Clitoria laurifolia. The varieties chapadensis (Malme) Fantz and macrocleistogama Fantz are localized endemics in Brazil (Mato Grosso). C. guianensis var. guianensis. —Generala. Savannas, 200–800 m; Bolívar (widespread), Amazonas (Río Ventuari basin). Widespread in northern Venezuela; other distribution as in species. ŠFig. 251. Clitoria javitensis (H.B.K.) Benth., J. Proc. Linn. Soc., Bot. 2: 42. 1858. —Neurocarpum javitense H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 409. 1823 [1824], emend. Fantz, Sida 9: 162. 1981. —Ternatea javitensis (H.B.K.) Kuntze, Revis. Gen. Pl. 1: 210. 1891. Liana. Tropical forests, usually associated with sandy soil, 50–700 m; Costa Rica, Panama, Colombia, Venezuela, Peru, Brazil; 4 varieties, 3 in Venezuela, 2 of these in the flora area. Variety portobellensis (Beurl.) Fantz is distributed in Central America, Colombia, and Venezuela (Carabobo, Miranda, Yaracuy). Variety longiloba Fantz is endemic to Peru (Loreto). Key to the Varieties of C. javitensis 1. Inflorescence 4–18 cm long, many-flowered; petiole commonly 10–25 cm long, petiolule 6–9 mm long; stipe 19–24 mm long; leaflets ovate, to 28 cm long, 15 cm wide ........................... var. grandifolia 1. Inflorescence 0.5–6 cm long, few-flowered; petiole commonly 4–10 cm long, petiolule 4–6 cm long; stipe 25–37 mm long; leaflets oblong-elliptic, to 18 cm long, 7(–9) cm wide ............... var. javitensis C. javitensis var. grandifolia (Ducke) Fantz, Sida 9: 170. 1981. —Clitoria grandifolia Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 141. 1930. 100–500 m; Amazonas (Cerro Sipapo, Río Cunucunuma). Colombia, Peru, Brazil. C. javitensis var. javitensis 50–700 m; Bolívar (Río Caroní, Río Caura, Río Paragua, Río Parguaza), Amazonas (wide-

284

F ABACEAE

spread). Amazonian Colombia, Brazil (Amazonas, Rondônia). ŠFig. 250. The typical form and f. bracteosubtenda Fantz are known from the flora area. The typical form has bracteoles 2–3 mm long, subopposite to alternate on the pedicel, inserted 2.5–5 mm below the base of the calyx, the apex below the calyx base or barely reaching it. Forma bracteosubtenda has bracteoles 3–5 mm long, subopposite, subtending calyx base, inserted 1–2 mm below the base of the calyx, the apex reaching the swollen calyx base (represented internally by the disk).

Neurocarpum javitense auct. non H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 409. 1823 [1824]: sensu Clitoria javitensis (H.B.K.) Benth., J. Proc. Linn. Soc., Bot. 2: 42. 1858. Liana. Climber in forests or thickets, or occasionally as trailing vine in open areas, associated with sandy soil. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 3 varieties, 2 in Venezuela, both in the flora area. Variety sagotii is endemic to French Guiana. Key to the Varieties of C. sagotii

Clitoria laurifolia Poir. in Lam., Encycl. supp. 2: 301. 1811. Neurocarpum cajanifolium C. Presl., Symb. Bot. 1: 17, t. 9. 1830. —Clitoria cajanifolia (C. Presl.) Benth. in Mart., Fl. Bras. 15(1): 121. 1862. Clitoria parvifolia Pittier, Bol. Técn. Minist. Agric. 5: 149. 1944, nom. nud. Subshrub. Sandy soil in savannas and along riverbanks, near sea level to 1300 m; Delta Amacuro (Isla Las Cidras near Misión San Francisco de Guayo), Bolívar (El Palmar, Gran Sabana, Puerto Ordaz), Amazonas (Río Negro). Anzoátegui, Apure, Zulia; West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, introduced and established in eastern Africa and southeastern Asia. Clitoria laurifolia is not known from Central America as has been reported in the literature. That material is usually misidentified Clitoria guianensis. Three forms are found in the flora area: (1) f. parvifolia with leaflets oblong-linear, 2.5–3.5 × 0.7–1.3 cm, petiole 1–1.5 mm long, and rachis 7–13 mm long; (2) f. glabrior with leaflets oblong to elliptic-oblong, 4–13 × 1.5– 3.5 cm, petioles 2–5 mm long, rachis 4–9 mm long, and a legume with weakly raised costa, extending 20–60% of the length of the valve, or ecostate (often a mixture of ecostate and costate legumes on same plant); and (3) f. laurifolia with leaflets similar to f. glabrior, but legume with prominently raised costa, extending ca. 75–90% of the length of the valve (otherwise all legumes similar). Clitoria sagotii Fantz, Sida 8: 94. 1979. Clitoria javitensis var. glabra Sagot, Ann. Sci. Nat. Bot. sér. 6, 13: 299. 1882.

1. Petiole ± canaliculate adaxially; bracteoles 4–6 mm long, inserted 2–4 mm below the calyx; petiolules 4–6 mm long; main veins of leaflets 6–8 pairs .............. ................................. var. canaliculata 1. Petiole ± flattened adaxially; bracteoles 7– 10 mm long, inserted 1–2 mm below the calyx; petiolules 6–9 mm long; main veins of leaflets 8–10 pairs ....... var. sprucei C. sagotii var. canaliculata Fantz, Sida 9: 351. 1982. 100–200 m; Bolívar (Río Cuyuní). Guyana, Suriname. ŠFig. 249. C. sagotii var. sprucei Fantz, Sida 9: 353. 1982. 100–200 m; Amazonas (Río Casiquiare, Río Guainía, Río Negro, San Antonio on upper Río Orinoco). Colombia (Amazonas), Brazil (Amazonas). Clitoria simplicifolia (Kunth) Benth., J. Proc. Linn. Soc., Bot. 2: 40. 1858. —Neurocarpum simplicifolium Kunth, Mimoses 213, t. 59. 1819 [1824]. —Ternatea simplicifolia (Kunth) Kuntze, Revis. Gen. Pl. 1: 120. 1891. Subshrub to perennial suffrutescent herb, trailing to erect, 10–40 cm tall. Riparian forests, disturbed areas, 50–200 m; Amazonas (Isla Ratón, around Puerto Ayacucho, Río Orinoco). Brazil (Goiás, Mato Grosso, Pará), Paraguay. ŠFig. 252. Clitoria steyermarkii Fantz, Ann. Missouri Bot. Gard. 76: 1165. 1989. Slender shrub to 2 m tall. Upland scrub, ca. 700–800 m; Bolívar (Cerro Guaiquinima). Endemic.

Clitoria 285

Fig. 248. Clitoria dendrina

Fig. 249. Clitoria sagotii var. canaliculata

286

F ABACEAE

Fig. 250. Clitoria javitensis var. javitensis

Fig. 251. Clitoria guianensis var. guianensis

Coursetia 287

Fig. 252. Clitoria simplicifolia

Fig. 253. Clitoria falcata var. falcata

18. COURSETIA DC., Ann. Sci. Nat. (Paris) 4: 92. 1825. by Nidia L. Cuello A. Small trees or shrubs. Stems usually spindly, erect to scandent. Leaves pulvinate, odd- or even-pinnate, or unifoliolate; petiolules shorter than the leaflets; leaflets orbicular or narrowly to widely elliptic, mucronate, upper surfaces glabrate to silky, lower surfaces pilose to silky or woolly, caducous; stipules small, subulate, persistent; stipels present or absent. Inflorescences of pendent to erect, sessile or pedunculate, axillary racemes, rarely panicles. Flowers ebracteolate; floral bracts broadly lanceolate to linear-lanceolate, readily caducous or sometimes persistent. Calyx campanulate, rounded or rarely attenuate at base, strigose to silky or tomentose, or sometimes stipitate-glandular, the lobes ligulate, lanceolate to triangular, acute to rounded; petals all yellowish brown, reddish brown, or purple; standard orbicular, emarginate, usually whitish to yellow and/or reddish, calluses and inflexed auricles usually well developed; wing petals commonly protruding beyond the keel at anthesis, parallel to the keel, whitish yellow to reddish; keels rostrate, the tip acute. Ovary granuliferous to silky, villous, or woolly, rarely glabrous, sessile or nearly so on the receptacle, 7–30-ovulate. Legumes glabrous to pubescent. Seeds uniformly brownish, with darker purple mottling. Neotropics (most diverse in Mexico and Central America); ca. 39 species, 4 in Venezuela, 1 of these in the flora area.

288

F ABACEAE

Fig. 254. Coursetia ferruginea

Coursetia ferruginea (H.B.K.) Lavin in Stirton, Adv. Legume Syst. 3: 63. 1987. —Robinia ferruginea H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 395. 1823 [1824]. —Humboldtiella ferruginea (H.B.K.) Harms, Repert. Spec. Nov. Regni Veg. 19: 12. 1923. —Majomo sabanero. Coursetia arborea Griseb., Fl. Brit. W. I. 2: 183. 1859. Tree or shrub 2–3 m tall; flowers white to pale lavender. Deciduous to evergreen lowland forests or savannas, 100–400 m; Delta Amacuro (Los Castillos), northern Bolívar, Amazonas (Caño Asisa, Río Ocamo). Widespread elsewhere in Venezuela; Panama, Colombia, Trinidad, Guyana, Brazil (Roraima). ŠFig. 254.

Crotalaria 289

19. CROTALARIA L., Sp. Pl. 714. 1753; Gen. Pl. ed. 5. 320. 1754. by Nidia L. Cuello A. Herbs or shrubs, erect to prostrate. Stems pubescent or glabrous, smooth or striate. Leaves ternately compound or unifoliolate by reduction; stipules varying greatly in size and shape or lacking; leaflets entire, glabrous or pubescent. Inflorescences racemose, terminal or opposite the leaves; bracts setaceous to foliaceous; bracteoles paired, similar in shape to the bracts, borne on the pedicel or the calyx. Flowers sometimes showy, bisexual, 5-merous. Calyx green, bilabiate, the tube usually campanulate, glabrous or pubescent, the lobes usually longer to much longer than the tube; corolla papilionaceous, usually yellow; standard orbicular or obovate, the apex rounded or retuse, the base clawed, the adaxial surface and veins near the base frequently becoming red-tinged; wing petals oblong, attached to a stalk continuous with the lower margin (cleaver-shaped), the blades usually puckered between some of veins; keel petals usually twisted at the tip, woolly-ciliate. Stamens 10, monadelphous, the staminal tube open on the upper side; anthers dimorphic with basifixed, long anthers alternating with medifixed short anthers, the short anthers positioned above the elongate ones at anthesis. Ovary terminated by a geniculate style. Fruit inflated, subcylindric, sessile or short-stipitate, glabrous or pubescent, the fruits of some species black at maturity. Seeds 7–46, oblique-cordiform; testa smooth and mostly impermeable to water. Tropics and subtropics, a few species in temperate areas; ca. 600 species, ca. 20 in Venezuela, 10 of these in the flora area. Key to the Species of Crotalaria 1. 1. 2(1).

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

5. 6(5).

Leaves 3-foliolate ....................................................................................... 2 Leaves 1-foliolate ....................................................................................... 4 Petioles equal to or longer than the terminal leaflet; bracts absent; fruits villous to spreading-pubescent, trichomes often ca. 2 mm long ............. ....................................................................................................... C. incana Petioles shorter than the terminal leaflet; bracts present; fruit appressedpubescent to strigulose, trichomes usually < 0.5 mm long .................. 3 Leaflets narrow or linear; racemes elongate; bracts 1–3 mm long, almost straight ................................................................................ C. maypurensis Leaflets oblong to ovate-lanceolate; racemes short; bracts 5–10 mm long, curled, silky-pubescent ................................................................ C. micans Stipules usually absent, inconspicuous when present.............................. 5 Stipules present and conspicuous, or caducous ........................................ 7 Leaflets linear-lanceolate to lanceolate, 0.2–0.9 cm wide; stipules usually absent, when present consisting of an inconspicuous, narrow (0.2–0.5 mm wide) decurrent wing terminating at the base of the leaf without visible lobes .......................................................................................... C. sagittalis Leaflets obovate, obovate-elliptic, lanceolate, or oblanceolate, 0.5–5 cm wide; stipules absent.............................................................................. 6 Leaflets obovate to obovate-elliptic, 1.5–5 cm wide; densely appressedpubescent on both sides; inflorescences 6–12 cm long; flowers 1–1.5 cm long ................................................................................................. C. nitens

290

F ABACEAE

6.

Leaflets lanceolate to oblanceolate, 0.5–1 cm wide; sparsely appressed pubescent on both sides; inflorescences 1.5–5 cm long; flowers 0.5–0.8 cm long ....................................................................................... C. aff. velutina 7(4). Stipules free; leaves oblanceolate .............................................................. 8 7. Stipules decurrent; leaves elliptic, obovate, lanceolate, or narrow-lanceolate ..................................................................................................... 9 8(7). Stipules and bracts persistent, ovate, 5–15 mm long; calyx glabrous ................................................................................................ C. spectabilis 8. Stipules and bracts caducous, narrow, 2–3 mm long; calyx strigulosepubescent .......................................................................................C. retusa 9(7). Stipules decurrent, longer than the length of one internode (3–8 mm wide, the width constant), this occurring throughout the length of the stem and thus the entire stem appearing winged; stipule lobes usually not spreading; inflorescences terminal or terminal and leaf-opposed .......... C. pilosa 9. Stipules decurrent, shorter than the length of one internode, usually only along the uppermost internodes of the stem; stipule lobes spreading or recurved; inflorescences not terminal, always leaf-opposed .............. 10 10(9). Leaflets obovate to ovate-elliptic, the lowermost usually obovate, the uppermost ovate to ovate-elliptic; stipule lobes falcate or recurved ................................................................................................. C. stipularia 10. Leaflets all elliptic-lanceolate or linear-lanceolate; stipule lobes spreading, sagittate, apiculate ........................................................... C. sagittalis Crotalaria incana L., Sp. Pl. 716. 1753. Herb or small shrub to 1.3 m tall; flowers yellow. Roadsides, ca. 50 m; Delta Amacuro (between San José and Clavellina, near Tucupita), Bolívar (Caicara). Anzoátegui, Apure, Aragua, Distrito Federal, Guárico, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Zulia; Mexico, Central America, Colombia, Ecuador, Peru, Brazil (Bahia, Pará), Bolivia, Argentina, Africa, Asia (China, Philippines, Malaysia).

C. maypurensis var. depauperata (Mart.) Windler & S.G. Skinner, Phytologia 50: 186. 1982. —Crotalaria depauperata Mart., Fl. Bras. 15(1): 30. 1859. —Generala. Herb with ascending stem to 1 m tall; flowers pale yellow. Swampy savannas, 50– 100 m; Bolívar (near Caicara, near Maripa, near San Juan de Manapiare), Amazonas (Río Ocamo, near San Juan de Manapiare). Brazil (Minas Gerais). ŠFig. 255.

Crotalaria maypurensis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 403. 1823 [1824]. —Generala. Crotalaria lectophylla Benth., Ann. Nat. Hist. 3: 430. 1839. Crotalaria anagyroides var. pauciflora Griseb., Cat. Pl. Cub. 69. 1866. Neotropics; 2 varieties, both in the flora area.

C. maypurensis var. maypurensis Shrub or suffrutescent herb; flowers yellow. Savannas, sandy open river banks, 50– 1000 m; Bolívar (widespread), Amazonas (near Canaripó, Culebra, Puerto Ayacucho to Samariapo, Santa Bárbara). Monagas, Zulia; Mexico, Central America, Colombia, British Guiana, Peru, Brazil, Paraguay, Argentina. ŠFig. 256.

Key to the Varieties of C. maypurensis 1. Leaflets linear, the terminal leaflet usually 0.8–3.5 × 0.1–0.25 cm ...................... ................................. var. depauperata 1. Leaflets oblanceolate, lanceolate, or elliptic, the terminal leaflet usually 4.5–5.5 × 0.7–1.5 cm .............. var. maypurensis

Crotalaria micans Link, Enum. Hort. Berol. Alt. 2: 228. 1822. Crotalaria anagyroides H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 404. 1823 [1824]. Suffrutescent herb or shrub to 3 m tall; flowers yellow. Sandy savannas, 50–800 m; Bolívar (Altiplanicie de Nuria, lower Río

Crotalaria 291

Caroní, near Santa Elena), Amazonas (Isla Ratón, upper Río Orinoco). Barinas, Carabobo, Distrito Federal, Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira, Sucre, Trujillo, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, widespread in Brazil, Bolivia, Paraguay, Argentina, Africa, Sri Lanka. Crotalaria nitens H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 399. 1824. Crotalaria bracteata Schltdl. & Cham., Linnaea 5: 575. 1830. Crotalaria schiedeana Steud., Nomencl. Bot. ed. 2, 1: 445. 1840. Herb 0.5–2 m tall; flowers yellow. Disturbed areas, 100–200 m; Amazonas (Tayari along upper Río Orinoco). Barinas, Mérida, Táchira; Mexico, Guatemala, Honduras, Costa Rica, Colombia, Ecuador, Peru, Bolivia. Crotalaria pilosa Mill., Gard. Dict. ed. 8. no. 2. 1768, non Crotalaria pilosa Thunb. 1800, nec Crotalaria pilosa Roxb. ex Mart. 1820. —Wareka marakae-yó (Panare). Crotalaria pterocaula Desv., J. Bot. Agric. 3: 76. 1814. Crotalaria pilosa var. skutchii Senn, Rhodora 41: 331. 1939. Suffrutescent to 60 cm tall; flowers yellow. Savannas, 100–400 m; Bolívar (Altiplanicie de Nuria, near Ciudad Piar, near Corozal on Río Maniapure, lower Río Caroní, near Santa Elena de Uairén, Serranía de los Pijiguaos), Amazonas (between Maguari and Río Parucito, Río Negro). Barinas, Mérida, Portuguesa; Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia, Paraguay. ŠFig. 259. The fruit of Crotalaria pilosa is used as a rattle by the Panare Amerindians. Crotalaria retusa L., Sp. Pl. 715. 1753. —Anilillo, Maraca de cascabel. Dolichos cuneifolius Forssk., Fl. Aegypt.Arab. 134. 1775. —Crotalaria cuneifolia (Forssk.) Schrank, Syll. Pl. Nov. 2: 78. 1828. Crotalaria retusifolia Stokes, Bot. Mat. Med. 3: 516. 1812. Herb or suffrutescent herb to 1 m tall; flowers yellow with reddish brown on outside of standard. Roadsides, sandy savannas, 50–

400 m; Delta Amacuro (Pedernales, near Tucupita), Bolívar (lower Río Caroní, middle Río Caura, Tumeremo, near Upata). Anzoátegui, Apure, Barinas, Distrito Federal, Falcón, Guárico, Miranda, Monagas, Portuguesa, Zulia; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Africa, India, Sri Lanka, China, Malesia, Philippines. Crotalaria sagittalis L., Sp. Pl. 714. 1753. Herb or suffrutescent herb 10–60 cm tall; flowers yellow. Savannas, ca. 100 m; Amazonas (La Esmeralda, near Puerto Ayacucho). Apure, Barinas, Yaracuy; U.S.A., Mexico, Central America, Colombia, Ecuador, Peru, Brazil (Goiás), Bolivia, Paraguay. Crotalaria sagittalis is a polymorphic species with the stipules varying in size and shape, 2–10 mm long or more, decurrent, sagittate, apiculate, pubescent or glabrate, or lacking. Crotalaria spectabilis Roth, Nov. Pl. Sp. 341. 1821. Erect herb; flowers yellow. Disturbed areas, 50–200 m; Bolívar (vicinity of El Callao and Tumeremo, lower Río Caroní). Barinas, Monagas, Portuguesa; Panama, Colombia, Trinidad, Peru, Brazil (Pará, São Paulo), native to India, naturalized in many tropical (or occasionally temperate) localities in the Old and New Worlds. Crotalaria stipularia Desv., J. Bot. (Desvaux) 3: 76. 1814, spelling variant: Crotalaria stipularis. —Ajonjolí sabanero. Erect to creeping herb with silvery green stems and leaves; flowers yellow. Savannas with igneous outcrops, slopes, 50–300 m; northern Bolívar. Anzoátegui, Aragua, Distrito Federal, Guárico, Mérida, Miranda, Monagas, Portuguesa, Sucre, Yaracuy; Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 257. Crotalaria aff. velutina Benth., Ann. Nat. Hist. 3: 429. 1839. Erect herb to 60 cm long; flowers yellow. Savannas, 50–300 m; Bolívar (road between Caicara and Puerto Ayacucho). Apure; Brazil. ŠFig. 258. This taxon may represent a new species, but more fruits and flowers are needed to determine this with certainty.

292

F ABACEAE

Fig. 255. Crotalaria maypurensis var. depauperata

Fig. 256. Crotalaria maypurensis var. maypurensis

Crotalaria 293

Fig. 257. Crotalaria stipularia

Fig. 258. Crotalaria aff. velutina

Fig. 259. Crotalaria pilosa

294

F ABACEAE

20. CYMBOSEMA Benth., J. Bot. (Hooker) 2: 60. 1840. by Richard H. Maxwell Vines to 10 m long. Leaves alternate, 3-foliolate; stipules acute; leaflets broadly lanceolate to oblong, entire. Inflorescence axillary, erect, pseudoracemose, unbranched; bracts ovate, persistent. Flowers ca. 4 cm long, several on each node. Calyx 4-lobed; corolla reddish, drying purplish, petals with claws 4–7 mm long; standard obovate; wing petals oblanceolate, free; keel petals oblanceolate, fused distally. Stamens 10, diadelphous, the vexillar one free. Ovary sessile, ca. 6-ovulate; stigma and style glabrous. Fruit a legume, ca. 44 × 18 × 4 mm, oblong, usually ± falcate, with a long, down-curved beak, glabrescent, ca. 4-seeded. Seeds ca. 10 × 6 × 4 mm, hard, dark brown, the hilum nearly halfway encircling the seed. Mexico, Costa Rica, Panama, Colombia, Venezuela, Amazonian Peru and Brazil; 1 species.

Fig. 260. Cymbosema roseum

Dalbergia 295

Cymbosema roseum Benth., J. Bot. (Hooker) 2: 60. 1840. —Raudal Bobadilla. Cymbosema apurense (H.B.K.) Pittier: sensu Pittier, Bol. Soc. Venez. Ci. Nat. 7: 154. 1941, non Dioclea apurensis H.B.K. 1823 [1824]. Vine; flowers reddish (may change to

purple when dried). Mostly river banks and forests, near sea level to 200 m; scattered in Delta Amacuro, northern Bolívar, and Amazonas. Apure, Miranda, Yaracuy; extra-Venezuelan distribution as in genus. ŠFig. 260. The standard of the flowers is not reflexed, which suggests hummingbird pollination.

21. DALBERGIA L. f., Suppl. Pl. 52. 1781 [1782], nom. cons. Amerimnon P. Browne, Civ. Nat. Hist. Jamaica 288. 1756. Ecastaphyllum P. Browne, Civ. Nat. Hist. Jamaica 299. 1756. by Gerardo A. Aymard C. Trees or shrubs, sometimes with scandent branches, or woody lianas. Leaves alternate, odd-pinnate, some species 1-foliolate; stipules ovate to subulate, small, deciduous; stipels absent; leaflets usually alternate, 3–several. Inflorescences paniculate, sometimes racemose, terminal or axillary. Flowers small, on short pedicels; bracts and bracteoles deciduous. Calyx campanulate, the teeth 5, the 2 upper ones broadest, the lowest one usually longest; petals subequal, clawed; standard glabrous, ovate, obovate, or orbicular; wing petals oblong; keel petals coherent along the lower margin. Stamens 9 or 10, monadelphous or disposed in 2 fascicles of 5, diadelphous, or absent; anthers minute, basifixed, dehiscing by 2 horizontal slits. Ovary stipitate, few-ovulate; style subulate, glabrous; stigma capitate or indeterminate. Fruits stipitate, reniform to oblong or linear, occasionally constricted medially, wingless or with the wing surrounding the seminiferous area, lightly reticulate, the margin not thickened, indehiscent. Seeds 1 or rarely 2, reniform and compressed. Pantropics; ca. 100 species, 16 in Venezuela, 15 of these in the flora area. Key to the Species of Dalbergia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

5(4). 5.

6(1). 6.

Leaves with 17–41 leaflets ......................................................................... 2 Leaves with 1–15 leaflets ........................................................................... 6 Leaves with 25–41 leaflets ........................................................... D. inundata Leaves with 17–21 leaflets ......................................................................... 3 Leaflets 0.8–1 × 0.4–0.6 cm .......................................................... D. inundata Leaflets 2–8 × 1–5 cm ................................................................................ 4 Leaflets oblong, the base rounded, apex obtuse; fruit with obvious reticulate venation ........................................................................... D. spruceana Leaflets ovate, ovate-elliptic, or circular, the base acute, apex acuminate; fruit without venation or with barely discernible reticulate venation ................................................................................................................ 5 Lower surface of leaflets appressed-pubescent; flowers dark violet; fruit 3–6 cm long, ovate, woody, not winged ........................................ D. foliosa Lower surface of leaflets glabrescent or sparsely pilose; flowers purple with white spots; fruit 7–10 cm long, oblong-elliptic, papery, winged ................................................................................................. D. spruceana Leaves 1-foliolate ....................................................................................... 7 Leaves 2–15-foliolate ................................................................................. 9

296

F ABACEAE

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

9(6). 9. 10(9). 10. 11(10).

11.

12(9). 12. 13(12).

13.

14(12). 14. 15(14). 15.

16(14). 16. 17(16). 17.

Leaflets orbicular; pedicels 3.5–4 mm long ........................................ D. sp. A Leaflets ovate or ovate-lanceolate; pedicels 1–2.5 mm long ..................... 8 Lower surface of leaflets not glaucous; calyx lobes unequal, the upper 2 fused; fruit oblong, glabrous .................................................. D. amazonica Lower surface of leaflets glaucous; calyx lobes equal, the upper 2 free or fused basally; fruit orbicular to ovate, sparsely pubescent .................... .......................................................................................... D. ecastophyllum Branches and branchlets glabrous or puberulent; lower surface of leaflets glabrous or sparsely pilose .................................................................. 10 Branches and branchlets ferruginous-tomentose or yellowish-tomentose; lower surface of leaflets pilose or densely pubescent ......................... 12 Leaflets orbicular or broadly elliptic; fruit orbicular ................. D. monetaria Leaflets ovate, ovate-lanceolate, ovate-oblong, or obovate; fruit reniform or oblong-elliptic .................................................................................. 11 Leaflets ovate, glabrous on upper surface, sparsely pubescent on lower surface, the apex retuse; inflorescence not fasciculate and clustered in the leaf axil, terminal or axillary; fruit oblong-elliptic, winged, membranaceous .............................................................................. D. frutescens Leaflets ovate-lanceolate, glabrous on both surfaces, the apex acuminate; inflorescence ± fasciculate, clustered in the leaf axil; fruit reniform, coriaceous, not winged, not membranaceous ........................... D. hygrophila Lower surface of leaflets appressed-pubescent or densely ferruginous-tomentose ................................................................................................ 13 Lower surface of leaflets pilose, or, if pubescent, pubescence not ferruginous-tomentose .................................................................................... 14 Lower surface of leaflets densely ferruginous-tomentose; calyx teeth equal, 4–5 mm long; fruit reniform, suborbicular, or quite orbicular, not winged, densely ferruginous-tomentose ..................................... D. riedelii Lower surface of leaflets appressed-pubescent; calyx teeth unequal, 1.5–2 mm long; fruit oblong, winged, glabrous, or sparsely pubescent ........... ................................................................................................ D. intermedia Leaflets widely ovate or orbicular ........................................................... 15 Leaflets oblong, obovate, ovate-elliptic, or obovate-oblanceolate ........... 16 Leaflets 3–6 cm wide, ovate, the upper surface shiny, the lower surface appressed-pubescent ....................................................................... D. sp. B Leaflets 7–12 cm wide, widely ovate or orbicular, chartaceous, the upper surface dull, the lower surface completely appressed-pubescent .......... ......................................................................................................... D. sp. C Lower surface of leaflets completely appressed-pubescent; calyx 1.2 mm long; ovary densely pubescent ........................................................ D. sp. D Lower surface of leaflets glabrous or sparsely appressed-pubescent; calyx 5–6 mm long; ovary glabrescent or pilose ........................................... 17 Leaflets elliptic-lanceolate; petals white; fruit reticulate ......... D. subcymosa Leaflets ovate-elliptic, oblong, or obovate; petals lilac or purple; fruit not reticulate ....................................................................................... D. foliosa

Dalbergia amazonica (Radlk. ex Köpff) Ducke, Leguminosas Amaz. Brasil. 122. 1939. —Ecastaphyllum amazonicum

Radlk. ex Köpff, Anat. Charakt. Dalberg. 40. 1892. —Anicillo, Caraota rebalsera, Kamatata (Yekwana).

Dalbergia 297

Scandent shrub or small tree; flowers white. Evergreen riparian lowland to lower montane forests, 50–500 m; Bolívar (Río Acanán, mouth of Río Botanamo, lower Río Caroní, upper Río Caura, Río Cuyuní, Río Erebato, mouth of Río Nichare, mouth of Río Paragua), Amazonas (Puerto Ayacucho, Raudal de Atures, lower Río Ventuari). Apure; Colombia, Guyana, French Guiana, Brazil (Amapá, Amazonas, Pará).

ests, 100–500 m; Bolívar (El Foco near Upata, Isla de Anacoco, La Escalera, Raudal Cotua on Río Asa, Río Caroní, Río Nichare), Amazonas (La Esmeralda, Río Casiquiare, Río Pasimoni, Río Siapa, Río Sipapo). Anzoátegui, Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amazonas, Bahia, Pará, Rio de Janeiro), Bolivia, Paraguay, northern Argentina. ŠFig. 261.

Dalbergia ecastophyllum (P. Browne ex L.) Taub. in Engl. & Prantl, Nat. Pflanzenfam. 3(3): 335. 1894. —Hedysarum ecastaphyllum P. Browne ex L., Syst. Nat. ed. 10, 2: 1169. 1759. —Chinay-yek (Arekuna), Bejuco saliva, Caraota rebalsera. Dalbergia brownei (Jacq.) Urb., Symb. Ant. 4: 295. 1905. —Amerimnon brownei Jacq., Enum. Syst. Pl. 27. 1760, “brownii.” Climbing shrub, liana, or small tree; petals white. Evergreen lowland to montane forests, flooded forests, 50–1300 m; Delta Amacuro (Caño Güiniquina, near Pedernales), Bolívar (45 km north of Cerro Impacto, Ciudad Bolívar, Puerto Ordaz, Río Botanamo, Río Caura, Río Cuyuní, Río Paragua, Río Tirica to Urimán, Sierra de Maigualida), Amazonas (Isla Ratón, ca. 9 km south of Puerto Ayacucho). Apure, Carabobo, Guárico, Mérida, Miranda, Monagas, Sucre, Táchira, Zulia; U.S.A. (southern Florida), Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amazonas, Bahia, Distrito Federal, Espirito Santo, Paraná, Rio de Janeiro), West Africa.

Dalbergia frutescens (Vell.) Britton, Bull. Torrey Bot. Club. 16: 324. 1889. —Pterocarpus frutescens Vell., Fl. Flumin. 283. 1825 [1829]. —Sangrito. Dalbergia variabilis Vogel, Linnaea 11: 196. 1837. Neotropics; 2 varieties, 1 in Venezuela.

Dalbergia foliosa (Benth.) A.M. Carvalho, Brittonia 49: 100. 1997. —Ecastaphyllum foliosum Benth., J. Bot. (Hooker) 2: 64. 1840. —Arepillo, Jadewa-jadakwadiyo (Yekwana), Uaypeu-roy-yek (Pemón). Ecastaphyllum glaucum Desv., Ann. Sci. Nat. (Paris) sér. 1, 9: 423. 1826. —Dalbergia glauca (Desv.) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 50. 1939, non Roxb. 1831, nec Kurz 1876. Shrub with climbing branches; petals violet. Evergreen lowland to lower montane for-

D. frutescens var. frutescens. —Bejuco de estribo, Bejuco roblito, Sangrito. Liana; flowers white. Deciduous to evergreen lowland or lower montane forests, 50– 300 m; Bolívar (Puerto Ordaz, Represa Guri, mouth of Río Paragua, Upata). Anzoátegui, Apure, Falcón, Miranda, Guárico, Sucre, Zulia; Peru, Brazil, Bolivia, Paraguay, northern Argentina. ŠFig. 265. Dalbergia hygrophila (Mart. ex Benth.) Hoehne, Fl. Brasilica 25(3): 21. 1941. —Ecastaphyllum hygrophilum Mart. ex Benth., Comm. Legum. Gen. 29. 1837. —Bejuco colorado. Scandent shrub, small tree, or liana; flowers white. Gallery forests, evergreen lowland to lower montane forests, 50–500 m; Delta Amacuro (Caño Araguabisi, Caño Güiniquina), Bolívar (Río Acanán, lower Río Caura), Amazonas (Caño Campana in Río Atabapo basin, Caño Ucata southeast of Síquita, Maroa, Río Baría, Río Cataniapo, Río Mawarinuma, Río Orinoco). Anzoátegui, Apure; Colombia, Peru, Brazil (Amazonas, Pará). ŠFig. 264. Dalbergia intermedia A.M. Carvalho, Brittonia 49: 101. 1997. —Ecastaphyllum tomentosum Spruce ex Benth, J. Linn. Soc. 4(supp.): 51. 1860. —Dalbergia tomentosa (Spruce ex Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 144. 1982, non Vogel 1837, nec (Benth.) Taub. 1891.

298

F ABACEAE

Liana; flowers white. Evergreen lowland forests, 50–500 m; Bolívar (Río Zariapo on upper Río Caura, Sierra de Maigualida), Amazonas (Isla Ratón, Puerto Ayacucho). Apure; Guyana, Brazil, Bolivia. ŠFig. 263. Dalbergia inundata Spruce ex Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 49. 1860. —Drepanocarpus paludicola Standl., Trop. Woods 33: 12. 1933. —Uña de gato. Dalbergia aturensis Pittier, Bol. Soc. Venez. Ci. Nat. 8: 260. 1943. Shrub with climbing branches, small tree, or shrub; flowers white and purple. Gallery and evergreen lowland forests, flooded forests, forests bordering white-water and black-water rivers, 100–200 m; Amazonas (widespread). Anzoátegui; Colombia, Guyana, Peru, Brazil (Amazonas, Minas Gerais, Pará). ŠFig. 262. Dalbergia monetaria L. f., Suppl. Pl. 317. 1781 [1782]. —Eburu, Oteratana, Otero, Realito, Tu-basabosa. Climbing shrub, small tree, or liana; petals white. Semideciduous to evergreen lowland forests, lower montane forests, gallery forests, flooded forests along black-water rivers, 50–1000 m; Delta Amacuro (widespread), Bolívar (Canaima, Cerro Guaiquinima, Río Karaurín, upper Río Paragua, Río Parupa, San Felíx), Amazonas (Caño San Miguel, Maroa, Puerto Ayacucho, Río Atabapo, Río Baria, Río Casiquiare, Río Mawarinuma, upper Río Orinoco, Río Yudi). Falcón, Miranda, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. ŠFig. 268. This species has also sometimes been called Dalbergia volubilis Urb. (Repert. Spec. Nov. Regni Veg. 16: 136. 1919), but that name was incorrectly based on Securidaca volubulis L. (Species Plantarum 707. 1753), Polygalaceae. Dalbergia riedelii (Benth.) Sandwith, Bull. Misc. Inform. 1931: 358. 1931. —Ecastaphyllum monetaria var. riedelii Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 50. 1860. —Ecastaphyllum riedeli (Benth.) Radlk. in Köpff, Anat. Charakt. Dalberg. 41. 1892.

Dalbergia pachycarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 145. 1922. Liana or climbing shrub; petals white. Evergreen lowland to montane forests, 50–1000 m; Bolívar (Caño Chiviripa near Río Maniapure, La Escalera), Amazonas (Cerro Cariche, La Esmeralda, Macuruco, Río Asisa, Río Atabapo, Río Casiquiare, Santa Barbara del Orinoco). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Acre, Amazonas, Pará, Roraima). ŠFig. 266. Dalbergia spruceana Benth., J. Proc. Linn. Soc. Bot. 4(suppl.): 35. 1860. Tree or climbing liana; flowers white. Evergreen lowland or seasonally dry forests, 100–300 m; Bolívar (El Pilón near Cerro Marimarota, Río Parguaza), Amazonas (Cerro Yutajé, Río Coro Coro). Brazil (Acré, Amapá, Amazonas), Bolivia. ŠFig. 267. Dalbergia subcymosa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 144. 1922. Liana. Evergreen lowland forests, near sea level to 100 m; Delta Amacuro (San Victor near Guyana border). Suriname, French Guiana, Peru, Brazil. Dalbergia sp. A Liana; flowers white. Granitic outcrops, evergreen lowland forests, ca. 100 m; Bolívar (Jabillal on Río Caura). Dalbergia sp. A is based on Fernández 5267 (MO, PORT). Dalbergia sp. B Liana; flowers white. Deciduous forests, 50–100 m; Bolívar (Puerto Ordaz to San Felíx). Dalbergia sp. B is based on Aristeguieta 5314 (MO, VEN). Dalbergia sp. C Liana; flowers white. Gallery forests, 100–200 m; Amazonas (Río Mawarinuma). Dalbergia sp. C is based on Davidse & Miller 27422 (MO, NY, VEN). Dalbergia sp. D. —Wakado (Yekwana). Tree; flowers yellow. Evergreen forests, 1200–1300 m; Amazonas (upper Río Ventuari). Dalbergia sp. D is based on Chaviel 417 (MO, PORT).

Dalbergia 299

Fig. 261. Dalbergia foliosa

Fig. 262. Dalbergia inundata

Fig. 263. Dalbergia intermedia

300

F ABACEAE

Fig. 264. Dalbergia hygrophila

Fig. 266. Dalbergia riedelii

Fig. 265. Dalbergia frutescens var. frutescens

Dalbergia 301

Fig. 267. Dalbergia spruceana

Fig. 268. Dalbergia monetaria

302

F ABACEAE

22. DERRIS Lour., Fl. Cochinch. 432. 1790, nom. cons. by Hans-Helmut Poppendieck Woody vines. Leaves alternate, odd-pinnate; stipules caducous, stipels absent, leaflets opposite, entire. Inflorescence a lateral or terminal pseudoraceme composed of short flower-bearing shoots to 3 mm long (to 10 mm long and secund in D. amazonica); bracts caducous; bracteoles 2, borne on the middle of the pedicel. Calyx dentate to truncate, cup-shaped; corolla white to light yellow (purple in D. amazonica); standard ovate, clawed, not reflexed, the base attenuate, without callosities, the apex obtuse (reflexed and truncate with basal callosities in D. amazonica); wing petals adhering to keel by pressure and structural conformity; keel petals coherent or connate; wing and keel petals about equal. Vexillar stamen free at base; anthers uniform; nectary glands indistinct. Ovary pubescent; ovules 2–4; style glabrous. Fruit indehiscent, flat, 1- or 2-seeded, with submarginal wings on either side close to the vexillar margin. Colombia, Venezuela, Guyana, Suriname, French Guiana, northern Amazonian Brazil, Peru; 3 species, all in the flora area. See Lonchocarpus for a discussion of the taxonomic problems between these two genera. Key to the Species of Derris 1.

1.

2(1). 2.

Corolla deep lavender, the standard with a light green to white spot at its base and with basal callosities; short flower-bearing shoots 5–10 mm long, secund; indumentum when present shortly sericeous, petiolule usually whitish-sericeous ...................................................... D. amazonica Corolla white to light yellow, the base of standard narrowed and without callosities; short flower-bearing shoots 1–3 mm long, indumentum when present brownish-rufescent ......................................................... 2 Legumes ovate .............................................................................. D. negrensis Legumes oblong to narrowly oblong ........................................ D. pterocarpus

Derris amazonica Killip, J. Wash. Acad. Sci. 24: 48. 1934. —Lonchocarpus negrensis Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 98. 1860, non Derris negrensis Benth. 1860. Vine to 30 m high. Lower montane forests on poor soil, 600–800 m; Amazonas (northern slopes of Cerro Duida). Guyana, Suriname, French Guiana, northern Amazonian Brazil. ŠFig. 270. The isolated position of this species within Derris is evident from the generic description and has already been noted by G. Amshoff (Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 59. 1939). It agrees with Lonchocarpus subgenus Phacelanthus Pittier in most characters except for the winged fruit. For further discussion, see Lonchocarpus.

Derris negrensis Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 98. 1860. Lonchocarpus killipii Ducke, Trop. Woods. 69: 5. 1942. High-climbing, robust liana to 20 m high; leaves coriaceous; flowers white, fragrant. Flooded forests, occasionally in disturbed but nonflooded evergreen lowland forests, 50– 300 m; Delta Amacuro (Sacupana), Bolívar (lower Río Caura, Río Parguaza), Amazonas (Río Casiquiare, Río Cunucunuma, Río Negro, Río Orinoco, Río Ventuari). Amazonian Brazil (Rio Negro basin). ŠFig. 269. This species is very similar to Derris pterocarpus when in flower, but differs from it by the fruit shape character presented in the key.

Derris 303

Fig. 269. Derris negrensis

Fig. 270. Derris amazonica

304

F ABACEAE

Derris pterocarpus (DC.) Killip, J. Wash. Acad. Sci. 26: 360. 1936. —Lonchocarpus pterocarpus DC., Prodr. 2: 260. 1825. Deguelia scandens Aubl., Hist. Pl. Guiane 750, t. 300. 1775. —Derris scandens (Aubl.) Pittier, Contr. U.S. Natl. Herb. 20: 41. 1917; Pittier, Legumin. Venez. 1: 105. 1944, non Derris scandens Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 106. 1860. Derris guyanensis Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 106. 1860. Glabrous liana, reaching high into the forest canopy. Flooded forests along rivers, disturbed forests on higher ground, 50–300 m; Bolívar (El Manteco, Puerto Ordaz, Serranía de Imataca). Apure; Guyana, Suriname, French Guiana, northern Amazonian Brazil. ŠFig. 271.

Fig. 271. Derris pterocarpus

23. DESMODIUM Desv., J. Bot. Agric. 1: 112, pl. 5. 1813, nom. cons. Meibomia Heist. ex Fabr., Enum. 168. 1759. by Nidia L. Cuello A. Herbs, subshrubs, or shrubs, prostrate, decumbent or scrambling to erect. Leaves 1–3(–5)-foliolate, petiolate, stipulate; leaflets petiolulate and stipellate. Inflorescences axillary or terminal (fasciculate and leaf-opposed in D. triflorum), racemose or racemose-paniculate, or extremely dense and capitate; primary bracts striate and ciliate, each subtending 1 flower or a fascicle of 2–several flowers; secondary bracts often present and similar to the primary bracts or depauperate, each subtending a single flower; bracteoles rarely if ever present in American species. Flowers pedicellate. Calyx 2-lobed, the upper lobe almost entire to bifid, the lower 3toothed with the central tooth longer than the laterals, or the calyx almost equally 5lobed; corolla exceeding the calyx; standard short-clawed and the wing petals subsessile; wing petals often attached to keel petals by a small appendage; keel petals long-clawed, partially fused above. Vexillar stamen free or partially fused; anthers elliptic. Ovary sessile or stipitate; ovules 2–many; style slender; stigma terminal. Fruit a loment, transversely septate into 1–many articles, stipitate, indehiscent, or tardily dehiscent, the articles nearly linear to almost saccate or circular and notched at the apex, sometimes folded on each other in accordion fashion, the surfaces glabrous to densely pubescent with straight or hooked trichomes, or pubescent only on the sutures, 1-seeded. Seeds oblong or subquadrate. North America, tropics and subtropics (most diverse in eastern Asia, Mexico, and Brazil), temperate Asia; ca. 300 species, ca. 25 in Venezuela, 17 of these in the flora area.

Desmodium 305

Key to the Species of Desmodium 1. 1.

Leaves all 1-foliolate, appearing simple .................................................... 2 Leaves all 3-foliolate, or both 3-foliolate and 1-foliolate on the same plant ................................................................................................................ 5 2(1). Leaflets linear to lance-ovate, narrowed to ± acute apex, rounded at base, 4–9.5 × 0.4–1.4 cm, prominently reticulate and soft-puberulent on lower surface ............................................................... D. sclerophyllum 2. Leaflets obovate, ovate, or elliptic, the blades wider, shorter or longer than above .............................................................................................. 3 3(2). Leaflets 4–8 cm wide, obovate to broadly ovate; inflorescence (2–)manybranched; erect shrub > 1 m tall ............................................ D. distortum 3. Leaflets 1–3.5 cm wide, ovate to elliptic-lanceolate; inflorescence 1–3branched; creeping to ascending herb or suffrutescent, < 1 m tall ...... 4 4(3). Leaflets mostly ovate, the lowermost sometimes oblong to oblong-lanceolate, the uppermost linear-lanceolate, the lower surface pilose to glabrescent; articles of loment 3–3.5 mm long, ovate-oblong to suborbiculate, hirtellous-pilose, the dorsal suture convex; isthmi (areas between articles of loment) central ................................. D. pachyrrhizum 4. Leaflets all elliptic, the lower surface strigose; articles of loment 4–5 mm long, ovate, uncinulate, pubescent, dorsal suture straight; isthmi subcentral ........................................................................... D. hickenianum 5(1). Leaves both 3-foliolate and 1-foliolate on the same plant ........................ 6 5. Leaves all 3-foliolate .................................................................................. 7 6(5). Herb or shrublet to 50 cm tall; leaflets 0.8–3 × 0.6–2.1 cm, ovate, oblong or elliptic, apex rounded to emarginate; stipules linear, acuminate, 4–7 mm long; frequent on granitic outcrops ....................................... D. orinocense 6. Herb or shrub to 3 m tall; leaflets 3.7–17 × 2–11 cm, elliptic to ovate, apex mostly obtuse and apiculate; stipules clasping stem, obliquely ovate, 6–15 × 3.5–6 mm; widespread in other habitats .................... D. distortum 7(5). Leaflets < 1 cm long; inflorescence (1)2–4 flowers, fasciculate, axillary or leaf-opposed .............................................................................. D. triflorum 7. Leaflets mostly > 1 cm long; inflorescence of many more than 4 flowers, not fasciculate, terminal or axillary ...................................................... 8 8(7). Inflorescence a short, dense raceme, ca. 1–2.5(–3.5) cm long, borne above the uppermost leaf; calyx densely long silky-pilose and ciliate, the trichomes to 2 mm long .............................................................. D. barbatum 8. Infloresecence a lax raceme, usually longer than 3.5 cm, or a sparingly to many-branched panicle; calyx variously puberulent or pilose but not as above ....................................................................................................... 9 9(8). Articles with dorsal suture straight or only slightly concave or notched at the center ............................................................................................. 10 9. Articles with dorsal suture curved, convex, sinuate, or obtusely to acutely angled ................................................................................................... 14 10(9). Loments with (1)2 or 3 articles, the articles with dorsal suture slightly concave or notched at the center ......................................................... 11 10. Loments with 4–many articles, the articles with dorsal suture straight .............................................................................................................. 12

306

F ABACEAE

11(10). Leaflets chartaceous to subcoriaceous, short- to long-pilose on lower surface; terminal leaflet rhombic-ovate or ovate to elliptic-ovate, obtuse to rounded at base, the margin not undulate; articles of loment 4–5.5 mm; loment stipe 3–8 mm long .......................................................... D. axillare 11. Leaflets thin, membranous, sparsely pilose to glabrescent on lower surface; terminal leaflet ovate, acuminate, truncate at base, the margin undulate; articles of loment 2.5–3 mm wide; loment stipe 1–1.5 mm long ..................................................................................... D. wydlerianum 12(10). Petioles usually slender, sparsely pilose to glabrescent; leaflets orbicular, obcordate, or elliptic, secondary and tertiary veins slightly visible but not prominent on lower surface; loments with 1–4 articles ................... ............................................................................................... D. adscendens 12. Petioles stout, with both uncinulate trichomes and moderately long spreading trichomes; leaflets generally ovate to ovate-rhombic, secondary and tertiary veins easily visible and prominent on lower surface; loments with 4–many articles ............................................................. 13 13(12). Stipules free from each other; petioles often equaling or longer than the lateral leaflets, (2.5–)3–7 cm long; leaflets thin; loments with to 5 articles, isthmi (area between articles) usually 1/4 or less of the width of the articles ..................................................................................... D. affine 13. Stipules fused basally or overlapping; petioles usually much shorter than the lateral leaflets, 1–2.5 cm long; leaflets thick; loments with to 8 articles, isthmi usually 1/4–1/2 the width of the articles ............... D. incanum 14(9). Plants prostrate or semiprostrate; articles of loment elliptic or rhombic in outline .................................................................................................. 15 14. Plants mostly erect, or erect to sprawling, usually becoming shrubby to 3 m or more; articles of loment orbicular or triangular in outline ..... 16 15(14). Articles of loment narrowly elliptic, sutures scarcely constricted at isthmi; loments with 7 or 8 articles ............................................. D. scorpiurus 15. Articles of loment mainly rhombic in outline, appearing slightly to muchtwisted, sutures markedly constricted at isthmi; loments with 5 articles ..................................................................................... D. procumbens 16(14). Articles of loment mostly orbicular in outline, but sometimes with the margins alternately revolute; isthmi between articles central .......... 17 16. Articles of loment triangular in outline; isthmi between articles excentric .............................................................................................................. 18 17(16). Pedicels > 1 cm long, spreading; articles of loment 3–3.5(-4) × 2.5-3 mm; lower surface of leaflets sparsely pilose mainly on the veins, margins sparsely ciliate ........................................................................ D. tortuosum 17. Pedicels < 1 cm long; articles of loments 1.5–2.5(–3) × 0.8–1.5(–2) mm; lower surface of leaflets densely soft-pilose throughout, margins densely ciliate ......................................................................... D. distortum 18(16). Inflorescences terminal and axillary, densely and many-flowered, racemose to racemose-paniculate, and many-branched; pedicels 2.5–6 mm long, erect at maturity; upper surface of leaflets usually lustrous, sparsely puberulent ............................................................ D. cajanifolium 18. Inflorescences axillary, racemose, lax and few-flowered; pedicels mostly 12–20 mm long, slender, flexuous; upper surface of leaflets not lustrous, appressed-pilose on both surfaces ....................... D. campyloclados

Desmodium 307

Desmodium adscendens (Sw.) DC., Prodr. 2: 332. 1825. —Hedysarum adscendens Sw., Prodr. 106. 1788. —Pega-pega. Repent herb, subshrub, or shrub to 1 m tall. Weedy areas, widespread in savannas, forest borders, along rivers, 50–900 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, Africa, Asia, Melanesia. Desmodium affine Schltdl., Linnaea 12: 312. 1838. Procumbent herb; flowers white or pale lilac with white. Roadsides, near sea level to 300 m; Delta Amacuro (Caño Mánamo, Pedernales), Bolívar (Isla Anacoco, Río Caura). Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.

Uairén), Amazonas (Isla Ratón). Aragua, Barinas, Carabobo, Lara, Mérida, Miranda, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. D. axillare var. stoloniferum (Rich. ex Poir.) B.G. Schub., J. Arnold Arbor. 44: 289. 1963. —Hedysarum stoloniferum Rich. ex Poir. in Lam., Encycl. 6: 421. 1804 [1805]. Desmodium axillare var. sintenisii Urb., Symb. Antill. 2: 303. 1900. Herb rooting at the nodes; flowers deep pink. Shaded places along forest borders, 50– 200 m; Bolívar (lower Río Caura). Cojedes, Monagas; Central America, Antilles, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia.

1. Stems with long, dense pubescence of straight trichomes; leaflets ovate to elliptic-ovate, the lower surface long-pilose, the apices acute to gradually acuminate ...................... var. acutifolium 1. Stems with rather inconspicuous pubescence of short, hooked trichomes; leaflets ovate, the lower surface short-pilose, the apices abruptly short-acuminate ............ ................................. var. stoloniferum

Desmodium barbatum (L.) Benth. in Miq., Pl. Jungh. 224. 1852. —Hedysarum barbatum L., Syst. Nat. ed. 10, 1170. 1759. —Këmëhkë, Kuhpëkwamën (Panare). Herb with ascending to erect stems or shrub 0.2–1 m tall; flowers purple-pink, white with some pink stripes, or white to light purple. Frequent in sandy savannas and along roadsides, 50–1000 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Old World Tropics. ŠFig. 273. Desmodium barbatum is a polymorphic species with variable habit. The leaves are sometimes unifoliolate to trifoliolate on the same plant, the leaflets are sometimes very small and look like those of D. triflorum, and the pubescence density on the lower leaf surface is also quite variable.

D. axillare var. acutifolium (Kunze) Urb., Symb. Antill. 4: 292. 1905. —Meibomia axillaris var. acutifolia Kunze, Revis. Gen. Pl. 1: 195. 1891. —Pega-pega. Herb with repent or trailing stems; flowers pink, pale-purple, or rose-purple. Shaded places along forest borders, Trachypogon savannas, cultivated fields, 50–800 m; Bolívar (lower Río Caura, near Santa Elena de

Desmodium cajanifolium (H.B.K.) DC., Prodr. 2: 331. 1825. —Hedysarum cajanifoliun H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 525. 1823 [1824]. Suffrutescent to 3 m tall or erect herb to 1.5 m; flowers rose turning blue-purple. Frequent in Mauritia palm swamps, 50–400 m; Bolívar (Ciudad Piar, Los Pijiguaos, Río Chicanán, Río Parguaza, lower Río Sua-

Desmodium axillare (Sw.) DC., Prodr. 2: 333. 1825. —Hedysarum axillare Sw., Prodr. 107. 1788. —Meibomia axillaris (Sw.) Kuntze, Revis. Gen. Pl. 1: 195. 1891. Neotropics; 4 varieties, 3 in Venezuela, 2 of these in the flora area. Key to the Varieties of D. axillare

308

F ABACEAE

pure), Amazonas (near Puerto Ayacucho). Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia. Desmodium campyloclados Hemsl., Biol. Cent.-Amer., Bot. 1: 276. 1880. —Meibomia campyloclada (Hemsl.) Kuntze, Revis. Gen. Pl. 1: 197. 1891. Desmodium flexuosum Pittier, Bol. Técn. Minist. Agric. 5: 26. 1944. Desmodium dubium Pittier, Bol. Soc. Venez. Ci. Nat. 11: 17. 1947. Vine, sprawling herb, or slender straggling shrub to 3 m tall or more; flowers lavender. Tepui slopes, 1500–1800 m; Amazonas (west of Cerro Yutajé). Barinas, Lara, Mérida, Táchira, Trujillo, Yaracuy; Central America, Colombia, Ecuador, Peru, Bolivia. Desmodium distortum (Aubl.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 8: 101. 1939. —Hedysarum distortum Aubl., Hist. Pl. Guiane 774. 1775. —Ihchan (Panare). Desmodium asperum (Poir.) Desv., J. Bot. Agric. 1: 122. 1813. —Hedysarum asperum Poir. in Lam., Encycl. 6: 408. 1804 [1805]. Herb to subshrub 1–3 m tall; flowers fragrant, purple or rose-lavender. Rocky or sandy savannas, sandy floodplains, ca. 50– 900 m; Bolívar (Los Pijiguaos, Río Parguaza), Amazonas (Puerto Ayacucho). Widespread elsewhere in Venezuela except the Andes; Mexico, Costa Rica, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay. Desmodium hickenianum Burkart, Darwiniana 3: 217. 1937. Herb to suffrutescent to 1 m tall; flowers purple. Savannas, 100–200 m; Bolívar (near El Manteco, near Moitaco). Aragua; Bolivia, Paraguay, Argentina, Uruguay. Desmodium incanum DC., Prodr. 2: 332. 1825. —Hedysarum incanum Sw., Prodr. 107. 1788. Hedysarum canum J.F. Gmel., Syst. Nat. 1124. 1792. —Desmodium canum (J.F. Gmel.) Schinz & Thell., Mém. Soc. Sci. Nat. Neuchâtel 5: 371. 1914 [1913].

Stems procumbent to ascending or erectascending; flowers rose, bluish gray, or lavender-rose. Weedy areas near towns, along roadsides, near sea level to 300 m; Delta Amacuro (Pedernales, Tucupita), Bolívar (El Dorado, northwest of El Manteco, lower Río Caura, Upata), Amazonas (Río Puruname). Widespread elsewhere in Venezuela; U.S.A., Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, widespread in the paleotropics. Desmodium orinocense (DC.) Cuello, Novon 4: 98. 1994. —Desmodium adscendens var. orinocense DC., Prodr. 2: 332. 1825. —Nicolsonia orinocensis (DC.) Schindl., Repert. Spec. Nov. Regni Veg. 23: 358. 1927. —Chapere (Guahibo). Hedysarum adscendens auct. non Sw. 1788: sensu H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 520, t. 597. 1823 [1824]. Shrublet to 50 cm tall, or prostrate shrub to 1.3 m tall, or spreading herb with divaricate stems; flowers purplish blue, lavender, or lilac. Herbaceous vegetation around igneous outcrops, 50–300 m; Amazonas (near Puerto Ayacucho, Río Atabapo, Río Baría, Río Casiquiare, Río Cuao, Río Pasimoni). Colombia (Vaupés). ŠFig. 272. Desmodium pachyrrhizum Vogel, Linnaea 12: 97. 1838. —Pega-pega. Herb, creeping to ascending. Savannas, 50–300 m; Bolívar (Altiplanicie de Nuria, north of El Manteco, near Represa Guri, lower Río Caura, Río Paragua). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Guárico, Miranda, Monagas, Portuguesa, Yaracuy; Colombia, Brazil, Bolivia, Paraguay, Argentina. Desmodium procumbens (Mill.) A. Hitchc., Annual Rep. Missouri Bot. Gard. 4: 76. 1893. —Hedysarum procumbens Mill., Gard. Dict. ed. 8, no. 10. 1768. Pantropics; 3 varieties, all in Venezuela, 1 in the flora area. D. procumbens var. procumbens Stems procumbent; flowers white or rose. Savannas, near sea level to 200 m; Delta

Desmodium 309

Amacuro (Tucupita), Bolívar (lower Río Caroní). Anzoátegui, Aragua, Carabobo, Distrito Federal, Guárico, Lara, Miranda, Portuguesa, Sucre, Trujillo, Zulia; U.S.A., Mexico, Central America, Antilles, Colombia, Guyana, Suriname, Ecuador, Brazil, Africa. Desmodium sclerophyllum Benth. in Mart., Fl. Bras. 15(1): 102. 1859. Suffrutescent to 1 m tall; flowers redpurple. Wet savannas dominated by Curatella americana, 100–400(–600) m; Bolívar (La Paragua, Río Chiguao, Río Paragua, base of Roraima-tepui), Amazonas (Río Parucito). Aragua; Colombia, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay. Desmodium scorpiurus (Sw.) Desv., J. Bot. Agric. 1: 122. 1813. —Hedysarum scorpiurus Sw., Prodr. 107. 1788. —Amjadi (Panare), Pega-pega. Subprostrate herbaceous vine; stems procumbent; flowers pinkish white or lavender. Savannas, near sea level to 300 m; Delta Amacuro (Tucupita), Bolívar (Ciudad Bolívar, La Paragua, around Represa Guri, lower Río Caroní), Amazonas (Puerto Ayacucho, Río Ocamo). Widespread elsewhere in Venezuela; U.S.A., Mexico, Central America, Antilles, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil (Rondônia), Bolivia, Africa. Desmodium tortuosum (Sw.) DC., Prodr. 2: 332. 1825. —Hedysarum tortuosum Sw., Prod. 107. 1788. —Arestín, Pega-pega. Woody herb or shrub to 1 m tall or a vine; flowers purple. Roadsides, borders of forests, 50–300 m; Delta Amacuro (Los Castillos), Bolívar (Kilómetro 88, Maripa, lower Río Caroní, Upata), Amazonas (Puerto Ayacucho). Aragua, Distrito Federal, Falcón, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; U.S.A., Mexico, Guatemala, Belize, Panama, Antilles, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia, Africa. Desmodium triflorum (L.) DC., Prodr. 2: 334. 1825. —Hedysarum triflorum L., Sp. Pl. 749. 1753. Low creeping herb; flowers blue. Savannas, 50–200 m; Bolívar (Isla Anacoco, La Vergareña), Amazonas (Puerto Ayacucho). Anzoátegui, Apure, Barinas, Carabobo, Cojedes,

Fig. 272. Desmodium orinocense

310

F ABACEAE

Miranda, Monagas, Portuguesa, Sucre; U.S.A., Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Asia. Desmodium wydlerianum Urb., Symb. Antill. 2: 302. 1900. Slender creeping herb, often a subshrub; flowers purple. Lower montane forests, in patches on ground, 300–700 m; Bolívar (Altiplanicie de Nuria, Río Chiguao), Amazonas (Río Ugueto). Distrito Federal, Sucre; Guatemala, Costa Rica, Panama, Antilles, Colombia, Guyana, Suriname, Peru, Brazil.

Fig. 273. Desmodium barbatum

24. DIOCLEA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 437, fig. 576. 1823 [1824]. by Richard H. Maxwell Woody vines or lianas, occasionally small, erect subshrubs. Leaves alternate, 3foliolate; leaflets ovate, oval, elliptic, to broadly lanceolate; stipules produced below insertion (i.e., extending above and below point of attachment) or not, rarely absent; leaflets ovate, oval, elliptic, to broadly lanceolate, entire, unlobed, stipellate. Inflorescence axillary, erect, usually single, pseudoracemose; bracts and bracteoles persistent to caducous. Flowers numerous. Calyx tube 4-lobed, the upper lobe entire or emarginate; corolla usually shades of purple, occasionally white; standard reflexed or occasionally straight, mostly emarginate, basally biauriculate, usually bicallose; wing petals free, occasionally with a spur; keel petals fused distally, beaked or unbeaked. Stamens 10, pseudomonadelphous; anthers dimorphic or monomorphic. Pistil geniculate or sigmoidal; ovary usually villous, 1–many-ovulate; style flat or terete distally; stigma terminal and capitate or somewhat subterminal. Fruit a legume, oblong to obovate, flat or turgid, dry or fleshy, dehiscent or indehiscent, mostly pubescent. Seeds mostly oval-oblong or cuboid, variable in size, soft or hard, the hilum linear, nearly 1/2–3/4 encircling the seed or short-oblong. Pantropics, mostly New World; ca. 55 species, ca. 20 species in Venezuela, 13 of these in the flora area. The genus is currently undergoing revision. Key to the Species of Dioclea 1.

1.

Stipules produced below insertion (extending above and below point of attachment); bracts 6–12 mm long, mostly caducous or semipersistent; anthers dimorphic (5 + 5) ...................................................................... 2 Stipules not produced below attachment; bracts to ca. 3 mm long, mostly persistent; anthers 10, monomorphic or dimorphic (5 + 5) .................. 3

Dioclea 311

2(1).

Leaflets with secondary veins in ca. 8 pairs, oval or ovate, the apices rounded or abruptly acute; legume compressed; seeds hard, the hilum 2/3–3/4 encircling; bracts ca. 10 mm long, reflexed, semipersistent or caducous; flowers buds somewhat oblong, the calyx lobes straight .......... ....................................................................................................... D. reflexa 2. Leaflets with secondary veins in ca. 12 pairs, broadly ovate to somewhat oval, the apices with abruptly acuminate tips; legume turgid; seeds soft, the hilum 1/2 encircling; bracts ca. 6 mm long, erect; flower buds slender, the calyx lobes upcurved ...................................... D. malacocarpa 3(1). Legume 6–16-seeded; seeds hard, the hilum linear, nearly 1/2 encircling; anthers 10, monomorphic; standard and keel petals mostly puberulent apically; keel petals oblong or obliquely oblong, the upper margin partially toothed .......................................................................................... 4 3. Legume 1–6(–8)-seeded; seeds soft, the hilum oblong, much less than 1/2 encircling; anthers 10, dimorphic (5 + 5); petals glabrous; keel petals triangular or somewhat semilunar or rostrate, the upper margin entire ................................................................................................................ 9 4(3). Calyx tube pubescent, sometimes basally or only on the midribs; bracteoles 1.5–3 mm long, ovate, acute or lanceolate, mostly persistent, occasionally caducous ................................................................................... 5 4. Calyx tube glabrous; bracteoles 6–16 mm long, broadly ovate or elliptic, mostly caducous ..................................................................................... 8 5(4). Flowers 1.2–2.5(–3) cm long; standard as wide as or almost as wide as long; calyx lobes about equal the tube length ....................... D. guianensis 5. Flowers 2.6–3.5 cm long; standard longer than wide; calyx lobes exceeding the tube length ................................................................................. 6 6(5). Legume oblong, 2–2.5 cm wide; calyx tube glabrous or glabrescent, except along the midribs; bracteoles ovate-orbicular ......................... D. apurensis 6. Legume linear-oblong or oblong, ca. 1.3–2 cm wide; calyx tube pubescent; bracteoles ovate, acute, or lanceolate .................................................... 7 7(6). Legume linear-oblong, ca. 1.5 cm wide; leaflets oval, elliptic, or broadly ovate, the apices rounded, both sides canescent-velutinous; calyx sparsely strigulose, occasionally almost glabrous, except along the midribs; flowers 3–3.5 cm long, mostly white, occasionally lilac; each wing petal with a spur ....................................................................... D. albiflora 7. Legume oblong, ca. 2 cm wide; leaflets mostly ovate, oval, to broadly lanceolate, the apices obtuse or acute, the lower surface villous; calyx sparsely pubescent basally as well as along the midribs; flowers 2.5–3.5 cm long, mostly purple, lilac, rosy white, or occasionally white; wing petal without a spur ................................................................... D. holtiana 8(4). Flowers > 4 cm long; keel oblanceolate, the upper margin weakly serrate; legume 10–16-seeded; standard straight ............................. D. macrantha 8. Flowers 2–3 cm long; keels somewhat oblong, the upper margin basally fimbriate; legume 7–10-seeded; standard reflexed ..................... D. virgata 9(3). Leaflets glabrous; mostly high-climbing forest lianas ............................ 10 9. Leaflets pubescent; lianas, woody vines, or shrublets of open woods and savannas ............................................................................................... 11 10(9). Anthers 5 + 5; ovary 2-ovulate; stems with conspicuous raised elliptic len-

312

F ABACEAE

ticels; calyx lobes strongly upcurved; legumes obovate-oblanceolate, 2-seeded; seeds compressed; leaflets coriaceous .......................... D. scabra 10. Anthers 10, monomorphic; ovary 4–8-ovulate; stems without raised lenticels; calyx lobes mostly straight; legumes mostly oblong, ca. 5-seeded; seeds compressed or rarely overgrown; leaflets papyraceous to somewhat membranaceous .......................................................... D. macrocarpa 11(9). Legume turgid, oblong, the beak upcurved; seeds cuboid; leaflets subcoriaceous, with drip tips; mostly forest lianas; flowers to ca. 2 cm long; anthers monomorphic; inflorescences occasionally branched ... D. ruddiae 11. Legume compressed, elliptic or oblanceolate, the beak downcurved; seeds compressed, oval-elliptic; leaflets rigidly coriaceous; small vines or erect shrublets of open woods and savannas; flowers ca. 1.5 cm long or unknown; anthers monomorphic or unknown; inflorescences single .............................................................................................................. 12 12(11). Leaflets subconduplicate, the bases cordate; woody vine to 1 m long; flowers unknown; known only from west rim of Cerro Parú, Amazonas ....................................................................................................... D. rigida 12. Leaflets unfolded, the bases mostly round; erect shrublets; flowers ca. 1.5 cm long; plants of open woods and savannas ............... D. steyermarkii Dioclea albiflora R.S. Cowan, Mem. New York Bot. Gard. 10(3): 150. 1958. Vine sprawling over low shrubs. Open forests, river banks, 100–200 m; Bolívar (Río Orinoco at Piedra Marimare), Amazonas (Isla Ratón on Río Orinoco, near Puerto Ayacucho). Endemic. Dioclea apurensis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 438. 1823 [1824]. Small to medium-sized vine. Mostly open areas, ca. 100 m; Bolívar (along Río Orinoco). Apure; Suriname, Amazon basin. This species is poorly known. The syntype has fruit but lacks flowers. Its legume is larger than those of the Dioclea guianensis complex. Collections with flowers that have been assigned to D. apurensis have small orbicular or acute bracteoles similar to D. guianesis s. lat., but frequently the calyx is pubescent only along the dorsal and ventral midlines, or the calyx is glabrous. Dioclea guianensis Benth., Comm. Legum. Gen. 70. 1837. —Frijolillo. Dioclea broadwayana Pittier, Bol. Técn. Minist. Agric. 5: 86. 1944, pro parte, nom. nud. Vine. Open areas, extending into forests, 50–800 m; Delta Amacuro (near Los Castillos de Guayana), widespread in Bolívar and Amazonas. Widespread in the rest of Venezuela;

Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. Pittier’s invalid Dioclea broadwayana was published without Latin. Pittier continued to segregate species from his concept of Dioclea guianensis as well as from that of his concept of D. broadwayana within the Guayana area, but he did not validly publish the names. These taxa are primarily segregates of D. guianensis s. lat., and since their significance extends beyond the Guayana area, their elucidation has been deferred pending further study. Dioclea holtiana Pittier ex R.H. Maxwell, Ann. Missouri Bot. Gard. 77: 584. 1990. —Dioclea holtiana Pittier, Bol. Técn. Minist. Agric. 5: 84, fig. 36. 1944, nom. nud. Vine or liana climbing over small trees. Lowland forests, along rivers and roads, also associated with granitic outcrops, 50–200 m; Amazonas (near Puerto Ayacucho, Río Cataniapo, near the mouth of Río Vichada on the Río Orinoco). Trujillo; Central America, possibly Mexico, Colombia. This species is similar in many respects to Dioclea albiflora and is found in the same localities in the flora. Dioclea macrantha Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 408. 1909.

Dioclea 313

Climbing, twining vine over shrubs. Secondary growth, 100–200 m; Bolívar (near El Palmar on the Río Grande). Colombia, Guyana, Suriname, French Guiana, Brazil (Amapá, Pará). Dioclea macrocarpa Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 410. 1909. —Bejuco de jabón. High-climbing forest liana. Flooded lowland riparian forests, 50–500 m; Delta Amacuro (vicinity of Curiapo, mouth of Río Amacuro, Río Grande), Bolívar (middle Río Chiguao, Serranía de Imataca), Amazonas (Río Mawarinuma). Colombia, Guyana, Suriname, Amazonian Ecuador, Peru, Brazil. Some collections contain legumes with twisting dehiscence and overgrown cuboid seeds that do not match the seeds of the type. Further investigation may warrant the naming of a variety of Dioclea macrocarpa or D. ruddiae, an additional variety of D. scabra, or possibly a new species. Dioclea malacocarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 170. 1922. —Ojo de zamuro, Yawade ansajüdü (Yekwana). Sprawling, coarse vine or forest liana. Flooded, riparian, and rain forests, 50–500 m; Delta Amacuro (northeast of El Palmar), Bolívar (Caño Adaua west of Cerro Guanacoco, southeast of El Dorado, Río Caura, upper Río Paragua), Amazonas (Río Atabapo, Río Casiquiare, Río Ventuari, Río Yaciba in Río Yatúa basin). Trinidad, Guyana, Suriname, French Guiana, Amazonian Brazil. Reports of this species from Central America, Colombia, western Venezuela, Ecuador, and Peru should probably be referred to the closely related Dioclea pulchra Moldenke. Dioclea reflexa Hook. f. in Hook., Niger Fl. 306. 1849. —Bejuco des amni, Bejuco de zamuro, Ojo de cari cari, Ojo de zamuro, Tangranta-moi (Taurepán). Coarse climbing or sprawling vine. Lowland wet forests, near sea level to 400 m; Delta Amacuro (Río Orinoco basin), northern Bolívar, Amazonas (upper Río Orinoco). Pantropics (distribution above the ocean drift line and upriver due to a flotation adaptation in the seed). ŠFig. 274.

The name Dioclea hexandra (Ralph) Mabberly was published (David Mabberley. Taxon 29: 605. 1980) based on Mucuna hexandra Ralph (Icon. Carpolog. 30, t. 34, fig. 5. 1849). Mabberley placed the name D. reflexa in synonymy, but it is retained here, because it is believed that the Asian D. hexandra (= D. javanica Benth.) is not the same species as the western African, northern South American, and Central American D. reflexa. Dioclea rigida R.S. Cowan, Mem. New York Bot. Gard. 10(3): 150. 1958. Woody vine to 1 m long. Occasional along upper ridges of summits, ca. 2000 m; Amazonas (Cerro Parú). Endemic. The species was described without flowers from a single collection only, but fits into Dioclea section Platylobium Benth. Cowan notes, “the rigidly coriaceous, venose, cordate leaflets are distinctive.” Dioclea ruddiae R.H. Maxwell, Ann. Missouri Bot. Gard. 75: 730. 1988. —Paronta-yek (Arekuna). High-climbing forest liana. Roadsides, savannas, wooded hills, slopes to upland, humid forests, 800–1600 m; Bolívar (near Cerro Venamo and La Escalera, Gran Sabana), Amazonas (Cerro Huachamacari, Simarawochi). Endemic. Dioclea scabra (Rich.) R.H. Maxwell, Ann. Missouri Bot. Gard. 77: 578. 1990. —Dolichos scabra Rich., Actes Soc. Hist. Nat. Paris 1: 111. 1792. Usually a high-climbing forest liana, woody vine, or shrublet. Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 3 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of D. scabra 1. Flowers 2.3–3 cm long ............ var. scabra 1. Flowers ca. 2 cm long ............ var. brownii D. scabra var. brownii R.H. Maxwell, Ann. Missouri Bot. Gard. 77: 579. 1990. High-climbing liana. Evergreen forests, ca. 100 m; Amazonas (known from a single collection from Caño Yagua). Endemic.

314

F ABACEAE

Fig. 274. Dioclea reflexa

Diplotropis 315

D. scabra var. scabra. —Bejuco de sapo, Bejuco de zamuro. Dioclea elliptica R.H. Maxwell, Ann. Missouri Bot. Gard. 77: 578. 1990, as synonym. Dioclea glabra auct. non Benth. 1837: sensu Pittier, Bol. Técn. Minist. Agric. 5: 79. 1944. Mostly high-climbing liana. Forests, extending into open areas, 100–400 m; Bolívar (Río Cuyuní near border with Guyana), Amazonas (widespread). Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, Pará). Dioclea steyermarkii R.H. Maxwell, Ann. Missouri Bot. Gard. 77: 585. 1990. Shrublet or vine to ca. 1 m tall. Savannas of hills and ridges, 100–600 m; Amazonas (south and southeast of Cerro Camani). Endemic. Dioclea steyermarkii is close to D. coriacea Benth., but with 10 perfect anthers instead of 5 perfect and 5 imperfect. It also shows an affinity to the imperfectly known D. rigida. Dioclea virgata (Rich.) Amshoff, Meded. Bot. Mus. Herb. Rijks. Univ. Utrecht 52: 59. 1939. —Dolichos virgatus Rich., Actes Soc. Hist. Nat. Paris 1: 111. 1792. Trailing or climbing roadside vine, or subshrub, sometimes forming dense clumps.

Neotropics, introduced into Paleotropics; 2 varieties, both in the flora area. Key to the Varieties of D. virgata 1. Keel petals with part of the upper margin crenate; bracteoles to ca. 6 mm long, semipersistent; fruit exocarp with short, curly, mostly canescent pubescence ........ ......................................... var. crenata 1. Keel petals with part of the upper margin long-fimbriate; bracteoles to ca. 10 mm long, caducous; fruit exocarp with long, stiff ferruginous, fugaceous pubescence .......................................... var. virgata D. virgata var. crenata R.H. Maxwell, Ann. Missouri Bot. Gard. 77: 585. 1990. Woody vine or liana. Margins of humid forests, 100 m; Amazonas (Río Casiquiare). Suriname, Brazil (Amapá, Amazonas, Pará). D. virgata var. virgata. —Ero-cuaja. Vine. Open areas, river banks, forest edges, near sea level to 200 m; Delta Amacuro (Caño Araguabisi between Caño Araguao and Caño Güiniquina, northeast of Tucupita), Bolívar (Río Parguaza), scattered in Amazonas. Throughout the rest of Venezuela; Neotropics, introduced into Paleotropics through botanical gardens.

25. DIPLOTROPIS Benth., Comm. Legum. Gen. 24. 1837. by Haroldo Cavalcante de Lima and Gerardo A. Aymard C. Trees. Leaves alternate, odd-pinnate; stipules small, caducous; stipels none. Inflorescence a terminal or corymbose panicle or terminal raceme. Flower buds clubshaped, curved. Calyx 5-dentate, the uppermost 2 joined higher up; petals rose to violet, plicate-corrugated; standard nearly hastate and auriculate above the claw; keel petals free. Stamens 10 and dimorphic, or 5 with 5 staminodes; filaments free, alternately long and short. Ovary subsessile or short-stipitate; ovules 3–6. Fruit indehiscent, samaroid or nut-like, oblong to oblong-orbiculate, membranous to woody. Seeds 1–few; radicle straight. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 12 species, 7 in Venezuela, all of these in the flora area. Key to the Species of Diplotropis 1.

1.

Leaves 12–20-foliolate, leaflets 1–3 cm wide, oblong to oblong-elliptic; inflorescences terminal racemes; androecium with 5 stamens and 5 staminodes .................................................................................... D. racemosa Leaves 5–16-foliolate, leaflets 4–9 cm wide, ovate, obovate, elliptic, or combinations of these; inflorescences terminal panicles; androecium with 10 stamens ..................................................................................... 2

316

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

3.

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

FABACEAE

Pedicels 4–5 mm long; calyx rusty-villose or appressed-ferruginous-pubescent .................................................................................................... 3 Pedicels 0.2–3 mm long; calyx sericeous or tawny-tomentose ................. 5 Branches and branchlets strigose; leaves 8–11-foliolate; leaflets strigose on the lower surface; calyx rusty-villose; standard 11–13 mm long ................................................................................................. D. strigulosa Branches and branchlets glabrescent; leaves 5–7-foliolate; leaflets glabrous on the lower surface; calyx appressed-ferruginous-pubescent; standard 7–9 mm long ........................................................................... 4 Leaflets with dull black on the lower surface; ovary sericeous; fruit woody, without wings .................................................................................. D. sp. A Leaflets without dull black on the lower surface; ovary ferrugineous; fruit a samara ....................................................................................... D. triloba Leaflets rigid-coriaceous; fruit woody, without wings ................. D. martiusii Leaflets membranaceous to coriaceous; fruit a flat samara, membranous to papery ................................................................................................ 6 Secondary and tertiary venation evident on the lower surface; calyx 6–10 mm long; fruits 8–12 × 3–5 cm ....................................... D. purpurea Secondary and tertiary venation strongly reticulate on the lower surface; calyx 4–6 mm long; fruits 5–6 × 1.5–2 cm ................... D. brasiliensis

Diplotropis brasiliensis (Tul.) Benth. in Mart., Fl. Bras. 15(1): 320. 1862. —Dibrachion brasiliensis Tul., Ann. Sci. Nat. Bot. sér. 2, 20: 139. 1843. —Bowdichia brasiliensis (Tul.) Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 31. 1915. Tree 6–18(–25) m tall; flowers pink; fruit a samara. Flooded forests, 50–200 m; Amazonas (San Carlos de Río Negro). Brazil (Acre, Amapá, Amazonas, Pará, Rondônia). Diplotropis martiusii Benth., Comm. Legum. Gen. 24. 1837. —Bowdichia martiusii (Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 131. 1922. —Congrio, Iyoynaji (Yanomami). Dibrachion riparium Spruce ex Benth. in Mart., Fl. Bras. 15(1): 321. 1861. Tree 6–18(–25) m tall; flowers pink; fruit woody. Flooded forests, 50–200 m; Amazonas (Río Atacavi, Río Baría, Río Casiquiare, Río Pamoni, Río Siapa, Río Sipapo, Río Yatúa). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 276. Diplotropis purpurea (Rich.) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 44. 1939. —Tachigali purpurea Rich., Actes Soc. Hist. Nat. Paris 1: 108. 1792, "Tachigalia." Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 2 varieties, 1 in the flora area.

D. purpurea var. purpurea. —Amiña, Amuña (Yekwana), Aru-ca-yu-dek (Arekuna), Cachicamo montañero, Congrio, Congrio blanco, OruKaiyet (Arekuna). Tree 20–40 m tall; flowers pink. Evergreen lowland to montane forests, 100–1100 m; Delta Amacuro (Río Toro), Bolívar (widespread), Amazonas (Puerto Ayachucho, Sierra Parima, Simarawochi, Río Coro Coro, upper Río Cunucunuma, Río Jenita on Río Ocamo, Yutajé). Sucre; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia. ŠFig. 275. Diplotropis racemosa (Hoehne) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 43. 1939. —Bowdichia racemosa Hoehne, Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 8: 55. 1919. Venezuela, Guyana, Suriname, Brazil; 2 varieties, 1 in Venezuela. D. racemosa var. racemosa Diplotropis racemosa var. kaieteurensis Amshoff, Bull. Torrey Bot. Club 75: 393. 1948. Tree 6–10(–30) m tall; flowers pinkish white. Borders of evergreen lowland forests, seasonally flooded forests near black-water rivers, 100–200 m; Amazonas (Piedra Guanare, Raudal Ceguera on Río Autana, Río Casiquiare). Guyana, Suriname, Brazil.

Diplotropis 317

Fig. 275. Diplotropis purpurea var. purpurea

318

FABACEAE

Fig. 276. Diplotropis martiusii

Diplotropis strigulosa R.S. Cowan, Mem. New York Bot. Gard. 10(1): 151. 1958. Tree to 20 m tall; flowers pink. Montane forests, 1800–2000 m; Amazonas (Cerro Parú). Endemic. Diplotropis triloba Gleason, Bull. Torrey Bot. Club 60: 355. 1933. Tree to 20 m tall; fruit membranous. Evergreen lowland forests, 100–200 m; Amazonas

(San Carlos de Río Negro). Brazil (Amazonas, Mato Grosso, Rondônia). Diplotropis sp. A Tree 6–8 m tall; flowers pink. Evergreen lowland forests, seasonally flooded forests near black-water rivers 100–200 m; Amazonas (Raudal Ceguera on Río Autana). Endemic.

26. DIPTERYX Schreb., Gen. Pl. 2: 485. 1791, nom. cons. Coumarouna Aubl., Hist. Pl. Guiane 740. 1775. Oleiocarpus Dwyer, Ann. Missouri Bot. Gard. 52: 51. 1965. by Haroldo Cavalcante de Lima Trees. Leaves alternate, even-pinnate; leaflets opposite or alternate, often pellucid-punctate; rachis flattened or winged, with apex projected beyond the leaflets; stipules small, caducous; stipels none. Inflorescence a terminal panicle. Calyx pellucid-punctate, bilabiate, upper lip with 2 large lobes and lower lip very short, entire, or 3-dentate; petals violet to whitish mauve; standard emarginate; keel petals adnate on lower side. Stamens 10, monadelphous. Ovary stalked; ovule 1. Fruit drupaceous,

Dipteryx 319

ovoid to oblong-fusiform; mesocarp thick-fibrous; endocarp bony or woody, tardily opening on ground. Seed cylindric-fusiform; radicle short, straight. Central America, South America; 9 or 10 species, 4 in Venezuela, all in the flora area. Key to the Species of Dipteryx 1. 1. 2(1). 2. 3(2). 3.

Leaflets 10–18, thin-coriaceous; calyx membranous, glabrous or pubescent; fruit ovoid, with sparse mesocarp and bony endocarp ... D. magnifica Leaflets 2–8, coriaceous; calyx coriaceous, tomentose; fruit ovoid-oblong to oblong-fusiform, with abundant mesocarp and woody endocarp .......... 2 Upper lip of calyx 18–25 mm long; leaflets mostly 14–20 × 6–8 cm ...... D. rosea Upper lip of calyx 10–15 mm long; leaflets mostly 7–15 × 3–7 cm ........... 3 Calyx rusty-tomentose with upper lip 10–15 mm long; petals whitish mauve .......................................................................................... D. odorata Calyx tawny-tomentose (violet-tomentose when fresh) with upper lip 6–9 mm long; petals violet ........................................................ D. punctata

Dipteryx magnifica (Ducke) Ducke, Trop. Woods 61: 8. 1940. —Coumarouna magnifica Ducke, Rev. Int. Bot. Appl. Agric. Trop. 14: 405. 1934. —Maikonaji (Yanomami), Sarrapia mona. Taralea casiquiarensis Pittier, Bol. Soc. Venez. Ci. Nat. 8: 262. 1943. Tree 20–45 m tall. Forests on white sand or on granitic outcrops, 100–300 m; Amazonas (Caño Caname, Río Casiquiare, Río Cunucunuma, Río Padamo, Santa Bárbara del Orinoco). Brazil.

Dipteryx odorata (Aubl.) Willd., Sp. Pl. 3(2): 910. 1802. —Coumarouna odorata Aubl., Hist. Pl. Guiane 740. 1775. —Sarrapia. Tree 10–30 m tall. Forests on river banks, 100–400 m; Delta Amacuro (Serranía de Imataca), cultivated in Bolívar and Amazonas. Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 277.

Fig. 277. Dipteryx odorata

320

FABACEAE

Dipteryx punctata (S.F. Blake) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 60. 1939. —Coumarouna punctata S.F. Blake, Contr. U.S. Natl. Herb. 20: 525. 1924. —Sarrapia, Sarrapia real. Dipteryx trifoliolata Ducke, Trop. Woods 61: 7. 1940. Tree 8–20 m tall. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (Ciudad Bolívar, El Palmar, Río Caura), Amazonas (Río Guainía, San Carlos de Río Negro). Widely cultivated in Venezuela; Colombia,

Guyana, Suriname, French Guiana, Brazil. A cultivated form of Dipteryx punctata with edible fruits and 2 or 3 leaflets was described as D. trifoliolata Ducke. The fruits are used to treat stomach pain. Dipteryx rosea Spruce ex Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 125. 1860. —Sarrapia, Sarrapia mona. Tree 10–30 m tall. Riparian forests, 50– 300 m; Bolívar (Serranía de Imataca), Amazonas (near San Carlos de Río Negro). Colombia, Peru, Brazil.

27. ERIOSEMA (DC.) Desv., Ann. Sci. Nat. (Paris) 9: 421. 1826. —Rhynchosia sect. Eriosema DC., Prodr. 2: 388. 1825, spelling variant: Euriosma. by Renée H. Fortunato Suffrutescent herbs or subshrubs, erect, ascending to prostrate. Roots woody, turnip-shaped or spindle-shaped. Stems simple or few-branched, glabrous or pubescent with various pubescence types, glandular or eglandular. Leaves alternate, unifoliolate or pinnate-trifoliolate (commonly the 1 or 2 older ones near base of stem unifoliolate); stipules free or connate, persistent, rarely deciduous; petioles canaliculate, short to ± lacking; leaflets membranous, chartaceous, or coriaceous, glabrous to pubescent, glandular on both surfaces or only on lower surface, seldom eglandular, glandular trichomes usually yellowish or reddish brown; stipels lacking. Inflorescences racemose, lax or congested, axillary, solitary, 1–many-flowered; peduncles surpassing or not surpassing the main cauline leaves; bracts small, persistent or deciduous; bractlets lacking. Calyx 5-lobed, externally pubescent and glandular, the lobes deltoid, triangular, or subulate, not exceeding corolla, the keel lobe frequently longer; corolla yellow, sometimes with the standard veins reddish purple; standard obovate, elliptic or roundish, emarginate, biauriculate, clawed, externally pubescent and glandular; wing petals narrowly oblong, ± pubescent and glandular at apex; keel petals falcate, incurved and glandular-pubescent at apex. Stamens 10, diadelphous; anthers 2-locular, ellipsoid, dorsifixed, uniform. Gynoecium sessile to subsessile; ovary 2-ovulate, pubescent; style incurved, glabrous; stigmas small, subcapitate. Legumes 2-seeded, obovoid, ellipsoid, compressed, dehiscent, obliquely beaked. Seeds reniform or ellipsoid, ± oblique in the legumes, lustrous, brown to black, with strophiole, the hilum long, linear; funicular attachment at extreme apex of the hilum. Pantropics; ca. 100 species, 6 in Venezuela, 4 of these in the flora area. Key to the Species of Eriosema 1. 1. 2(1). 2.

Leaves all unifoliolate ........................................................... E. simplicifolium Leaves trifoliolate (usually 1 or 2 leaves unifoliolate at base of stems) ... 2 Middle leaflets < 5 times longer than wide, elliptic to obovate; stipules mostly free, ovate ........................................................................... E. rufum Middle leaflets > 5 times longer than wide, oblong to narrowly elliptic; stipules connate, rarely free, lanceolate ............................................... 3

Eriosema 321

3(2).

3.

Inflorescence pseudocorymbose, congested, when fully expanded equal to or shorter than the main cauline leaves; flowers > 9 mm long ............. .................................................................................................... E. crinitum Inflorescence racemose, lax, when fully expanded equaling or exceeding the main cauline leaves; flowers 4.5–7.5 mm long ................ E. violaceum

Eriosema crinitum (H.B.K.) G. Don, Gen. Hist. 2: 348. 1932. —Glycine crinita H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 421. 1823 [1824]. —Rhynchosia crinita (H.B.K.) DC., Prodr. 2: 389. 1825. Erect to decumbent subshrub; stems and leaflets densely reddish-, yellow-, or white-pilose, puberulent, or glabrate; inflorescences usually in roundish clusters at end of peduncles, 1–8-flowered. Sandy and rocky savannas, wet forests along rivers, forest edges, 50–1200 m; Delta Amacuro, Bolívar, Amazonas. Mexico, Central America, Greater Antilles, tropical and subtropical South America; 4 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of E. crinitum 1. Stems and leaflets densely reddish-pilose or yellow-pilose, or glabrate; inflorescences 2–8-flowered ....... var. crinitum 1. Stems and leaflets white-pilose; inflorescences 1–4-flowered ...... var. stipulare E. crinitum var. crinitum Ca. 20–40 cm tall. Savannas, edge of forests in open grassland, 50–1200 m; Bolívar (widespread), Amazonas (near Puerto Ayacucho, Sierra Parima). Widespread elsewhere in Venezuela; Mexico, Guatemala, Belize, Honduras, Greater Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. E. crinitum var. stipulare (Benth.) Fortunato, Novon 3: 25. 1993. —Eriosema stipulare Benth., Linnaea 22: 519. 1849. 20–100 cm tall. Savannas, 50–300 m; Delta Amacuro (between Los Castillos de Guayana and Piacoa), northern Bolívar, central and northern Amazonas. Apure, Cojedes, Guárico, Monagas, Portuguesa, Zulia; Costa Rica, Colombia, Guyana, Suriname, Brazil. ŠFig. 279. Eriosema rufum (H.B.K.) G. Don, Gen. Hist. 2: 347. 1823. —Glycine rufa H.B.K., Nov.

Gen. Sp. (quarto ed.) 6: 423, t. 574. 1823 [1824]. —Rhynchosia rufa (H.B.K.) DC., Prodr. 2: 384. 1825. Subshrub 50–100 cm tall. West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay; 2 varieties, 1 in Venezuela. E. rufum var. rufum Erect stems reddish-pubescent; stipules commonly free; leaflets short-pilose on both surfaces. Dry savannas, 50–400 m; Delta Amacuro (between San Félix and Los Castillos), common in northern Bolívar. Aragua, Apure, Cojedes, Guárico, Lara, Monagas, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Antilles, Colombia, Trinidad, Guyana, Suriname, Peru, Brazil, Bolivia. ŠFig. 278. Eriosema simplicifolium (H.B.K.) G. Don, Gen. Hist. 2: 348. 1832. —Glycine simplicifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 419. 1823 [1824]. —Rhynchosia simplicifolia (H.B.K.) DC., Prodr. 2: 389. 1825. Eriosema simplicifolium var. micranthum Grear, Mem. New York Bot. Gard. 20(3): 88. 1970. Decumbent to prostrate, rarely erect; stems simple or few-branched, 50–80 cm long; stipules connate. Savannas, edges of gallery forests, 50–400(–1200); Delta Amacuro (between Los Castillos and Piacoa), Bolívar (scattered), northern and central Amazonas. Anzoátegui, Apure, Barinas, Guárico, Monagas, Portuguesa, Zulia; Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Eriosema violaceum (Aubl.) G. Don, Gen. Hist. 2: 347. 1832. —Cytisus violaceous Aubl., Hist. Pl. Guiane 766. 1775. —Rhynchosia violacea (Aubl.) DC., Prodr. 2: 388. 1825. Stems erect to ascending; stipules connate or the older ones often free; leaflets appressed-villous or tomentose on both sur-

322

FABACEAE

Fig. 278. Eriosema rufum var. rufum

faces. Wet savannas, edges of forests along rivers, 50–900 m; scattered in Bolívar and Amazonas. Anzoátegui, Apure, Barinas,

Fig. 279. Eriosema crinitum var. stipulare

Monagas; Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil, Paraguay.

28. ERYTHRINA L., Sp. Pl. 706. 1753. by David A. Neill Trees, shrubs, or (outside the flora area) herbaceous perennials with woody rootstocks, often deciduous and flowering when leafless; bark usually armed with conical prickles. Leaves pinnately 3-foliolate, with glandular stipels on petiole below lateral leaflets; lower surface of leaflets glabrous, pubescent, or glaucous with epicuticular wax. Inflorescence racemose, terminal or pseudo-terminal, rarely axillary, erect or horizontal, flowers in fascicles of (2)3(–6). Flowers showy, red, orange, or pink. Calyx tubular or campanulate, thick and fleshy or chartaceous, the apex truncate or 1–5-dentate, sometimes cleft or bilabiate; standard larger than wing and keel petals, sometimes conduplicate and completely enclosing the latter; keel petals coherent or free. Stamens usually monadelphous, the vexillar one free for its upper 3/4

Erythrina 323

and the other 9 free for the upper 1/4. Ovary stipitate, usually tomentose; stigma capitate. Fruit linear, sometimes constricted between the seeds, dehiscent. Seeds red, bicolored (red and black) or brown, often persistent on pods long after dehiscence. Tropics, subtropics, and warm-temperate regions worldwide; ca. 115 species, 8 in Venezuela, 4 of these in the flora area. Erythrina pallida Britton & Rose, known from coastal Venezuela, Trinidad, and Tobago, is closely related to E. mitis. It may occur in drier forests of the flora area but flowering material or mature seeds, necessary for positive identification, have not been collected. Key to the Species of Erythrina 1.

1.

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

3.

Inflorescence erect; calyx tubular; standard tubular, at least 3 times longer than wide, conduplicately folded with wing and keel petals and sexual organs concealed within the standard; seeds red .................................. 2 Inflorescence horizontal or arching; calyx campanulate; standard spreading, revealing the keel petals, not conduplicate, < 2 times longer than wide; seeds brown .................................................................................. 3 Corollas scarlet, blades of the keel petals free from each other and obtusely rounded at apex .................................................................... E. mitis Corollas dark pink to orange-red, blades of the keel united by their exterior margins or, if free, then acute to acuminate at tip ....... E. rubrinervia Evergreen trees, usually growing in swampy or riparian habitats; leaf with small (ca. 1 mm diameter) glandular, not cupular, stipel; leaflets elliptic with revolute margins; corollas relatively thick and fleshy, the standard with a claw about 1/3 the length of the blade, the wings relatively large, about 1/2 the length of the ovate keel; standard pale orange or salmoncolored, the wing and keel petals ivory at base, red at apex ......... E. fusca Deciduous trees, usually growing on alluvial terraces but not in swampy habitats; leaf with large cupular stipel to 4 mm long and 3 mm diameter on petiole below lateral leaflets; leaflets rhombic to rhombic-ovate; corollas thin-chartaceous, the standard with a barely discernible claw, the wing petals minute, barely exceeding the calyx; petals uniformly deep orange .................................................................................... E. poeppigiana

Erythrina fusca Lour., Fl. Cochinch. ed. 1, 427. 1790. —Bucare, Oparu. Erythrina glauca Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: 428. 1801. Tree to 25 m tall. Usually swampy or riparian habitats, sometimes forming extensive monospecific stands, near sea level to 200 m; Delta Amacuro (road between Caño Guará and La Horqueta, Caño Rico between Caño Macareo and Río Grande, Río Orocoima), Bolívar (La Paragua). Anzoátegui, Apure, Aragua, Carabobo, Distrito Federal, Guárico, Lara, Mérida, Zulia; Central America, West Indies, Colombia, coastal Ecuador, eastern

Peru, Brazil, Bolivia, widespread in tropical southeast Asia, Mascarene Islands, western Pacific islands, New Guinea. ŠFig. 281. This is the only species of Erythrina known from both the Old World and New World tropics. Erythrina mitis Jacq., Pl. Hort. Schoenbr. 2: 47. 1797. —Peniolilla, Pionía. Deciduous tree to 8 m tall, flowering when leafless; legumes strongly constricted between the seeds. Semideciduous and lower montane forests, 50–500 m; Bolívar (Altiplanicie de Nuria, Caño Maracapra on lower Río Paragua, Río Botanamo basin), Ama-

324

FABACEAE

zonas (Salto Salas on upper Río Orinoco). Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia. Erythrina poeppigiana (Walp.) Cook, U.S.D.A. Div. Bot. Bull. 25: 57. 1901. —Micropteryx poeppigiana Walp., Linnaea 23: 740. 1850. —Pericoco. Tree to 30 m tall. Semideciduous lowland forests and seasonally flooded riparian forests, 100–400 m; Bolívar (Río San Pedro 200 km south of Caicara), Amazonas (Raudal Arata on Río Ocamo). Anzoátegui, Barinas, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Panama,

along base of Andes in Colombia, Ecuador, eastern Peru, and eastern Bolivia, widely planted as shade tree for coffee and cacao (and naturalized in places) in Nicaragua, Costa Rica, western Panama, and West Indies. ŠFig. 280. Erythrina rubrinervia H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 434. 1823 [1824]. Deciduous tree to 25 m tall, flowering when leafless; branchlets spinose, pith chambered; legumes strongly constricted between the seeds. Semideciduous lowland forests, ca. 200 m; Amazonas (Raudal Arata on Río Ocamo). Barinas, Lara, Mérida, Táchira, Trujillo; Belize, Panama, Colombia, Ecuador, Peru, Bolivia.

Fig. 280. Erythrina poeppigiana

Fig. 281. Erythrina fusca

Etaballia 325

29. ETABALLIA Benth., J. Bot. (Hooker) 2: 99. 1840. by Nidia L. Cuello A. Trees. Young stems glabrous, unarmed. Leaves alternate, unifoliolate, the leaflet large, entire, pinnately veined, ovate to ovate-oblong, acute to acuminate, rounded or subcordate at the base; stipules small, deciduous; petiolule very short. Inflorescences axillary spikes; bracts and bracteoles small, deciduous. Flowers 10–15 mm long, not papilionaceous. Calyx tubular, closed in bud, subregularly 2–5-toothed; petals 5(6), free, subequal, linear, imbricate. Stamens 10, rarely 11; filaments alternately long and short, connate at the base into a tube free or shortly adnate to the petals; anthers uniform, short, didymous, opening lengthwise. Ovary subsessile, 1–3 ovulate; style 1 mm long; stigma truncate, oblique. Fruit obovoid, 2-valved but drupelike, indehiscent. Seed usually 1; cotyledons fleshy. Venezuela, Guyana, Brazil; 1 species.

Fig. 282. Etaballia dubia

Etaballia dubia (H.B.K.) Rudd, Phytologia 20: 427. 1970. —Hecastophyllum dubium H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 388. 1823 [1824]. —Brasil, Sangrito. Etaballia guianensis Benth., J. Bot. (Hooker) 2: 99. 1840. Tree to ca. 30 m tall; inner bark with red sap; petals linear, yellow to orange. Semideciduous to evergreen lowland forests, 50–200 m; Bolívar (Caicara, Las Bonitas, Puerto Ordaz to San Félix), Amazonas (Caño Yapacana, Puerto Ayacucho, Río Ventuari). Anzoátegui, Apure, Guárico; Guyana, Brazil (Amazonas, Pará). ŠFig. 282.

326

F ABACEAE

30. FISSICALYX Benth., J. Proc. Linn. Soc., Bot. 5: 79. 1861. by Gerardo A. Aymard C. Trees. Leaves alternate, odd-pinnate; stipules deciduous; leaflets opposite or subopposite, without stipels. Inflorescences terminal panicles; bracts and bracteoles small, the latter persistent. Calyx tube turbinate, 1-lipped, the limb acuminate, entire or shortly 2-dentate, splitting in flower and spathaceous; petals yellow to orange, inserted with the stamens at the top of the calyx tube; standard ovate; wing petals obliquely oblong, free; keel petals ± similar to the wing petals, free. Stamens connate into a sheath split above; anthers versatile, opening by 2 apical pores. Ovary shortly stipitate, 2-ovulate; style filiform; stigma minute, terminal. Fruit flat, ovateelliptic including the broad wings, indehiscent. Seed pendent, the hilum small. Panama, Venezuela, Guyana, Brazil, Paraguay; 1 species. Fissicalyx fendleri Benth., J. Proc. Linn. Soc., Bot. 5: 79. 1861. Monopteryx jahnii Pittier, Bull. Torrey Bot. Club 42: 626. 1915. Tree to 20 m tall; petals orange to yellow. Semideciduous to evergreen lowland and lower montane forests, 100–500 m; Bolívar (Cerro Bolívar, La Paragua, lower Río Caroní), Amazonas (Río Manapiare). Aragua, Anzoátegui, Barinas, Carabobo, Cojedes, Guárico, Lara, Miranda, Portuguesa, Táchira; Panama, Guyana, Brazil, Paraguay. ŠFig. 283.

Fig. 283. Fissicalyx fendleri

Galactia 327

31. GALACTIA P. Browne, Civ. Nat. Hist. Jamaica 298. 1756. by Richard H. Maxwell Trailing, twining, or climbing, woody or herbaceous perennials, occasionally small, erect subshrubs. Leaves 3-foliolate or simple; stipules and stipels present; leaflet blade with entire, unlobed margins. Inflorescence axillary or terminal, erect, pseudoracemose; bracts and bracteoles present. Calyx 4-lobed; corolla pink, purplish, or white; standard reflexed; wing petals free; keel petals fused distally. Stamens 10, monadelphous or the vexillar one free. Ovary many-ovulate; style frequently exserted during anthesis; stigma minute, capitate. Fruit a legume, oblong to linear, dehiscent, somewhat compressed. Seeds ovoid, the hilum small, oval. Pantropics, extending into temperate areas; 50 species, ca. 7 in Venezuela, 3 of these in the flora area. Key to the Species of Galactia 1. 1. 2(1). 2.

Leaves digitately 3-foliolate; leaflets linear .............................. G. gracillima Leaves pinnately 3-foliolate; leaflets variable, but never linear .............. 2 Herbaceous or somewhat woody twining, climbing, or prostrate vines; leaflets elliptic-ovate ..................................................................... G. striata Woody erect shrubs or subshrubs; leaflets oval, ovate, oblong, or narrowly elliptic ..................................................................................... G. jussiaeana

Galactia gracillima Benth. in Mart., Fl. Bras. 15(1): 142. 1859. —Chinak guaiquin-chinaten (Pemón). Herbaceous, slender twiner to ca. 50 cm long. Moist savannas, 1000–1100 m; Bolívar (Río Kukenán). Suriname, southern Brazil, Paraguay, Argentina, Uruguay.

Glycine tenuiflora Klein ex Willd., Sp. Pl. 3(2): 1059 1802. —Galactia striata var. tenuiflora (Klein ex Willd.) Burkart, Darwiniana 16: 721. 1971. —Galactia tenuiflora (Klein ex Willd.) Wight & Arn. 1834, pro parte: sensu Benth. in Mart., Fl. Bras. 15(1): 143. 1859.

Galactia jussiaeana Kunth, Mimoses 196, t. 55. 1819 [1824]. —Alcornoquillo, Juan Zamora. Galactia angustifolia Kunth, Mimoses 201, t. 56. 1819 [1824]. —Galactia jussiaeana var. angustifolia (Kunth) Burkart, Darwiniana 16: 714. 1971. Galactia camporum Sprague, Trans. & Proc. Bot. Soc. Edinburgh 22: 430. 1905 [1904]. Shrub; pubescence sericeous to tomentose. Savannas, 50–600 m; Delta Amacuro (between San Felíx and Los Castillos), northern Bolívar, northern Amazonas. Sucre, widespread in the Llanos; West Indies, Colombia, Guyana, Suriname, French Guiana, northern Brazil. ŠFig. 284. Galactia striata (Jacq.) Urb., Symb. Antill. 2: 320. 1900. —Glycine striata Jacq., Hort. Bot. Vindob. 1: 32. 1770, “Glicine.”

Fig. 284. Galactia jussiaeana

328

F ABACEAE

Prostrate, trailing, or low twining vine. Open disturbed areas, 100–500 m; Bolívar (near Caicara, Moitaco), Amazonas (El Porvenir 54 km south of Puerto Ayacucho, Santa Rosa de Ucata). Throughout much of Venezuela; U.S.A. (Florida), Central America, West

Indies, tropical South America, Paraguay, Argentina. This species has been introduced worldwide as a forage crop for tropical pasture improvement and has become naturalized, extending its distribution.

32. GLIRICIDIA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 393. 1823 [1824], spelling variants: Glyricidia, Gliciridia. Hybosema Harms, Repert. Spec. Nov. Regni Veg. 19: 66. 1923. by Nidia L. Cuello A. Shrubs or medium-sized trees. Leaves alternate, sometimes partly subopposite or opposite, odd-pinnate; leaflets 5–19, entire, often drying with purple mottling beneath, without stipels; stipules small, caducous. Inflorescences axillary and/or cauline, of clustered racemes, appearing before or with the leaves; bracts inconspicuous, caducous; bracteoles absent. Flowers often rose or rose-tinged, pedicellate, hypanthium distinctively cup-shaped. Calyx teeth short and broad, or absent; petals 5, of similar length, free except the basally connate keel petals, short-clawed; standard nearly orbicular, erect or reflexed. Stamens 10, diadelphous, the vexillar stamen free; anthers uniform. Ovary short-stipitate, slender, straight, flattened, glabrous; ovules several to many; style glabrous, ca. 1/2 the length of the ovary; stigma capitate. Fruit a legume, dehiscent, stipitate, flattened, often wider toward the apex, the margins sometimes slightly thickened, nonseptate, exocarp smooth or only faintly indented between the seeds, glabrous, the valves hard and often coiling in dehiscence. Seeds to 10 mm long, nearly round to oblong, compressed. Neotropics; ca. 6 species, 1 in Venezuela. Gliricidia sepium (Jacq.) Kunth ex Walp., Repert. Bot. Syst. 1: 679. 1842. —Robinia sepium Jacq., Enum. Syst. Pl. 28. 1760. —Mata ratón. Tree to 6 m tall. Cultivated and perhaps a locally naturalized escape, 100–300 m; northeastern Bolívar, Amazonas (Puerto Ayacucho). Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Portuguesa, Sucre, Yaracuy; native to Mexico, Central America, and Colombia, introduced and naturalized in West Indies, Guyana, Ecuador, Brazil, and Paleotropics. ŠFig. 285. Gliricidia sepium is cultivated for forage. It is also used as a living fence or as a mouse poison.

Fig. 285. Gliricidia sepium

Hymenolobium 329

33. HYMENOLOBIUM Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 84. 1860. by Haroldo Cavalcante de Lima and Gerardo A. Aymard C. Large or small trees or shrubs. Leaves alternate, crowded at the end of the branches, deciduous; leaflets 15–55, opposite on the rachis; stipules linear or lanceolate, caducous; stipels minute. Inflorescence a terminal panicle. Calyx campanulate, apex truncate, margin obscurely 5-dentate; petals rose or rose-violet; standard orbiculate and emarginate; keel petals adnate on lower side. Stamens 10, connate into a sheath open on the upper side. Ovary with many ovules. Fruit indehiscent, orbicular to oblong-elliptic, samaroid or nut-like, reticulate-veined, with 2 parallel prominent veins near the margins, membranous or coriaceous; seed chambers central. Seeds 1(2), with short straight radicle. Central America, South America; 12 species, 2 in Venezuela, both in the flora area. Key to the Species of Hymenolobium 1.

1.

Leaflets pilose to densely pubescent on lower surface, the apex apiculate; calyx 8–9 mm long; fruit orbicular to oblong, nut-like, coriaceous, 4–7 × 3–4 cm ............................................................................... H. heterocarpum Leaflets glabrous on both surfaces, the apex emarginate; calyx 4–6 mm long; fruit oblong-elliptic, samaroid, membranous, 5–12 × 1.5–2.5 cm .................................................................................................. H. petraeum

Fig. 286. Hymenolobium heterocarpum

330

F ABACEAE

Hymenolobium heterocarpum Ducke, Trop. Woods 47: 6. 1936. —Anzuelito, Caraota montañera, Mañu, Siñate (Yekwana). Tree 10–30 m tall; flowers pinkish red. Riparian forests, 50–200 m; Bolívar (Río Tabaro near Río Nichare), Amazonas (Maroa to Yavita, Río Cuao, Río Mawarinuma, Río Sipapo, near San Fernando de Atabapo). Suriname, Ecuador, Brazil. ŠFig. 286.

Hymenolobium petraeum Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 36. 1915. —Alcornoque, Alcornoque montañero, Atebrino, Pelote, Tanyeechemen (Panare). Tree 20–40 m tall; flowers pink; fruit red. Evergreen lowland and semideciduous forests, granitic slopes, ecotone of savannas, 100–400 m; Bolívar (Río Maniapure, Río Parguaza, Río Toro), Amazonas (Caicara to Puerto Ayacucho road, Río Cataniapo). Guárico; Colombia, Guyana, Suriname, French Guiana, Brazil.

34. INDIGOFERA L., Sp. Pl. 751. 1753. Anila Mill., Gard. Dict. Abr. ed. 4. 1754. by Gerardo A. Aymard C. Perennial herbs or small woody shrubs, strigose with pale, appressed, T-shaped (malpighiaceous) trichomes. Leaves odd-pinnate, pinnately 3-foliolate, or 1-foliolate; leaflets entire; stipels present, often inconspicuous; stipules setaceous, deciduous, apparently adnate to the petioles. Inflorescences axillary racemes or spikes; pedicels short or apparently absent; bracteoles absent. Flowers usually reddish or white; hypanthium campanulate. Calyx 5-dentate, the teeth subequal; standard broad, circular, subsessile, strigose without; wing petals oblong, somewhat adherent to the keel petals, auriculate; keel petals coherent, laterally spurred. Stamens 10, diadelphous, vexillar stamen free, anthers with an apical projection. Ovary slender, sessile, usually strigose; ovules few to many; style short, glabrous; stigma capitate. Fruits oblong or linear, terete or 4-angled, curved or straight, dehiscent or apparently indehiscent, septate. Seeds globose to cylindric and truncate, attached at the middle. Pantropics (most diverse in Africa and Asia); ca. 700 species, 11 in Venezuela, 6 of these in the flora area. Key to the Species of Indigofera 1. 1. 2(1). 2. 3(2). 3.

4(2). 4. 5(4). 5.

Leaves 1-foliolate, linear-lanceolate, 1.3–5 mm wide ............. I. bongardiana Leaves with 3–20 leaflets, rarely 1-foliolate, oblong-elliptic, leaflets 6–15 mm wide ................................................................................................. 2 Leaflets acute at the base; fruits curved ................................................... 3 Leaflets cuneate at the base; fruits straight ............................................. 4 Fruits 1–1.5 cm long, strongly curved, finely pubescent; leaflets pubescent on both surfaces .................................................................... I. suffruticosa Fruits 3–3.5 cm long, slightly curved, glabrous or sparsely pubescent; leaflets usually glabrous on upper surface, pubescent on lower surface ..................................................................................................... I. tinctoria Prostrate herb; leaflets 0.2–0.8 cm long; flowers purple; fruits 0.4–1 cm long ......................................................................................... I. microcarpa Erect or procumbent herb or shrub; leaflets 1.5–5 cm long, eglandular; flowers red or pink; fruits 1.5–4 cm long .............................................. 5 Stems, petioles, inflorescence rachis, and fruits appressed-sericeous to glabrescent ......................................................................... I. lespedezioides Stems, petioles, inflorescence rachis, and fruits densely hirsute with long brown-red, erect trichomes ........................................................... I. hirsuta

Indigofera 331

Indigofera bongardiana (Kuntze) Burkart, Darwiniana 4: 171. 1942. —Anila bongardiana Kuntze, Revis. Gen. Pl. 2: 938. 1891. Suffrutescent ca. 60 cm tall; flowers purple. Periodically flooded savannas, 100–200 m; Amazonas (Río Parucito). Southern Brazil, Paraguay, northern Argentina, Uruguay. Indigofera hirsuta L., Sp. Pl. 1062. 1753. —Aristín. Perennial herb or shrub, sometimes with procumbent branches; flowers pink. Savannas, roadsides, 100–200 m; Bolívar (Maripa). Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Guyana, Brazil, Bolivia, Paraguay. Indigofera lespedezioides H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 457. 1823 [1824]. —Añilito, Juan Zambrano, Juan Zamora, Mato zabandu, Raíz de zamuro, Romero de sabana. Indigofera pascuorum Benth., Ann. Nat. Hist. 3: 431. 1839. Perennial herb or shrub; flowers pink. Wet savannas, roadsides, 50–400 m; northern Bolívar, Amazonas (near Puerto Ayacucho). Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Guyana, Brazil, Bolivia, Paraguay. ŠFig. 287. This species is locally cultivated for its roots, which are used to treat diarrhea. Indigofera microcarpa Desv., J. Bot. (Desvaux) 3: 79. 1814. —Pega pega rosada. Prostrate herb; flowers purple. Open savannas, 50–200 m; Bolívar (northeast of Cerro Cuchivero, El Tigre on Río Cuchivero). Guárico, Monagas, Zulia; Peru, Brazil, Bolivia, Paraguay, Uruguay. Indigofera suffruticosa Mill., Gard. Dict. ed. 8, no. 2. 1768. —Añilito, Inchanpichanemen (Panare). Suffrutescent or shrub to 3 m tall; flowers pink. Open and disturbed areas, near sea level to 600 m; Delta Amacuro (Isla Iduburojo near Misión San Francisco de Guayo, Pedernales) Bolívar (Ciudad Bolívar, Icabarú, Kilómetro 88, Río Caura, Río Suapure, Upata). Widespread elsewhere in Venezuela; U.S.A., Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.

Fig. 287. Indigofera lespedezioides

332

F ABACEAE

Indigofera suffruticosa is used as a fish poison in the lower Río Caura basin. Indigofera tinctoria L., Sp. Pl. 751. 1753. Erect or sprawling shrub to 1.5 m tall; flowers pink. Open and disturbed areas, ca.

50 m; Bolívar (Ciudad Bolívar). Monagas; U.S.A. (Florida), Mexico, Central America, Antilles, native of the Old World (India) introduced and naturalized in Neotropics as the source of blue indigo dye.

35. LECOINTEA Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 128. 1922. by Basil Stergios Trees. Leaves 1-foliolate, variously serrate to crenate, glabrous; stipules caducous. Flowers not papilionaceous, rather small, pedicellate, borne on axillary, often branched racemes; bracts persistent; bracteoles small, persistent. Calyx gamosepalous, tube campanulate to cotyliform (disk-shaped but with raised or ascending border) with 5 obscure lobes; petals 5, separate, spatulate, clawed, imbricate, soon deciduous, outer one much wider than inner 4. Stamens 9–13, free, ± unequal; anthers basifixed, longitudinally dehiscent, pilosulous at tip. Ovary stipitate; ovules 4– 6; style straight or slightly bent, stout and exerted; stigma small, oblique. Fruit thick, globose to oval or broadly turbinate, indehiscent. Seeds 1 or 2, thick and semicompressed, ovoid to orbicular. Central America, Venezuela, Suriname, Ecuador, Peru, Brazil, Bolivia; 6 species, 2 in Venezuela, 1 of these in the flora area.

Fig. 288. Lecointea amazonica

Lonchocarpus 333

Lecointea amazonica Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 129, pl. 8. 1922. —Níspero, Palo de arco, Pilón. Tree 15–25 m tall; flowers white. Evergreen lowland and flooded forests, 50–600 m;

Delta Amacuro (upper Río Toro), Bolívar (Represa Guri, Río Nichare), Amazonas (southern Río Manapiare basin). Guatemala, Belize, Costa Rica, Suriname, Ecuador, Peru, Brazil. ŠFig. 288.

36. LONCHOCARPUS H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 383. 1823 [1824], nom. cons. by Hans-Helmut Poppendieck Trees, shrubs, or vines. Leaves alternate, odd-pinnate; leaflets opposite, entire; stipules caducous; stipels absent (present in L. densiflorus). Inflorescence a lateral or terminal pseudoraceme, the flowers either in pairs on distinct or short, lateral peduncles, or numerous on lateral short shoots, or arising directly from the rachis and appearing verticillate, paired, or solitary; peduncular and pedicellar bracts caducous; bracteoles 2, borne on the pedicel or at the base of the calyx. Calyx dentate to truncate, cup-shaped; corolla purple or white; standard blade broadly ovate to obovate, clawed, the base either narrow and without callosities, or truncate to cordate with callosities; wing petals adhering to the keel petals by pressure and structural conformity; keel petals touching or connate. Vexillar stamen free at base; anthers uniform; nectary glands indistinct. Ovary pubescent; ovules 2–12; style glabrous. Fruit generally indehiscent, occasionally dehiscent along the vexillar suture, or separating into 1-seeded loments, papery to thin or thick woody, 1–12-seeded, vexillar margin occasionally winged, but wings never submarginal. Mexico, Central America, West Indies, South America south to Peru and northern Argentina, 1 species in western Africa; ca. 150 species, 21 in Venezuela, 16 of these in the flora area. The circumscription adopted here for Derris and Lonchocarpus is conventional, but far from satisfactory. It is maintained following earlier treatments by Benth. (in Carl Friedrich Philipp von Martius, Flora Brasiliensis 15(1): 275–287. 1862), G. Amshoff (in A. Pulle, Flora of Suriname 2(2): 141–147. 1939), and H. Pittier (Bol. Tec. Minist. Agric. Cría, Caracas 5: 96–106. 1944), and because it is consistent with some recent chemical and other surveys and avoids nomenclatural changes with farreaching consequences. There is a close connection between Lonchocarpus subgenus Phacelanthus Pittier and the controversial American species of Derris. The winged fruit, which is the diagnostic character differentiating the two genera, has certainly evolved more than once in a parallel manner. Lonchocarpus utilis is a rotenone-yielding, cultivated species that is often found in cultivation or escaped. It has never been collected in flower or fruit and therefore is not included in the key. It has oblong-elliptic leaflets with a gradually acuminate apex and a round to acute base, whereas the other known rotenone-containing species, L. urucu, has obovate-oblong leaflets with an abruptly acuminate apex and a roundish to obtuse base. Key to the Species of Lonchocarpus (except L. utilis; see above) 1. 1.

Flowers numerous in short lateral shoots; generally lianas, rarely trees or shrubs (subgenus Phacelanthus) ........................................................... 2 Flowers paired on a common peduncle, or directly arising from the spike; trees or shrubs, rarely vines (subgenus Lonchocarpus) ....................... 6

334

F ABACEAE

2(1). 2.

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

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

10.

11(10).

11.

12(9).

12.

13(12). 13.

Deciduous trees; legumes flattened, lanceolate to linear, fragile and sublomentaceous, 1–8-seeded ........................................................ L. pictus Evergreen plants, initially shrubby but then becoming large lianas; legumes either flattened and 1- or 2-seeded, or strongly thickened, 1–12seeded ..................................................................................................... 3 Stipels present; ovules 10–12; legumes 7–15 × 4–5 cm, to 12-seeded, lomentaceous .......................................................................... L. densiflorus Stipels absent; ovules 2–6; legumes flattened and different from above ................................................................................................................ 4 Calyx and corolla without latex ducts; standard with ± distinct auriculae or basal callosities, base truncate, claw glabrous ................ L. floribundus Calyx and corolla with latex ducts; standard without auriculae or basal callosities, base narrowed, claw pilose .................................................. 5 Lower surface of leaflets distinctly spreading-pilose, trichomes ca. 0.2 mm long; margins of legume only slightly thickened ...................... L. martynii Lower surface of leaflets appressed-pilose, appearing subglabrous, trichomes 0.25–0.5 mm long; margins of legume distinctly thickened ........................................................................................................ L. urucu Flowers arising directly from the spike and appearing verticillate, paired, or solitary; peduncle of fruit not jointed ............................................... 7 Flowers paired on a common peduncle; peduncle of fruit jointed ............ 9 Flowering spikes 14–18 cm long ..................................................... L. fendleri Flowering spikes ≤ 11 cm long ................................................................... 8 Leaflets obtuse ..................................................................... L. crucisrubierae Leaflets acuminate ........................................................................ L. tubicalyx Calyx denticulate ..................................................................................... 10 Calyx entire, or at most slightly undulate .............................................. 12 Spikes dense, flowers touching each other, often at least partly dark violet to blackish when dry; leaflets 11–20 × 6.8–8.5 cm, without pellucid dots; legumes to 7.5 × 2.3 cm .............................................. L. heptaphyllus Spikes lax, flowers not touching each other and light-colored when dry; leaflets 3.5–11 × 1.5–6.5 cm, with or without pellucid dots; legumes at least 7.5 × 2.8 cm ................................................................................. 11 Leaflets rugose, the margin involute, the main veins of lower surface distinctly pilose; legumes 3.5–4 cm wide, to 10 mm thick, prominently and abruptly thickened at the seeds, the seeds nearly spherical ................. ........................................................................................... L. crassispermus Leaflets flat, the margin flat, tomentose on both surfaces; legumes 2.8–3.8 cm wide, to 2 mm thick, not abruptly thickened at the seeds, the seeds not spherical ........................................................................... L. hedyosmus Leaves pellucid-punctate, glabrous; spikes 11–19 cm long; peduncles 7–10 mm long; pedicels 5–7 mm long; legumes 7–17 × 3–3.2 cm, glabrous .................................................................................................. L. punctatus Leaves not pellucid-punctate, glabrous to pubescent; spikes 9.5–15 cm long; peduncles to 3 mm; pedicels to 3 mm long; legumes 4–11 × 1.3–2.5 cm, pubescent or glabrous ................................................................... 13 Vexillar margin of legume conspicuously winged ................ L. dipteroneurus Vexillar margin of legume without wings ............................................... 14

Lonchocarpus 335

14(13). Legumes (1–)3–7-seeded, 5.5–11 × 1.3–1.8 cm, bullate, dark brown, glabrous; leaves glabrous .......................................................... L. imatacensis 14. Legumes 1- or 2-seeded, 4–6.5 × 2–2.5 cm, woody, dark gray, pubescent; leaves pubescent on lower surface, scabrid to pubescent on upper surface .............................................................................................. L. sericeus Lonchocarpus crassispermus Poppend., Novon 2: 53. 1992. —Jebe, Mahomo chino. Tree to 20 m tall. Semideciduous forests, savanna edges, 200–300 m; Bolívar (15 km southeast of El Callao, between La Encrucijada and El Pao, 7 km north of mouth of Río Paragua, 17 km from Upata toward San Félix). Endemic. Lonchocarpus crucisrubierae Pittier, Arb. Arbust. Venez. 6–8 [reprinted from Bol. Minist. RR. EE. no. 8/9]: 100. 1927. —Majomo rebalsero, Menudito, Tocorito. Tree or shrub 2–8 m tall; trunk to 40 cm diameter, mostly branched from the base, occasionally with elongate vining branches; flowers precocious. Semideciduous and seasonally flooded forests, 50–200 m; Bolívar (Ciudad Bolívar, Puerto Ordaz, lowlands of Río Orinoco basin and tributaries). Barinas, Guárico. Lonchocarpus densiflorus Benth., Ann. Nat. Hist. 3: 433. 1839. Lonchocarpus glabrescens Benth., Hooker’s J. Bot. Kew Gard. Misc. 2: 233. 1850. —Derris glabrescens (Benth.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 261. 1943. Lonchocarpus boliviensis Pittier, Contr. U.S. Natl. Herb. 20: 93, fig. 43. 1917. Small shrub, flowering when 3 m tall, becoming a high-climbing liana; young branches reddish brown, lenticellate. Riparian forests, 50–200 m; Delta Amacuro (Caño Carisal), Bolívar (Río Orinoco), Amazonas (Río Negro). Apure; Colombia, Peru, northern Amazonian Brazil, Bolivia. The 1-seeded segments of the peculiar lomentaceous fruits are dispersed by water. Lonchocarpus dipteroneurus Pittier, Contr. U.S. Natl. Herb. 20: 90. 1917. Lonchocarpus stenopteris Pittier, Arb. Arbust. Venez. 6–8 [reprinted from Bol. Minist. RR. EE. no. 8/9]: 102. 1927

Small tree 3–10 m tall, often manystemmed or -branched from the base; twigs gray to dark brown, lenticellate. Deciduous forests, 100–300 m; Bolívar (Represa Guri, Villa Lola). Aragua, Distrito Federal, Guárico, Miranda. The collections from Bolívar state are either widely disjunct from the populations in northern Venezuela, or else the species has not been collected in intermediate areas. Lonchocarpus fendleri Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 94. 1860. Lonchocarpus fendleri subsp. pubescens Pittier, Contr. U.S. Natl. Herb. 20: 91. 1917. Lonchocarpus sanctae-marthae Pittier, Contr. U.S. Natl. Herb. 20: 92, fig. 42, pl. 1. 1917. Lonchocarpus stenurus Pittier, Arb. Arbust. Venez. 6–8 [reprinted from Bol. Minist. RR. EE. no. 8/9]: 102. 1927. Deciduous tree 6–25 m tall, often lowbranching, with dense, rounded to almost flat crown, bark light gray; flowers appearing before the leaves. Gallery forests, near sea level to 100 m; Delta Amacuro (Caño Arature). Frequent in northern Venezuela; Panama, Colombia. Lonchocarpus floribundus Benth., Ann. Nat. Hist. 3: 432. 1839. —Derris floribundus (Benth.) Ducke, Bol. Técn. Inst. Agron. N. 18: 197. 1949. Lonchocarpus nitidulus Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 98. 1860. Shrublet, flowering when 1 m tall, becoming a large liana in closed forests. Locally frequent in clearings, mostly on sandy soil, 100–200 m; Bolívar (Serranía Baraguán, Río Parguaza), Amazonas (Río Negro, Río Orinoco and tributaries). Guyana, Suriname, French Guiana, Amazonian Brazil. Lonchocarpus floribundus is used as a fish poison in Brazil. Lonchocarpus hedyosmus Miq., Linnaea 18: 564. 1844. —Derris hedyosma (Miq.)

336

F ABACEAE

J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 262. 1943. —Jebe, Majomo. Lonchocarpus margaritensis Pittier, Contr. U.S. Natl. Herb. 20: 91, fig. 41. 1917. Lonchocarpus ernestii Harms, Repert. Spec. Nov. Regni Veg. 17: 321. 1921, “ernesti.” —Derris ernestii (Harms) Ducke, Bol. Técn. Inst. Agron. N. 18: 196. 1949, “ernesti.” Lonchocarpus paniculatus Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 161. 1922. Tree 8–30 m tall; trunk to 1 m diameter, often branched from the base; bark gray. Semideciduous and evergreen lowland forest, forested slopes along savannas, 50–500 m; widespread in northern Bolívar, Amazonas (San Fernando de Atabapo). Venezuelan Coastal Cordillera and Andes; Guyana, Suriname, Brazil. The hard and durable wood of Lonchocarpus hedyosmus is used for construction. Lonchocarpus heptaphyllus (Poir.) DC., Prodr. 2: 259. 1825. —Dalbergia heptaphylla Poir. in Lam., Encycl. suppl. 2: 445. 1811 [1812]. —Tocorito. Dalbergia pentaphylla Poir. in Lam., Encycl. suppl. 2: 445. 1811 [1812]. —Lonchocarpus pentaphyllus (Poir.) DC., Prodr. 2: 259. 1825. Lonchocarpus latifolius H.B.K. ex DC., Prodr. 2: 260. 1825. —Derris latifolia (H.B.K. ex DC.) Ducke, Bol. Técn. Inst. Agron. N. 18: 195. 1949. Lonchocarpus discolor Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr. 3: 421. 1902. Tree 7–26 m tall; trunk to 38 cm or more diameter, occasionally vining; hollow branches ant-inhabited; bark reddish brown to grayish. Evergreen lowland forests, 100– 300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Serranía de Imataca). Apure, Barinas, Carabobo, Distrito Federal, Miranda, Monagas, Sucre; Central America, West Indies, Colombia, Peru, Brazil (Amapá, Pará). Lonchocarpus imatacensis Poppend., Novon 2: 53. 1992. —Jebe, Majomo. Tree to 26 m tall; trunk to 40 cm diameter. Evergreen lowland forests, 50–400 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Salto Pará, Serranía de Imataca). Endemic. ŠFig. 291.

This species has been misidentified as Lonchocarpus nitidus Benth., but can be distinguished by the bullate seed region of the fruit. The wood is used for railroad ties. Lonchocarpus martynii A.C. Sm., Amer. J. Bot. 26: 577. 1937. Scandent shrub or liana eventually reaching large dimensions; young branches dark brown, pubescent. Evergreen lowland to lower montane forests, gallery forests, forest borders, 200–900 m; Bolívar (Río Nichare basin, Santa Elena de Uairén). Guyana, adjacent Amazonian Brazil. Lonchocarpus pictus Pittier, Arb. Arbust. Venez. 2/3 [reprinted from Bol. Comerc. Industr. Venez. no. 34]: 26. 1923. —Majomo, Tocorito, Tocorito blanco. Slender or branched deciduous tree or shrub 8–15 m tall; flowers present as or rarely before the leaves expand. Semideciduous to evergreen lowland forests, gallery forests, 50–200 m; Delta Amacuro (near Río El Toro), Bolívar (Ciudad Bolívar, Río Suapure to La Paragua). Venezuelan Coastal Cordillera; Guyana, Suriname, French Guiana. ŠFig. 290. Like Hibiscus tiliaceus L., the bark of Lonchocarpus pictus is used locally against erysipelas, a febrile disease producing deep red color of the skin. Lonchocarpus punctatus H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 383. 1823 [1824]. —Jebe. Robinia violacea Jacq., Enum. Syst. Pl. 28. 1760, nom. dubium. —Lonchocarpus violaceus (Jacq.) DC., Prodr. 2: 259. 1825. Lonchocarpus benthamianus Pittier, Contr. U.S. Natl. Herb. 20: 86. 1917. Spreading, branched tree or shrub 1–15 m tall; bark grayish. Deciduous to semidecidous forests, 50–300 m; Bolívar (Río Caroní, mouth of Río Paragua, Río Upata, Río Yuruari). Venezuelan Coastal Cordillera; West Indies, Colombia, Ecuador, Peru, Brazil. Lonchocarpus sericeus (Poir.) H.B.K. ex DC., Prodr. 2: 260. 1825. —Robinia sericea Poir. in Lam., Encycl. 6: 226. 1804. —Majomo.

Lonchocarpus 337

Fig. 289. Lonchocarpus sericeus

Fig. 290. Lonchocarpus pictus

338

F ABACEAE

Fig. 291. Lonchocarpus imatacensis

Tree 12–25 m tall, the wood hard and durable; leaf rachis, branchlets, and inflorescences ferruginous-pubescent; bark brownish or grayish, lenticellate, yielding resinous fluid when cut. Semideciduous to evergreen lowland or lower montane forests, 50–400 m; Delta Amacuro (Río Acure), Bolívar (near El Dorado, near La Paragua). Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, western Africa. ŠFig. 289. Lonchocarpus tubicalyx Pittier ex Poppend., Novon 2: 57. 1992. Shrub or tree 4–5 m. Habitat not known, near sea level to 100 m; Delta Amacuro (Caño Mánamo south of Tucupita). Apure. Lonchocarpus tubicalyx is currently known from just two specimens with very short pedicels, making the flowers appear verticillate. Lonchocarpus urucu Killip & A.C. Sm., J. Wash. Acad. Sci. 20: 81, fig. 4, 1930. —Derris urucu (Killip & A.C. Sm.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 266. 1943. Erect shrub becoming a huge woody vine, often found in large clumps of several acres. Well-drained, fertile soils in nonflooded forests, ca. 100 m; Amazonas (La Esmeralda). Amazonian Brazil. Lonchocarpus urucu is used in Brazil as a fish poison and a source of commercially ex-

ploited rotenone. It is very difficult to differentiate from L. utilis, which is used in the same manner. The leaf shape affords the only, and often difficult, way to distinguish the species. Lonchocarpus utilis A.C. Sm., Amer. J. Bot. 24: 580. 1937. —Derris utilis (A.C. Sm.) Ducke, Bol. Técn. Inst. Agron. N. 18: 197. 1949. —Barbasco, Barbasco blanco, Barbasco blanco caicareño, Barbasco bravo, Timbo. Lonchocarpus nicou auct. non (Aubl.) DC. 1825: sensu Killip & A.C. Sm., J. Wash. Acad. Sci. 20: 74. 1930. —Derris nicou (Aubl.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 263. 1943, non Robinia nicou Aubl., Hist. Pl. Guiane 771. 1775. Erect shrub becoming a woody vine. Secondary forests at sites of old cultivated areas, near sea level to 300(–1300) m; scattered throughout Delta Amacuro, Bolívar, and Amazonas. Elsewhere in Venezuela in the adjacent Río Orinoco basin; widely cultivated throughout the Amazon basin, Guyana, Suriname, French Guiana, and eastern Brazil. The rotenone-yielding roots of Lonchocarpus utilis are used as a fish poison. The flowers and fruits are unknown; it is propagated vegetatively and spreads in cultivation by means of its roots.

Machaerium 339

37. MACHAERIUM Pers., Syn. Pl. 2: 276. 1807, nom. cons. Drepanocarpus G. Mey., Prim. Fl. Esseq. 236. 1818. by Velva E. Rudd Lianas, trees, or shrubs, scandent with spine-like prehensile shoots; sap usually reddish. Leaves alternate, odd-pinnate, 3–many-foliolate; leaflets alternate, entire; stipules usually indurated, spinescent or sometimes caducous; stipels lacking. Inflorescence terminal or axillary, racemose, sometimes paniculate; bracts small, caducous or stipule-like, spiny but smaller; bracteoles small, usually broadly ovate, obtuse, paired at base of calyx. Flowers papilionaceous, ca. 4–18 mm long. Calyx campanulate or briefly tubiform, 5-lobed to subtruncate; petals white to yellow, pink, bluish, or purple. Stamens 10, monadelphous or diadelphous (5:5 or 9:1); anthers dorsifixed. Ovary 1- or 2-ovulate; style filiform; stigma minute, capitate, terminal. Fruit indehiscent, stipitate, usually samaroid with a basal seminiferous body and a sterile, terminal wing, or sometimes lunate, falcate, or subreniform with the wing reduced or lacking. Seeds compressed, ovate, reniform, or orbicular. Tropical and subtropical America, 1 species in coastal West Africa; ca. 130 species, ca. 35 in Venezuela, 22 of these in the flora area. Key to the Species of Machaerium 1. 1. 2(1). 2. 3(2).

3.

4(3).

4.

5(2). 5. 6(5).

Leaflets with the secondary veins approximately parallel, usually well defined, extending to the margin or nearly so ...................................... 2 Leaflets with the secondary veins branching, not always parallel, usually not extending to the margin, sometimes weakly defined ..................... 9 Secondary veins ca. 3–25 mm apart; leaves 5–15-foliolate; leaflets 1.5– 11 cm wide; fruit winged ........................................................................ 3 Secondary veins ca. 1 mm apart; leaves 5–35-foliolate; leaflets 0.5–3 cm wide; fruit winged or unwinged ............................................................ 5 Lower surface of leaflets with trichomes lax or curled; venation conspicuous; fruit ferruginous-tomentulose, glabrescent with age; flowers 8– 17 mm long .............................................................................. M. quinatum Lower surface of leaflets subglabrous or with trichomes appressed and usually minute; secondary veins usually evident but lesser veins inconspicuous; fruit puberulent to tomentulose, usually subglabrous by maturity; flowers 5–10 mm long ................................................................ 4 Leaflets predominantly ovate to elliptic, acute to acuminate, sometimes obtuse; flowers 6–10 mm long on pedicels ca. 1 mm long; calyx 3–4 × 2– 2.5 mm; fruit with stipe 3–5(–10) mm long ...................... M. floribundum Leaflets predominantly obovate to elliptic, obtuse; flowers 5–8 mm long, essentially sessile; calyx 2.5–3 × 1.5–2 mm; fruit with stipe 2–3 mm long .................................................................................. M. macrophyllum Fruit with a well-developed terminal wing ............................................... 6 Fruit lunate or falcate, lacking a terminal wing ....................................... 7 Leaflets ca. 1–2 × 0.2–0.6 cm, linear to oblong or elliptic-oblong, the surfaces puberulent with lax trichomes, somewhat glabrescent with age; flowers (7–)8–10 mm long, standard puberulent on the outer face; fruit essentially straight ........................................................................ M. affine

340

F ABACEAE

6.

7(5).

7.

8(7).

8.

9(1). 9. 10(9).

10.

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

13.

14(13). 14.

Leaflets 1–5(–9) × (0.5–)1–2(–3.5) cm, elliptic to oblong, the upper surface glabrous or nearly so, the lower surface puberulent with lax trichomes or finely appressed-pubescent, glabrescent with age; flowers (8–)10–15 mm long, the standard glabrous or subglabrous on the outer face; fruit usually bent at about a 90° angle ..................................... M. robiniifolium Leaflets acute to obtuse or emarginate, aristate at the apex, the arista (0.5–)1–5 mm long, rarely lacking, the secondary veins diverging from the midvein at angles of about 90°; flowers 10–14 mm long; calyx 5–7 × 4 mm .................................................................................... M. aristulatum Leaflets obtuse or retuse, not apiculate, the secondary veins diverging from the midvein at angles of about 45°; flowers (7–)8–13 mm long; calyx 3.5–5(–7) × 2–2.5 mm ....................................................................... 8 Flowers ca. 10–13 mm long; calyx ferruginous-pubescent; fruit falcate or lunate, not curved into a circle, puberulent, glabrescent with age, sessile, or stipe to ca. 5 mm long; leaves ca. 15–35-foliolate; lower surface of leaflets puberulent with lax trichomes ............................... M. ferox Flowers ca. 10 mm long; calyx glabrous to moderately pubescent, not ferruginous; fruit usually curved into a circle, glabrous or subglabrous at maturity, the stipe 5–10 mm long; leaves 5–13-foliolate; lower surface of leaflets glabrous to moderately pubescent with minute appressed trichomes ....................................................................................... M. lunatum Leaflets ca. (0.2–)0.3–3(–4) × 0.1–1.1 cm, predominantly elliptic to linear or oblong; fruit winged ......................................................................... 10 Leaflets ca. 2–20 × 1.5–8.5 cm, predominantly ovate to ovate-elliptic or oblong; fruit with or without a terminal wing .................................... 12 Flowers (8–)10–12 mm long; calyx 3.5–5 × 2.5–3 mm; petals white to purple, the standard pubescent on the outer face; leaflets elliptic to oblong, 5–30 × 3–11 mm ........................................................ M. altiscandens Flowers 4–6(–7) mm long; calyx 1–2.5 × 1–1.5 mm; petals white to creamcolored or yellowish, sometimes with a few purplish or reddish markings, the standard glabrous on the outer face; leaflets linear or linearoblong, 5–30(–40) × 1–10 mm .............................................................. 11 Leaves 37–90(–100)-foliolate; leaflets linear or linear-oblong, 5–13 × 1– 2 mm .............................................................................. M. multifoliolatum Leaves ca. 35–51-foliolate; leaflets linear-oblong, 10–30(–40) × (2–)3– 5(–10) mm .......................................................................... M. myrianthum Fruit lunate or subreniform, lacking a terminal wing ........................... 13 Fruit with a well-developed terminal wing ............................................. 15 Leaflets coriaceous, usually blackening on drying, sometimes pubescent on lower surface along the midvein otherwise glabrous; fruit ferruginous- or fulvous-velutinous; flowers white to pinkish ........ M. inundatum Leaflets subcoriaceous, usually not much darkening on drying, minutely appressed-pubescent to subglabrous on lower surface; fruit fulvoustomentulose or puberulent, glabrescent; flowers dark red to purple .... 14 Fruit ca. 7–9 cm long, the stipe 5–7 mm long; pubescence generally whitish or gray; trees ........................................................................ M. dubium Fruit ca. 3.5 cm long, the stipe 1–3 mm long; pubescence generally brownish or tawny; small trees, shrubs, or lianas ....................... M. leiophyllum

Machaerium 341

15(12). Flowers 13–18 mm long; calyx 6–10 × 4–5 mm; fruit ferruginoustomentulose, sometimes setose, usually glabrous at maturity, the stipe 5–10 mm long ............................................................................... M. kegelii 15. Flowers 5–11 mm long; calyx 2–4 × 1–2.5 mm; fruit sericeous to puberulent or glabrous, the stipe 2–15 mm long ............................................ 16 16(15). Lianas; flowers 8–11 mm long; calyx 3.5–4 × 2–3 mm ............................ 17 16. Lianas or trees; flowers ca. 5–8 mm long; calyx 2–3.5 × 1–2 mm .......... 18 17(16). Flowers 9–11 mm long; petals white but drying black, rusty-brownvillous; leaves 11–17-foliolate; fruit sometimes puberulent at the base, otherwise glabrous, the stipe 10–12 mm long ................... M. amazonense 17. Flowers 8–10 mm long; petals bluish; leaves 11- or 13-foliolate; fruit fulvousto feruginous-velutinous, the stipe 5–15 mm long ................. M. tovarense 18(16). Fruit fulvous-sericeous, glabrescent, not darkening on drying, the stipe 2– 5 mm long; flowers 5–7 mm long; calyx 2.5–3 × 2 mm, subsericeous scarcely striate; petals purplish or white, not darkening on drying, the standard sericeous on the outer face; upper surface of leaflets glabrous, the lower surface sericeous or subsericeous to glabrous; usually lianas .............................................................................................. M. madeirense 18. Fruit glabrous to puberulent, usually darkening on drying, the stipe 4– 15 mm long; flowers 5–8 mm long; calyx 2–3.5 × 1–2 mm; petals white or greenish, usually darkening on drying; leaflets glabrous on both surfaces or puberulent to glabrous on upper surface, appressed-pubescent to subsericeous on lower surface; trees or shrubs .............................. 19 19(18). Calyx ca. 2 × 2 mm, scarcely striate; fruit essentially glabrous at maturity, the stipe (6–)10–15 mm long; leaves 11–17-foliolate; upper surface of leaflets puberulent to glabrous, the lower surface appressed-pubescent to subsericeous ..................................................................... M. acutifolium 19. Calyx ca. 2–3.5 × 1–2 mm, with prominent longitudinal striations; fruit puberulent to glabrous at maturity, the stipe 4–10 mm long; leaves 3– 9-foliolate; leaflets glabrous on both surfaces ..................................... 20 20(19). Flowers ca. 5–6.5 mm long; calyx 2–2.5 × 1–1.5 mm; leaflets acuminate, the acumen ca. 1.5–2 cm long; fruit with stipe 7–10 mm long ............... ............................................................................................ M. acuminatum 20. Flowers ca. 8 mm long; calyx 3–3.5 × 2 mm; leaflets acuminate, the acumen to ca. 1 cm long; fruit with stipe 5.5–7 mm long..... M. guaremalense Machaerium acuminatum H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 391. 1823 [1824]. —Nissolia acuminata (H.B.K.) DC., Prodr. 2: 258. 1825. —Cumariche. Tree to ca. 20 m tall. Deciduous or semievergreen forests, 100–1100 m; Amazonas (near Puerto Ayacucho). Aragua, Guárico, Lara, Miranda, Trujillo, Yaracuy. ŠFig. 293. The only collection from the flora area (Morillo 6741, VEN) is somewhat disjunct from the rest of the range of Machaerium acuminatum. It has slightly smaller fruit and is tentatively assigned here.

Machaerium acutifolium Vogel, Linnaea 11: 187. 1837. Tree to 10 m tall. Venezuela, Brazil, Bolivia; 3 varieties, 1 in Venezuela, 1 in the flora area. M. acutifolium var. pseudacutifolium (Pittier) Rudd, Phytologia 25: 399. 1973. —Machaerium pseudacutifolium Pittier, Bol. Soc. Venez. Ci. Nat. 7: 148. 1941. —Tucurito. Semideciduous forests, savannas, 50–300 m; Bolívar (Represa Guri, Río Asa, Río

342

F ABACEAE

Paragua). Falcón, Guárico; northern Brazil, Bolivia. Machaerium affine Benth., Comm. Legum. Gen. 34. 1837. —Bainepá, Vainepá. Machaerium rectipes Pittier, Bol. Soc. Venez. Ci. Nat. 9: 121. 1944. Tree to ca. 17 m tall. Seasonally flooded savannas and gallery forests, 50–300 m; Bolívar (near La Paragua). Guyana. Machaerium altiscandens Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 75. 1925. Liana or tree to ca. 25 m tall. Evergreen lowland forests, 50–200 m; Bolívar (Río Parguaza). Suriname, Amazonian Brazil. The leaflets of this species show considerable variation in width. Machaerium amazonense Hoehne, Arq. Bot. Estado São Paulo, n.s. 1: 46, t. 56. 1939. High-climbing liana. Evergreen lowland to montane primary and secondary forests and savannas, 100–900 m; Bolívar (near Santa Elena de Uairén), Amazonas (slope of Río Cataniapo). Brazil (Amazonas). Machaerium aristulatum (Spruce ex Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 311. 1925. —Drepanocarpus aristulatus Spruce ex Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 69. 1860. —Robasesina. Tree, shrub, or liana, to ca. 10 m tall. Moist savannas, river banks, 50–200 m, Bolívar (near Ciudad Bolívar, San Rafael), Amazonas (Isla Ratón, Río Orinoco). Apure, Barinas; Colombia, Peru, northern Brazil, Bolivia. Machaerium dubium (H.B.K.) Rudd, Phytologia 24: 122. 1972. —Drepanocarpus dubius H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 390. 1823 [1824]. —Almendro. Machaerium caicarense Pittier, Bol. Soc. Venez. Ci. Nat. 9: 120. 1944. Tree to ca. 15 m tall. Seasonally flooded evergreen gallery forests, 50–100 m; Bolívar (Río Orinoco at Caicara). Apure, Guárico.

Machaerium ferox (Mart. ex Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 311. 1925. —Drepanocarpus ferox Mart. ex Benth., Comm. Legum. Gen. 32. 1837. —Bejuco de murciélago, Uña de gavilán. Drepanocarpus ferox ß macrophyllum Benth. in Mart., Fl. Bras. 15(1): 256. 1862. Liana, shrub, or small tree, to ca. 20 m tall. Flooded or nonflooded riparian forests, near sea level to 200 m; Delta Amacuro (Caño Guara near Isla Guara), Amazonas (above La Esmeralda, Río Orinoco near mouth of Río Atabapo, Río Pamoni). Colombia (Amazonas), Guyana, Suriname, Brazil. ŠFig. 294. Machaerium floribundum Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 68. 1860. —Chaperno, Sangre de toro, Uña de gavilán. Drepanocarpus venezuelensis Pittier, Contr. U.S. Natl. Herb. 20: 122, fig. 59. 1918. —Machaerium venezuelense (Pittier) Hoehne, Fl. Brasilica 25(3): 53. 1941. Machaerium decorticans Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 150. 1922. Machaerium woodworthii Standl., Contr. Arnold Arbor. 5: 81. 1933. Machaerium rosescens Standl., Publ. Carnegie Inst. Wash. 461: 24. 1935. Drepanocarpus ?ovalifolium Pittier, Bol. Soc. Venez. Ci. Nat. 7: 149. 1941. —Machaerium longistipitatum Pittier, Bol. Técn. Minist. Agric. 5: 119. 1944, based on Drepanocarpus ovalifolium Pittier, non Machaerium longistipitatum Hoehne 1939. Southern Mexico, Central America, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, northern Brazil, Bolivia; 3 varieties, all in Venezuela, 1 in the flora area. Variability in size and shape of leaflets seems to be the chief reason for the several synonyms. M. floribundum var. floribundum Tree, shrub, or liana to ca. 25 m tall. Moist forests, near sea level to 200 m; Delta Amacuro (Caño Araguao, Caño Daudacana),

Machaerium 343

Amazonas (between Cerro Yapacana and Santa Bárbara, upper Río Atacavi, Río Orinoco). Northern Venezuela; southern Mexico, Central America, Colombia, Guyana, French Guiana, Peru, northern Brazil. ŠFig. 295. Machaerium guaremalense Pittier, Arb. Legum., part 3, Trab. Mus. Comercial Venezuela 4 [reprinted from Bol. Minist. R.R. E.E. no. 4–7]: 213. 1928. Small tree or liana to ca. 5 m tall. Semideciduous to evergreen lowland forests, savannas, 50–300 m; Bolívar (island in Lago Guri, Río Orinoco east of mouth of Río Horeda and Cerro Gavilán). Anzoátegui, Carabobo, Portuguesa. This species is similar to and possibly synonymous with Machaerium darienensis Pittier, a species from Panama and is closely related to M. striatum J. Johnst. from Colombia and coastal Venezuela.

piare). Anzoátegui, Barinas; Mexico, Central America, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 298. Machaerium leiophyllum (DC.) Benth., Comm. Legum. Gen. 36. 1837. —Nissolia leiophylla DC., Prodr. 2: 258. 1825. Colombia, Venezuela, Guyana, French Guiana, Peru, Amazonian Brazil, Bolivia; 4 varieties, 2 in Venezuela, 1 of these in the flora area.

Machaerium inundatum (Mart. ex Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 311. 1925. —Drepanocarpus inundatus Mart. ex Benth., Comm. Legum. Gen. 32. 1837. —Jarizo, Urapo. Liana or tree to 20 m tall. Flooded forests along rivers, near sea level to 200 m; Delta Amacuro (Río Acure, Serranía de Imataca), northern Bolívar, Amazonas (widespread). Apure, Monagas, Guárico; Colombia, Guyana, Suriname, French Guiana, Peru, northern Brazil. ŠFig. 296.

M. leiophyllum var. latifolium (Benth.) Rudd, Phytologia 22: 56. 1971. —Drepanocarpus crista-castrensis var. latifolium Benth. in Mart., Fl. Bras. 15(1): 258. 1862. Drepanocarpus frondosus Mart. ex Benth., Comm. Legum. Gen. 32. 1827. —Machaerium frondosum (Mart. ex Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 151. 1922. Liana, sometimes a tree or shrub. River banks, flooded forests, 50–100 m; Bolívar (Río Horeda, Río Parguaza), Amazonas (mouth of Río Samariapo, mouth of Río Sipapo). Apure; Guyana, French Guiana, Amazonian Brazil. ŠFig. 297. Without fruit this variety is virtually indistinguishable from var. leiophyllum, which has much larger, less lunate fruit than var. latifolium.

Machaerium kegelii Meisn., Linnaea 21: 257. 1848. Machaerium bracteatum Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 65. 1860. Machaerium pachyphyllum Pittier, Contr. U.S. Natl. Herb. 20: 469. 1922. Machaerium steinbachianum, Hoehne, Arq. Bot. Estado São Paulo, n.s. 1: 48. 1939. Liana, shrub, or tree to ca. 20 m tall. Primary, semi-evergreen, and semideciduous forests, 200–1300 m; Delta Amacuro (southeast of Los Castillos de Guayana, southeast of Piacoa), Bolívar (Caño Pablo near Salto Pará, road between Tumeremo and Bochinche), Amazonas (Río Mana-

Machaerium lunatum (L. f.) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 310. 1925. —Pterocarpus lunatus L. f., Suppl. 317. 1781. —Drepanocarpus lunatus (L. f.) G. Mey., Prim. Fl. Esseq. 238. 1818. —Arepillo, Olvidanovia, Siete conchas. Shrub or small tree to ca. 8 m tall, sometimes scandent. Brackish coastal marshes, sandy soil, mangrove swamps, near sea level to 50 m; Delta Amacuro (widespread). Monagas, Sucre; Nicaragua, Costa Rica, Panama, Hispaniola, Puerto Rico, Lesser Antilles, coastal Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil, western coast of Africa from Senegal to Angola. ŠFig. 301.

344

F ABACEAE

Machaerium macrophyllum Benth., Comm. Legum. Gen. 35. 1837. Colombia, Venezuela, Guyana, Suriname, Peru, Brazil; 2 varieties, 1 in Venezuela. M. macrophyllum var. macrophyllum Liana, sometimes a tree, to ca. 18 m tall. Evergreen forests, 100–800 m; Bolívar (Caño Iguapo, Río Erebato, mouth of Río Nichare), Amazonas (Cucurital near middle Caño Yagua, Culebra, Mavaca, San Carlos de Río Negro, Sierra de la Neblina, Simarawochi, Yavita). Colombia (Amazonas), Guyana, Suriname, Peru, Brazil (Amazonas). Machaerium madeirense Pittier, Contr. U.S. Natl. Herb. 20: 119. 1918. Machaerium compressicaule Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 76. 1925. Machaerium compressicaule var. manoense Ducke, Arq. Inst. Biol. Veg. 2: 45. 1935. —Machaerium latifolium var. manoense (Ducke) Hoehne, Fl. Brasilica 25(3): 65. 1941. Liana, sometimes a tree to ca. 10 m tall. Gallery forests, evergreen lowland forests, 100–500 m; Bolívar (west side of Amaruaytepui, southwest of El Dorado, Río Acanán, Río Asa), Amazonas (below mouth of Caño Yapacana, Culebra, El Porvenir, Río Orinoco between Tamatama and Esmeralda). Southeastern Colombia, Guyana, Suriname, Peru, Amazonian Brazil, northern Bolivia. ŠFig. 302. Variation in leaflet shape seems to be chiefly responsible for the synonymy. Machaerium multifoliolatum Ducke, Bull. Mus. Hist. Nat. (Paris) sér. 2, 4: 734. 1932. Liana to ca. 30 m long. Evergreen lowland and lower montane forests, 100–1000 m; Amazonas (slopes of Cerro Duida, San Carlos de Río Negro). Colombia (Vaupés), Brazil (Amazonas), Bolivia. ŠFig. 299. Machaerium myrianthum Spruce ex. Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 59. 1860. Liana or scandent tree to ca. 10 m tall. Lowland to lower montane forests, 200–800 m; Bolívar (Altiplanicie de Nuria, Cerro Jaua, northwest of El Manteco, Santa Elena

de Uairén, Sierra de Lema). Colombia (Vaupés), Guyana, Suriname, northern Brazil. ŠFig. 300. Machaerium quinatum (Aubl.) Sandwith, Bull. Misc. Inform. Kew 1931: 359. 1931. —Nissolia quinata Aubl., Hist. Pl. Guiane 743, t. 4. 1775. Nissolia ferruginea Willd., Sp. Pl. 3(2): 900. 1802. —Machaerium ferrugineum (Willd.) Pers., Syn. Pl. 2: 276. 1807. Tree, shrub, or liana to ca. 16 m tall. Lowland to upland riparian forests, savannas, usually on sandy soil. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 varieties, both in the flora area. Key to the Varieties of M. quinatum 1. Flowers ca. 8–12 mm long; calyx 2.5–3 × 2– 3 mm; bracteoles 1–2 × 1.5–2.5 mm ................................. var. parviflorum 1. Flowers ca. 12–17 mm long; calyx 4–6 × 3–4 mm; bracteoles 2–3 × 3–4 mm ...................................... var. quinatum M. quinatum var. parviflorum (Benth.) Rudd, Phytologia 24: 121. 1972. —Machaerium ferrugineum ß parviflorum Benth. in Mart., Fl. Bras. 15(1): 253. 1862. —Acoi-yerí-yek (Arekuna), Chaparillo, Maripa-yen-ya-pupen-yek (Arekuna), Uña de murciélago. Machaerium nervosum Vogel, Linnaea 11: 186. 1837. 100–1200 m; Bolívar (Arabopó, Roraimatepui, base of Sororopán-tepui), Amazonas (San Carlos de Río Negro, Yavita). Southeastern Colombia, French Guiana, Brazil. ŠFig. 304. M. quinatum var. quinatum 100–900 m; Bolívar (Río Cuyuní, Río Parguaza, near Santa Elena de Uairén), Amazonas (Simarawochi). Guyana, Suriname, French Guiana, Brazil (Amazonas). ŠFig. 303. Machaerium robiniifolium (DC.) Vogel, Linnaea 11: 190. March 1837. —Nissola robiniifolia DC., Prodr. 2: 258. 1825. Machaerium sieberi Benth., Comm. Legum. Gen. 34. June 1837.

Machaerium 345

Machaerium eggersii Hoehne, Arq. Bot. Estado São Paulo, n.s. 1: 47, t. 58. 1939. Tree to ca. 20 m tall, with stipular spines. Semideciduous to evergreen lowland forests, 50–800 m; northern Bolívar. Widespread in northern Venezuela; Colombia, Trinidad, Guyana, Brazil (Rio Branco area). ŠFig. 292. There has been confusion with Machaerium moritzianum Benth. and many specimens so determined are actually M. robiniifolium. Machaerium tovarense Pittier, Contr. U.S. Natl. Herb. 20: 121. 1918. Liana. Semideciduous forests on hills, 200–300 m; Bolívar (island in Lago Guri 40

km south of Represa Guri). Aragua, Distrito Federal, Falcón, Miranda, Zulia. Machaerium sp. A Tree ca. 25 m tall. Montane riparian forests, 700–900 m; Amazonas (Sierra Parima along Río Matacuni). The single collection, Steyermark 107084 (US), previously annotated as Machaerium acuminatum, bears flower buds but it cannot be reliably identified. There appears to be some relationship to M. acuminatum, M. amazonense, and M. guaremalense. However, it does not compare well with any of those species and until better material is available it is better to maintain it as Machaerium sp. A.

Fig. 292. Machaerium robiniifolium

346

F ABACEAE

Fig. 293. Machaerium acuminatum

Fig. 294. Machaerium ferox

Machaerium 347

Fig. 295. Machaerium floribundum var. floribundum

348

F ABACEAE

Fig. 296. Machaerium inundatum

Fig. 297. Machaerium leiophyllum var. latifolium

Machaerium 349

Fig. 298. Machaerium kegelii

Fig. 299. Machaerium multifoliolatum

350

F ABACEAE

Fig. 300. Machaerium myrianthum

Fig. 301. Machaerium lunatum

Machaerium 351

Fig. 302. Machaerium madeirense

352

F ABACEAE

Fig. 303. Machaerium quinatum var. quinatum

Fig. 304. Machaerium quinatum var. parviflorum

Macroptilium 353

38. MACROPTILIUM (Benth.) Urb., Symb. Antill. 9: 457. 1928. —Phaseolus sect. Macroptilium Benth. in Mart., Fl. Bras. 15(1): 189. 1859. by Gerardo A. Aymard C. Vines or sometimes erect or sprawling herbs, sometimes perennial from a thick rootstock. Leaves pinnate, trifoliolate or rarely unifoliolate; stipules veined, not prolonged below the insertion; stipels ciliate to tomentose; leaflets glabrous or pubescent, lacking hooked trichomes. Inflorescences with stiff, elongate peduncles, the flowers mostly congested at the apex; bracteoles narrow, at least distally, veined, caducous; rachis with small swellings at the nodes, lacking extrafloral nectaries; pedicels equaling or shorter than the calyx. Flowers purplish, violet, or white, the interior often vivid. Calyx campanulate, the teeth free, the upper 2 teeth sometimes reduced; standard orbicular, emarginate, with 2 small basal auricles, lacking medial thickenings; wing petals longer than the standard and the keel petals, long-stipitate; keel petals apically spiraled, basally adnate to the staminal tube. Style apically recurved and thickened, caducous. Fruit linear, turgid or compressed, nonseptate. Seeds numerous, small, with a short hilum. Pantropics, cultivated in Africa and Asia as forage; ca. 20 species, 6 species in Venezuela, 4 of these in the flora area. Key to the Species of Macroptilium 1. 1. 2(1). 2. 3(2). 3.

Leaves unifoliolate ............................................................... M. monophyllum Leaves trifoliolate ....................................................................................... 2 Leaflets linear, 2–7 mm wide ........................................................... M. gracile Leaflets ovate, elliptical, rhombic, oblong, or linear, 1–4 cm wide .......... 3 Stipels glabrous, ca. 3 mm long; stipules lanceolate, striate; calyx with lanceolate teeth; fruit 7–12 cm long .......................................... M. lathyroides Stipels long-ciliate, ca. 1 mm long; stipules lanceolate-subulate, veined; calyx with deltoid teeth; fruit 3–6 cm long ............. M. longepedunculatum

Macroptilium gracile (Poepp. ex Benth.) Urb, Symb. Antill. 9: 457. 1928. —Phaseolus gracilis Poepp. ex Benth., Comm. Legum. Gen. 77. 1837. Perennial herb or vine; flowers red, purple, or pink. Savannas, edges of towns, 50–400 m; Delta Amacuro (between Los Castillos and Piacoa), Bolívar (Altiplanicie de Nuria, Caicara, Ciudad Bolívar, La Paragua, La Urbana, lower Río Caroní), Amazonas (Puerto Ayacucho, Río Parucito). Anzoátegui, Apure, Carabobo, Distrito Federal, Monagas, Portuguesa; Mexico, Central America, Antilles, Colombia, Guyana, Brazil (Amapá, Maranhão, Roraima), Argentina. ŠFig. 307. Macroptilium lathyroides (L.) Urb., Symb. Antill. 9: 457. 1928. —Phaseolus lathyroides L., Sp. Pl. ed. 2, 2: 1018. 1762 [1763].

Neotropics, cultivated as forage in Asia; 2 varieties, 1 in Venezuela. M. lathyroides var. lathyroides Erect herb; flowers purple-red to black. Savannas, edges of towns, 50–200 m; Delta Amacuro (Tucupita to La Horqueta road), Bolívar (Ciudad Bolívar, El Palmar), Amazonas (Puerto Ayacucho, Río Manapiare). Barinas, Guárico, Miranda, Portuguesa; Mexico, Central America, Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 306. Macroptilium longepedunculatum (Mart. ex Benth.) Urb., Symb. Antill. 9: 457. 1928. —Phaseolus longepedunculatus Mart. ex Benth., Comm. Legum. Gen. 77. 1837. Phaseolus campestris Mart. ex Benth., Comm. Legum. Gen. 77. 1837.

354

F ABACEAE

Fig. 306. Macroptilium lathyroides var. lathyroides Fig. 305. Macroptilium monophyllum Fig. 307. Macroptilium gracile

Monopteryx 355

Perennial vine; flowers red to violet. Disturbed places, Trachypogon savannas, 50–500 m; Bolívar (Altiplanicie de Nuria, Ciudad Bolívar, middle Río Orinoco), Amazonas (Laguna El Sillón 78 km northeast of Puerto Ayacucho, lower Río Ventuari). Lara, Portuguesa; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amapá, Goiás, Mato Grosso), Argentina.

Macroptilium monophyllum (Benth.) Maréchal & Baudet, Bull. Jard. Bot. Belg. 47: 257. 1977. —Phaseolus monophyllus Benth., Comm. Legum. Gen. 76. 1837. Perennial herb or vine; flowers yellow to orange. Savannas, 50–200 m; Bolívar (Maripa, Río Ore in Río Parguaza basin), Amazonas (Maypures, Puerto Ayacucho, Rincones de Chachorro 30 km north of Puerto Ayacucho). Monagas; Colombia, Brazil, Paraguay. ŠFig. 305.

39. MONOPTERYX Spruce ex Benth. in Mart., Fl. Bras. 15(1): 307. 1862. by Charles H. Stirton and Gerardo A. Aymard C. Large trees to 50 m tall. Stipules small, elliptic-obovate, caducous. Leaves 3– 11-foliolate, odd-pinnate. Leaflets coriaceous, inserted opposite or alternate, small and many or few and large. Inflorescences terminal panicles or racemose in upper axils. Flowers pink, bisexual, zygomorphic, pedicellate, bracteolate, bracteate. Calyx lobes united in 2 lips, the larger covering the flower in bud and the lower small with 3 very short teeth; petals sessile; standard obovate to suborbiculate; keel petals subfalcate-oblong, fused along base and at apex, longer than the free wing petals. Stamens subequal, kinked, free; anthers linear-oblong. Ovary stipitate, glabrous or sparsely pilose, 1-ovulate; style short conical, arcuate, stigma laterally introrse. Fruit flattened, arcuate, tapering at both ends, finely transversely ridged, with apex acute. Colombia, Venezuela, French Guiana, Brazil; 4 species, all in the flora area. Key to the Species of Monopteryx 1.

1.

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

Rachis of the leaves canaliculate, ± winged at the base; petiolules 0.2–2 mm long; leaflets oblong, obovate, or obovate-elliptic, drying brownish on the lower surface, apex rounded; fruit falcate ................................. 2 Rachis of the leaves not canaliculate, not subwinged at the base; petiolules > 4 mm long; leaflets ovate, elliptic, ovate-lanceolate, not drying brownish on the lower surface, apex acuminate; fruit not falcate ....... 3 Leaves 3- or 5-foliolate; terminal leaflets 3–4 cm wide ..................... M. sp. A Leaves 7–15-foliolate; terminal leaflets 1–2 cm wide ....................... M. inpae Leaflets glaucous, densely minutely tomentulose and not reticulate on the lower surface; calyx 9–12 mm long .............................. M. angustifolia Leaflets glabrous, not glaucous, strongly reticulate on the lower surface; calyx 6–9 mm long ........................................................................ M. uaucu

Monopteryx angustifolia Spruce ex Benth. in Mart., Fl. Bras. 15(1): 307, t. 122. 1862. —Barbasco. Tree 20–30 m tall, with buttressed trunks and hard wood; leaves (5)7- or 9-foliolate, ovate-lanceolate, coriaceous, the lower surface densely minutely tomentulose, the upper surface nitid; petals dark pink; fruit

woody, flat, dehiscent. Nonflooded evergreen lowland forests, 50–200 m; Amazonas (Río Atabapo, Yavita). Colombia, Brazil (Amazonas: Rio Negro). ŠFig. 308. Monopteryx inpae W.A. Rodrigues, Acta Amazon. 5: 153, fig. 1. 1975. Tree to 35 m tall; lower surface of leaflets

356

F ABACEAE

Fig. 308. Monopteryx angustifolia

drying brownish; fruits narrowly falcate, with a winged border on both sides of legume. Evergreen lowland forests on sandy soil, 100–200 m; Amazonas (Río Mawarinuma). French Guiana, Brazil (Amapá, Amazonas). Monopteryx uaucu Spruce ex Benth. in Mart., Fl. Bras. 15(1): 308. 1862. —Guaco. Tree to 40 m tall, with buttressed trunks. Evergreen lowland forests on ultisols, 50– 200 m; Amazonas (San Carlos de Río Negro to Solano, Yavita to Maroa). Colombia, Brazil (Amazonas). Monopteryx sp. A Tree ca. 15 m tall; fruits dark brown, falcate, finely transversely ridged near maturity. Lower montane forests, 200–300 m; Bolívar (Minas de Manaima in middle Río Paragua). Endemic. The only specimen of this species, Stergios 10259 (MO, PORT), is in fruit only. It appears most closely related to Monopteryx inpae.

Mucuna 357

40. MUCUNA Adans., Fam. Pl. 2: 325. 1763, nom. cons. Stizolobium P. Browne, Civ. Nat. Hist. Jamaica 290. 1756. by Gerardo A. Aymard C. Herbs to climbing woody vines. Leaves alternate, pinnately 3-foliolate; lateral leaflets oblique; stipules and often stipels deciduous. Inflorescences axillary on leafy shoots or on old branches, subumbellate, condensed-paniculate, or falsely racemose through reduction of lateral branches, mostly pendent; bracts and bracteoles deciduous, often subfoliaceus, enclosing the bud. Flowers showy, purple, reddish, orange, greenish, yellow, or white. Calyx campanulate, 4- or 5-toothed, the upper teeth connate and forming an entire or bifid lip, the lower 3 usually unequal, often with irritating trichomes; standard rounded with median claw and pair of inflexed lateral auricles at base, usually much shorter than other petals; wing and keel petals narrowed into basal claw with small dorsal auricle, margins basally ciliate, the keel petals partially connate along lower margin, usually horny, apically falcate and indurate. Stamens 10, diadelphous, the filaments alternately thick and thin, long and short, the vexillar stamen free; anthers barbate. Ovary tomentose, the short stipe surrounded by a glandular disk; style slender, glabrous or pubescent; stigma small, capitate, sometimes with a tuft of trichomes. Fruit ovoid, oblong, or linear, thick or flattened, the margins often winged, undulate between the seeds and ± compressed laterally between the seeds, the surface sometimes layered with parallel or irregularly raised thin plates which may form elongated outgrowths or 1 or 2 transverse ribs, usually densely covered with stiff irritating trichomes, usually dehiscent, occasionally indehiscent. Seeds globose or oblong, the hilum narrow, encircling more than 1/2 of the periphery. Pantropics; 120 species, 5 in Venezuela, 3 of these in the flora area. Key to the Species of Mucuna 1.

1. 2(1). 2.

Fruit 4–9 × 1–1.5 cm, without prominent transverse thin plates, but with 2 horizontal ribs on each valve; flowers purple with white tips ............ ................................................................................................... M. pruriens Fruit ≥ 10 × 2–7 cm, with prominent transverse thin plates; flowers reddish orange, yellow-green, or pale greenish white ............................... 2 Flowers 6–9 cm long, reddish orange; inflorescences 6–13(–15) cm long; peduncle 1–1.2 cm long; anthers pilose .................................... M. rostrata Flowers ≤ 5 cm long, pale greenish white or yellow-green; inflorescences > 15 cm long; peduncle > 15 cm long; anthers basally barbate ....... M. urens

Mucuna pruriens (L.) DC., Prodr. 2: 405. 1825. —Dolichos pruriens L., Herb. Amb. 23. 1754. —Stizolobium pruriens (L.) Medik., Vorles. Churpfälz. Phys.Öcon. Ges. 2: 399. 1787. —Caraota, Nescafé, Pica Pica, Toddy. Paleotropics, introduced in the Neotropics for food and forage; 3 varieties, 2 in Venezuela, 1 of these in the flora area.

M. pruriens var. utilis (Wall. ex Wight) Baker ex Burck, Ann. Jard. Bot. Buitenzorg 11: 187. 1893. —Mucuna utilis Wall. ex Wight, Icon. Pl. Ind. Orient. 1: 280. 1840. Stizolobium aterrimun Piper & Tracy, U.S.D.A. Bur. Pl. Industr. Bull. 179: 18. 1910. Stizolobium hassjo Piper & Tracy,

358

F ABACEAE

U.S.D.A. Bur. Pl. Industr. Bull. 179: 17. 1910. —Mucuna hassjoo (Piper & Tracy) Mansf., Kulturpflanze 7: 204. 1959. Vine, occasionally high-climbing; flowers purple. Cultivated or escaped on disturbed ground, 50–200 m; Bolívar (near El Dorado, Tumeremo), Amazonas (Isla Ratón). Widespread elsewhere in Venezuela; Mexico, Central America, West Indies, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia. ŠFig. 309. Mucuna rostrata Benth. in Mart., Fl. Bras. 15(1): 171. 1859. —Guaraguao, Ojo de burro, Ojo de zamuro, Jijije. High-climbing woody vine; flowers orange to red. Evergreen lowland to lower montane flooded or nonflooded forests, near sea level to 500 m; Delta Amacuro (Caño Arature, Caño Curiapo, Caño Guará, La Horqueta, Punta Barima), Bolívar (base of Auyántepui, Río Uruyen). Widespread elsewhere in Venezuela; Guatemala, Belize, Honduras, Panama, Colombia, French Guiana, Ecuador, Peru, Brazil (Amazonas, Pará), Bolivia. ŠFig. 310.

Mucuna urens (L.) Medik., Vorles. Churpfälz. Phys.-Öcon. Ges. 2: 399. 1787. —Dolichos urens L., Syst. Nat. ed. 10, 2: 1162. 1759. —Bejuco de zamuro, Calabazín, Jeamo, Ojo de zamuro, Pepa de zamuro. Dolichos altissmus Jacq., Enum. Syst. Pl. 27. 1760. —Stizolobium altissimum (Jacq.) Pers., Syn. Pl. 299. 1807. —Mucuna altissima (Jacq.) DC., Prod. 2: 405. 1825. High-climbing vine; flowers pale greenish white with dull purple. Humid or gallery forests, 50–200 m; Delta Amacuro (Caño Araguabisi between Caño Araguao and Caño Güiniquina, Caño Joba-Suburu east of Caño Sacupana), Bolívar (widespread), Amazonas (Caño Yureba, near La Esmeralda, Río Casiquiare, Río Mawarinuma, Río Padamo, Río Parú). Widespread elsewhere in Venezuela; Mexico, Guatemala, Costa Rica, Panama, West Indies, Suriname, Peru, Brazil, Argentina. ŠFig. 311.

Fig. 309. Mucuna pruriens var. utilis

Mucuna 359

Fig. 310. Mucuna rostrata

Fig. 311. Mucuna urens

360

F ABACEAE

41. MUELLERA L. f., Suppl. Pl. 53, 329. 1781 [1782], nom. cons. Coublandia Aubl., Hist. Pl. Guiane 937. 1775. by Nidia L. Cuello A. Shrubs or small trees, branches glabrous to sparsely pilose. Leaves alternate, odd-pinnate; leaflets 5 or 7, subopposite, upper surfaces glabrous, lower surfaces puberulent; stipules minute; stipels lacking. Inflorescences in axillary or lateral racemes; bracts and bracteoles small, inconspicuous, caducous. Flowers in groups on short peduncles along an axillary rachis, violet or white. Calyx campanulate, truncate, shortly 5-dentate or with very short or obsolete teeth; petals clawed, glabrous; standard broadly ovate or suborbicular, auricle callosities absent; wing petals falcate-oblong, auriculate, slightly adherent to the keel petals; keel petals incurved, obtuse, shorter than the wings. Stamens 10, monadelphous, connate into a tube closed at both sides, vexillar stamen free at the base but connate from the middle with the others into a closed tube; anthers versatile. Ovary subsessile; ovules numerous; style filiform, incurved; stigma, small, terminal. Fruit thick, fleshy, hard, indehiscent, subterete, constricted between the seeds and moniliform, or by abortion 1-seeded and subglobose. Seeds ovoid-subglobose, slightly compressed, with a lateral hilum. Mexico, Central America, Antilles, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Argentina; 2 species, 1 in Venezuela.

Fig. 312. Muellera frutescens

Myrocarpus

Muellera frutescens (Aubl.) Standl., Tropical Woods 34: 41. 1933. —Coublandia frutescens Aubl., Hist. Pl. Guiane 937, pl. 356. 1775. Muellera moniliformis L. f., Suppl. Pl. 59, 329. 1781 [1782]. Tree 2–7 m tall; flowers purple. Flooded and wet forests, mangrove swamps, near sea

361

level to 50 m; Delta Amacuro (Angosturita, Caño Araguao, Caño Bagre, Caño JobaSuburu east of Caño Sacupana, Pedernales). Sucre; Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil (Amapá, Maranhão, Pará). ŠFig. 312.

42. MYROCARPUS Allemão, Pl. Novas. Brasil pl. 5. post 26 Oct. 1847. by Gerardo A. Aymard C. Large trees, with hard wood and resinous bark. Leaves odd-pinnate, (1–)3–10foliolate, alternate; leaflets chartaceous, alternate, with pellucid dots; stipules small, stipels lacking. Inflorescences axillary or pseudoterminal, many-flowered spicate racemes (almost catkin-like); bracts minute, deltoid. Flowers small, subactinomorphic, pedicellate. Calyx turbinate-campanulate, with 5 short, subequal lobes, the vexillar lobes sometimes connate; petals (3–)5, free, whitish, imbricate, subequal, linear, clawed. Stamens 6, 8, or 10; filaments free, subequal, mostly exceeding the petals; anthers subcordiform, basifixed, dehiscent by lateral slits. Ovary shortly stipitate, usually 3–5-ovulate; style short, inflexed; stigma minute, terminal. Fruits elongate, laterally compressed, samaroid with marginal wings, resinous. Seeds 1–5, oblong or compressed-fusiform, the hilum apical. Venezuela, Brazil, Bolivia, Paraguay, Argentina; 4 species, 1 in Venezuela. Myrocarpus venezuelensis Rudd, Phytologia 23: 404. 1972. —Cereipo, Kamakari (Yekwana), Nosamo de casa (Yekwana). Large tree to 20 m tall; outer bark corky, with strong lenticels; fruits elongate, ca. 9 × 2 cm. Evergreen lowland forests, 50–400 m;

Bolívar (mouth of Río Nichare), Amazonas (Río Oinoquito in upper Río Ofinoco basin). Apure (San Camilo). ŠFig. 313. This species has purple heartwood and is extremely resistant to termites; it is used locally in house construction (center posts) and in wood carving.

Fig. 313. Myrocarpus venezuelensis

362

F ABACEAE

43. MYROSPERMUM Jacq., Enum. Syst. Pl. 4: 20. 1760. by Charles H. Stirton Large shrubs or small trees. Stipules minute or lacking. Leaves 10–25foliolate, odd-pinnate, alternate, deciduous. Leaflets chartaceous, alternate to subopposite, with pellucid dot and linear balsam glands. Inflorescences axillary or terminal racemes. Flowers white, zygomorphic, bracteate, bracteolate. Calyx turbinate-campanulate with 5 subequal lobes, valvate in bud; petals 5, unequal, free, glabrous, clawed; standard white or yellow with a nectar patch, elliptic to obovate; wing petals smaller and narrower than keel petals. Stamens 10, free, persistent; anthers

Fig. 314. Myrospermum frutescens

Myroxylon 363

equal, uniform, ovoid or ellipsoid, dorsifixed on elongated filaments. Ovary stipitate, 5–7-ovulate, pubescent or glabrous; style minute or distinct; stigma small, terminal, capitate. Fruits stipitate, compressed, indehiscent, samaroid (apically 1-seeded) and distinctly winged, or oblong (1–3-seeded) and scarcely winged. Seeds oblong-reniform, the hilum subapical, elliptic. Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Puerto Rico, Colombia, Trinidad, Venezuela; 3 species, 1 in Venezuela. Myrospermum frutescens Jacq., Enum. Syst. Pl. 4: 20. 1760. —Cereipo. Large shrub or small tree to 12 m tall; leaves 10–21-foliolate, oblong to elliptic, secondary veins inconspicuous; fruits flat, winged, indehiscent. Deciduous forests and shrublands, granitic outcrops, 100–300 m; Bolívar (50 km southwest of Caicara in Serranía La Encaramada, islands in Lago Guri, Puerto Ordaz). Anzoátegui, Falcón, Guárico, Portuguesa, Yaracuy; southern

Mexico, Central America, Colombia. ŠFig. 314. Myrospermum frutescens is the emblem species of Anzoátegui state. It is often confused with Myrospermum secundum Klotzsch, which is found in the West Indies, north-central Venezuela, and Trinidad, but this is a much larger fruited species. Myrospermum frutescens has also been confused with the genus Myroxylon, which has similar fruits but quite different flowers.

44. MYROXYLON L. f., Suppl. Pl. 34, 233. 1781., nom. cons. —Myrospermum sect. Myroxylon (L. f.) DC., Prodr. 2: 95. 1825. Toluifera L., Sp. Pl. 384. 1753. by Charles H. Stirton Trees to 40 m tall. Stipules minute or lacking. Leaves 5–15-foliolate, odd-pinnate, alternate, deciduous. Leaflets chartaceous, alternate, lanceolate to elliptic, acute to acuminate, with pellucid glandular dots and lines, margins undulate. Inflorescences axillary or in terminal racemes. Flowers white, bisexual, zygomorphic;

Fig. 315. Myroxylon balsamum

364

F ABACEAE

bracteate and bracteolate, caducous. Calyx turbinate-campanulate, with 5 subequal lobes, valvate in bud; petals 5, free, unequal, the standard much larger, the other 4 ± equal. Stamens 10, free, persistent, exserted; anthers uniform oblong, acuminate, sagittate, dorsifixed on short filaments. Ovary stipitate, 2-ovulate near apex, smooth; style short, subulate; stigma terminal, minute. Fruits stipitate, compressed, indehiscent, samaroid (apically 1-seeded), distinctly winged. Seeds reniform, resinous, the hilum subapical, elliptic. Mexico, Central America, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil; 2 variable species, 1 in Venezuela. Myroxylon balsamum (L.) Harms, Notizbl. Königl. Bot. Gart. Berlin 5: 94. 1908. —Toluifera balsamum L., Sp. Pl. 384. 1753. —Kamabadek (Pemón). Large tree with buttressed trunks; leaves 5–15-foliolate, elliptic, one half usually broader than the other, arcuate; young branchlets with white lenticels. Cultivated in the flora area, and possibly escaping. Carabobo, Miranda, Yaracuy; Mexico, Cen-

tral America, Colombia, Suriname, Ecuador, Peru, Bolivia. ŠFig. 315. This species has been used for hundreds of years as incense and as a medicinal plant. It is still used in the perfume trade and as a medicine in other countries. A sterile specimen from Río Paragua 13 km north of Karún, A. Fernández 4618 (MO, PORT), appears to be this species.

45. ORMOSIA Jack., Trans. Linn. Soc. London 10: 358, t. 25. 1811. nom. cons. by Charles H. Stirton and Gerardo A. Aymard C. Shrubs to very tall trees. Clusters of pearl bodies usually present in axils of leaflets, pulvinus, bracts, bracteoles or inflorescence branches, few to many. Leaves 1–21-foliolate, odd-pinnate, alternate, persistent, stipellate but early caducous; leaflets chartaceous or coriaceous, alternate to opposite; stipules small or lacking, deltoid to linear, caducous. Inflorescences axillary or terminal racemes. Flowers from white, yellow, pink, mauve, or blackish purple, with variously colored nectar patches, bisexual, zygomorphic, bracteate, bracteolate. Calyx campanulate, hypanthoid, with equal or subequal teeth or lobes, vexillar teeth fused for half their length and with a convex sinus; petals 5, unequal, free, glabrous, clawed; standard suborbicular or broadly ovate, emarginate, darker in color than other petals, broadly ovate to obovate; wing petals oblique, obovate-oblong, longer and broader than keel petals, petal sculpturing arching, transcostal; keel petals often imbricate along lower margin. Stamens 10, free, subequal in 2 or 3 ranks; filaments all thin or variously thickened at the base, inserted at the apex of the hypanthium; anthers equal, dorsifixed. Ovary sessile to stipitate, 2–7-ovulate, pubescent; style elongate, recurved; stigma introrse, bilobed. Fruits usually dehiscent, a few indehiscent, glabrous to densely velutinous, compressed to turgid, with or without septa, oblong to elongate, apex usually acute. Seeds ellipsoid, globose, or lenticular, 1–6, monochromatic (red, scarlet, yellow, black) or dichromatic (red and black or orange and black), usually polymorphic, the hilum terminal or subterminal, linear or elliptic. Neotropics, southeast Asia to northeast Australia; 145 species, 23 in Venezuela, 18 of these in the flora area. A collection (Gentry & Stein 46858, MO) from Río Mawarinuma, Amazonas state, may represent a new species, but fruits and flowers are needed to determine this with certainty.

Ormosia 365

Key to the Species of Ormosia 1.

Lower surface of leaflets sparsely minutely appressed-pubescent, glabrescent, or glabrous ..................................................................................... 2 1. Lower surface of leaflets densely sericeous, tomentose, or velutinous .............................................................................................................. 10 2(1). Apex of leaflets conspicuously acuminate; secondary veins on lower surface inconspicuous or scarcely raised; fruits black; seeds bright red ............................................................................................... O. grandiflora 2. Apex of leaflets acute, obtuse, shortly acuminate, emarginate, or truncate; secondary veins on lower surface slightly to conspicuously raised; fruits vermillion, dark brown, or chestnut brown; seeds dull red or red and black ................................................................................................ 3 3(2). Apex of leaflets retuse or emarginate; fruits 3–4 × 1.5–2 cm, not woody ................................................................................................................ 4 3. Apex of leaflets acute, obtuse, or shortly acuminate; fruits > 5 × 3 cm, woody ...................................................................................................... 7 4(3). Leaflets ovate, oblong, lanceolate, or ovate-lanceolate; fruits indehiscent or very tardily dehiscent, broadly beaked, fulvous-tomentose .............. ................................................................................................. O. williamsii 4. Leaflets elliptic to obovate; fruits dehiscent, acutely beaked, glabrate ................................................................................................................ 5 5(4). Upper surface of leaflets smooth; valves of fruit ± woody or coriaceous, 1– 2 mm thick ................................................................................ O. costulata 5. Upper surface of leaflets rugose; fruit woody, 2–7 mm thick ................... 6 6(5). Leaflets with secondary veins 3–10 mm apart; inflorescences ferruginoustomentulose; beak of fruit straight ........................................... O. coccinea 6. Leaflets with secondary veins 10–25 mm apart; inflorescences fulvoustomentulose; beak of fruit arcuate ....................................... O. subsimplex 7(3). Leaflets elliptic-oblong, lower surface with slightly raised secondary veins, upper surface with major veins flush with the surface; flowers 9– 10 mm long ............................................................................... O. paraensis 7. Leaflets ovate to ovate-oblong, elliptic, or obovate, lower surface with distinctly raised secondary veins, upper surface with major veins deeply impressed on the surface; flowers 10–25 mm long ............................... 8 8(7). Leaflets with secondary veins 3–10 mm apart, midvein on lower surface tomentulose; inflorescences ferruginous-tomentulose; beak of fruit straight ....................................................................................... O. coccinea 8. Leaflets with secondary veins 10–25 mm apart, midvein on lower surface puberulent or appressed-pubescent; inflorescences fulvous-tomentulose; beak of fruit arcuate .................................................................. 9 9(8). Leaflets rigid-coriaceous, elliptic to obovate; bracts deltoid or linear-deltoid, 2–3.5 mm long; calyx ferrugineos-tomentulose, 7–9 mm long ............................................................................................... O. subsimplex 9. Leaflets subcoriaceous to coriaceous, ovate to ovate-oblong; bracts linear, 3–10 mm long; calyx gray-pubescent, 10–15 mm long ........ O. macrocalyx 10(1). Lower surface of leaflets aureo-velutinous ............................................. 11 10. Lower surface of leaflets sericeous or tomentose .................................... 14

366

F ABACEAE

11(10). Secondary veins of leaflets terminating in a series of prominent arches forming a distinct intramarginal vein ............................... O. macrophylla 11. Secondary veins of leaflets upturned and gradually diminishing apically inside the margin, connected to superadjacent secondaries by a series of cross veins without forming a distinct intramarginal vein ............ 12 12(11). Flowers 10–14 mm long, calyx 3–8 mm long; fruits obliquely elliptic or oblong, sutures not ridged; seeds 1(2) ........................................ O. discolor 12. Flowers 15–25 mm long, calyx 10–15 mm long; fruits moniliform, sutures distinctly ridged; seeds 2–6 ................................................................. 13 13(12). Lower surface of leaflets golden yellow, the secondary and tertiary venation prominent; fruits rugose, dehiscent along both sutures, the valves recurved ....................................................................................... O. nobilis 13. Lower surface of leaflets silvery gray, the secondary and tertiary venation weakly developed; fruits smooth, tardily dehiscent along lower suture, the valves not recurved ....................................................... O. bolivarensis 14(10). Leaflets tomentose, secondary veins, almost parallel ............................ 15 14. Leaflets sericeous, secondary veins arcuate ........................................... 17 15(14). Leaflets elliptic or elliptic-oblong; fruits woody, 7–8 × 2.5–4 cm, to 25 mm thick; seeds scarlet or scarlet and black ............................... O. lignivalvis 15. Leaflets elliptic-ovate, ovate, or obovate; fruits subwoody, 2–5 × 1–1.5 cm, to 1.1 mm thick .................................................................................... 16 16(15). Young stems fulvo-velutinous; leaf base obtuse, secondary veins 18– 25; calyx externally and fruit velutinous.. ....................... O. maguireorum 16. Young stems cinerous- to fulvous-tomentulose; leaf base rounded to cordate, secondary veins 10–15; calyx externally fulvous- to ferruginoustomentulose; fruit velutinous essentially glabrous .............. O. amazonica 17(14). Fruits chartaceous, finely brown-velutinous, 15–25 × 11 mm, 1–2 mm thick ..................................................................................... O. steyermarkii 17. Fruits woody, fulvous-velutinous, 25–40 × 15–25 mm, 2–5 mm thick ... 18 18(17). Leaflets ovate to elliptic, the apex acuminate to attenuate .............. O. sp. A 18. Leaflets oblong-elliptic or obovate, the apex acute to shortly acuminate .............................................................................................................. 19 19(18). Trees to 30 m tall, pubescence tawny .......................................... O. coarctata 19. Shrubs to 3 m tall, pubescence ferruginous ....................................... O. sp. B Ormosia amazonica Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 139. 1922. Ormosia euneura Harms, Notizbl. Bot. Gard. Berlin-Dahlem 9: 972. 1926. Tree to 20 m tall; leaves 7–11-foliolate, leaflets coriaceous, the lower surface finely and tightly crisp-pubescent. Lowland wet forests, 200–300 m; Amazonas (Río Mawarinuma). Colombia, Ecuador, Peru, Brazil. Ormosia bolivarensis (Rudd) Stirton, comb. nov. —Ormosia nobilis var. bolivarensis Rudd, Contr. U.S. Natl. Herb. 32: 345. 1965. —Guanacoco, Wanaca-có (Yekwana).

Tree to 10–25 m tall; leaves 5–9-foliolate, leaflets elliptic-oblong, coriaceous, the lower surface minutely fulvous-sericeous. Whitesand and forested hills, 400–900 m; Bolívar (Cerro Camarón, Cerro Ichún, Río Canaracuni, upper Río Caura, Río Curutu, Río Icabarú, Río Salto) Guyana, Brazil. ŠFig. 316. Ormosia bolivarensis is distinguished from O. nobilis by its smaller, pale orange to orange-yellow seeds and distinctive fulvoussericeous lower surfaces of leaves. Ormosia coarctata Jacks., Trans. Linn. Soc. London 10: 363, t. 27. 1811. —Pericoca.

Ormosia 367

Ormosia cuneata Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 64. 1925. Tree to 30 m tall; leaves 5–11-foliolate, leaflets obovate, coriaceous, the lower surface densely pubescent with laxly crispate to subpatent trichomes. Humid forests, 200– 300 m; Bolívar (53 km northeast of Los Rosos, Reserva Forestal Imataca, Río Botanamo, Río Cuyuní). Mérida; Trinidad, Guyana, Suriname, French Guiana, Brazil, Bolivia. Ormosia coarctata is little collected in the region. Flowering material is needed from Venezuela. Ormosia coccinea (Aubl.) Jacks., Trans. Linn. Soc. London 10: 360, t. 25, 1811. —Robinia coccinea Aubl., Hist. Pl. Guiane 773. 1775. —Gateado, Macure, Palo macure, Peonía, Peonío, Pionilla, Pionina, Tento, Too’o (Yanomami), Ya’u balé. Large shrub or tree to 30 m tall; leaves 9 (–14)-foliolate; leaflets oblong, coriacious, large. Forest margins and islands, shrubby white-sand savannas, 50–200 m; Bolívar (Piedra Marimare in middle Río Orinoco), Amazonas (widespread). Apure, Monagas; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas). Ormosia costulata (Miq.) Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 392. 1925. —Leptolobium costulatum Miq., Stirp. Surinam. Select. 17. 1850 [1851]. —Peonía. Tree to 13 m tall; leaves 1–7-foliolate; leaflets obovate to elliptic, coriaceous, tertiary veins scarcely visible. Sandy savanna areas; 50–400 m; Bolívar (Río Asa, Río Paragua). Guyana, Suriname, French Guiana, Brazil (Amazonas). ŠFig. 318. Ormosia discolor Spruce ex Benth. in Mart., Fl. Bras. 15(1): 318. 1862. —Peonía. Ormosia micrantha Ducke, Arq. Inst. Biol. Veg. 4: 21. 1938. Tree to 19 m tall; leaves (3)5- or 7foliolate, ovate to oblong, coriaceous, lower surface fulvous-sericeous; flowers small. Nonflooded evergreen lowland forests, 100– 200 m; Amazonas (Río Casiquiare, Río Ne-

gro, Río Pasimoni, Río Yatúa). Amazon basin of Colombia and Brazil. Ormosia grandiflora (Tul.) Rudd, Contr. U.S. Natl. Herb. 32: 301. 1965. —Diplotropis grandiflora Tul., Arch. Mus. Hist. Nat. 4: 109. 1844. Tree or shrub to 10 m tall; leaves (1)3- or 5-foliolate; leaflets ovate, elliptic, or obovate, subcoriaceous, glabrescent, shiny. Evergreen lowland forests, ca. 300 m; Amazonas (Río Sipapo). Costa Rica, Ecuador, Peru, Brazil, Bolivia. Ormosia lignivalvis Rudd, Contr. U.S. Natl. Herb. 32: 331. 1965. —Chaparillo, Kaguaitariyek (Arekuna), Pericosa, Wanaekoko (Yekwana). Tree to 50 m tall; leaves 5–11-foliolate; leaflets elliptic or oblong, coriaceous, venation well developed. Forest margins, 200– 500 m; Bolívar (El Dorado, Río Asa, Río Paragua at mouth of Río Karún, Río Tabaro, Santa María de Erebato, Serranía Pia-Zoi, east-southeast of Villa Lola). Guyana, French Guiana, Peru, Brazil (Amazonas). Ormosia macrocalyx Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 137. 1922. Ormosia toledana Standl., Publ. Carnegie Inst. Wash. 461: 64. 1935. Ormosia chlorocalyx Ducke, Bol. Técn. Inst. Agron. N., 2: 23. 1944. Tree to 40 m tall; leaves 7–11-foliolate; leaflets ovate to ovate-oblong, sparsely pubescent to glabrescent. Evergreen lowland forests, 100–200 m; Amazonas (Río Sipapo). Barinas, Cojedes, Falcón, Portuguesa, Zulia; Central Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil. Ormosia macrophylla Benth., Ann. Wiener Mus. Naturgesch 2: 88. 1838. —Kabaituri, Palo macure, Peonía, Tento. Tree to 10 m tall; leaves 5–9-foliolate; leaflets broadly ovate or elliptic, tertiary veins inconspicuous. Shrublands, whitesand savanna edges, scrubby forests, 50–600 m; Amazonas (Caño Chimoni of Río Casiquiare, slopes of Cerro Aracamuni, base of Cerro Yapapana, Río Barí, Río Guasacavi, Río Pasimoni, Río Siapa, San Carlos de Río

368

F ABACEAE

Negro, Santa Rosa de Ucata). Colombia (Amazonas, Vaupés), Ecuador, Brazil (Amazonas). ŠFig. 317. Branches of Ormosia macrophylla often become hollowed out and are occupied by ants. Ormosia maguireorum Rudd, Contr. U.S. Natl. Herb. 32: 351. 1965. Tree 15–25 m tall; leaves 9-foliolate; leaflets elliptic-ovate, coriaceous, venation prominent, densely hairy. Montane forests, 1000–1200 m; Amazonas (Río Yatúa on Sierra de la Neblina). Endemic. No collections with flowers have yet been obtained of this species and are needed to better characterize it. Ormosia nobilis Tul., Arch. Mus. Hist. Nat. 4: 106. 1844. Neotropics; 2 varieties, 1 in Venezuela. O. nobilis var. nobilis Shrub or tree to 10 m tall; leaves 7- or 9foliolate, elliptic to elliptic-oblong, coriaceous, finely velutinous, glabrescent. Savannas, scrubby forests, 100–300 m; Bolívar (upper Río Caura), Amazonas (Caño Caname, Río Aracamuni, Río Pasimoni, Río Siapa). Colombia, Guyana, French Guiana, Suriname, Brazil (Amazonas, Maranhão, Pará), Bolivia. Ormosia paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 62. 1925. —Kapakun, Metari-yek, Mureyenu-yek (Arekuna), Mutare-yek, Peonía, Peonilla, Peonío, Pionío. Ormosia crassicarpa Pierce ex Pittier, Bol. Soc. Venez. Ci. Nat. 10: 108. 1944. Ormosia heterophylla Pires, Bol. Técn. Inst. Agron. N. 38: 24. 1960. Tree to 20 m tall; leaves (1–)7–15foliolate; leaflets oblong-elliptic, coriaceous, secondary veins inconspicuous. Lowland and upland forests, 50–1500 m; Bolívar (Gran Sabana, Ptari-tepui, Río Asa, lower Río Caura, Río Paragua, Río Turiba), Amazonas (Caño Cucurital in Río Ventuari basin, 25 km northeast of Puerto Ayacucho, Río Matacuni, Serranía Batata 55 km southeast of Puerto Ayacucho). Mérida, Zulia; Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 320.

Ormosia steyermarkii Rudd, Contr. U.S. Natl. Herb. 32: 352. 1965. —Ormosia microsperma Pittier, Bol. Soc. Venez. Ci. Nat. 10: 109. 1945, non Baker 1868 [1867]. —Mutare-yek (Arekuna), Pronilla. Tree to 20 m tall; leaves 3–7-foliolate; leaflets ovate, coriaceous, the lower surface densely hairy, secondary veins prominent. Nonflooded montane forests, 800–1600 m; Bolívar (Salto de Pacairao near Kavanayén, Sororopántepuí). Brazil (Amazonas, Roraima). A collection in fruit from the Río Ayaiche, Steyermark 89475, may be conspecific with O. steyermarkii Rudd. or may represent a new taxon. The resinous wood of Ormosia steyermarkii is said to be used to kindle fires. Ormosia subsimplex Spruce ex Benth. in Mart., Fl. Bras. 15: 316. 1862. —Ormosia coccinea var. subsimplex (Spruce ex Benth.) Rudd, Contr. U.S. Natl. Herb. 32: 328. 1965. —Peonía. Tree to 13 m tall; leaves 7–11-foliolate; leaflets ovate, elliptic or oblong, coriaceous, secondary veins broadly spaced, tertiary veins weak. Riparian forests in and around savanna areas, 50–200 m; Bolívar (Río Parguaza), Amazonas (lower Río Ventuari, San Fernando de Atabapo, Solano). Apure; Panama, Brazil (Amazonas: Rio Aracá). ŠFig. 319. Ormosia subsimplex is related to O. coccinea but has differently shaped fruits and corollas, larger flowers, widely spaced secondary veins and scarcely developed tertiary venation. Included here are Fariñas et al. 331 (VEN) from Amazonas state and Davidse & González 12429 (MO) from Apure state. These are trees to 8 m tall, with strongly beaked fruits, and may represent a new species. Ormosia williamsii Rudd, Contr. U.S. Natl. Herb. 32: 313. 1965. —Peonía rebalsera, Peonío, Tento. Tree to 15 m tall; leaves 5–9-foliolate; leaflets ovate or oblong, coriaceous, glabrescent. Riparian black-water forests, 100–200 m; Amazonas (Caño Magua, Piedra Cocuy, Río Atabapo, Samariapo, near San Carlos de Río Negro). Colombia, Brazil (Amazonas: Rio Negro).

Ormosia 369

Fig. 316. Ormosia bolivarensis

Fig. 317. Ormosia macrophylla

370

F ABACEAE

Fig. 318. Ormosia costulata

Fig. 319. Ormosia subsimplex

Ormosia 371

Fig. 320. Ormosia paraensis

372

F ABACEAE

Ormosia sp. A Tree to 25 m tall; leaves 5- or 7-foliolate; leaflets ovate to elliptic, subcoriaceous, the lower surface densely tawny; flowers not known. Montane forests, 1300–1400 m; Bolívar (La Escalera, Sierra de Lema). Endemic. Ormosia sp. B Shrub to 3 m; leaves 7-foliolate, the up-

per base of petiole flattened; leaflets to 20 cm long, elliptic-oblong, coriaceous, densely ferruginous-pubescent with crispate trichomes, tertiary veins visible, the upper surface dark green, nitid, and rugulate, the lower surface tawny-brown. Secondary forests, 1100–1300 m; Amazonas (35 km northeast of Parima B). Endemic.

46. PACHYRHIZUS Rich. ex DC., Prodr. 2: 402. 1825, nom. cons. by Nidia L. Cuello A. Twining annual, rarely semierect perennial herbs to somewhat woody vines. Roots tuberous. Leaves alternate, pinnately 3-foliolate; stipules linear; petioles with prominent pulvinus; leaflets entire, lobed, or deeply dissected, with linear, caducous stipels. Inflorescences axillary pseudoracemes; bracts progressively of reduced size from main axis to lateral axis; peduncle often longer than the petiole, the flowers somewhat congested apically; bracteoles subulate, shorter than and subtending the calyx; pedicels slender, slightly longer than the calyx. Flowers blue, floral prophylls caducous and silky. Calyx campanulate-tubular, 4-lobed with upper lobe larger and emarginate; standard suborbicular, with green basal spot, emarginate and with 2 basal incurved auricles enclosing the auricles of the wings; wing petals medially adherent to keel petals; keel petals obliquely oblanceolate, recurved with a hump, equaling the wings, blades connate, not auriculate. Stamens 10, diadelphous; anthers elliptical, dorsifixed. Ovary subsessile, with a crenulate disc round its base;

Fig. 321. Pachyrhizus erosus

Phaseolus 373

ovules numerous; style incurved, sometimes forming a circle, ventrally ciliate, dorsally glabrous; distal part of stigma subglobose on the vertical surface, widening at apex. Legume linear-oblong, internally septate between seeds, externally contracted between seeds. Seeds flat and suborbicular to flat and square, or plump and reniform. Neotropics; 5 species, 2 in Venezuela, 1 of these in the flora area. The genus is widely cultivated for the edible tuberous roots. Pachyrhizus erosus (L.) Urb., Symb. Antill. 4: 311. 1905. —Dolichos erosus L., Sp. Pl. 726. 1753. —O’ mochahcho (Panare). Herbaceous climbing or trailing vine. Semideciduous forests bordering savannas,

50–100 m; Bolívar (El Corozal, lower Río Parguaza). Aragua, Distrito Federal, Portuguesa; Mexico, Central America, West Indies, French Guiana, Brazil, Paraguay, Argentina, introduced and naturalized in the Paleotropics. ŠFig. 321.

47. PHASEOLUS L., Sp. Pl. 723. 1753; Gen. Pl. ed. 5, 323. 1754. by Gerardo A. Aymard C. Vines, herbs, or subshrubs. Leaves trifoliolate; stipules acute, veined, often pubescent, not prolonged below the insertion; stipels often oblong, thin, glabrous; leaflets mostly entire, pubescent or glabrate but minute, hooked trichomes present. Inflorescences axillary, the flowers congested in fascicles along the rachis, the nodes not swollen, not glandular; rachis eglandular; bracteoles ovate or greatly reduced and much shorter than the calyx, veined, puberulent, persistent at least until anthesis; bracts ovate or lanceolate, veined, puberulent; pedicels mostly longer than the calyx. Flowers blue, purple, violet, yellow, or white. Calyx mostly 2-lipped, the upper teeth partly united; standard symmetrical, orbicular, basally appendaged; wing pet-

Fig. 322. Phaseolus lunatus

374

F ABACEAE

als partly spiralled, apically hooded; keel petals in 2 or 3 spirals. Vexillar stamen free, the free part of the others elongate; anthers nearly uniform. Ovary 1–manyovulate; style apically thickened, curved in 1.5–2.5 spirals, hairy inside, distally caducous. Fruit linear-oblong, straight or slightly curved, not septate, compressed or turgid, sometimes beaked. Seeds 1–many, oblong to reniform, the hilum short and central. Neotropics, widely cultivated in temperate and tropical countries; ca. 50 species, 4 in Venezuela, 2 of these in the flora area. Key to the Species of Phaseolus 1. 1.

Stipels 1–1.5 mm long; peduncle 2–3 cm long; standard pubescent outside; fruits curved ........................................................................ P. lunatus Stipels 2–4 mm long; peduncle < 1 cm long; standard sparsely pilose to glabrate on the midvein outside; fruits straight ........................ P. vulgaris

Phaseolus lunatus L., Sp. Pl. 724. 1753. Climbing or trailing vine or erect herb; flowers yellow or purplish. Escaped from cultivation, 100–200; Amazonas (Río Cataniapo). Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. ŠFig. 322. Phaseolus vulgaris L., Sp. Pl. 723. 1753. Neotropics, but cultivated in almost every country around the world; 2 varieties, 1 in Venezuela.

P. vulgaris var. vulgaris Climbing or trailing vine or erect herb; flowers white or purplish. Northern Bolívar (cultivated). Widely cultivated elsewhere in Venezuela, but grows wild in Lara, Mérida, Portuguesa, Táchira, Trujillo; widely cultivated as the source of black beans in almost every temperate and tropical country. This species was domesticated in the New World probably some 8000–10,000 years ago from a wild ancestral form distributed in the highlands between northern Mexico and northern Argentina.

48. PISCIDIA L., Syst. Nat. ed. 2, 1151, 1155, 1376. 1759, nom. cons. by Gerardo A. Aymard C. Trees or shrubs, unarmed. Leaves alternate, odd-pinnate, 5–27-foliolate; stipules (bud scales) obliquely ovate, semiorbicular, or reniform, early caducous; stipels absent; leaflets opposite. Inflorescences axillary or lateral, usually racemose, sometimes spicate. Flowers white with pink to purplish markings; bracts at base of pedicels, ovate, elliptic, or lanceolate, early caducous; bracteoles paired at base of calyx, ovate, oblong, or linear, caducous. Calyx campanulate with 5 short lobes; standard suborbicular, usually pubescent on the outer face, but glabrous in one species; wing petals falcate, oblong, commonly a little longer than the standard, adherent to the keel; keel petals connate at the base. Stamens 10, monadelphous but the vexillar filament free at the base; anthers oblong, dorsifixed. Ovary sessile, many-ovulate; style glabrous above; stigma minutely penicillate. Fruit indehiscent, 1–10-seeded, compressed, with 4 longitudinal wings. Seeds reniform, tan to reddish or dark brown, laterally compressed, the hilum lateral, elliptic to suborbicular. Neotropics; ca. 7 species, 1 in Venezuela. Piscidia carthaginensis Jacq., Enum. Syst. Pl. 27. 1760. Piscidia guaricensis (Pittier) Pittier, Mesa Guanipa 49. 1942. —Lonchocarpus gua-

ricensis Pittier, Arb. Legum., part 3, Trab. Mus. Comercial Venezuela 4 [reprinted from Bol. Minist. R.R. E.E. no. 4–7.] 4: 231. 1928.

Platymiscium 375

Fig. 323. Piscidia carthaginensis

Small tree; calyx pink; petals white and pink. Dry to gallery forests, 50–100 m; Bolívar (Las Bonitas on the middle Río Orinoco). Anzoátegui, Guárico, Miranda, Mona-

gas, Nueva Esparta, Portuguesa; Mexico, Central America, Antilles, Colombia, Ecuador, Peru. ŠFig. 323.

49. PLATYMISCIUM Vogel, Linnaea 11: 198. 1837. by Nereida Xena de Enrich Trees; bark dark, rough, fissured. Leaves opposite or verticillate, compound, odd-pinnate, with 5 or 7(9) opposite or subopposite leaflets; stipules coriaceous, ovate-orbicular or lanceolate, caducous or subpersistent; stipels absent. Inflorescence axillary, paniculate, glabrous or variously pubescent. Flowers small, yellow, numerous; hypanthium small or almost absent. Calyx campanulate, dark to pale green, glabrous or pubescent, teeth small and unequal, the vexillar ones almost totally connate; corolla yellow; standard ovate-orbicular, glabrous, yellow, with ± conspicuous brown nectar guides, without auricles, with a short claw; wing petals oblong-elliptic, glabrous, free, auriculate; keel petals slightly connate at the apex, upper margin ciliate. Stamens 10, frequently the vexillar one free or partially connate

376

F ABACEAE

to the staminal column; anthers monomorphic, ovate-hastate, versatile with longitudinal dehiscence, extrorse. Pistil glabrous, stipe elongate; ovary with 1 ovule; style curved, filiform; stigma terminal, inconspicuous. Fruit indehiscent, flat, stipitate, membranous, oblong-elliptic or narrowly elliptic, with seed in central chamber. Seed large, reniform, flattened, brown; embryo curved, the cotyledons thin, foliaceous, the radicle curved, inflexed. Neotropics; ca. 12 species, 3 in Venezuela, 2 of these in the flora area. This genus is frequently cultivated as a shade and ornamental tree. The wood is sold under the name “Panama redwood” in the lumber industry. Key to the Species of Platymiscium 1.

1.

Inflorescences glabrous or scarcely pubescent; flowers pale yellow, 10–14 mm long; calyx pale green, generally glabrous; leaflets 5 or 7, ovate-elliptic, ± abruptly acuminate at apex ....................................... P. pinnatum Inflorescences densely pubescent; flowers yellow-orange, 15–17 mm long; calyx externally dark, golden-pubescent; leaflets 5, ovate-oblong, obtuse at apex ................................................................................ P. trinitatis

Fig. 324. Platymiscium pinnatum

Platypodium 377

Platymiscium pinnatum (Jacq.) Dugand, Contr. Hist. Nat. Colomb. 1: 7. 1938 —Amerimnon pinnatum Jacq., Select. Stirp. Amer. Hist. 200. 1763. —Roble, Roble blanco, Roble colorado. Platymiscium polystachyum Benth. in Seem., Bot. Voy. Herald 111, t. 21. 1853. Platymiscium polystachyum var. fendleri Benth., J. Proc. Linn. Soc., Bot. 4: 83. 1860. Tree to 20 m tall. Deciduous forests, woody savannas, 100–500 m; Bolívar (island in Danto Manchado sector of Lago Guri, 30 km south of El Manteco, near El Palmar, La Paragua, Puerto Ordaz, Río Icabarú, northeast of Upata). Anzoátegui, Apure, Barinas, Carabobo, Cojedes, Guárico, Lara, Monagas, Portuguesa, Zulia; Central America, Colombia, Brazil. ŠFig. 324.

Platymiscium trinitatis Benth., J. Proc. Linn. Soc., Bot. 4: 82. 1860. —Roble, Roble blanco, Roble colorado. Platymiscium duckei Huber, Bol. Mus. Paraense Hist. Nat. 6: 83. 1909. Platymiscium nigrum Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 157. 1922. —Platymiscium duckei var. nigrum (Ducke) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 87. 1925. Tree to 20 m tall. Riparian, montane, and premontane semideciduous or deciduous forests, 100–500 m; Bolívar (northern deciduous zone, Cerro Cotorra between La Paragua and mouth of Río Paragua, Temblador on lower Río Caura, northwest of Tumeremo). Anzoátegui, Apure, Aragua, Barinas, Falcón, Lara, Mérida, Miranda, Sucre, Yaracuy; Trinidad, Brazil (Pará).

50. PLATYPODIUM Vogel, Linnaea 11: 420. 1837. by Gerardo A. Aymard C. Trees 20–30 m tall; branches pilose-pubescent. Leaves odd- or even-pinnate, alternate; rachis pilose-pubescent; leaflets 1.5–2.5 × 2–7 cm, in 5–18 pairs, alternate or irregularly opposite, upper surface glabrous, lower surface puberulent to pubescent, apex emarginate; stipules linear, deciduous; stipels absent. Inflorescences racemose in the upper leaf axils; bracts and bracteoles small, caducous. Flowers showy, yellow or yellow-orange, pedunculate; bracts 2, acute, deciduous. Calyx turbinate at the base, glabrous or laxly pilose, lobes pubescent within, the upper 2 teeth broader, connate high up; standard glabrous, obovate, emarginate, not appendaged; wing petals obliquely obovate or oblong; keel petals oblong or obovate, obtuse, connate at the apex (dorsally). Stamens 10, divided equally among 2 fascicles or sometimes the vexillar stamen free; anthers versatile. Ovary long-stipitate, glabrous or laxly pilose; ovules 3–5; style filiform; stigma small, suborbicular terminal. Fruit stipitate, 4–8 mm long, samaroid, indehiscent, woody at the apex, narrowed at the base into an obliquely oblong venose wing. Seeds 1(2), oblong to reniform. Panama, Colombia, Venezuela, Brazil, Bolivia, Paraguay; 2 species, 1 in Venezuela. Platypodium elegans Vogel, Linnaea 11: 421. 1837. Panama, Colombia, Venezuela, Brazil, Bolivia, Paraguay; 2 subspecies, both in Venezuela, 1 in the flora area. P. elegans subsp. maxonianum (Pittier) H.C. Lima, comb. nov. —Platypodium

maxonianum Pittier, Contr. U.S. Natl. Herb. 18: 234. 1917. —Canalete amarillo. Tree 20–30 m tall; flowers yellow. Semideciduous to wet forests, 100–300 m; Bolívar (Túriba). Barinas, Falcón, Portuguesa, Trujillo, Zulia; Panama, Colombia, Brazil, Bolivia, Paraguay. ŠFig. 325.

378

F ABACEAE

Fig. 325. Platypodium elegans subsp. maxonianum

51. POECILANTHE Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 80. 1860. by Nidia L. Cuello A. Small trees or shrubs. Leaves odd-pinnate, sometimes unifoliolate; leaflets alternate or opposite; stipels minute or absent; stipules caducous or inconspicuous. Flowers in short axillary or lateral racemes or panicles; bracts and bractlets small. Calyx turbinate at the base, the upper 2 lobes connate into one ± 2-toothed lip; petals clawed; standard orbiculate, not appendaged; wing petals falcate-oblong or obovate; keel petals incurved, subrostrate, connate at the back. Stamens all connate into a sheath split above, sometimes nearly diadelphous; anthers alternately longer and basifixed or shorter and versatile. Ovary subsessile or shortly stipitate; ovules several; style filiform, glabrous, incurved; stigma small, terminal. Fruit flat-com-

Poecilanthe 379

pressed, woody-coriaceous, 2-valved. Seed obovate, hard and shining, compressed, with a basal hilum, estrophiolate. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; ca. 10 species, 2 in Venezuela, both in the flora area. Key to the Species of Poecilanthe 1. 1.

Pedicels 0.5–1 mm long; flowers 8–10 mm long, purple or red; fruit 4–6.5 cm long .................................................................................... P. amazonica Pedicels 3–4 mm long; flowers 12–15 mm long, brown; fruit 10–13 cm long ................................................................................................ P. hostmannii

Poecilanthe amazonica (Ducke) Ducke, Bull. Mus. Hist. Nat. (Paris) sér. 2, 4: 734. 1932. —Cyclolobium amazonicum Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 146. 1922. —Arepito. Small tree or shrub; flowers purple or red. Evergreen lowland and flooded forests, 100– 200 m; Amazonas (Caño Morocoto below San Fernando de Atabapo, Río Sipapo, San Carlos de Río Negro). Brazil (Amapá, Amazonas). ŠFig. 326. Poecilanthe hostmannii (Benth.) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 61. 1939. —Cyclolobium hostmannii Benth., J. Proc. Linn. Soc., Bot. 4: 52. 1860. Tree; flowers deep maroon. Evergreen lowland forests, 100–200 m; Amazonas (base of Cerro Yapacana, Río Baría). Suriname, French Guiana, Brazil (Amapá, Amazonas).

Fig. 326. Poecilanthe amazonica

380

F ABACEAE

52. POIRETIA Vent., Mém. Cl. Sci. Math. Inst. Natl. France 1807(1): 4. July 1807, nom. cons. by Nidia L. Cuello A. Herbs or suffrutescents, erect or scandent, 1–4 m tall. Leaves alternate, 4foliolate; leaflets chartaceous to coriaceous, linear to ovate, obovate, or suborbicular, glandular-punctate on lower surface, glands sometimes visible on upper surface, opposite; stipules lanceolate, usually glandular, caducous; stipels present at base of lower pair of leaflets, linear to lanceolate or ovate, usually glandular, caducous. Inflorescences axillary or terminal, racemose, sometimes paniculate or spicate, few- or many-flowered; bracts stipule-like; bracteoles lacking. Flowers small to mediumsized, 4–20 mm long. Calyx glandular-punctate, campanulate with 5 subequal lobes shorter than the tube, the vexillar lobes smaller than the others and somewhat connate; petals yellow, glandular-punctate; standard reflexed, glabrous or nearly so. Stamens 10, monadelphous; anthers dimorphic, alternately oblong and basifixed or elliptic and dorsifixed. Ovary sessile, glabrous or pubescent; ovules 1–8; stigma terminal, minutely capitate. Fruit sessile, oblong, laterally compressed, with 1–8 articles, glandular-punctate, sometimes verruculose. Seeds brown, reniform, compressed, hilum present, elliptic. Mexico, Central America, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil (most diverse in east-southeast part), Bolivia; ca. 6 species, 1 in Venezuela. Poiretia punctata (Willd.) Desv., J. Bot. Agric. 1: 122, pl. 5, fig. 17. 1813. —Glycine punctata Willd., Sp. Pl. 3(2): 1066. 1802. Herbaceous vine 1–3 m long; flowers bright yellow. Open areas, forest edges, roadsides, 100–200 m; Bolívar (La Paragua). Aragua, Carabobo, Distrito Federal, Miranda; Mexico, Central America, Antilles, Colombia, Ecuador, Peru, eastern and southeastern Brazil, Bolivia. ŠFig. 327.

Fig. 327. Poiretia punctata

Pterocarpus 381

53. PTEROCARPUS Jacq., Select. Stirp. Amer. Hist. 283. 1763, nom. cons. by Nereida Xena de Enrich, Celia Gil, and Paul E. Berry Trees; bark smooth or slightly to strongly fissured, exuding a red resin; crown usually umbrella-shaped; young branches terete, glabrous to densely pubescent, frequently with lenticels. Leaves alternate, compound, odd-pinnate, nonstipulate; stipels linear or narrowly triangular, caducous, rarely persistent; petiole and petiolules terete, 2–5(–6) cm long, glabrous or pubescent; leaflets (3)5–11(13), alternate or subopposite, occasionally opposite, varying in shape and size along the rachis, glabrous or densely pubescent, the base rounded, obtuse, or sometimes acute, the apex acuminate or emarginate and sometimes mucronate, margins entire. Inflorescences axillary, paniculate or racemose, glabrescent or densely pubescent; bracts and bracteoles small, linear or triangular, early caducous or sometimes persistent, glabrous or pubescent. Flowers small, 11–20 mm long, numerous, zygomorphic, pedicellate or sessile; hypanthium turbinate, frequently basally incurved, 1–4 mm long. Calyx 5-dentate, teeth small, unequal, the vexillar ones almost joined, apex rounded or acute; corolla yellow-orange, nectar guide usually whitish or violet at the middle of the standard; standard 10–19 mm long, orbicular-ovate or obovate-elliptic, glabrous, basally attenuate and ending in a small or medium-sized claw, apex slightly emarginate; wing petals 10–18 mm long, oblong-elliptic or elliptic-obovate, free, glabrous, auriculate, basally clawed; keel petals 10–15 mm long, glabrous, auriculate, connate at the middle and basal parts, basally clawed. Stamens 10, the vexillar one frequently joined up to 1/4 of the staminal column or shortly joined basally; anthers versatile, isomorphic, elliptic, longitudinally dehiscent. Ovary 1-locular, stipitate to sessile, glabrous or pubescent; ovules 2–7; style incurved, filiform, glabrous; stigma terminal, inconspicuous. Fruit flattened, indehiscent, winged, orbicular or suborbicular, sometimes asymmetric, stipitate or sessile, glabrous or glabrescent, membranous or coriaceous; seed chamber hard, central, distal, or lateral, sometimes partitioned. Seeds 1–3, reniform; testa smooth or slightly rugose, the hilum conspicuous. Pantropics; ca. 20 species, 5 in Venezuela, all of these in the flora area. The wood from species in this genus is used for construction, musical instruments, crafts, and toothpicks. The resin is called “sangre de drago” and is widely used in popular medicine. Key to the Species of Pterocarpus 1.

1.

2(1).

Fruit with a prominent, membranous or chartaceous wing; seed chamber in central or proximal position; inflorescence densely pubescent, a simple raceme or else with 1–3 short branches at the base of the main axis; calyx pubescent on outside ........................................................... 2 Fruit with a coriaceous or much-reduced wing; seed chamber in a central or more often a lateral position; inflorescence glabrous or pubescent, a raceme or a much-branched panicle; calyx glabrous or pubescent on outside .................................................................................................... 3 Fruit stipitate, the stipe 8–25 mm long, the wing with a small outgrowth distally; seeds 1–3, the testa smooth, pale brown or brown-pink; pedicel

382

2.

3(1).

3.

4(3).

4.

F ABACEAE

(6–)7–13(–17) mm long; leaflets (7)9–11(13), apex emarginate, mucronate, or briefly acuminate-mucronate (usually the mucro caducous) .............................................................................................. P. acapulcensis Fruit estipitate, the wing without small outgrowth distally; seed solitary, the testa rugose, pale brown to dark brown; pedicel (2.5–)4–9(–12) mm long; leaflets (3)5–9(11), apex short or long obtuse-acuminate ..... P. rohrii Fruit stipitate, the stipe 2.5–6 mm long; inflorescence a much- and usually laxly branched axillary panicle; calyx glabrous on the outside; fruit glabrous, the seed-bearing part variously thickened, usually distinctly veined ....................................................................................... P. officinalis Fruit estipitate; inflorescence a simple, axillary raceme, or if branched, only 1–3 short and few-flowered branches near the base; calyx densely brownish pubescent on the outside; fruit usually minutely brownish tan-pubescent, or glabrescent, the seed-bearing part generally not distinctly veined ......................................................................................... 4 Fruit smooth to rugulose, the seed-bearing part slightly or rather evenly thickened, gradually grading into the wing, glabrescent and usually golden tan; inflorescence racemose, sometimes spindle-shaped; stems generally hollow and ant-inhabited ...................................... P. amazonum Fruit wrinkled or deeply rugose, the seed-bearing part rather abruptly thickened from the small surrounding wing, usually yellowish to rusty brown with a fine pubescence or else glabrescent; inflorescence racemose or with 1–3 branches at the base; stems solid, generally not antinhabited ............................................................................ P. santalinoides

Pterocarpus acapulcensis Rose, Contr. U.S. Natl. Herb. 5: 143. 1897. —Drago, Sangre de drago, Sangredrago. Pterocarpus podocarpus S.F. Blake, Contr. U.S. Natl. Herb. 20: 524. 1924. Pterocarpus vernalis Pittier, Arb. Arbust. Venez. 6–8 [reprinted from Bol. Minist. RR. EE. no. 8/9]: 100. 1927. Deciduous tree 5–25(–30) m tall; fruits persisting for several months. Deciduous, semideciduous, and gallery forests, 50–500 m; northern Bolívar. Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Lara, Miranda, Portuguesa, Táchira, Zulia; Mexico, Central America, Colombia. ŠFig. 329. Pterocarpus amazonum (Benth.) Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 56. 1939. —Phellocarpus amazonum Mart. ex. Benth., Comm. Legum. Gen. 42. 1837. Pterocarpus ulei Harms, Verh. Bot. Vereins Prov. Brandenburg 48: 171. 1907.

Evergreen tree 5–15 m tall; branchlets thickened, hollow and ant-inhabited. Riparian and swamp forests, 50–300 m; Delta Amacuro (Caño Jota-Sabuca near Caño Mariusa), Bolívar (Isla Anacoco, Río Nichare basin, Serranía de Imataca), Amazonas (Río Atabapo). Apure; Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Pterocarpus officinalis Jacq., Select. Stirp. Amer. Hist. 283, t. 183, fig. 92. 1763. —Cacú, Drago, Iburu (Warao), Lagunero, Mucanana, Mucunana, Mucutena, Sangredrago, Sangrito. Pterocarpus draco L., Sp. Pl. ed. 2, 1662. 1763. Tree 5–30 m tall; lower trunk with sinuous buttresses to 3 m tall and 5 m wide. Swamp and flooded riparian forests, occasionally nonflooded lowland forests, near sea level to 300(–600) m; widespread in lower Delta Amacuro, Bolívar (Altiplanicie de Nuria, Serranía de Imataca, east of Túriba). Anzoátegui, Monagas, Sucre; Mexico, Central America, West Indies, Colombia, Trinidad, Ecuador, Brazil. ŠFig. 328.

Pterocarpus 383

Pterocarpus officinalis is common along water courses in Delta Amacuro state and is easily recognizable by the characteristic buttress roots. The light wood is locally used in artisanry to make carved wooden animals. Pterocarpus rohrii Vahl, Symb. Bot. 2: 79. 1791. —Sangredrago, Sangrito alado. Pterocarpus rupestris Pittier, Arb. Arbust. Venez. 6–8 [reprinted from Bol. Minist. RR. EE. no. 8/9]: 23. 1927. Pterocarpus magnicarpa Schery, Fieldiana, Bot. 28: 261. 1952. Pterocarpus rohrii var. rubiginosus Schery, Fieldiana, Bot. 28: 261. 1952. Tree 6–30 m tall; trunk narrowly buttressed. Semideciduous to evergreen lowland forests, often along watercourses, 50–200 m; Delta Amacuro (Río Toro area along Bolívar border), Bolívar (middle Río Caura, Río Paragua, Río Toro area), Amazonas (La Esmeralda, Río Cunucunuma, around San Carlos de Río Negro). Anzoátegui, Apure, Barinas, Carabobo, Falcón, Lara, Miranda, Yaracuy, Zulia; Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 330.

Pterocarpus santalinoides L’Hér. ex DC., Prodr. 2: 419. 1825. —Cuai-eneru, Guajenera, Para-para, Sangrito. Pterocarpus amazonicus Huber, Bol. Mus. Paraense Hist. Nat. 5: 402. 1909. Pterocarpus grandis R.S. Cowan, Mem. New York Bot. Gard. 10: 86. 1961. Usually evergreen tree 5–15 m tall; trunk buttressed. Swamp, riparian, and semideciduous forests, near sea level to 300 m; Delta Amacuro (widespread), Bolívar (Serranía de Imataca), Amazonas (lower Río Cipapo, Río Yatúa). Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, western Africa. The collections from Amazonas state (including the type of Pterocarpus grandis) have unusually large leaflets and fruits and are questionably referred to this species.

Fig. 328. Pterocarpus officinalis

384

F ABACEAE

Fig. 329. Pterocarpus acapulcensis

Fig. 330. Pterocarpus rohrii

Rhynchosia 385

54. RHYNCHOSIA Lour., Fl. Cochinch. 425, 460. 1790, nom. cons. Dolicholus Medik., Vorles. Churpfälz. Phys.-Ökon. Ges. 2: 354. 1787, nom. rejic. Arcyphyllum Elliott, J. Acad. Nat. Sci. Philadelphia 1: 371. 1818. by Renée H. Fortunato Suffrutescent herbs, lianas, or subshrubs, erect, ascending, or twining to prostrate. Roots woody, turnip-shaped, spindle-shaped, or club-shaped. Stems terete to angular, simple or branched. Leaves unifoliolate or pinnate-trifoliolate, subsessile to long-petiolate; stipules free, persistent or deciduous; leaflets 3-veined, pubescent, glabrous, glandular on both surfaces or only the lower one, glandular trichomes yellowish, sometimes reddish brown when dried; stipels small, persistent (sometimes early caducous) or lacking. Inflorescence racemose, axillary, simple or few-branched, 1–many-flowered; peduncles surpassing the main cauline leaves or not; bracts small, persistent or deciduous; bracteoles lacking. Calyx campanulate, 5-lobed, the 2 lateral lobes subequal to 3 or more times the length of the tube, the 2 vexillar lobes connate; corolla yellow, often striped or flushed with purple or reddish brown in the standard veins; standard obovate to orbiculate, emarginate to rounded, sometimes apiculate, biauriculate at the base, clawed, externally pubescent and glandular or glabrous; wing petals narrowly oblong, uniauriculate, glandular-pubescent or glabrous at the apex, clawed; keel petals falcate, incurved and glandular-pubescent to glabrous at apex, clawed. Stamens 10, diadelphous; anthers ellipsoid, dorsifixed, uniform. Ovary sessile to subsessile, villous; ovules (1)2; style incurved, filiform, glabrous; stigma small. Legumes 2-seeded, ovoid, obovoid, ellipsoid, falcate, or dumbbell-shaped, deeply to shallowly constricted between seeds, usually compressed, pubescent and glandular, dehiscent, beaked. Seeds subreniform to suborbicular, brown, black, mottled, or red and black, lustrous; strophiole inconspicuous to prominent, the hilum ovoid to linear; funicular attachment distal or central to subcentral to the hilum. Widely distributed in warm-temperate and tropical areas of both hemispheres, but absent from Europe and northern and central Asia; ca. 190 species, 6 in Venezuela, all of these in the flora area. Key to the Species of Rhynchosia 1. 1. 2(1).

2.

3(1).

3.

Lateral calyx lobes > 2 times as long as the tube ..................................... 2 Lateral calyx lobes ± equal to the length of the tube ................................ 3 Standard pubescent and gland-dotted externally, 6–10 × 4–6 mm; stems twining or creeping; legumes puberulous to glabrate, trichomes uniform in length .......................................................................... R. reticulata Standard glabrous externally, 12–20 × 10–17 mm; stems erect to ascending; legumes viscid-hirsute and puberulous, trichomes of 2 lengths ........................................................................................... R. schomburgkii Legumes deeply to shallowly constricted between the seeds (dumbbellshaped); seeds red and black; leaflets and standard with glands not turning black when dried ...................................................................... 4 Legumes ellipsoid or falcate; seeds black, brown, or mottled; leaflets and standard usually with glands turning black when dried ..................... 5

386

4(3).

4.

5(3). 5.

F ABACEAE

Legumes glabrous or puberulous, glabrescent and black with age; seeds predominantly black, red only in the hilum area; stipels present; pedicels 2–5 mm long ......................................................... R. melanocarpa Legumes densely brown- or yellowish brown-puberulous, not glabrescent with age; seeds with nearly equal areas of red and black almost equal; stipels usually absent; pedicels 0.5–1.5 mm long .............. R. phaseoloides Legumes ellipsoid; funicle attached distally to hilum; flowers 7.5–12 mm long ................................................................................................. R. edulis Legumes ± falcate; funicle attached centrally or nearly so to hilum; flowers 3–6(–6.5) mm long ................................................................ R. minima

Rhynchosia edulis Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 123. 1874. —Eriosema edule (Griseb.) Burkhart, Darwiniana 6: 261. 1943. Rhynchosia melanosticta Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 123. 1874. Eriosema volubile Micheli, Mém. Soc. Phys. Genève 28(7): 36. 1883. Herbaceous or suffrutescent vine; stems erect, suberect, prostrate, and twining; stipules persistent; leaflets ovate, deltate, or rhombic, glands on both surfaces; inflorescences elongate, usually equaling or exceeding the leaves. Savannas, edges of forests in open grassland, along riverbanks, 100–400 m; Bolívar (near Upata and Guasipati). Aragua, Carabobo, Distrito Federal, Mérida, Miranda, Portuguesa, Trujillo; southeastern U.S.A., Central America, Colombia, Peru, Brazil, Paraguay, Argentina. Rhynchosia melanocarpa Grear, Mem. New York Bot. Gard. 31(1): 43. 1978. Rhynchosia phaseoloides var. erecta Micheli, Mém. Soc. Phys. Genève 28(7): 33. 1883. Twining, suffrutescent vine; stems fewbranched; stipules caducous; leaflets rhombic, ovate, or deltate; stipels linear; inflorescences elongate, equaling or exceeding the leaves. Thickets, margins of gallery forests along rivers, disturbed areas, 400–500 m; Bolívar (near Kamarata). Anzoátegui, Apure, Guárico, Monagas; Peru, Brazil, Bolivia, Paraguay, northeastern Argentina. ŠFig. 333. Rhynchosia minima (L.) DC., Prodr. 2: 385. 1825. —Dolichos minimus L., Sp. Pl. 726. 1753.

Glycine reflexa Nutt., Gen. N. Amer. Pl. 2: 115. 1818. Suffrutescent to herbaceous vine; stems twining or prostrate; stipules persistent; leaflets rhombic-circular, ± rhombic, or ovate; inflorescences elongate, equaling or exceeding the leaves. Weed in disturbed and cultivated land in different soil types, near sea level; Delta Amacuro (Pedernales). Widespread elsewhere in Venezuela; tropics and subtropics worldwide. ŠFig. 334. Rhynchosia phaseoloides (Sw.) DC., Prodr. 2: 385. 1825. —Glycine phaseoloides Sw., Prodr. 105. 1799. —Dolicholus phaseoloides (Sw.) DC. in Kuntze, Revis. Gen. Pl. 3(2): 62. 1898. Woody vine; stems angular, becoming terete; stipules caducous; racemes elongate, usually equaling or exceeding the leaves, branched. Margins of woods bordering savannas, clearings in evergreen forests, disturbed areas, 50–1100 m; Delta Amacuro (Río Orocoima), Bolívar (Altiplanicie de Nuria, Río Grande, Río Suapure, west of Santa Elena de Uairén). Distrito Federal, Guárico, Miranda, Sucre, Yaracuy, Zulia; southern Panama, Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ŠFig. 331. Rhynchosia reticulata (Sw.) DC., Prodr. 2: 385. 1825. —Glycine reticulata Sw., Prodr. 2: 105. 1788. Dolicholus kuntzei Harms ex Kuntze, Revis. Gen. Pl. 3(2): 61. 1898. —Rhynchosia kuntzei Harms ex Kuntze, Revis. Gen. Pl. 3(2): 61. 1898, as synonym. —Rhynchosia reticulata var. kuntzei (Harms ex Kuntze) Grear, Mem. New York Bot. Gard. 31(1): 116. 1978.

Rhynchosia 387

Fig. 331. Rhynchosia phaseoloides

Fig. 333. Rhynchosia melanocarpa

Fig. 332. Rhynchosia schomburgkii

Fig. 334. Rhynchosia minima

388

F ABACEAE

Suffrutescent herb or prostrate or twining vine; stems 3-angled; stipules lanceolate to widely ovate, base truncate or cordate, caducous, rarely some persistent; leaflets ovate, rhombic, or round-ovate; racemes congested to elongate, shorter than or exceeding the leaves. Thickets, margin forests, along rivers, grasslands, shrub savannas, 100–200 m; Bolívar (El Cuchivero). Barinas, Carabobo, Distrito Federal, Falcón, Monagas, Sucre, Táchira, Yaracuy; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

Rhynchosia schomburgkii Benth. in Mart., Fl. Bras. 15(1): 203. 1859. Dolicholus pittieri Standl., Contr. U.S. Natl. Herb. 17: 433. 1914. Rhynchosia caliensis Harms, Notizbl. Bot. Gard. Berlin-Dahlem 11: 783. 1933. Subshrub to suffrutescent herb; stems erect to twining at the tips; stipules caducous; leaflets discolorous; flowers 1–2.2 cm long; calyx only exceeding the corolla by the keel lobes. Sandy savannas, patches of forests, along rivers, 200–1200 m; Bolívar (widespread). Zulia; Colombia, Guyana, northern Brazil. ŠFig. 332.

55. SESBANIA Scop., Intr. Hist. Nat. 308. 1777, nom. cons. by Gerardo A. Aymard C. Annual or perennial herbs, shrubs, or small trees. Leaves alternate with numerous leaflets; leaflets entire, even-pinnate, obtuse at the apex, rounded at the base, often glaucous, short-petiolate; stipules small, deciduous; stipels minute or apparently absent. Inflorescences short and few-flowered axillary racemes; bracts and bracteoles small, apparently deciduous. Flowers with the hypanthium broadly campanulate, as broad as long or broader, the 5 teeth equal, often short, truncate to triangular with the tip acute to acuminate; petals white, purplish yellow, red, or variegated; standard longer than other petals, mostly orbicular or nearly so, reflexed, short-clawed and usually appendaged at the base; wing petals free; keel petals curved, joined below, with long claws. Stamens diadelphous, the vexillar stamen free, geniculate near the base, the united stamens equal or 5 alternate ones somewhat longer. Ovary stipitate, linear; stigma small, capitate. Fruit linear, short-stipitate, beaked, septate between the seeds within, the exocarp occasionally indented between the seeds in some species, 2-valved, mostly dehiscent, occasionally indehiscent, the calyx often not persistent in fruit. Seeds many, oblong, slightly compressed, smooth, brown. Pantropics; ca. 50 species, 5 in Venezuela, 2 of these in the flora area. Key to the Species of Sesbania 1.

1.

Pedicels 0.8–1 mm wide; calyx teeth 1–1.5 mm long; standard suborbicular, 1–1.5 cm long; wing petals oblong, 1.5 cm long; fruit 2–3.5 mm wide ...................................................................................................... S. emerus Pedicels 1.5–2 mm wide; calyx teeth 3–4 mm long; standard ovate, 3 cm long; wing petals oblong to obovate, to 2.5 cm long; fruit 4–6 mm wide ................................................................................................ S. exasperata

Sesbania emerus (Aubl.) Urb., Repert. Spec. Nov. Regni Veg. 16: 149. 1919. —Aeschynomene emerus Aubl., Hist. Pl. Guiane 775. 1775. —Maguey. Erect herb or shrub to 3 m tall; flowers

yellow with purple spots. Savannas, wet places, near sea level to 200 m; Delta Amacuro (Caño Araguao, Isla Cocuina, Sacupana), Bolívar (El Manteco, Kilómetro 88, Represa Guri). Aragua, Falcón, Guárico,

Sesbania 389

Fig. 335. Sesbania exasperata

390

F ABACEAE

Lara, Miranda, Portuguesa, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Sesbania exasperata H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 534. 1823 [1824]. Erect herb or shrub to 3 m tall; flowers

yellow. Wet places, edges of rivers and creeks, near sea level to 200 m; Delta Amacuro (Caño Güiniquina), Amazonas (Río Cataniapo). Apure, Barinas, Cojedes, Portuguesa, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 335.

56. SOEMMERINGIA Mart., Soemmeringia Nov. Pl. Gen. 27. 1828. by Nidia L. Cuello A. Small, creeping herbs from a central tap root, sometimes prostrate or somewhat shrubby. Leaves alternate, odd-pinnate, sensitive; leaflets 7–9, bluish green, oblanceolate, or obovate, shallowly emarginate at apex, glabrous; blades asymmetric about the midvein, the secondary venation parallel and prominent on both surfaces, extending to the leaflet margin and forming small teeth; stipules linear-lanceolate, sharply acuminate, striate, papery, persistent; stipels absent. Flowers 1 or 2, axillary; bracts small, subulate; bracteoles ovate to lanceolate, striate, persistent at calyx base. Calyx green, 2-lipped, upper lip 2-toothed, the teeth much longer than the tube, the upper 2 broader and fused almost to their apex; corolla yellow; standard subsessile, reticulate-venose, becoming brownish and scarious and persisting as a delicately veined protective covering completely enclosing the green plicate fruit. Stamens in 2 lateral bundles; anthers uniform. Ovary stipitate; ovules several; style incurved. Fruit stipitate, joints plano-convex, reticulate. Venezuela, Brazil, Bolivia; 1 species. Soemmeringia semperflorens Mart., Diss. de Soemmer 28. 1828. Prostrate shrub or subprostrate herb; petals light yellow. Open savannas, disturbed areas, 50–300 m; Bolívar (near Caicara, La

Paragua, lower Río Caroní). Anzoátegui, Apure, Cojedes, Guárico, Monagas, Portuguesa; Brazil (Amapá, Bahia, Ceará, Maranhão, Pará, Roraima), Bolivia. ŠFig. 336.

Fig. 336. Soemmeringia semperflorens

Spirotropis 391

57. SPIROTROPIS Tul., Arch. Mus. Hist. Nat. 4: 113. 1844. by Charles H. Stirton and Gerardo A. Aymard C. Trees or lianas. Stipules spatulate-oblanceolate. Leaves odd-pinnate, 1–4-jugate, opposite or subalternate, coriaceous, the upper surface glabrous, the lower surface velvety, apex sharply acuminate. Inflorescence terminal, racemose-paniculate. Flowers purple to purplish red. Calyx split into 1 or 2 parts at maturity, reflexed, upper half 2-lipped, lower half 3-lipped; petals short-clawed; standard elliptic to orbicular, glabrous; wing petals equal to keel petals but shorter than standard. Filaments free or scarcely united at their base. Ovary with 3–6 ovules, densely pubescent; style filiform; stigma capitate. Fruit flattened, reticulate, hard, oblong, tapering at both ends, apex acute, indehiscent. Venezuela, Guyana, Suriname, French Guiana; 2 species, both in the flora area.

Fig. 337. Spirotropis longifolia

392

F ABACEAE

Key to the Species of Spirotropis 1.

1.

Leaves and flowering axis finely puberulous, tawny or silvery; calyx split into 2 halves, recurved, quite twisted; filaments fused near the base .................................................................................................. S. longifolia Leaves and flowering axis coarsely pubescent, ferruginous; calyx split once into a broad, recurved, slightly twisted flap; filaments nearly free .......................................................................................................... S. sp. A

Spirotropis longifolia (DC.) Baill., Hist. Pl. 2: 364. 1870. —Swartzia longifolia DC., Mém. Légum. 406. 1806. —Turäyek (Arekuna). Dipteryx phaeophylla Steyerm., Ann. Missouri Bot. Gard. 71: 313. 1984. Tree 4–15 m tall. Lowlands, montane forests, 100–1000 m; Bolívar (Río Carapo on Cerro Guaquinima). Guyana, Suriname, French Guiana. ŠFig. 337.

The Venezuelan collections could belong to two undescribed species; more flowering material is needed to be certain. Spirotropis sp. A Small tree 8–10 m tall; petals purple. Gallery forests, along river margins and islands, 300–400 m; Bolívar (Río Canaracuni, Río Ichún). Endemic.

58. STYLOSANTHES Sw., Prodr. 108. 1788. by Gerardo A. Aymard C. Herbs, rarely subshrubs. Stems erect, ascending, or sprawling-procumbent, diffusely branched, terete, finely striate, unarmed. Leaves alternate, pinnately 3foliolate; leaflets linear-lanceolate, elliptic, or oblong, apices obtuse to acute, occasionally mucronate, the base attenuate, the costas prominent; petioles 1.5–15 mm long; stipules amplexicaul, pubescent, 2-lobed, 3–11-veined; stipels absent. Inflorescences spicate, terminal or axillary, 1–several-flowered, each flower surrounded by a series of bracts and bracteoles, the outermost bracts similar to stipules and giving rise to 1–3 reduced leaflets, bracts subtended by a smaller outer bracteole, usually 3veined, ciliate along the margin. Flowers 5-merous. Calyx 4–15 mm long, tubular, 5lobed; petals yellow or yellow-orange with purple stripes. Stamens 10, monadelphous; 5 versatile anthers alternating with 5 subbasifixed ones. Ovary subsessile, placentation marginal; ovules 2(3); style long, filiform; stigma minute, terminal. Fruit a loment, with 2 articles, distally fertile, proximally abortive or fertile, reticulate or muricate, beaked. Seeds ovate or lenticular, compressed, light brown to black, lustrous. U.S.A., Mexico, Neotropics (most diverse), introduced and naturalized in Malaysia and Australia; ca. 50 species, 10 in Venezuela, 6 of these in the flora area. The taxonomy of some of the taxa included here probably will not be resolved until the genus has been revised. Key to the Species of Stylosanthes 1. 1. 2(1).

2.

Leaflets linear to linear-lanceolate ............................................................ 2 Leaflets lanceolate to elliptic or oblong ..................................................... 3 Spike densely capitate; bracts hispid with golden trichomes; loment 1.5– 2.5 cm wide, with sessile glands, glabrous or with very short pubescence (tuberculate) ................................................................. S. guianensis Spike elongate; bracts ciliate, otherwise nearly glabrous; loment 0.8– 1.5 cm wide, minutely pubescent ........................................ S. angustifolia

Stylosanthes 393

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

6.

Branches densely pubescent to laxly pilose with short, viscid, sometimes glandular black trichomes ........................................................... S. viscosa Branches glabrous to puberulent or pilose, without glandular black trichomes .................................................................................................... 4 Stems usually ascending or sometimes prostrate; beak of fruit 1–3.5 mm long, strongly hooked ............................................................................. 5 Stems erect, rarely scandent and prostrate; beak of fruit 0.2–0.8 mm long, erect or shortly hooked .......................................................................... 6 Leaflets glabrous to lax-pilose on both surfaces, apex obtuse to subacute; flowers with 2 inner bracteoles .................................................. S. hamata Leaflets short-bristly-ciliate to nearly glabrous on both surfaces, apex acute; flowers with 1 inner bracteole ......................................... S. humilis Spikes capitate to elongate, bracts 3–9-veined, glabrous to puberulent with bristly golden trichomes, outer bracteole glabrous to pilose; standard suborbiculate ................................................................. S. guianensis Spikes capituliform, bracts 11–17-veined, copiously soft-pubescent, outer bracteole ciliate; standard obovate ............................................ S. capitata

Stylosanthes angustifolia Vogel, Linnaea 12: 63. 1838. Erect herb, much-branched from the base; flowers yellow. Wet savannas, 100–300 m; Bolívar (Caicara, Cerro San Borja along lower Río Suapure, near Ciudad Bolívar, Maripa, lower Río Caroní, lower Río Parguaza); Apure, Aragua, Guárico, Sucre; Guyana, Suriname, French Guiana, Brazil. Stylosanthes capitata Vogel, Linnaea 12: 70. 1838. Stems erect, branched, to 1 m tall; flowers yellow. Open places, savannas, 50–100 m; Bolívar (near Ciudad Bolívar). Anzoátegui, Apure, Guárico, Monagas, Zulia; Brazil, Bolivia. Stylosanthes guianensis (Aubl.) Sw., Kongl. Vetensk. Acad. Nya Handl. 11: 296. 1789. —Trifolium guianense Aubl., Hist. Pl. Guiane 776, pl. 309. 1775. Widespread in Neotropics; 7 varieties, 3 in Venezuela, 2 of these in the flora area. Key to the Varieties of S. guianensis 1. Leaflets linear-lanceolate; inflorescences usually terminal; spikes densely capitate; loment with sessile glands (minutely tuberculate) or glabrous .............. ......................................... var. gracilis 1. Leaflets elliptic to lanceolate; inflorescences terminal or sometimes axillary; spikes loosely capitate; loment usually glabrous or with very short pubescence .................................... var. guianensis

Fig. 338. Stylosanthes guianensis var. gracilis

394

F ABACEAE

S. guianensis var. gracilis (H.B.K.) Vogel, Linnaea 12: 66. 1838. —Stylosanthes gracilis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 507, t. 596. 1823 [1824]. —Cadillo. Erect herb; flowers yellow. Savannas, meadows, 100–1100 m; Bolívar (widespread), Amazonas (Puerto Ayacucho, Sierra Parima). Anzoátegui, Guárico, Monagas, Sucre, Trujillo, Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 338. S. guianensis var. guianensis Semierect to erect herb; flowers yellow. Savannas, open places, 200–1100 m; Bolívar (Ciudad Bolívar, Gran Sabana, lower Río Caroní, Río Cuyuní, near Tumeremo), Amazonas (near Puerto Ayacucho, Río Ocamo, Sierra Parima). Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Mérida, Monagas, Portuguesa, Sucre, Trujillo, Zulia; Mexico, Central America, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. Stylosanthes hamata (L.) Taub., Verh. Bot. Vereins Prov. Brandenburg 32: 22. 1890. —Hedysarum hamatum L., Syst. Nat. ed. 10, 2: 1170. 1759. Ascending herb, or sometimes prostrate, often branched; flowers yellow. Savannas, open places, 50–400 m; Bolívar (Ciudad Piar, La Escalera, near Puerto Ordaz). Anzoátegui, Aragua, Falcón, Guárico, Lara, Nueva

Esparta, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; U.S.A. (Florida), West Indies, Guatemala, Panama, Colombia, Brazil. Stylosanthes humilis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 506, t. 594. 1823 [1824]. —Cola de Zorro. Herb, usually ascending, sometimes prostrate; flowers yellow. Open savannas with Trachypogon, 50–200 m; Bolívar (Caicara, near Ciudad Bolívar, Ciudad Piar, Maripa, lower Río Caroní), Amazonas (near Puerto Ayacucho). Apure, Barinas, Carabobo, Cojedes, Guárico, Mérida, Monagas, Nueva Esparta, Portuguesa, Sucre, Trujillo, Zulia; Mexico, Central America, West Indies, Colombia, Brazil, Bolivia, introduced and naturalized in Malaysia and Australia. Stylosanthes viscosa (L.) Sw., Prod. 108. 1788. —Hedysarum hamatum var. viscosum L., Pl. Jamaic. Pug. 20. 1759. —Cola de zorro, Oregano canaotero, Oregano montañero, Peludita. Ascending and spreading or prostrate herb; flowers yellow. Open savannas, 100– 900 m; Bolívar (Ciudad Bolívar, El Manteco, El Pao, Gran Sabana, near Maripa, Río Caroní, near Tumeremo, Upata). Anzoátegui, Barinas, Falcón, Guárico, Monagas, Nueva Esparta, Sucre, Trujillo, Zulia; U.S.A. (Texas, introduced in Illinois), Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina.

59. SWARTZIA Schreb., Gen. Pl. 2: 518. 1791, nom. cons. Possira Aubl., Hist. Pl. Guiane 934. 1775. Tounatea Aubl., Hist. Pl. Guiane 549. 1775. Rittera Schreb., Gen. Pl. 1: 364. 1789. Hoelzelia Neck., Elem. Bot. 3: 62. 1790. Riveria H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 266. 1825. Fairchildia Britton & Rose, N. Amer. Fl. 23: 348. 1930. Huertia Mutis ex Alva in Mutis, Diario de Observ. 1: 215. 1957. by Nidia L. Cuello A. and Richard S. Cowan Shrubs, trees, rarely lianas; stipules minute and needle-like to large and foliaceous. Leaves odd-pinnate, 1–many-foliolate; petiole and rachis winged to terete, often stipellate at insertion of leaflets. Inflorescence racemose to paniculate-racemose, borne on trunk, branches, or branchlets, axillary, or supra-axillary, solitary to fasciculate; bracts usually present at base of pedicels, bracteoles often present at base of or on pedicels. Calyx entire in bud, opening in 2–5 regular or irregular segments;

Swartzia 395

petal 1 or none, usually clawed. Stamens dimorphic (except for 2 species), larger ones 2–25, with larger filaments and anthers than the many smaller stamens. Gynoecium 1(–3)-pistillate; gynophore usually well developed; style usually present and either terminal on ovary or lateral to its long axis, occasionally absent. Fruit 1–manyseeded, oval, moniliform, or elliptic, thin- to thick-walled, coriaceous to woody. Seeds generally ovoid with crustose or chartaceous testa, mostly arillate, the aril white, yellow, or red. Neotropics; ca. 200 species, ca. 55 species in Venezuela, 52 of these in the flora area. Key to the Species of Swartzia 1. 1. 2(1). 2. 3(2).

3.

4(2). 4.

5(4).

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

7.

8(5). 8. 9(8). 9.

Leaves with a single leaflet, appearing simple ......................................... 2 Leaves with 3 or more leaflets ................................................................. 10 Stipules conspicuously developed, at least 4 mm long, persistent ........... 3 Stipules not conspicuous or caducous ....................................................... 4 Lower leaflet surface and inflorescence pilose, primary veins of the leaflets interconnected intramarginally; stipules foliaceous, lanceolate, 6.5–28 × 1.5–9 mm .................................................................... S. buntingii Lower leaflet surface glabrous, primary veins of the leaflets not interconnected in an intramarginal vein; stipules nonfoliaceous, 4–5 × 1 mm ................................................................................................... S. stipellata Inflorescence axes and flower buds glabrous; blades of leaflets lanceolate .......................................................................................................... S. sp. F Inflorescence axes and flower buds minutely strigulose, malpighiaceousstrigulose, sericeous, or densely pilose; blades of leaflets elliptic, oblong elliptic, broadly elliptic, or ovate ........................................................... 5 Inflorescence axes and flower buds sparingly minutely strigulose to sericeous, or malpighiaceous-strigulose, flower buds globose to elliptic in outline .................................................................................................... 6 Inflorescence axes and flower buds densely pilose-velutinous, flower buds oval in outline ........................................................................................ 8 Lower surface of leaflets golden-silky with appressed, contiguous and parallel trichomes at least along midrib ........................................ S. palustris Lower surface of leaflets glabrous or sparingly micro-strigulose or glaucous ......................................................................................................... 7 Blades of leaflets coriaceous, the lower surface glaucous or yellowish, the apex rounded and bluntly apiculate; calyx segment deciduous; style 1–2 mm long, not persistent in fruit .................................... S. vaupesiana Blades of leaflets chartaceous, the lower surface not glaucous or yellowish, the apex bluntly acute to acuminate; calyx segment persistent with the fruits; style 4.5–6 mm long, persistent in fruit ................... S. conferta Gynoecium densely villose marginally ......................................... S. maguirei Gynoecium glabrous ................................................................................... 9 Leaflets narrowly oblong to elliptic-oblong, 8–18(–22) × 3.5–6(–9) cm, margin of the leaflets usually narrowly revolute ................. S. floribunda Leaflets broadly oblong (18–)24–34 × 9–14 cm, margin of leaflet plane .......................................................................................................... S. sp. E

396

FABACEAE

10(1). 10. 11(10). 11. 12(11). 12. 13(12). 13. 14(13).

14.

15(13).

15.

16(12). 16. 17(16). 17. 18(17). 18. 19(18).

19.

20(17).

20. 21(11). 21.

22(21).

Lateral leaflets alternate .......................................................... S. microcarpa Lateral leaflets opposite ........................................................................... 11 Inflorescences axillary, simple racemes .................................................. 12 Inflorescences borne on trunk, branches, or branchlets, simple or panicled racemes ................................................................................................ 21 Lateral leaflets ≥ 2 pairs .......................................................................... 13 Lateral leaflets 1 or 2 pairs ...................................................................... 16 Gynoecium and young fruits pubescent; stipules linear-lanceolate or foliaceous, 5–15 mm long ............................................................................ 14 Gynoecium and young fruits glabrous or partly sericeous; stipules triangular or linear-attenuate, not foliaceous, < 5 mm .............................. 15 Leaflets 3 or 4 pairs, the blades 4–12 × 2.5–5.5 cm; base of leaflets rounded; gynoecium 2-pistillate, stipules deciduous on old leaves and present on new shoots, linear, pubescent, 5–10.5 mm long ......................... S. dipetala Leaflets 2 or 3 pairs, the blades 10.5–21 × 4.5–9.5 cm; base of leaflets acute to obtuse; gynoecium 1-pistillate; stipules persistent on old leaves, foliaceous, 10–15 mm long .................................................. S. sp. C Leaflets 2 or 3 pairs, often even-pinnate; petioles, leaf rachis, and inflorescense axes minutely strigulose to glabrescent; pedicels 12–23 mm long, minutely strigulose; ovary laterally sericeous ............... S. parvifolia Leaflets 4 or 5 pairs, odd-pinnate; petioles, leaf rachis and inflorescence axes densely brownish-sericeous; pedicels stout, 8–12 mm long, densely silky-pilose, clavately thickened upward; ovary glabrous ..................................................................................................... S. oedipus Leaf rachis with a distinct narrow wing .................................. S. arborescens Leaf rachis lacking a wing ....................................................................... 17 Inflorescence axes densely tomentose ..................................................... 18 Inflorescense axes glabrous or strigulose ................................................ 20 Lower surface of leaflets densely subappressed-pilose over entire surface ................................................................................................... S. roraimae Lower surface of leaflets glabrescent, or strigose on midrib .................. 19 Flower buds 6–10 mm wide, densely brownish silky-pilosulous, pedicels thick 2–4 mm wide, 12–25 mm long; lower surface of leaflets glanddotted, thinly strigose on midrib .............................................. S. aymardii Flower buds 4–5 mm wide, golden-sericeous, pedicels slender < 1 mm wide, 7–10 mm long; lower surface of leaflets glabrous over entire surface, not gland-dotted .................................................... S. brachyrachis Leaflets 1–3, oblong-elliptic, obtuse to rounded at base, lower surface of leaflets glaucous or yellowish, secondary veins not prominent on upper surface ................................................................................... S. vaupesiana Leaflets always 3, obovate-elliptic, angustate at base, lower surface not glaucous, secondary veins prominent on upper surface ................. S. sp. A Style terminal, i.e., continuing long axis of ovary; fruit moniliform, rarely elliptic-oblong ....................................................................................... 22 Style lateral, i.e., perpendicular to long axis of ovary or appearing so by abrupt curvature of ovary apex; fruit variously shaped, but never moniliform ............................................................................................ 39 Gynoecium densely pubescent or sericeous on all surfaces ................... 23

Swartzia 397

22. 23(22). 23. 24(23).

24. 25(24). 25.

26(23). 26. 27(26). 27.

28(26). 28. 29(28). 29. 30(28).

30.

31(30). 31. 32(22). 32. 33(32). 33. 34(32). 34. 35(34).

Gynoecium sparingly strigulose, only partly pubescent or glabrous ...... 32 Rachis marginate to winged ..................................................................... 24 Rachis terete, canaliculate, or lacking stipels ......................................... 26 Leaflets rigid, bullulate by the impressed venation on the upper surface, the veins strongly salient on the lower surface; ovary tomentose, gynophore glabrous ..................................................................... S. pachyphylla Leaflets chartaceous to thinly coriaceous, not at all bullulate; ovary sericeous; the gynophore pubescent .......................................................... 25 Mature buds verruculose; petal surface obscured dorsally by the dense pubescence; fruit ± oblong, ca. 4 cm wide, obliquely costate .... S. laxiflora Mature buds costulate longitudinally or smooth; petal sericeous on the back, at least on the veins; fruit moniliform, ca. 2 cm broad, ± verrucose but not obliquely costate ....................................................... S. grandifolia Young branchlets strongly sulcate, the stipules very broad, conspicuous .............................................................................................................. 27 Young branchlets not sulcate, the stipules narrower and not conspicuous .............................................................................................................. 28 Stipules acute at apex; rachis of leaves terete, stipellate at each pair of leaflets, densely light brown to golden-tomentose ................... S. laxiflora Stipules rounded at apex; rachis of leaves canaliculate on upper surface, not stipellate at each pair of leaflets, densely dark brown-pannose ................................................................................................... S. panacoco Lower surface of leaflets densely puberulent microscopically and buff-colored ....................................................................................................... 29 Lower surface of leaflets glabrous or minutely and sparingly striguloseglabrescent ........................................................................................... 30 Lower surface of leaflets with 2 types of trichomes, peltate, short-stalked ones and much longer, subulate, simple ones ........................ S. jenmannii Lower surface of leaflets uniformly micro-puberulent, trichomes peltate, short-stalked ...................................................................... S. schomburgkii Rachis of leaves slightly canaliculate on upper surface; anther connectives apiculate, at least on the larger stamens; fruit moniliform, each section 1–2 × 1.5 cm .................................................. S. cardiosperma Rachis of leaves terete, marginate or shortly winged; anther connectives obtuse, not apiculate; fruit elliptic to oblong in outline, 4.5–9 × 2– 2.5 cm, if moniliform, each section 2.5–4.5 × 1.5–2 cm ...................... 31 Pedicels > 15 mm long; leaflets 1- or 2-jugate; filaments of larger stamens 17–22 mm long ............................................................................ S. pinnata Pedicels mostly 7–10 mm long; leaflets 3- or 4-jugate; filaments of larger stamens 5–9 mm long ............................................................ S. leptopetala Lateral leaflets 2 or 3 pairs ...................................................................... 33 Lateral leaflets 4–8 pairs ......................................................................... 34 Leaflets obovate to obovate-elliptic; stipules nonfoliaceous .......... S. leiogyne Leaflets elliptic; stipules foliaceous ......................................... S. macrocarpa Young branchlets strongly ribbed; lateral leaflets 5–8 pairs .................. 35 Young branchlets not strongly ribbed; lateral leaflets 4 or 5 pairs ........ 36 Leaflets 6–8 pairs, blades narrowly lanceolate or linear lanceolate, 2.5– 6 × 0.8–1.8 cm; ovary laterally and ventrally densely pale sericeous, dor-

398

FABACEAE

35. 36(34). 36. 37(36). 37. 38(36).

38. 39(21). 39. 40(39). 40. 41(40).

41. 42(40).

42.

43(39). 43. 44(43). 44. 45(44). 45. 46(43). 46. 47(46). 47. 48(47). 48. 49(48).

sally glabrescent .......................................................................... S. cowanii Leaflets 5 or 6 pairs, blades oval to elliptic to narrowly elliptic to oblongelliptic 4–18.5 × 2.5–8 cm; ovary glabrous over entire surface ..... S. sprucei Leaflets glabrous on lower surface .......................................................... 37 Leaflets strigulose or strigose to pilose, not glabrescent on lower surface .............................................................................................................. 38 Leaflets ± minutely warty; fruits woody, markedly white-lenticellate ................................................................................................. S. polyphylla Leaflets not at all warty; fruits neither woody nor white-lenticellate .................................................................................................. S. cuspidata Blades of the lowermost pair of leaflets usually oval 7.7–10(–15) × 3.5– 6 cm, the blades of the other pairs elliptic to elliptic-oblong to oblong or obovate-oblong, 13.5–23 × 4.5–9 cm; petal blade sinuate-dentate .......................................................................................................... S. picta Blade of all leaflets elliptic to obovate-elliptic, 3–9.5 × 1.5–2.5 cm; petal blade not sinuate-dentate ......................................................... S. piarensis Rachis of leaves distinctly winged ........................................................... 40 Rachis of leaves unwinged, terete, marginate or stipellate at each pair of leaflets .................................................................................................. 43 Lateral leaflets 2 or 3 pairs ...................................................................... 41 Lateral leaflets 6–11 pairs ........................................................................ 42 Calyx densely pubescent on inner surfaces; apex of leaflets bluntly acuminate; filaments of larger stamens glabrous or slightly strigulose ................................................................................................ S. guianensis Calyx glabrous on inner surfaces; apex of leaflets abruptly acute to acuminate; filaments white-villose .............................................. S. steyermarkii Lateral leaflets 6 or 7 pairs; wing segment of rachis herbaceous, with a deltate and sharply mucronate tip at the insertion of each pair of leaflet; valves of pods smooth and coriaceous .......................... S. alato-sericea Lateral leaflets 10 or 11 pairs; wing segments coriaceous and revolute, mostly with a rounded, slightly mucronate tip; valves of pods corrugated and woody .............................................................................. S. sp. B Leaf rachis stipellate ................................................................................ 44 Leaf rachis lacking wings and stipels ..................................................... 46 Inner surfaces of calyx silky; filaments of larger stamens pubescent ....................................................................................................... S. sericea Inner surfaces of calyx glabrous .............................................................. 45 Leaflets usually 2–4-jugate, elliptic to ovate ............................ S. tessmannii Leaflets 6- or 7-jugate, narrowly oblong .............................. S. angustifoliola Gynoecium glabrous ................................................................................. 47 Gynoecium pubescent .............................................................................. 50 Lateral leaflets 1 pair........................................................................... S. sp. D Lateral leaflets 2 or more pairs ............................................................... 48 Leaflets 2 pairs, blades conspicuously reticulate-venose and elevated on upper surface .......................................................................... S. wurdackii Leaflets (2)3–5 pairs; veins of blades on upper surface plane or obscure or slightly impressed ................................................................................ 49 Bracteoles lacking; blades of leaflets bluntly acute to rounded-obtuse at apex ............................................................................................... S. pittieri

Swartzia 399

49. 50(46). 50. 51(50). 51. 52(51). 52. 53(52).

53.

54(53).

54.

Bracteoles present, foliaceous; blades of leaflets caudate-cuspidate at apex, the cauda linear ................................................................ S. caudata Principal lateral veins of leaflets joined submarginally into a ± conspicuous vein; leaflets golden- or silvery-silky on lower surface ...... S. argentea Principal lateral veins not forming a submarginal vein; leaflets not silky on lower surface, strigose, strigulose, or glabrescent ......................... 51 Median leaflets 27 × 12 cm, strongly venose on both sides; petioles ca. 8 mm diameter at base ...................................................... S. benthamiana Median leaflets smaller, venation less marked; petioles more slender .............................................................................................................. 52 Mature buds warty; fruit with anastomosing, strongly salient, nearly transverse ridges ............................................................... S. benthamiana Mature buds not warty; fruit warty or smooth ....................................... 53 Buds oval, densely golden-silky, ca. 10 mm diameter; leaflets densely strigulose on lower surface; inflorescence axes densely yellowish or golden appressed-pubescent; valves of pods golden-sericeous in inmature pods, papillose-verrucose to lenticellate in mature pods ........... S. cupavenensis Buds globose or obnapiform, sparingly strigulose, 5–7 mm diameter; leaflets usually glabrous; inflorescence axes thinly minutely silky-strigulose or glabrescent, never yellowish or golden-pubescent, valves of pods glabrous and smooth or glabrous and minutely warty but not verrucose nor lenticellate in mature pods ................................................ 54 Ovary mostly glabrous except for few trichomes at base and along sutures; valves of mature pods smooth, at middle shallowly sulcate lengthwise and the sutures developed into wings, 2 on the adaxial side and 1 on the abaxial side ............................................................ S. triptera Ovary densely pubescent in all surfaces; valves of mature pods minutely warty, not sulcate lengthwise and the sutures unmodified ..... S. laevicarpa

Swartzia alato-sericea Barneby, Ann. Missouri Bot. Gard. 78: 177. 1991. Tree ca. 18 m tall. Lowland nonflooded forests, 200–400 m; expected in Amazonas. Brazil (Amazonas: known only from the southwestern base of Serra da Neblina). Swartzia angustifoliola Schery, Fieldiana, Bot. 28: 262. 1952. —Chamanare de tierra firme. Tree ca. 10 m tall. Nonflooded, riparian forests, 100–200 m; Amazonas (Capibara, San Carlos de Río Negro, Yavita to Maroa road). Endemic. ŠFig. 339. Swartzia arborescens (Aubl.) Pittier, J. Wash. Acad. Sci. 11: 157. 1921. —Possira arborescens Aubl., Hist. Pl. Guiane 934. 1775. —Guamo, Kajehai-detidi (Yekwana), Sibara-koni. Possira triphylla Sw., Prodr. 82. 1788. Rittera dodecandra Vahl, Symb. Bot. 2:

60. 1791. —Swartzia dodecandra (Vahl) Willd., Sp. Pl. 2: 1220. 1799. Swartzia triphyllata Willd., Sp. Pl. 2: 1220. 1799. Swartzia parviflora DC., Prod. 2: 423. 1825. Swartzia bifida Steud., Flora 26: 757. 1843. Swartzia rariflora Hoehne, Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 12: 16. 1922. Shrub or tree 3–10 m tall. Riparian forests, secondary forests, 50–700 m; Bolívar (mouth of Río Antavari, Río Nichare, Santa María de Erebato), Amazonas (widespread). Carabobo; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 342. Swartzia argentea Spruce ex Benth. in Mart., Fl. Bras. 15(2): 31. 1870. Colombia, Venezuela, Brazil; 2 varieties, 1 in Venezuela.

400

FABACEAE

S. argentea var. argentea. —Chamanare orillero, Jabón de raya, Macuca grande, Palo de chamanare. Tree 3–20 m tall; lower surface of leaves silver-brown; petal yellow. Permanently flooded forests along streams and rivers, 100–200 m; Amazonas (Caño San Antonio on upper Río Orinoco, Río Cuao, Río Negro-Río Casiquiare basin, Río Sipapo). Northwestern Brazil (Amazonas: Rio Negro basin). ŠFig. 345. The resin from the trunk is used to treat sting-ray wounds. Swartzia aymardii Barneby, Ann. Missouri Bot. Gard. 78: 178. 1991. Tree ca. 5–8 m tall; standard white. Brushy savannas, gallery forests, 700–800 m; Bolívar (northern slope of Sierra Pacaraima near source of Río Icabarú). Endemic. Swartzia benthamiana Miq., Stirp. Surinam. Select. 15. 1850 [1851]. Swartzia rosea Mart. ex Benth. in Mart., Fl. Bras. 15(2): 32. 1870. Occasional in riparian forests, 100–400 m. Southeastern Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, northwestern Brazil; 2 varieties, both in the flora area. Key to Varieties of S. benthamiana 1. Median leaflets smaller and venation less marked than below; petioles < 8 mm diameter at base; surface of fruits with anastomosing, strongly salient, nearly transverse ridges .. var. benthamiana 1. Median leaflets ca. 27 × 12 cm, strongly venose on both sides, third degree venules forming coarse reticulum; petiole ca. 8 mm diameter at base; surface of fruits golden-sericeous and verrucose, without ridges .............. var. yatuensis S. benthamiana var. benthamiana. —Chamanare. Tree 6–15 m tall. Riparian forests, 50–300 m; Amazonas (Guayapo on lower Río Caura, Río Autana, Río Guainía, Río Siapa, Río Sipapo, Río Ventuari, San Carlos de Río Negro). Southeastern Colombia, Guyana, Suriname, French Guiana, northwestern Brazil.

S. benthamiana var. yatuensis R.S. Cowan, Fl. Neotrop. Monogr. 1: 137. 1968. Tree 10–30 m tall. Riparian forests, 100– 200 m; Amazonas (Río Baría, Río Mawarinuma, Río Siapa, Río Yatúa). Endemic. Swartzia brachyrachis Harms, Notizbl. Königl. Bot. Gart. Berlin 6: 309. 1915. Western Colombia, Venezuela, Guyana, eastern Peru, Brazil; 5 varieties, 1 in Venezuela. S. brachyrachis var. glabrata R.S. Cowan, Fl. Neotrop. Monogr. 1: 192, figs. 39, 4a, 4b. 1968. Tree or shrub 3–15 m tall; fruit orange. Riparian forests, sometimes in dry secondary forests, forested slopes, 50–700 m; Bolívar (El Abismo, Río Samay). Guyana. Swartzia buntingii R.S. Cowan, Fl. Neotrop. Monogr. 1: 204. 1968. Tree 8–20 m tall; flowers yellowish; fruits green to orange. Riparian forests, 100–200 m; Amazonas (Río Cunucunuma, Río Mawarinuma, Río Paciba, Río Yatúa). Endemic. ŠFig. 341. Swartzia cardiosperma Spruce ex Benth. in Mart., Fl. Bras. 15(2): 33. 1870. —Chamanare, Guamo, Macuca montañero, Palo de morrocoy. Tree 6–20 m tall, cauliflorous; flowers yellow. Flooded and nonflooded rain forests, 100–300 m; Amazonas (Caño San Miguel, Cerro Huachamacari, La Esmeralda, Río Baría, Río Casiquiare, lower Río Guainía, Río Mawarinuma, Río Negro, upper Río Orinoco, Yavita to Maroa road). Southeastern Colombia, eastern Peru, northwestern Brazil. Swartzia caudata R.S. Cowan, Fl. Neotrop. Monogr. 1: 122. 1968. —Mudujai, Shi-ñateu. Tree 4–25 m tall. Forests along rivers, 100–400 m; Amazonas (near Cerro Duida, near Cerro Huachamacari, Ocamo, Río Cunucunuma, upper Río Orinoco, Río Padamo, Río Ventuari). Endemic. Swartzia conferta Spruce ex Benth., Fl. Bras. 15(2): 20. 1870.

Swartzia 401

Southeastern Colombia, Venezuela, northwestern Guyana, northwestern Brazil; 2 varieties, 1 in Venezuela.

pure), Amazonas (near Galipero, Isla Ratón, near Samariapo, San Juan de Manapiare). Apure; Guyana, Brazil.

S. conferta var. conferta Shrub or tree 1.5–15 m tall; flowers white; fruit orange. Riparian and secondary forests, 50–700 m; Bolívar (Río Cuyuní), Amazonas (Cerro Duida, Cerro Huachamacari, Río Casiquiare-Río Negro basin, Río Cunucunuma). Southeastern Colombia, northwestern Guyana, northwestern Brazil. ŠFig. 340.

Swartzia floribunda Spruce ex Benth. in Mart., Fl. Bras. 15(2): 21. 1870. —Anón, Guaquito, Palo de golondrona. Shrub or small tree 1.5–10 m tall; flowers white. Lowland forests, bushy savannas, secondary forests, 50–200 m; Amazonas (Caño Caname, Maroa, Río Atabapo, Río Atacavi, Río Casiquiare, Río Guainía, Río Negro, Río Temi, Yavita). Endemic.

Swartzia cowanii Steyerm., Brittonia 33: 33. 1981. Tree 4–15 m tall, cauliflorous; flowers yellow. Near rivers on tepui summits, 700–800 m; Bolívar (Cerro Guaiquinima). Endemic. ŠFig. 344. Swartzia cupavenensis R.S. Cowan, Fl. Neotrop. Monogr. 1: 139. 1968. —Dudu, Guayaparu, Guayapu. Tree 6–15 m tall; petal cream-white. Forests, 100–300 m; Amazonas (Caño Caname, near the mouth of Caño Cupaven, Caño Soromoni near La Esmeralda, Caño Yagua, Isla Cangrejo near Cerro Duida, Río Cuao, Río Guayapo, Río Orinoco, Río Pasimoni, Río Sipapo, Río Ventuari, Río Yatúa, near San Carlos Río Negro). Endemic. Swartzia cuspidata Spruce ex Benth. in Mart., Fl. Bras. 15(2): 36. 1870. —Chamanare. Shrub or tree 3–18 m tall, cauliflorous; flowers white. Riparian, roadside, and secondary forests, 100–700 m; Amazonas (Río Casiquiare, Río Negro basin, Sierra de la Neblina, Yavita to Maroa road). Eastern Peru, Amazonian Brazil. Swartzia dipetala Willd. ex J. Vogel, Linnaea 11: 173. 1837. —Miriyon (Panare), Panillo, Sowi (Yekwana). Swartzia dicarpa Moric. ex Meisn., Pl. Vasc. Gen., Commentarius 68. 1837. Swartzia microstylis Benth., J. Bot. (Hooker) 2: 89. 1840. Tree 5–30 m tall; fruit orange; petals yellow; flowers fragrant. Riparian forests, semideciduous forests, granitic slopes, savannas, 50–200 m; Bolívar (lower Río Caura, Río Maniapure, Río Parguaza, lower Río Sua-

Swartzia grandifolia Bong. ex Benth., J. Bot. (Hooker) 2: 85. 1840. —Akayuwajunai, Lawadema-eje-de-deu, Yawademo (Yekwana). Tree 6–18 m tall, cauliflorous; flowers creamy-white to yellow with purple veins. Lowland forests along rivers on clay, sandy clay, or rocky soils above level of seasonal flooding, 100–500 m; Bolívar (Río Caura, Río Suapure), Amazonas (Río Casiquiare, Río Cunucunuma, upper Río Orinoco, Río Ventuari). Guyana, Suriname, French Guiana, northcentral and northwestern Brazil. ŠFig. 346. Swartzia guianensis (Aubl.) Urb., Symb. Antill. 5: 365. 1908. —Tounatea guianensis Aubl., Hist. Pl. Guiane 550. 1775. —Dawadema, Majako shodo (Yekwana). Swartzia alata Willd., Sp. Pl. 2: 1220. 1799. Tree 6–15 m tall. Understory of forests on sandy or lateritic clay soil, riparian forests, forested slopes, 100–400 m; Amazonas (Cerro Huachamacari, Río Cunucunuma). Guyana, Suriname, French Guiana. The cortex of Swartzia guianensis is used by Yekwana Amerindians to treat skin infections. Swartzia jenmanii Sandwith, Bull. Misc. Inform. Kew 1934: 361. 1934. Large tree. Forested slopes, 800–1100 m; Bolívar (Uei-tepui). North-central Guyana. Swartzia laevicarpa Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 37. 1939. Tree 4–30 m tall; fruit gray-green. Widespread in lowland, high forests along creeks

402

FABACEAE

and rivers, usually on sandy soils but also on sandy clay, mostly in nonflooded locations, also in scrub forest on summit of tepui, forested slopes of tepui with to 20 m trees, 100– 1000 m; Bolívar (Amaruay-tepui), Amazonas (Isla Maracá in Río Negro, Río Mawarinuma). Southeastern Colombia, Guyana, Suriname, northwestern Brazil. Swartzia laxiflora Bong. ex Benth., J. Bot. (Hooker) 2: 86. 1840. —Tounatea laxiflora (Bong. ex Benth.) Taub., Bot. Centralbl. 47: 391. 1891. —Tounatea laxiflora (Bong. ex Benth.) Kuntze, Revis. Gen. Pl. 1: 211. 1891. —Chamanare. Swartzia polycarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 126. 1922. Tree 4–15 m tall; flowers orange. Frequent in nonflooded riparian lowland forests. 50– 100 m; Amazonas (Río Cuao, Río Guayapo, Río Sipapo). Brazil (Amazonas). Swartzia leiogyne (Sandwith) R.S. Cowan, Fl. Neotrop. Monogr. 1: 46. 1968. —Swartzia grandifolia var. leiogyne Sandwith, Bull. Misc. Inform. Kew 1937: 103. 1937. Tree 13–22 m tall. Lowland to lower montane forests, 50–400 m; expected to be found in Amazonas. Guyana, Brazil (Roraima). Swartzia leptopetala Benth., J. Bot. (Hooker) 2: 87. 1840. —Carrasposo, Chamanare, Congrio, Yawademo de rebalse. Swartzia discolor Poepp., Nov. Gen. Sp. Pl. 3: 62. 1845. Swartzia fugax Spruce ex Benth. in Mart., Fl. Bras. 15(2): 30. 1870. Swartzia melanoxylon Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 123. 1922. Swartzia rotundata R.S. Cowan, Mem. New York Bot. Gard. 8: 116. 1953. Tree 5–30 m tall; flowers yellow. Frequent in riparian forests, 50–300 m; Bolívar (Río Asa, Río Caroní, Río Caura, Río Erebato), Amazonas (Cerro Yutajé, Maroa, Río Casiquiare, Río Cataniapo, Río Coro Coro, Río Guainía, Río Matacuni, Río Ocamo, Río Orinoco). Anzoátegui, Apure, Miranda, Monagas, Yaracuy; southeastern Colombia, Brazil. ŠFig. 347. Swartzia macrocarpa Spruce ex Benth. in Mart., Fl. Bras. 15(2): 38. 1870.

Tree to 5 m tall. Riparian, seasonally or permanently flooded forests, 100–200 m; expected to be found in Amazonas. Northwestern Brazil (Amazonas: Rio Negro Basin). ŠFig. 350. Swartzia maguirei R.S. Cowan, Mem. New York Bot. Gard. 8: 115. 1953. —Guaquito. Tree 3–10 m tall; fruit orange. Evergreen lowland forests, 100–300 m; Amazonas (Cerro Sipapo, Culebra, El Gavilán near Puerto Ayacucho, Río Cunucunuma, Río Coromoto, Río Yureba, near San Carlos Río Negro). Brazil (Amazonas). Swartzia microcarpa Spruce ex Benth., Fl. Bras. 15(2): 35. 1870. —Chamanare negro. Tree to 25 m tall. Flooded plains, 50–200 m; Amazonas (Río Guayapo, Río Pasimoni, Río Sipapo). Colombia, Brazil. Swartzia oedipus Barneby, Ann. Missouri Bot. Gard. 78: 179, fig. 1. 1991. Tree to 20 m tall; trunk to 20 cm diameter. Nonflooded forests, 100–500 m; Amazonas (Río Negro near Piedra Cocuy). Brazil. Swartzia pachyphylla Harms, Notizbl. Königl. Bot. Gart. Berlin 6: 310. 1915. Tree 10–15 m tall. Gallery forests, ca. 1400 m; Bolívar (Roraima-tepui). Endemic. Swartzia palustris Barneby, Ann. Missouri Bot. Gard., 78: 179. 1991. —Wa-hú. Slender tree 3–25 m tall, when crowded sometimes producing long and lithe branches. Swamp forests, rain forests on low hills, riparian forests along black-water streams, 100–400 m; Amazonas (upper Río Baría, Río Cunucunuma, southwestern base of Sierra de la Neblina). Endemic. Swartzia panacoco (Aubl.) R.S. Cowan, Fl. Neotrop. Monogr. 1: 32. 1968. —Robinia panacoco Aubl., Hist. Pl. Guiane 768. 1775. Robinia tomentosa Willd., Sp. Pl. 3(2): 1134. 1802. —Swartzia tomentosa (Willd.) DC., Prod. 2: 423. 1825. Swartzia similis Benoist, Bull. Mus. Hist. Nat. (Paris) 25: 297. 1919. Tree. Upland and highland forests. Venezuela, Guyana, Suriname, French Guiana,

Swartzia 403

Brazil; 8 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of S. panacoco 1. Leaflets coriaceous, strigulose on lower surface .......................... var. cardonae 1. Leaflets rigid, tomentose on lower surface ..................................... var. tepuiensis S. panacoco var. cardonae (R.S. Cowan) R.S. Cowan, Fl. Neotrop. Monogr. 1: 34. 1968. —Swartzia tomentosa var. cardonae R.S. Cowan, Bol. Soc. Venez. Ci. Nat. 13: 99. 1954. —Aka-yek, Chamanare, Guamo terciopelo, Kajari, Leche cochino, Okoi-yek (Pemón), Tasajo, Terciopelo. Tree 10–30 m tall. Primary, riparian forests on sandy nonflooded soils, 400–1200 m; Bolívar (Río Canaracuni, Río Caura, Río Erebato, Río Icabarú, upper Río Paragua, Río Tonoró, Río Venamo), Amazonas (Río Cunucunuma). Guyana, Brazil (Amazonas: near junction of Rio Amazon and Rio Negro). S. panacoco var. tepuiensis (Schery) R.S. Cowan, Fl. Neotrop. Monogr. 1: 37. 1968. —Swartzia tepuiensis Schery, Fieldiana, Bot. 28: 265. 1952. —Ve-yek. Tree 10–20 m tall. Slopes of sandstone tepuis, dwarf open forests, 1600–2000 m; Bolívar (Amaruay-tepui, Carrao-tepui, Gran Sabana, Río Aícha near Los Caribes, Río Oris in Río Paragua basin). Endemic. ŠFig. 352. Swartzia parvifolia Schery, Fieldiana, Bot. 28: 263. 1952. Tree 10–25 m tall. Summits of sandstone tepuis, 700–1700 m; Bolívar (El Paují, near Kavanayén, Macizo del Chimantá), Amazonas (base of Cerro Cuao, Río Cuao). Endemic. ŠFig. 349. Swartzia piarensis R.S. Cowan, Fl. Neotrop. Monogr. 1: 45. 1968. —Uoki-yek. Tree 8 m tall. Riparian forests, 500–800 m; Bolívar (Cerro Bolívar, Río Caroní near mouth of Río Icabarú). Endemic. Swartzia picta Spruce ex Benth. in Mart., Fl. Bras. 15(2): 25. 1870. Tree 8–13 m tall. Riparian forests, forested slopes of tepuis, 100–900 m. Venezuela, Brazil; 2 varieties, both in the flora area.

Key to the Varieties of S. picta 1. Leaflets strigulose on lower surface, attenuate basally, the apex shortly and sharply acute; pedicels cylindric, 20–35 mm long; bracteoles arising below pedicel apex ............. var. bolivarensis 1. Leaflets usually pilose on lower surface, obtuse basally, the apex obtuse or shortly and bluntly acute; pedicels clavate, 12– 24 mm long; bracteoles apical on pedicels or on base of calyx ................ var. picta S. picta var. bolivarensis R.S. Cowan, Fl. Neotrop. Monogr. 1: 45. 1968. 400–900 m; Bolívar (quebrada Los Brasileros near Icabarú, Río Chaberu, Río Samay). North-central Brazil (Amazonas: near Río Branco). ŠFig. 353. S. picta var. picta. —Chamanare. 100–200 m; Bolivar (Río Acanán), Amazonas (Río Negro, Río Guainía). Northwestern Brazil. ŠFig. 354. Swartzia pinnata (Vahl) Willd., Sp. Pl. 2: 1220. 1799. —Rittera pinnata Vahl, Eclog. Amer. 2: 38. 1798. Tree 7–15 m tall. Dry forests, 50–200 m; Amazonas (Gavilán near Puerto Ayacucho, Río Orinoco, Río Sipapo). Coastal Venezuela; Trinidad. Swartzia pittieri Schery, Fieldiana, Bot. 28: 263. 1952. Tree 7–10 m tall. Granitic, rocky areas, 50–200 m; Bolívar (50 km southwest of Caicara in Serranía La Encaramada), Amazonas (Caño Chimoni tributary of Río Casiquiare, near Puerto Ayacucho, Río Orinoco). Anzoátegui, Barinas, Guárico, Portuguesa; West-central and southwestern Venezuela. ŠFig. 351. Swartzia polyphylla DC, Prod. 2: 424. 1825. —Cajario, Candilón, Canjilón de agua, Dau bagibagi (Waroa), Guaraba, Palo de raya. Swartzia acuminata Willd. ex J. Vogel, Linnaea 11: 173. 1837. —Tounatea acuminata (Willd. ex J. Vogel) Taub., Bot. Centralbl. 47: 390. 1891. Tounatea acuminata var. puberula Taub., Flora 75: 81. 1892. —Swartzia acu-

404

FABACEAE

minata var. puberula (Taub.) Glaz., Bull. Soc. Bot. France 53. Mem. 36: 155. 1906. Swartzia acuminata var. tridynamia Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr. 2: 506. 1898. Swartzia platygyne Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 127. 1922. Swartzia opacifolia J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 226. 1943. Swartzia urubuensis Ducke, Bol. Técn. Inst. Agron. N. 2: 21 1944. Tree 3–40 m tall; flowers white. Evergreen forests, permanently flooded areas along rivers, seasonally flooded river margins, near sea level to 800 m; Delta Amacuro (Caño Araguabisi between Caño Araguao and Caño Güiniquina, Caño Güiniquina, Caño JobaSuburu east of Caño Sacupana, Caño Jobure, near Curiapo, Misión San Francisco de Guayo), Amazonas (Río Casiquiare, Río Cuao, Río Mawarinuma, Río Negro, base of Sierra de la Neblina). Colombia, Guyana, French Guiana, eastern Peru, Amazonian Brazil. ŠFig. 343. The wood of Swartzia polyphylla is used for paddles. Swartzia roraimae Sandwith, Bull. Misc. Inform. Kew 1934: 362. 1934. Tree of unknown stature. Forested slope of tepui, 1500–1600 m; Bolívar (southwestern face of Roraima-tepui). Endemic. Swartzia schomburgkii Benth. in Mart., Fl. Bras. 15(2): 38. 1870. —Dudu, Kanaguarajodi (Yekwana), Níspero, Oliverio, Para-cutaco, Paretaura, Pore-taurai, Pretaurayese (Pemón), Sánamo, Shodocona (Yanomami), Wvaraba (Yeral). Swartzia schomburgkii var. guayanensis R.S. Cowan, Fl. Neotrop. Monogr. 1: 90. 1968. Tree 10–30 m tall; flowers white. Riparian forests, 50–800 m; Bolívar (Cerro Camarón, El Cácaro on upper Río Caura, between El Dorado and Kilómetro 88, Río Erebato, Río Ichún, Río Karún, Río Nichare, Río Paragua), Amazonas (Caño Ucata southeast of Síquita, Río Atabapo, Río Negro-Río Cunucunuma, upper Río Orinoco, Río Siapa near base of Cerro Aracamuni, Río Ugueto, base of Sierra de la Neblina). Southeastern Colombia, Guyana, Suriname, northwestern Brazil.

Swartzia schomburgkii is treated here as including var. guayanensis since the differences in fruiting material are not consistent. Also, some specimens from the southern part of Amazonas state have a pubescent gynoecium, which means that the var. guayanensis is not restricted to Bolívar state as was thought. The green wood of Swartzia schomburgkii is cut into fine pieces and used to start fires. Swartzia sericea J. Vogel, Linnaea 11: 176. 1837. Southeastern Colombia, Venezuela, French Guiana, Brazil (Amazonas); 2 varieties, 1 in Venezuela. S. sericea var. sericea. —Chamanare, Chimanare, Cupana, Cupana de chamanare, Maca, Macuca, Tucana guarana. Tree 4–15 m tall; fruits orange, with thick white aril along seed. Riparian forests, 100– 600 m; Bolívar (El Dorado to Santa Elena), Amazonas (Caño Cupueni, Pimichín, Río Atabapo, Río Atacavi, Río Baría, Río Guainía, Río Negro-Río Casiquiare, Río Siapa, Río Sipapo, Río Temi, Río Ventuari, Río Yatúa, San Juan). Southeastern Colombia, French Guiana, Brazil (Amazonas: Rio Negro basin). ŠFig. 355. The seeds of this species are used for fish bait. Swartzia sprucei Benth. in Mart., Fl. Bras. 15(2): 37. 1870. Tree 3–30 m tall. Guyana, Venezuela; 2 varieties, both in the flora area. Key to the Varieties S. sprucei 1. Filaments of larger stamens sparingly villose; leaflets strigose but glabrescent and not tessellate on lower surface; leaf rachis distinctly stipellate at each pair of leaflets .......................... var. sprucei 1. Filaments of larger stamens glabrous; sparingly strigulose and usually minutely tessellate on lower surface; leaf rachis not at all stipellate ...... var. tessellata S. sprucei var. sprucei Tree ca. 10 m tall, cauliflorous. Riparian forests, 50–300 m; Bolívar (Río Caura near Salto Pará), Amazonas (Río Pasimoni). Endemic.

Swartzia 405

S. sprucei var. tessellata R.S. Cowan, Fl. Neotrop. Monogr. 1: 40. 1968. Lower slope of ironstone hill, 100–500 m; Bolívar (road between El Dorado and Kilómetro 88). Guyana. Swartzia steyermarkii R.S. Cowan, Fl. Neotrop. Monogr. 1: 160. 1968. Tree 4–10 m tall. Lowland to montane forest, 100–1000 m; Bolívar (Río Ayaiche near base of Sierra de Lema, Río Uroi). Endemic. ŠFig. 358. Swartzia stipellata R.S. Cowan, Fl. Neotrop. Monogr. 1: 211. 1968. Tree 5–7 m tall. Riparian forests, 100–200 m; Amazonas (Río Puruname). Brazil (Amazon basin). Swartzia tessmannii Harms, Notizbl. Bot. Gart. Berlin-Dahlem 9: 971. 1926. —Okiyeu (Pemón), Wadi (Yekwana). Tree 8–24 m tall; flowers yellow; fruit orange. Riparian forests, 200–500 m; Bolívar (Cerro Cimarrón on upper Río Caura, Cerro Sarisariñama, El Cácaro between Río Caura and Río Paragua, El Paují, upper Río Caura, Río Kurutú on Río Paragua, Río UenanIkabarú), Amazonas (upper Río Cunucunuma). Amazonian Peru and Brazil. ŠFig. 356. Swartzia triptera Barneby, Ann. Missouri Bot. Gard. 78: 181, fig. 2. 1991. Tree 4–12 m tall; petal white; fruit green, with sticky red exudate. Lowland nonflooded forests, riparian forests, 100–200 m; Amazonas (Cerro Marahuaka, Río Mawarinuma). Endemic. Swartzia vaupesiana R.S. Cowan, Fl. Neotrop. Monogr. 1: 126. 1968. —Hoja del sol. Swartzia dolichopoda R.S. Cowan, Fl. Neotrop. Monogr. 1: 130. 1968. Swartzia vaupesiana var. glauca R.S. Cowan, Fl. Neotrop. Monogr. 1: 127. 1968. Tree 6–15 m tall; petals orange-yellow. Riparian forests, 100–300 m; Amazonas (Caño Cabeza Manteco tributary of Río Autana, Caño Cupueini east of San Fernando de Atabapo, Río Autana, upper Río Orinoco, Río Sipapo, Río Yatúa, Sierra de la Neblina). Southeastern Colombia. ŠFig. 357.

Swartzia vaupesiana is treated here as including S. vaupesiana var. glauca and S. dolichopoda, because of the great resemblance of all parts observed in the type specimens of these three taxa. Variation in number of leaflets (1 or 3) seems to be common in S. vaupesiana. Swartzia wurdackii R.S. Cowan, Fl. Neotrop. Monogr. 1: 148. 1968. Tree 15 m tall. Forests at base of escarpments, 1300–1400 m; Bolívar (Macizo del Chimantá [base of Amurí-tepui]). Endemic. Swartzia sp. A. —Macho. Tree 12–22 m tall, with reddish exudate; fruits yellow to orange. Lower montane forests, 200–300 m; Delta Amacuro (Río Grande). Guyana, Suriname. ŠFig. 348. This taxon is based on Blanco 164 (VEN), Zabala 126 (VEN), and Marcano-Berti 232 (MO). Swartzia sp. B. —Chamanare. Tree 15–20 m tall; fruits orange. Nonflooded lower montane forests, 200–300 m; Amazonas (Río Yatúa). This taxon is based on Velazco 1420 (MO, PORT). Swartzia sp. C Small tree; fruit orange, seed black with white aril. Low open forest on granitic outcrop, ca. 600 m; Amazonas (Cerro Aracamuni). This taxon is based on Liesner & Carnevali 22271 (MO). Swartzia sp. D Tree 10 m tall; fruit bright orange, seed black with white aril. Riparian forest, ca. 300 m; Amazonas (Río Mawarinuma). This taxon is based on Davidse & Miller 27210 (MO). Swartzia sp. E Vining tree; flowers white. Nonflooded forests, 500–600 m; Amazonas (Caño Colorado and Serranía Batata in Cuao-Sipapo massif). This taxon is based on Sanoja 3200 and 3371 (MO, PORT). Swartzia sp. F Small tree. Lowland riparian forest, 50– 100 m; Amazonas (Caño Monomi in Río Casiquiare basin). This taxon is based on Vareschi 7793 (VEN).

406

F ABACEAE

Fig. 339. Swartzia angustifoliola

Fig. 340. Swartzia conferta var. conferta

Fig. 341. Swartzia buntingii

Swartzia 407

Fig. 342. Swartzia arborescens

Fig. 343. Swartzia polyphylla

408

F ABACEAE

Fig. 344. Swartzia cowanii

Fig. 345. Swartzia argentea var. argentea

Swartzia 409

Fig. 346. Swartzia grandifolia

410

F ABACEAE

Fig. 347. Swartzia leptopetala

Fig. 348. Swartzia sp. A

Swartzia

Fig. 349. Swartzia parvifolia

Fig. 350. Swartzia macrocarpa

411

412

F ABACEAE

Fig. 351. Swartzia pittieri

Fig. 352. Swartzia panacoco var. tepuiensis

Swartzia 413

Fig. 353. Swartzia picta var. bolivarensis Fig. 354. Swartzia picta var. picta

414

F ABACEAE

Fig. 355. Swartzia sericea var. sericea

Fig. 356. Swartzia tessmannii

Swartzia

Fig. 357. Swartzia vaupesiana

Fig. 358. Swartzia steyermarkii

415

416

FABACEAE

60. TARALEA Aubl., Hist. Pl. Guiane 745. 1775. by Haroldo Cavalcante de Lima and Gerardo A. Aymard C. Trees or shrubs. Leaves alternate or opposite, odd-pinnate or even-pinnate; leaflets alternate or opposite, pellucid-punctate or not; rachis cylindric or flattened, tip often projected beyond the leaflets; stipules small, caducous; stipels none. Inflorescence a terminal panicle. Calyx pellucid-punctate, bilabiate, upper lip with 2 large lobes, lower lip very shortly 3-dentate; petals violet; standard emarginate; keel petals adnate on lower side. Stamens 10, connate in a sheath open on upper side. Ovary short-stalked; ovule 1. Fruit elastically dehiscent, 2-valved, flat. Seed compressed-ovoid; cotyledons fused, convolutely wrinkled; radicle central at base of a deep, closed sinus. Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil; 5 or 6 species, 4 in Venezuela, all in the flora area. Key to the Species of Taralea 1. 1. 2(1).

2.

3(2). 3.

Leaflets thick-coriaceous, appressed-puberulent on lower surface and midrib ...................................................................................... T. crassifolia Leaflets coriaceous to thin-coriaceous, glabrous ....................................... 2 Leaflets with venation prominent and reticulate on both surfaces, apex obtuse or retuse-acute; upper lip of calyx obovate-orbicular or orbicular, 7–11 mm wide .................................................................... T. reticulata Leaflets with venation inconspicuous on upper surface, apex acute or acuminate; upper lip of calyx oblong or oblong-lanceolate, 3–6 mm wide ................................................................................................................ 3 Upper lip of calyx coriaceous, oblong-lanceolate; ovary densely pilose; fruit 5–7 × 3–4 cm; seed 2–2.5 × 1.5–2 cm .......................... T. oppositifolia Upper lip of calyx membranous, oblong to round; ovary pilose only at the margins; fruit 3–4 × 1.5–2 cm; seed 1–1.5 × 0.5–0.8 cm ............. T. cordata

Taralea cordata Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 71. 1925. —Dipteryx cordata (Ducke) R.S. Cowan, Mem. New York Bot. Gard. 10(1): 152. 1958. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 varieties, both in the flora area. Key to the Varieties of T. cordata 1. Leaflets 4–6(7), ovate or orbicular, the base cordate; rachis tip acute or obtuse ......................................... var. cordata 1. Leaflets 1–3(–5), ovate-lanceolate, the base acute; rachis tip subulate ............... ........................................... var. rigida T. cordata var. cordata Shrub to slender tree 1–8 m tall. Wet lowland and montane forests, 200–1400 m;

Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Sipapo). Colombia, Guyana, Suriname, French Guiana, Brazil. ŠFig. 359. T. cordata var. rigida (Schery) H.C. Lima, comb. & stat. nov. —Taralea rigida Schery, Fieldiana, Bot. 28: 266. 1952. —Dipteryx rigida (Schery) R.S. Cowan, Mem. New York Bot. Gard. 10(1): 15. 1958. Slender shrub 0.5–3 m tall, with elongate stems. Savannas, 100–1500 m; Bolívar (Icabarú, Kavanayén), Amazonas (Cerro Cucurito, Cerro Yapacana, Río Temi, San Fernando de Atabapo). Brazil (Amazonas: Serra Aracá). Taralea crassifolia (Benth.) Ducke, Rev. Int. Bot. Appl. Agric. Trop. 14: 407. 1934. —Dipteryx crassifolia Benth., J. Bot. (Hooker) 2: 235. 1840.

Taralea 417

Fig. 359. Taralea cordata var. cordata

Fig. 360. Taralea crassifolia

418

FABACEAE

Fig. 361. Taralea oppositifolia

Fig. 362. Taralea reticulata

Tephrosia 419

Shrub or tree 2–20 m tall; flowers purple. Savannas on sandstone, evergreen forests, 500–1100 m; Bolívar (Auyán-tepui, Cerro Guacamaya, Cerro Guaiquinima, Cerro Mahedi, El Paují, 26.5 km east of Icabarú, Río Asa, upper Río Caroní, Río Paragua, upper Río Supamo), Amazonas (Caño Iguapo, Cerro Mahedi, Río Coro Coro). Guyana, Suriname, French Guiana, Brazil. ŠFig. 360. Taralea oppositifolia Aubl., Hist. Pl. Guiane 745. 1775. —Dipteryx oppositifolia (Aubl.) Willd., Sp. Pl. 3(2): 910. 1802. —Coumarouna oppositifolia (Aubl.) Taub., Bot. Centralbl. 47: 389. 1891. —Arepillo, Zapatero. Dipteryx applanata Benth., J. Bot. (Hooker) 2: 234. 1840.

Dipteryx oppositifolia var. parviflora Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 126. 1860. Tree 8–30 m tall; flowers purple. Forests on black-water river banks, 100–400 m; Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 361. Taralea reticulata (Benth.) Ducke, Rev. Int. Bot. Appl. Agric. Trop. 14: 407. 1934. —Dipteryx reticulata Benth., J. Bot. (Hooker) 2: 235. 1840. Taralea steyermarkii Schery, Fieldiana, Bot. 28: 268. 1952. Tree 5–30 m tall. Savannas, shrubby forests along savannas, 1000–1400 m; Bolívar (Ikabarú, Kavanayén, upper Río Aponguao, Río Karaurín). Guyana. ŠFig. 362.

61. TEPHROSIA Pers., Syn. Pl. 2: 328. 1807. by Gerardo A. Aymard C. Herbs or shrubs, erect or trailing; roots usually thick; pubescence usually dense and matted. Leaves alternate, odd-pinnate; leaflets (1 or 2)3–41, almost always pubescent, at least on lower surface; rachis usually grooved on upper surface; exstipellate but tufts of trichomes sometimes present in the axils; stipulate. Inflorescence terminal or axillary, racemose, elongate, the flowers in clusters of 2–6 or more at the nodes, each cluster usually with a primary bract at the base and each pedicel with a secondary bract at the base. Flowers red, purple, or white. Petals clawed; standard hairy outside, often densely so; wing petals about as long as the standard and usually basally adnate to the keel petals. Stamens usually diadelphous, the vexillar stamen frequently fused to the stamen tube above, free at the base. Ovary sessile, slender, usually hairy; style bearded above in many species, or glabrous. Fruit flat, linear or oblong, straight or slighty upcurved, often obliquely contracted distally, beaked, nonseptate or occasionally slightly septate, usually coiling in dehiscence. Seeds several to many, circular to oblong, flattened. Temperate North America, tropics and subtropics (most diverse in Mexico, Africa, and tropical Australia); ca. 400 species, 8 in Venezuela, 5 of these in the flora area. Many species produce rotenone and related compounds which are used as fish poisons and insecticides. The vernacular name “barbasco” is used for some of the fish-poisoning species throughout the Neotropics. The taxonomy of some of the taxa included here probably will not be resolved until the genus has been revised. Key to the Species of Tephrosia 1. 1.

Herbaceous plants, either erect annuals, prostrate to ascending herbs, or climbing herbaceous vines ..................................................................... 2 Erect perennial herbs with woody base, subshrubs, or shrubs to 3.5 m tall ................................................................................................................ 3

420

2(1).

2.

3(1).

3. 4(3).

4.

FABACEAE

Stems, inflorescences, and leaflets on both surfaces brownish yellow-pubescent; stipules 7–10 mm long; inflorescence 15–30 cm long; legume 12–15-seeded ................................................................................ T. adunca Stems, inflorescences, and lower surfaces of leaflets silvery-strigose or laxely pilose; stipules 3–5 mm long; inflorescence 5–13 cm long; legume 8–10-seeded .................................................................................. T. cinerea Erect shrubs to 3.5 m tall; leaves 15–30 cm long; leaflets 15–40; calyx 5–8 mm long, densely pubescent with long yellow-brown trichomes ..................................................................................................... T. sinapou Erect herbs or subshrubs to 1.5 m tall; leaves 3–10 cm long; leaflets 3–9; calyx 2–4 mm long, shortly sericeous, pubescent, or laxely pilose ...... 4 Leaflets 5–9, oblong-obovate, appressed-pubescent on both surfaces or glabrate; terminal leaflet 2.5–4 cm long; flowers purple or red-purple ......................................................................................................... T. senna Leaflets 3, lanceolate-obovate, the lower surface with silvery-silky pubescence; terminal leaflet 5–10 cm long; flowers yellow ........... T. sessiliflora

Tephrosia adunca Benth., Ann. Nat. Hist. 3: 431. 1839. Decumbent herb, stems ascending to 50 cm tall; flowers rose-pink. Open savannas, 50–200 m; Bolívar (Maripa, lower Río Caura). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Guárico, Monagas, Portuguesa, Sucre; Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina, Uruguay.

Galega cathartica Sessé & Moc., Fl. Mexic. ed. 2, 175. 1894. —Tephrosia cathartica (Sessé & Moc.) Urb., Symb. Antill. 4: 283. 1905. Erect suffruticose herb, much-branched; flowers red-purple. Open areas, edges of towns, ca. 50 m; Bolívar (near Ciudad Bolívar, El Pao). Falcón, Sucre, Zulia; Mexico, Central America, West Indies, Colombia, Brazil.

Tephrosia cinerea (L.) Pers., Syn. Pl. 2: 328. 1807. —Galega cinerea L., Syst. Nat. ed. 10, 2: 1172. 1759. Vicia littoralis Jacq., Enum. Syst. Pl. 27. 1760. —Tephrosia cinerea var. littoralis (Jacq.) Benth. in Mart., Fl. Bras. 15(1): 49. 1859. —Tephrosia littoralis (Jacq.) Pers., Syn. Pl. 2: 329. 1807. Prostrate to erect herb or herbaceous vine; flowers purplish blue. Open savannas, edges of towns, 50–1100 m; Delta Amacuro (near Tucupita), Bolívar (near Caicara, near Ciudad Bolívar, Ciudad Piar, El Manteco, El Miamo, Gran Sabana, Puerto Ordaz, Represa Guri, lower Río Paragua, base of Roraima-tepui). Amazonas (near Puerto Ayacucho). Widespread elsewhere in Venezuela; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ŠFig. 363.

Tephrosia sessiliflora (Poir.) Hassl., Repert. Spec. Nov. Regni Veg. 16: 162. 1919. —Cytisus sessiliflorus Poir. in Lam., Encycl. suppl. 2: 439. 1811 [1812]. —Generala. Erect herb or shrub to 1.5 m tall; flowers yellow. Trachypogon and Curatella savannas, 50–400 m; Bolívar (near Caicara, near Maripa, lower Río Caroní, Río Caura, Río Paragua, lower Río Suapure). Amazonas (Caño Guanay, Cerro Yutajé basin, near Puerto Ayacucho, Río Parucito, near San Juan de Manapiare). Apure, Aragua, Barinas, Cojedes, Guárico, Portuguesa, Zulia; West Indies, Guyana, Suriname, French Guiana, Colombia, Brazil, Paraguay. ŠFig. 364.

Tephrosia senna H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 458. 1823 [1824].

Tephrosia sinapou (Buc’hoz) A. Chev., Compt. Rend. Hebd. Séances Acad. Sci. 180: 1522. 1925. —Galega sinapou Buc’hoz, Hist. Univ. Règne Vég. pl. 994. 1775. —Barbasco, Barbasco caicareño, Damai (Yekwana).

Tephrosia 421

Fig. 363. Tephrosia cinerea

Fig. 364. Tephrosia sessiliflora

422

FABACEAE

Galega toxicaria Sw., Prod. 108. 1788. — Tephrosia toxicaria (Sw.) Pers., Syn. Pl. 2: 329. 1807. Shrub or erect subshrub to 3.5 m tall; flowers white to red. Edges of towns, open areas, 50–300 m; Bolívar (lower Río Caroní, lower Río Caura, Río Paragua, Santa María de Erebato), Amazonas (upper Río Cunu-

cunuma, Río Padamo, San Carlos de Río Negro). Anzoátegui, Carabobo, Cojedes, Lara, Sucre, Zulia; Mexico, Central America, West Indies, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Tephrosia sinapou is locally cultivated for its roots, which are used as a fish poison.

62. TERAMNUS P. Browne, Civ. Nat. Hist. Jamaica 290. 1756. by Nidia L. Cuello A. Slender, twining herbs, usually densely pubescent. Leaves pinnate, trifoliolate; stipules narrow, striate; stipels subulate; leaflets mostly broadly or narrowly ovate, mucronate. Inflorescences axillary racemes or fascicles; peduncles slender, severalflowered; bracteoles 2, subtending the calyx or just below the top of the pedicel; pedicels short, pubescent. Flowers inconspicuous, purplish; bracts small; bracteoles lanceolate to subulate, striate; hypanthium none. Calyx teeth 5, subequal, or the 2 upper ones connate; standard obovate, narrowed at base, not appendaged, emarginate, glabrous; wing petals narrow, adherent to and longer than the keel petals; keel petals almost straight, obtuse. Stamens 10, pseudomonadelphous; alternate anthers abortive or small and sterile. Ovary sessile, linear, pubescent; style incurved, glabrous; stigma capitate, thick, not bearded. Fruit linear, mostly straight, 2-valved,

Fig. 365. Teramnus uncinatus subsp. uncinatus

Vatairea

423

septate between the seeds, beaked by the persistent inflexed and hooked tip. Seeds numerous, septate, plump, shiny, lenticulate. Pantropics, ca. 8 species; 3 in Venezuela, 1 of these in the flora area. Teramnus uncinatus (L.) Sw., Prodr. 105. 1788. —Dolichos uncinatus L., Sp. Pl. ed. 2, 2: 1019. 1762 [1763]. Pantropics; 3 subspecies, 1 in Venezuela. T. uncinatus subsp. uncinatus Herb or vine; flowers purplish. Savannas, disturbed places, ca. 100 m; Bolívar (lower

Río Caroní). Aragua, Barinas, Cojedes, Distrito Federal, Falcón, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; Mexico, Central America, West Indies, Mexico, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, cultivated in Central Africa as forage. ŠFig. 365.

63. VATAIREA Aubl., Hist. Pl. Guiane 755. 1775. by Haroldo Cavalcante de Lima and Gerardo A. Aymard C. Trees. Leaves alternate, odd-pinnate, crowded at the end of the branches, deciduous; stipules small, caducous; stipels minute. Inflorescence a terminal panicle. Calyx campanulate or funnelform-turbinate, margin 5-dentate; petals purplish; standard orbiculate to ovate, emarginate; keel petals briefly joined at the back. Stamens 10, connate in a sheath open on upper side. Ovary with 1 or 2 ovules; style cylindric or flattened. Fruit indehiscent, samaroid or nut-like, winged at the top, transverse-veined or wing rudimentary, basal seed chambers with spongy-fibrous mesocarp. Seed 1, with straight radicle. Mexico, Guatemala, Belize, Honduras, Nicaragua, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 8 species, 2 in Venezuela, both in the flora area. Key to the Species of Vatairea 1.

1.

Leaflets with venation inconspicuous or discretely reticulate on the lower surface; calyx 1–2.5 cm long, campanulate, densely gray-appressedpubescent; gynoecium fusiform, style cylindric; fruit nut-like, orbicular to ovate, with rudimentary wing ............................................ V. guianensis Leaflets with venation prominent and reticulate on the lower surface; calyx 5–7 mm long, funnelform-turbinate, sparsely appressed-pubescent; gynoecium knife-like with flattened style; fruit samaroid, knifeshaped, with transverse-veined wing at the top ..................... V. paraensis

Vatairea guianensis Aubl., Hist. Pl. Guiane 755. 1775. —Guaboa, Mapadaro. Ormosia pacimonensis Spruce ex Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 119. 1890. Vatairea surinamensis Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 403. 1926. Tree 8–25 m tall; flowers purple. Seasonally flooded forests mostly along black-water rivers, 50–200 m; Delta Amacuro (Caño Atoiba north of Caño Araguao, Serranía de Imataca), Amazonas (Caño San Miguel, Río Baría, Río Casiquiare, Río Negro, Río Pasi-

moni, Río Sipapo, Río Temi). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 366. Vatairea paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 140. 1930. —Canelito negro, Pilón. Tree 30–40 m tall; flowers purple or blueviolet. Evergreen lowland to basimontane forests, 50–300 m; Delta Amacuro (Serranía de Imataca), Bolívar (El Palmar, mouth of Río Paragua, Sierra de Lema, Soledad). Suriname, Brazil. ŠFig. 367.

424

FABACEAE

Fig. 366. Vatairea guianensis

Fig. 367. Vatairea paraensis

Vataireopsis

Fig. 368. Vataireopsis speciosa

425

426

F ABACEAE

64. VATAIREOPSIS Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 473. 1932. by Haroldo Cavalcante de Lima and Gerardo A. Aymard C. Trees. Leaves alternate, odd-pinnate, crowded at the end of the branches, deciduous; stipules small, caducous; stipels minute. Inflorescence a terminal panicle. Calyx funnelform-turbinate, 5-dentate; petals blue-violet; standard orbiculate; keel petals mostly free or occasionally adnate on back. Stamens 9 or 10, connate in a sheath, except open 1/3 on basal side. Ovary with 1(2) ovules; style flattened. Fruit a samara, indehiscent, winged at top, transverse-veined; basal seed chambers with 2 small lateral wings. Seed 1; radicle curved. South America; 4 species, 2 in Venezuela, both in the flora area. Key to the Species of Vataireopsis 1.

1.

Leaflets membranaceous; rachis of inflorescences, bracteoles, and calyx externally and densely cinereous-appressed-pubescent; standard 7– 11 mm long .................................................................................. V. speciosa Leaflets subcoriaceous; rachis of inflorescences, bracteoles, and calyx ferruginous-appressed-pubescent externally; standard 14–17 mm long ............................................................................................ V. surinamensis

Vataireopsis speciosa Ducke, Notizbl. Bot. Gard. Berlin-Dahlem 11: 474. 1932. —Tarawuina (Yanomami). Tree 5–15 m tall; flowers purple. Evergreen lowland forests, 200–300 m; Amazonas (Río Matacuni). Guyana, Brazil, Bolivia. ŠFig. 368.

Vataireopsis surinamensis H.C. Lima, Rodriguésia 32): 30. 1980. —So-wo (Yanomami). Deciduous tree 12–30 m tall; flowers purple. Evergreen lowland and riparian forests, 200–300 m; Amazonas (Culebra, Río Cunucunuma). Suriname, Brazil.

65. VIGNA Savi, Nuovo Giorn. Lett. 8: 113. 1824 ser. 3, nom. cons. by Gerardo A. Aymard C. Vines or herbs, rarely shrubby, mostly with sturdy rootstocks, lacking hooked trichomes. Leaves pinnate or subdigitate-trifoliolate, rarely 1-foliolate, mostly entire; stipules sometimes produced below the insertion, sometimes bilobed; stipels veined, mostly blunt; glabrate. Inflorescences axillary or terminal, pseudoracemose or subcapitate; rachis contracted, the nodes glandular; bracts and bracteoles caducous; pedicels 1 or 2 per node, mostly shorter than the calyx. Flowers yellowish or white, sometimes with some purple or violet. Calyx 2-lipped, the upper lip emarginate, the lowest tooth of the lower lip longest; standard orbicular, auriculate, sometimes appendaged on the dorsal face; wing petals about equaling the standard and keel petals; keel petals apically curved to the first spiral, often oblique. Vexillar stamen free; anthers 10, uniform. Style apically thickened and barbate on the inner face, caducous, sometimes with a curved beak. Legume linear to oblong, turgid or compressed, straight or curved, not septate. Seeds reniform or quadrate, the hilum short or long, a well-developed aril sometimes present. Pantropics, mostly Paleotropics; ca. 150 species, 14 in Venezuela, 9 of these in the flora area. The taxonomy of some of the taxa included here may not be resolved until the genus has been revised.

Vigna 427

Key to the Species of Vigna 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(3). 5. 6(5). 6. 7(5). 7. 8(7). 8.

Leaflets linear or linear-oblong, 3–9 mm wide ............................... V. linearis Leaflets narrow-ovate, ovate, ovate-deltoid, or rhombic, > 10 mm wide ................................................................................................................ 2 Leaflets sometimes 3-lobed, densely whitish-tomentulose; calyx and pedicels sericeous .................................................................... V. lasiocarpa Leaflets unlobed, glabrous to pilose; calyx and pedicels glabrous to pilose, but not sericeous .................................................................................... 3 Leaflets ovate-deltoid, deltoid, or rhombic ................................................ 4 Leaflets narrowly ovate, ovate, oblong, lanceolate, or lanceolate-oblong ................................................................................................................ 5 Flowers white, sometimes with dull purple; fruits 5–7 mm wide .............. ..................................................................................................... V. candida Flowers purple; fruits 2–3 mm wide ...................................... V. peduncularis Stipules with a spur at the base, truncate or auriculate (auricle to 15 mm long) ........................................................................................................ 6 Stipules truncate or subauriculate (auricle to 1.5 mm long) at base ....... 7 Leaflets ovate; stipels glandular; inflorescences pseudoracemose ............. ..................................................................................................... V. juruana Leaflets oblong, lanceolate, or oblong-lanceolate; stipels eglandular; inflorescences subumbellate ........................................................... V. longifolia Petiolules glabrate; bracteoles and bracts 1–2 mm long ................. V. luteola Petiolules hispidulous or tomentose; bracteoles and bracts 3–4 mm long ................................................................................................................ 8 Leaflets 5–8 cm wide; flowers white; styles coiled; pods 8–10 mm wide ................................................................................................. V. adenantha Leaflets 2–3.5 cm wide; flowers blue-lavender; styles erect; pods 4–5 mm .................................................................................................... V. vexillata

Vigna adenantha (G. Mey.) Maréchal, Mascherpa & Stainier, Taxon 27: 202. 1978. —Phaseolus adenanthus G. Mey., Prim. Fl. Esseq. 239. 1818. Phaseolus truxillensis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 451. 1823 [1824]. Perennial vine; flowers purplish or pink. Disturbed and wet places, near sea level to 200 m; Delta Amacuro (southeast of Piacoa), Bolívar (Ciudad Bolívar, Piedra Marimare on the Río Orinoco, Tumeremo to Bochinche road), Amazonas (Puerto Ayacucho). Apure, Aragua, Cojedes, Distrito Federal, Guárico, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; Panama, Trinidad, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, cultivated in Central Africa and Asia. ŠFig. 373. Vigna candida (Vell.) Maréchal, Mascherpa & Stainier, Taxon 27: 201. 1978.

—Phaseolus candidus Vell., Fl. Flumin. 311. 1825 [1829]. Phaseolus appendiculatus Benth., Comm. Legum. Gen. 73. 1837. Perennial vine; flowers white and purple. Open places 100–1400 m; Bolívar (Jabillal on Río Caura, Represa Guri), Amazonas (Cerro Moriche, Cerro Yutajé, Cuao-Sipapo massif). Central America, Colombia, Brazil, Paraguay. Vigna juruana (Harms) Verdc., Kew Bull. 24: 540. 1970. —Phaseolus juruanus Harms, Notizbl. Bot. Gart. BerlinDahlem 7: 506. 1921. Slender vine; flowers yellow. Open and/or wet places, 100–400 m; Delta Amacuro (Caño Güiniquina, Curiapo, Río Toro), Bolívar (Isla Anacoco, Río Botanamo, upper Río Paragua, Salto Pará), Amazonas (Río Siapa). Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Central Africa.

428

F ABACEAE

Vigna lasiocarpa (Mart. ex Benth.) Verdc., Kew Bull. 24: 539. 1970. —Phaseolus lasiocarpus Mart. ex Benth., Comm. Legum. Gen. 76. 1837. —Frijol de sapo. Phaseolus diversifolius Pittier, Bol. Técn. Minist. Agric. 5: 56. 1944. Herbaceous vine; flowers yellow. Savannas, edges of towns, 50–1000 m; Bolívar (Ciudad Piar, Gran Sabana, Maripa, Río Pargueni), Amazonas (La Esmeralda, San Carlos de Río Negro, San Juan de Manapiare, Santa Barbara del Orinoco, Yavita). Apure, Barinas, Guárico, Portuguesa; Central America, Colombia, Guyana, Suriname, French Guiana, Brazil, Paraguay, Argentina. Vigna linearis (H.B.K.) Maréchal, Mascherpa & Stainier, Taxon 27: 202. 1978. —Phaseolus linearis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 445. 1823 [1824]. Neotropics; 2 varieties, 1 in Venezuela. V. linearis var. linearis. —Barbasquillo, Golondrina. Perennial herb or vine; flowers redpurple. Savannas, Mauritia palm swamps, open places, 50–900 m; widespread in Bolívar and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 369. Vigna longifolia (Benth.) Verdc., Kew Bull. 24: 541. 1970. —Phaseolus longifolius Benth., Comm. Legum. Gen. 75. 1837. Slender vine; flowers yellow. Wet places,

100–200 m; Delta Amacuro (Caño Acoimo), Amazonas (Minicio). Guárico, Lara; Panama, Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Vigna luteola (Jacq.) Benth. in Mart., Fl. Bras. 15(1B): 194. 1859. —Dolichos luteolus Jacq., Hort. Bot. Vindob. 1: 39. 1770. Vine; flowers yellow. Wet places, sandy beaches, secondary scrub, near sea level to 100 m; Delta Amacuro (Caño Araguao, Caño Güiniquina, Río Acure). Widespread in northern Venezuela; U.S.A. (Florida), Mexico, Central America, Antilles, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Brazil, Paraguay, Argentina, Uruguay, Central Africa, Southest Asia. ŠFig. 370. Vigna peduncularis (H.B.K.) Fawc. & Rendle, Fl. Jamaica 4: 68. 1920. —Phaseolus peduncularis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 447. 1823 [1824]. Neotropics; 3 varieties, 1 in Venezuela. V. peduncularis var. peduncularis Slender vine; flowers lavender and white. Savannas, disturbed open places, 50–900 m; Bolívar (Caicara, Río Ore in Río Parguaza basin), Amazonas (La Esmeralda, Puerto Ayacucho, Río Ocamo, lower Río Ventuari, San Pedro del Cataniapo, Simarawochi). Anzoátegui, Apure, Barinas, Distrito Federal, Guárico, Portuguesa, Sucre, Zulia; Central America, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ŠFig. 371.

Fig. 369. Vigna linearis var. linearis

Vigna 429

Fig. 370. Vigna luteola

Fig. 372. Vigna vexillata var. vexillata

Fig. 371. Vigna peduncularis var. peduncularis

Fig. 373. Vigna adenantha

430

F ABACEAE

Vigna vexillata (L.) A. Rich. in Sagra, Hist. Phys. Cuba, Bot. Pl. Vasc. 11: 191. 1845. —Phaseolus vexillatus L., Sp. Pl. 724. 1753. Pantropics; 6 varieties, 1 in Venezuela. V. vexillata var. vexillata Slender vine; flowers blue or lavender turning white or yellow. Open, wet places,

granitic outcrops, near sea level to 200 m; Delta Amacuro (Capure, Pedernales, Río Acure), Amazonas (Puerto Ayacucho, Tamatama). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Mérida, Miranda, Táchira, Yaracuy, Zulia; Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 372.

66. ZOLLERNIA Wied-Neuw. & Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 13(2), pref. 13, t. C & D. 1827. by Gerardo A. Aymard C. Trees or shrubs. Leaves simple; petiole developed as a rugose joint extending into the base of the blade; margin serrate-undulate; stipules free, rigid, borne on the enlarged node. Inflorescences terminal and axillary racemes, often forming a panicle; bracts small; bracteoles minute on upper half of pedicel. Calyx closed in bud, ovoid to lanceolate, acumimate, splitting in 2 valves to short conical or saucershaped, deciduous hypanthium at anthesis; petals 5(6), imbricate, slighty unequal, shorter than calyx, obtuse, clawed; standard broader than the other petals and exterior to them. Stamens (9)10–12(–15), uniform; filaments short, linear-lanceolate, sagittate at base; anthers many times longer than the filaments. Ovary subsessile or stipitate; ovules numerous; style short and subulate; stigma small, obliquely terminal. Fruit indehiscent, drupaceous, ellipsoid, laterally compressed, stipitate, apiculate; mesocarp fleshy; endocarp thin and tough; seeds 1–few. Neotropics; ca. 10 species, 2 in Venezuela, both in the flora area. Zollernia kanukuensis R.S. Cowan, known from the Kanuku mountains in Guyana, might eventually be found in the Venezuelan Guayana.

Fig. 374. Zollernia paraënsis

Zornia 431

Key to the Species of Zollernia 1. 1.

Inflorescence rachis, pedicels, and calyx ferruginous-pubescent; anthers sparsely pilose, with trichomes 1–1.5 mm long ................... Z. grandifolia Inflorescence rachis, pedicels, and calyx sericeous-canescent; anthers glabrous or with trichomes ca. 0.2 mm long ................................ Z. paraënsis

Zollernia grandifolia Schery, Fieldiana, Bot. 28: 270. 1952. Tree to 20 m tall; leaf blades elliptic, margins serrate; petals pink, 5–6 mm long; anthers pilose. Evergreen lowland forests, ca. 200 m; Bolívar (Río Caura, Río Nichare). Endemic.

Zollernia paraënsis Ducke, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 6: 81. 1910. —Masiri (Yekwana). Zollernia ulei Harms, Notizbl. Königl. Bot. Gart. Berlin 6: 309. 1915. Tree to 20 m tall. Evergreen lowland forests, 200–400 m; Amazonas (Río Ocamo, lower Río Orinoquito). Guyana, Brazil. ŠFig. 374.

67. ZORNIA J.F. Gmel., Syst. Nat. 2: 1076. 1791. Myriadenus Desv., J. Bot. Agric. 1: 121. 1813. by Gerardo A. Aymard C. Annual herbs or suffruticose perennials to 2 m tall. Stems diffusely branched, prostrate and spreading to erect and ascending. Leaves alternate, digitately 2- or 4foliolate; leaflets often pellucid-punctate, exstipellate; stipules subfoliaceous, paired, peltate, auriculate, 3–11-veined, often punctate. Inflorescence spicate, rarely racemose, terminal or axillary, interrupted to congested, 1–many-flowered; bracts paired, lateral, stipuliform, 3–15-veined, sometimes punctate, enclosing a sessile flower; bracteoles absent. Calyx subhyaline, the tube short, 5-lobed, usually 5–10veined, ciliate, the 2 upper lobes connate nearly to the apex, the 2 lateral lobes much shorter, the lowest oblong or lanceolate, shorter than the upper lip; corolla yellow, orange, or rarely white, frequently purple-striated; standard suborbicular, clawed, to 17 mm long; wing petals obliquely obovate or oblong, shorter than standard, usually clawed and auriculate; keel petals incurved, subrostrate, smaller than the wing petals. Stamens 10, monadelphous; filaments connate into a closed tube, splitting dorsally, persistent; anthers alternately longer, subbasifixed, versatile. Ovary sessile; ovules 2–several, campylotropous; styles filiform basally; stigmas minute, terminal. Fruit a loment of 2–15 articles, compressed, the upper suture nearly straight, the lower deeply sinuate, articles glabrous to puberulent or pilose, smooth or echinate, usually reticulate, indehiscent. Seeds black or dark brown, orbicular to subreniform, estrophiolate (lacking an appendage to the hilum). Tropical and subtropical regions worldwide (most diverse in the Neotropics); ca. 80 species, 15 in Venezuela, 11 of these in the flora area. The taxonomy of some of the taxa included here probably will not be resolved until the genus has been revised. Key to the Species of Zornia 1. 1. 2(1).

Annual herbs, very slender, to 20 cm tall ..................................... Z. herbacea Perennnial herbs, stout near base, 30–100 cm tall ................................... 2 Stems, petioles, petiolules, leaflets, bracts of inflorescences, and calyx densely sericeous .......................................................................... Z. sericea

432

F ABACEAE

2.

Stems, petioles, petiolules, leaflets, bracts of inflorescences, and calyx glabrous or strigose to villous, never sericeous ......................................... 3 3(2). Leaflets linear-filiform, 1–2 mm wide; loment with 3 segments ................ ................................................................................................. Z. lasiocarpa 3. Leaflets wider, 3–12 mm wide; loment with 4–15 segments .................... 4 4(3). Leaves 4-foliolate ....................................................................................... 5 4. Leaves 2-foliolate ....................................................................................... 8 5(4). Bracts of inflorescences elliptic, apex obtuse; loment densely glandular ................................................................................................................ 6 5. Bracts of inflorescences lanceolate-elliptic, apex acute; loment eglandular ................................................................................................................ 7 6(5). Leaflets lanceolate to broadly lanceolate, subacute; inflorescences manyflowered; calyx pilose; loment reticulate ............................. Z. guanipensis 6. Leaflets obovate, retuse; flowers solitary, axillary; calyx glabrous; loment without reticulation ............................................................... Z. myriadena 7(5). Leaflets oblanceolate, strigose; stipules 7-veined; loment erect, with hairy bristles ................................................................................... Z. brasiliensis 7. Leaflets ovate-lanceolate or lanceolate; stipules 5-veined; loment curved, without hairy bristles ................................................................. Z. curvata 8(4). Leaflets epunctate; loment glandular with numerous hairy bristles (never retrorse) ...................................................................................... Z. diphylla 8. Leaflets punctate or rarely epunctate; loment eglandular with numerous retrorse bristles ...................................................................................... 9 9(8). Bracts of inflorescences oblong-lanceolate or broadly lanceolate; loment without reticulation .................................................................... Z. gemella 9. Bracts of inflorescences linear, lanceolate to lance-ovate; loment with evident reticulation .................................................................................. 10 10(9). Petiole glabrous; inflorescences congested near tip; bracts of inflorescences linear, 1–2 mm wide, auricle to 2 mm; loment with retrorse bristles 0.6–1.5 mm long ............................................................. Z. latifolia 10. Petiole glabrous to pilose; inflorescences crowded; bracts of inflorescences lanceolate to lance-ovate, 4–8 mm wide, auricle 2–3.5 mm long; loment with retrorse bristles 0.2–0.4 mm long ................................... Z. reticulata Zornia brasiliensis Vogel, Linnaea 12: 62. 1838. Erect perennial herb to 60 cm tall; flowers yellow. Edges of towns, open areas, savannas, 50–300 m; Bolívar (Caicara, Puerto Ordaz, Río Aro, near Tumeremo). Monagas, Nueva Esparta; Brazil. Zornia curvata Mohlenbr., Webbia 16: 132. 1961. Erect herb to 50 cm tall; flowers yellow. Savannas, edges of towns, 50–400 m; Bolívar (Ciudad Bolívar, upper Río Paramichi). Amazonas (Galipero, Isla Ratón, near Puerto Ayacucho). Anzoátegui, Aragua, Distrito Federal, Lara, Mérida, Miranda, Monagas, Táchira, Trujillo; Mexico, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia, Uruguay, Argentina.

Zornia diphylla (L.) Pers., Syn. Pl. 2: 318. 1807. —Hedysarum diphyllum L., Sp. Pl. 747. 1753. Perennial prostrate to erect herb; flowers yellow. Edges of towns, savannas, open areas, 50–200 m; Bolívar (near Caicara), Amazonas (Río Sipapo, San Carlos de Río Negro, San Pedro del Orinoco). Mérida, Trujillo, Zulia; Colombia, Guyana, Brazil, Paraguay, Argentina, Zaire, Sri Lanka, Vietnam. Zornia gemella (Willd.) Vogel, Linnaea 12: 61. 1838. —Hedysarum gemellum Willd., Sp. Pl. 5(1): 1178. 1810. Erect herb to 60 cm tall; flowers yellow. Savannas, open areas, 50–400 m; Bolívar (Altiplanicie de Nuria, near Tumeremo).

Zornia 433

Anzoátegui, Monagas; U.S.A. (Texas), Mexico, Costa Rica, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Paraguay, Argentina, Uruguay.

jillo; Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, West Africa.

Zornia guanipensis Pittier, Bol. Soc. Venez. Ci. Nat. 6: 194. 1940. Zornia tenuifolia var. latifolia Benth. in Mart., Fl. Bras. 15(1): 81. 1859. Zornia marajoara Huber, Bol. Mus. Paraense Hist. Nat. 5: 150. 1908, nom. nud. Erect herb to 70 cm tall; flowers yellow. Savannas, Mauritia palm swamps, 50–200 m; Bolívar (Tumeremo). Anzoátegui, Monagas; Suriname, Brazil (Maranhão).

Zornia myriadena Benth. in Mart., Fl. Bras. 15(1): 85. 1859. Erect herb to 60 cm tall; flowers yellow. Savannas, ca. 100 m; Bolívar (lower Río Caroní). Cuba, Jamaica, Brazil (Bahia, Minas Gerais, Pernambuco).

Zornia herbacea Pittier, Bol. Soc. Venez. Ci. Nat. 6: 192. 1940. Annual herb to 20 cm tall; flowers yellow. Savannas, ca. 200 m; Bolívar (northeast of Upata). Anzoátegui, Zulia. Zornia lasiocarpa A.R. Molina, Ceiba 1: 257. 1951. Erect herb 20–40 cm tall; flowers yellow. Open areas, edges of towns, 50–200 m; Bolívar (Ciudad Bolívar, Los Pijiguaos). Mexico, Honduras, Brazil. Zornia latifolia Sm. in Rees, Cycl. 39: Zornia no. 4. 1819. Herb to subshrub, 30–70 cm tall; flowers yellow. Pantropics; 2 varieties, 1 in Venezuela. Z. latifolia var. latifolia. —Pega-pega. Zornia pubescens H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 515. 1823 [1824]. Zornia surinamensis Miq., Ann. Mag. Nat. Hist. 11: 14. 1843. Herb to subshrub 30–70 cm tall; flowers yellow. Savannas, edges of towns, 50–1200 m; widespread in Bolívar and Amazonas. Apure, Cojedes, Monagas, Portuguesa, Tru-

Zornia reticulata Sm. in Rees, Cycl. 39: Zornia no. 2. 1819. Erect herb to subshrub 30–70 cm tall; flowers yellow. Savannas, open areas, 50– 500 m; Bolívar (Ciudad Piar, Las Trincheras, Urimán), Amazonas (Caño San Miguel, Isla Ratón, Puerto Ayacucho, Salto Yureba in lower Río Ventuari basin, Yavita). Aragua, Barinas, Distrito Federal, Falcón, Guárico, Monagas, Portuguesa, Zulia; U.S.A. (Arizona, Texas), Mexico. Central America, Antilles, Colombia, Guyana, Ecuador, Brazil, Bolivia, Paraguay, Argentina. Zornia sericea Moric., Pl. Nouv. Amér. 126. 1844. —Crineja. Erect herb to 50 cm tall; flowers yellow. Curatella-Trachypogon savannas, edges of semideciduous forests, 100–200 m; Bolívar (Caicara, lower Río Caura near Maripa, between Upata and Villa Lola), Amazonas (San Fernando de Atabapo). Apure, Mérida; Brazil (Bahia, Maranhão, Mato Grosso, Minas Gerais, Pará, Piaui). ŠFig. 375.

Fig. 375. Zornia sericea

434

F LACOURTIACEAE

FLACOURTIACEAE by Mark Olson, Paul E. Berry, and Gerardo A. Aymard C. Shrubs or trees, rarely scrambling climbers, sometimes armed with axillary or trunk spines. Leaves alternate or rarely opposite (Abatia), often 2-ranked, simple, entire or glandular-crenate or -serrate, often pellucid-punctate, sometimes with pairs of glands at bases of leaves, pubescence usually simple, rarely stellate, pinnately or palmately veined; petioles short or with leaf bases decurrent to petioles, or petioles long, sometimes with pulvini at distal ends, often thickened or wrinkled at base and/or apex; stipules usually present, often deciduous. Inflorescences terminal or axillary, sometimes epiphyllous or on old wood, usually cymose, or in fascicles, spikes, racemes, panicles, corymbs, or cone-like pseudanthia, or flowers solitary and axillary; bracts and bracteoles small, scale-like. Flowers actinomorphic, usually small, bisexual or unisexual (plants monoecious, dioecious, or sometimes andromonoecious), usually hypogynous, rarely perigynous or epigynous. Sepals (2)3–8(–15), rarely more, distinct or connate toward the base, persistent or deciduous, sometimes accrescent, imbricate or occasionally valvate; petals (0)3–8(–15), free, imbricate or valvate, mostly alternating with the sepals and deciduous, sometimes persistent and accrescent, sometimes spirally arranged and poorly differentiated from the sepals, occasionally possessing basal scales on the adaxial surfaces. Disk often present, intrastaminal or extrastaminal, occasionally composed of discrete interstaminal glands. Stamens numerous or occasionally equal in number to the petals, sometimes some of them staminodal; filaments distinct, grouped into bundles opposite the petals alternating with glands, or united into a tube; anthers dithecal, dehiscing longitudinally, less often by terminal pores, sometimes with prolonged or glandular connective. Ovary of 2–10 united carpels (1 in paleotropical Aphloia), 1-locular with 2–9 parietal or virtually basal placentas, sometimes incompletely plurilocular by deeply intruded placentas, rarely distinctly plurilocular and with axile placentation (Prockia), superior or rarely inferior (only in Madagascar); ovules 1–many on each placenta, generally numerous; styles free or variously united, rarely absent; stigmas distinct, 2–5-lobed. Fruit a berry or a loculicidal capsule or drupe, sometimes winged, horned, or prickly. Seeds 1–many, often compressed, sometimes arillate or with cottony hairs; endosperm abundant, oily, and proteinaceous. Pantropics, with some representatives reaching temperate regions of South America (Aphaerema in southern Brazil and Berberidopsis in Chile), Africa, Asia, and Australia; 89 genera and ca. 875 species, 13 genera and 47 species in the flora area. A possible record of Hasseltia floribunda H.B.K. from the flora area is Steyermark et al. 115059 (MO, VEN). It is from the Caño Güiniquina basin in Delta Amacuro state, but the specimen is sterile and cannot be confirmed as this species. Hasseltia can be recognized by its flowers with 4 sepals and 4 petals borne in highly branched terminal racemes. Key to the Genera of Flacourtiaceae 1.

Petioles apically pulvinate, at least some flexed at the apex; petals more numerous than the sepals; fruit a capsule or a dry berry with bristly spines, longitudinal wings, or small conical projections; floral disk absent ......................................................................................................... 2

FLACOURTIACEAE 435

1.

2(1).

2.

3(2).

3.

4(1).

4.

5(4).

5.

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

7.

Petioles lacking an apical pulvinus, not flexed at apex; petals equaling the sepals in number or absent; fruit a fleshy berry or a dry to fleshy capsule, without spines, conical projections, or wings, though surface sometimes convolutedly papillate and with abundant stellate hairs (Ryania), or 3–sulcate (Zuelania); floral disk present or absent ......... 4 Fruit with 10–14 longitudinal papery wings, these sometimes highly crisped; flowers usually unisexual; styles 4–8; petals larger than the sepals; staminate flowers in few-flowered racemes (sometimes solitary), pistillate flowers solitary; leaves not resinous .... 2. Carpotroche Fruit with bristles, prickles, or conical projections, not winged; flowers bisexual or unisexual; styles 1, 3, or 4; petals larger than the sepals or equal in length; flowers in racemes, fascicles, or solitary; leaves resinous or not ................................................................................................ 3 Fruit with smooth or furrowed, glabrous or puberulent, conical projections 1–18 mm long; usually at least some flowers bisexual, style 1; petals equal to the sepals in length; flowers in axillary racemes, the rachis often persistent after many or sometimes all of the flowers or fruits have fallen; leaves often resinous and shiny, especially young ones ....................................................................................... 8. Lindackeria Fruit with curving, flattened, pubescent prickles 1–5 mm long; flowers unisexual, plants dioecious, styles 3 or 4; petals exceeding sepals in length; staminate flowers in axillary fascicles, pistillate flowers solitary in leaf axils; leaves not shiny and resinous .................. 10. Mayna Sepals 13–45 mm long, often showy; petals absent; disk urceolate to cupshaped; plants often with stellate pubescence; fruit a tardily dehiscent capsule, usually pubescent and often gently lobed toward the apex, the surface often minutely and complicatedly convoluted or covered with many papillae .............................................................................. 11. Ryania Sepals 1–9 mm long, not showy; petals present or absent; disk absent or composed of lobes or glands, if cup-shaped, then formed of disk lobes adnate to filaments; plants never with stellate pubescence; fruit a berry, or a dry to fleshy capsule, not covered with pubescent, convoluted papillae ......................................................................................... 5 Inflorescences spikes or panicles of spikes, the flowers sessile; petals absent; stamens 8, 4 long and 4 short; disk lobes 8, spatulate, pubescent, alternating with the stamens; stipules 3–22 mm long, ovate-lanceolate and enveloping the terminal bud, soon deciduous ................................................................................................... 4. Euceraea Inflorescences racemes or fascicles, or the flowers solitary, pedicellate or sessile; petals absent or present; stamens and disk varied; stipules, when present, smaller and not enveloping the terminal bud .............. 6 Petals present, often persistent with the calyx on the fruit; inflorescences 4–30 cm long .......................................................................................... 7 Petals absent; inflorescences < 8 cm long, usually very short .................. 8 Ovary semi-inferior, hairy; disk represented by pubescent glands opposite the sepals; sepals usually 5–8, stamens in fascicles of (2)3–7, alternating with the disk glands; stipules minute, deciduous ........... 6. Homalium Ovary superior, usually glabrous; disk absent; sepals usually 3 or 4; base of lamina or petiole apex usually with 1 or 2 glands; stamens numerous and free; stipules 2–4 mm long, ± persistent .............................. 1. Banara

436

F LACOURTIACEAE

8(6).

Trunk often with stout, branched spines; leaves never pellucid-punctate; plants generally dioecious (flowers sometimes appearing bisexual, but then anthers ± abortive) .......................................................... 12. Xylosma 8. Trunk unarmed; leaves often pellucid-punctate; plants with at least some flowers bisexual ..................................................................................... 9 9(8). Disk appendages absent (no staminodes); stamens 10–50, bracts and bracteoles free or united to form a cupule; sepals petaloid, pellucidpunctate, and -lineate, often reflexed at anthesis ........................ 7. Laetia 9. Disk appendages or staminodes present; stamens 5–100, bracts and bracteoles always free; sepals thick, not pellucid-punctate or -lineate, only occasionally reflexed at anthesis ................................................. 10 10(9). Stigma a thick, peltate disk, subsessile (the pistil therefore having the shape of a tiny, squat urn); flowers in rather sparsely distributed fascicles, usually occurring on leafless shoots; stamens 20–40; staminodes 15–20; fruit a fleshy capsule, depressed-globose, shallowly 3-sulcate, yellowish green to dark green when fresh, finally splitting into 3 valves .................................................................................................. 13. Zuelania 10. Styles present, entire or 3-lobed, stigma not a subsessile peltate disk; flowers in rather few-flowered cymes or in densely distributed fascicles commonly in the axils of leaves and on leafy shoots; stamens either ≤ 25 or > 80; staminodes ≤ 15; fruit a dry or fleshy capsule .................. 11 11(10). Flowers numerous in elongate, slender, pedunculate racemes; calyx shortly split into 2 or 3 valvate lobes; leaves with 3 or 5 main palmate veins ........................................................................................... 9. Lunania 11. Flowers either few in axillary cymes or else in short, congested inflorescences; calyx deeply 4-, 5-, or 6-lobed, the lobes mostly imbricate; leaves with pinnate venation .......................................................................... 12 12(11). Flowers in axillary, few-flowered cymes; stamens 80–100, arranged in 3 series external to about 15 distally hairy staminodes .........5. Hecatostemon 12. Flowers in axillary, sessile, or pedunculate fascicles or condensed cymes, often in the axils of leaves or on leafy shoots, inflorescences often occur densely, with each axil in a shoot bearing a tuft of flowers, in many species the bracts and bracteoles, along with the peduncles or the bases of the peduncles below the articulation, forming cushions in the axils of the leaves; staminodes usually equal in number to the stamens, usually in the same series and often adnate to them; stamens (5)6–10(–25) .................................................................................................... 3. Casearia 1. BANARA Aubl., Hist. Pl. Guiane 547, t. 217. 1775. Kuhlia H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 234, t. 652. 1825. Xyladenius Desv. ex Ham., Prodr. Pl. Ind. Occid. 41. 1825. Ascra Schott in Spreng., Syst. Veg. 4(cur. post.): 407. 1827. Boca Vell., Fl. Flumin. 232. 1825 [1829]. Shrubs or trees. Leaves alternate, not pellucid-punctate, often distichous, 3–5veined from the base or pinnately veined, entire or glandular-serrate or -crenate, sometimes with one or a pair of glands at the base of the blade or on the petiole (glands on the petiole are often cupped and borne on short stalks); petiolate; stipules deciduous or subpersistent. Inflorescences terminal, subterminal, or more rarely

Banara

437

axillary, paniculate, racemose, corymbose, or fasciculate. Flowers bisexual, usually yellow; pedicels articulate at or above the base; bracts and bracteoles deltoid, minute, generally deciduous. Sepals 3(4), connate basally, valvate or rarely subimbricate distally; petals same number as and similar to the sepals; perianth persistent in fruit. Disk absent. Stamens numerous, free, in several series, often conspicuous and rather showy; filaments filiform, usually glabrous; anthers small, subglobose or oblong, basifixed, introrse, longitudinally dehiscent. Ovary sessile, incompletely plurilocular by 3–8 filiform or lamelliform multiovulate placentas intruded far into the cavity, glabrous or rarely hairy; style simple, persistent; stigma capitate or subcapitate, minutely lobed according to the number of placentas. Fruit indehiscent, dry or fleshy; pericarp thin-coriaceous. Seeds numerous, oblong to ovoid-applanate; testa crustaceous, sometimes slightly foveolate. Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; 31 species, 4 in Venezuela, 3 of these in the flora area. Key to the Species of Banara 1. 1. 2(1).

2.

Lower surface of leaves glabrous; inflorescence axis glabrous to laxly puberulent ...................................................................................... B. nitida Lower surface of leaves and inflorescence axis noticeably pubescent ..... 2 Inflorescence 10–40 cm long, panicles many-flowered; flower buds 2–4 mm diameter; leaf margins with teeth mostly 20–25 per side, the sinuses between serrations mostly 1–2 mm deep; lower surfaces of leaves brownish tan, with loosely appressed tan or brownish trichomes .... B. guianensis Inflorescence < 10 cm long, panicles 5–10-flowered; flower buds 6–10 mm diameter; leaf margins with teeth mostly 20–40 per side, the sinuses between serrations 0.5–1 mm deep; lower surfaces of leaves grayish green with densely appressed, canescent trichomes ........... B. orinocensis

Banara guianensis Aubl., Hist. Pl. Guiane 548, t. 217. 1775. —Guanábana silvestre, Pan de acure, Pinito blanco, Rastrojero, Rastrojero blanco, Vara blanca, Ya’-ra paya yo’ (Panare). Xyladenius gladulosus Desv. ex Ham., Prodr. Fl. Ind. Occid. 41. 1825. —Banara glandulosa (Desv. ex Ham.) Speg., Revista Argent. Bot. 1: 210. 1926. Kuhlia mollis Poepp., Nov. Gen. Sp. Pl. 3: 74, pl. 285. 1845. —Banara mollis (Poepp.) Tul., Ann. Sci. Nat. Bot. sér. 3, 7: 288. 1847. —Banara guianensis var. mollis (Poepp.) Eichler in Mart., Fl. Bras. 13(1): 501. 1871. Shrub or small tree 4–10(–15) m tall. Semideciduous to evergreen lowland forests, secondary forests, forest-savanna ecotones, granitic outcrops, 50–600 m; widespread in northern Bolívar, Amazonas (Puerto Ayacucho). Widespread elsewhere in Venezuela;

Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 377. Banara nitida Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl 2): 93. 1861. —Cayenito. Banara guianensis auct. non Aubl. 1775: sensu Ll. Williams, Field Mus. Nat. Hist., Bot. Ser. 15: 351. 1936, pro parte; Steyerm., Acta Bot. Venez. 3: 140. 1968, pro parte. Tree to 25 m tall. Evergreen lowland forests, 200–300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Serranía de Imataca). Aragua, Carabobo; Amazonian Colombia, Ecuador, Peru, and Brazil (Acre). ŠFig. 376. Banara orinocensis (Cuatrec.) Sleumer, Blumea 24: 118. 1978. —Prockia

438

F LACOURTIACEAE

Fig. 376. Banara nitida Fig. 377. Banara guianensis

orinocensis Cuatrec., Trop. Woods 101: 25. 1955. —Caramacatillo. Banara sacupanensis Aristeg., Bol. Soc. Venez. Ci. Nat. 21: 299, fig. on p. 301. 1960. Shrubby tree 3–6 m tall. Evergreen lowland riparian forests, 0–200 m; Delta Ama-

curo (Caño Jota-Sabuca near Caño Mariusa, Caño Sacupana), Bolívar (mouth of Río Caura), Amazonas (Caño Yagua, Isla Ratón and scattered along Rio Orinoco to below mouth of Cano Yapacana). Apure; Colombia (Vichada). The fruit of Banara orinocensis is used as fish bait.

2. CARPOTROCHE Endl., Gen. Pl. 918. 1839. Shrubs or trees. Leaves alternate, often crowded toward the ends of the branches, persistent or rarely periodically deciduous, subentire to dentate or serrate, venation pinnate; petioles with an apical pulvinus, often flexed; stipules lanceolate to subulate, deciduous. Flowers scented, pedicellate, unisexual or bisexual (plants monoecious, dioecious, polygamomonoecious, or polygamodioecious). Inflorescences axillary, the staminate ones of short, few-flowered cymes, the pistillate ones usually 1-flowered, sometimes on leafless branches or cauliflorous. Sepals generally 3, imbricate, subpersistent; petals 4–12, imbricate, ± biseriate, generally larger than the sepals; pistillate flowers often larger than staminate flowers. Disk or staminodes absent. Stamens numerous; filaments pubescent; anthers basifixed, linear, pubescent. Ovary smooth or with 4–16 longitudinal ridges or wings, often densely, finely pubescent; placentas 4–8, multiovulate; styles 4–8(–10), short, persis-

Carpotroche

439

tent, connate at base; stigmas capitate or shortly lacerate-palmatifid. Fruit a winged or ridged capsule, with a thick, often fibrous pericarp, tardily dehiscent by 3–7 valves, with 4–16 vertical ridges or entire to crenate-lacerate wings. Seeds generally numerous; testa smooth; endosperm copious; embryo straight; cotyledons foliaceous. Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 11 species, 2 in Venezuela, both in the flora area. Key to the Species of Carpotroche 1.

1.

Fruit ca. 6 cm diameter with highly crisped wings not overlapping like shingles, sometimes so dense it is difficult to discern the individual wings; petiole 5–10(–15) mm long; leaves oblong-ovate, 8–20(–30) cm long; sepals 2–8 mm long; petals 2.5–18(–20) mm long; stamens ca. 40; filaments 1.5–2 mm long ............................................. C. crispidentata Fruit 2.5–5 cm diameter with ca. 12 longitudinal wings (5–)10–30 mm high appressed to the fruit surface and partly overlapping each other, not highly crisped; petiole 15–30(–50) mm long; leaves elliptic-oblong to -obovate, 15–37(–46) cm long; sepals 12–20 mm long; petals (20–)25– 30 mm long; stamens ca. 80; filaments 2–3 mm long .......... C. grandiflora

Fig. 378. Carpotroche grandiflora

440

F LACOURTIACEAE

Carpotroche crispidentata Ducke, Arq. Inst. Biol. Veg. 4: 55. 1938. —Palo de camarón. Small tree to 10 m tall. Evergreen and deciduous lowland forests on white sand, 100– 200 m; Amazonas (Aemoba in upper Río Orinoco, Río Cunucunuma). Zulia; French Guiana, Brazil (Amazonas). Carpotroche grandiflora Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl 2): 81. 1861. —Mayna grandiflora

(Spruce ex Benth.) R.E. Schult., Bot. Mus. Leafl. 15: 69. 1951. —Palo cachicamo, Palo de camarón, Sawe (Yekwana), Yará, Yariyala (Bale). Carpotroche amazonica Mart., Fl. Bras. 13(1): 437. 1871. Shrub or small tree 4–11 m tall. Evergreen lowland forests, 100–200 m; Amazonas (between La Esmeralda and Cerro Duida, Río Cunucunuma, Río Guayapo, Río Yatúa, San Carlos de Rio Negro, Yavita). Colombia (Vaupés), Brazil (Amazonas). ŠFig. 378.

3. CASEARIA Jacq., Enum. Syst. Pl. 4, 21. 1760. Shrubs or trees. Branchlets rarely thorny. Leaves alternate, distichous, pinnately veined, entire or glandular-crenate or -serrate, rarely spiny, membranaceous to coriaceous, often with pellucid dots and/or lines, petiolate, stipulate, without basal glands. Flowers bisexual, small, in axillary, sessile or pedunculate fascicles or glomerules, rarely in cymes, very rarely solitary; pedicels articulate; basal bracts free, scale-like, often numerous and forming a cushion. Sepals (4)5(6, 9), usually connate at the base or rarely to half their length, imbricate, subpersistent; petals absent. Stamens usually 8 or 10, to 25 in some species, ± perigynous, uniseriate; filaments free, or rarely adnate to the disk (C. spinescens); anthers globose to ovoid, sometimes apiculate by a glabrous or barbate gland-like connective. Disk lobes generally in the same row with the stamens and alternating with them, or else inserted inside the row of stamens. Ovary free, 1-locular, with usually 3 parietal multiovulate placentas; ovules anatropous; style simple or divided at apex into three branches; stigma(s) capitate. Capsule dry to succulent, often 3-angled, dehiscing by 3 or less often 4 valves. Seeds generally numerous, glabrous or pubescent, completely to partially enveloped by a soft, often colored and fimbriate aril; testa crustaceous, foveolate; albumen fleshy; cotyledons flat. Tropics and subtropics worldwide; ca. 180 species, ca. 27 in Venezuela, 21 of these in the flora area. Due to the difficulty of distinguishing some of the numerous species of Casearia in the flora area, two keys (one for flowering material, the other for fruiting material) are offered in the hope that one of them (or a combination of the two) will enable the user to identify material from the Venezuelan Guayana. The key to flowering material appears below, followed by a key to fruiting material on page 442. Key to the Species of Casearia based on flowering material 1. 1. 2(1). 2. 3(2).

Disk lobes intrastaminal, in their own row inside the row of stamens, not alternating with them nor adnate to them ........................................... 2 Disk lobes interstaminal, in the same row as the stamens and alternating with them, often adnate to them to varying degrees ........................... 5 Style unbranched bearing only 1 stigma ........................................ C. tremula Style 3-branched, each branch bearing a capitellate stigma .................... 3 Ovary and fruit with fine reddish pubescence inside and out; sepals usually reflexing as fruit develops ................................................ C. javitensis

Casearia 441

3.

4(3). 4. 5(1).

5.

6(5). 6. 7(6). 7. 8(7).

8.

9(8). 9. 10(7). 10. 11(10).

11.

12(10). 12. 13(12).

13.

Ovary and fruit glabrous inside or with grayish or golden trichomes inside and out; sepals usually remaining erect under the fruit or fertilized flower .............................................................................................. 4 Flowers (3–)5–8 mm long; stamens 10–20, drying brownish; leaves drying greenish .......................................................................... C. commersoniana Flowers 6–15 mm long; stamens 20–25, drying blackish, prominent against pale disk lobes; leaves drying brownish .................... C. spruceana Disk lobes and filaments adnate for most of their length, the anthers appearing to be inserted in the sinuses between the fused disk lobes; deciduous tree with divaricating, vining branches, the leaves often congested toward the branch tips ................................................ C. spinescens Both disk lobes and filaments free from each other their entire length or fused below only to their lower third; branches not vining or divaricately branched ...................................................................................... 6 Style 3-branched near the apex, each branch bearing a capitellate stigma ................................................................................................... C. sylvestris Style not branched, bearing only one stigma ............................................ 7 Stamens usually 8 ...................................................................................... 8 Stamens usually 10 .................................................................................. 10 Leaves generally drying light to olive green, usually glabrous or with scattered hairs on the midrib, especially on the lower surface; plants may bear spines on trunk or branches; flowers usually 3–4 mm long (occasionally longer) .................................................................. C. aculeata Leaves usually drying brownish or reddish brown, sometimes greenish brown and usually distinctly pubescent, at least on midrib on the lower surface; plants unarmed; flowers 4–6 mm long .................................... 9 Leaves glabrous or with usually appressed grayish pubescence ............... ................................................................................................ C. guianensis Leaves usually with erect golden pubescence ................................. C. hirsuta Flowers small, ≤ 2.5 mm long .................................................................. 11 Flowers > 2.5 mm long ............................................................................. 12 Leaves glabrous, often shiny, often drying greenish, usually entire; stipules very short, inconspicuous, ± triangular; filaments hairy, anthers without an apical gland ............................................... C. zizyphoides Leaves glabrous or pubescent, drying brownish, usually toothed; stipules linear, deciduous; filaments glabrous, anthers with a small apical gland ................................................................................................... C. ulmifolia Lower surface of leaves hairy to densely minutely tomentose ............... 13 Lower surface of leaves glabrous, or, if pubescent, then only on the midrib .............................................................................................................. 16 Leaves ovate, apex obtuse, acute, or bluntly acuminate, the lower surface often with veins little raised but bearing hairs the same color or lighter than the rest of the lower blade; flowers pedicellate; inflorescence sessile ............................................................................................. C. mollis Leaves narrowly oblong, apex often narrowly long-acuminate, the lower surface often with veins prominently raised and darker than the rest of the blade; flowers pedicellate or sessile; inflorescence often pedunculate ........................................................................................................ 14

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F LACOURTIACEAE

14(13). Flowers on 2–4 mm long pedicels, puberulent to glabrous ........... C. arborea 14. Flowers sessile, finely tomentose ............................................................ 15 15(14). Lower surface of leaves tan in dried specimens; flowers 4–6 mm long ............................................................................................... C. grandiflora 15. Lower surface of leaves brown in dried specimens; flowers 6–8 mm long .................................................................................................. C. rusbyana 16(12). Disk lobes glabrous ..................................................................... C. singularis 16. Disk lobes laxly to densely hairy ............................................................. 17 17(16). Inflorescence on a pedicel 3–4 mm long; Serra Pirapucú, Brazil .... C. neblinae 17. Inflorescence sessile ................................................................................. 18 18(17). Flowers 2.5–3 mm long, sepals not reflexed; leaves generally drying olive green; anthers with apical gland ............................................ C. prunifolia 18. Flowers 4–6 mm long, sepals reflexed or not; leaves often drying brownish or blackish; anthers with apical gland or not ..................................... 19 19(18). Anthers with apical gland covered with stiff hairs; sepals not reflexed .................................................................................................... C. pitumba 19. Anthers lacking an apical gland; sepals usually reflexed ...................... 20 20(19). Pedicels usually filiform, ca. 0.1 mm diameter; leaves 5–18 cm long ........................................................................................... C. mariquitensis 20. Pedicels more robust, usually ca. 0.5 mm diameter; leaves 3–9(–10) cm long ........................................................................................... C. decandra Key to the Species of Casearia based on fruiting material 1. 1. 2(1). 2. 3(2).

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

7(6).

Fruit a dry capsule enclosed at least halfway up by the persistent sepals at maturity ............................................................................................. 2 Fruit a dry capsule enclosed less than halfway to not at all within the sepals at maturity, or else a fleshy capsule or berry ................................ 6 Mature fruit 10–15 mm long; leaves glabrous or nearly so, usually entire or nearly so .............................................................................. C. spruceana Mature fruit < 10 mm long; leaves glabrous to tomentose, especially on the lower surface, serrate to (sub)entire ............................................... 3 Leaves usually oblong to elliptic, with or without an acuminate tip, glabrous on the lower surface or with some pubescence on the larger veins ................................................................................................... C. ulmifolia Leaves usually narrowly oblong, often with a long-acuminate apex, usually minutely tomentose on the lower surface ...................................... 4 Fruits on pedicels 2–4 mm long; lower surface of leaves pubescent to subglabrous ......................................................................................... C. arborea Fruits sessile; lower surface of leaves tomentellous ................................. 5 Lower surface of leaves drying tan ........................................... C. grandiflora Lower surface of leaves drying brown ......................................... C. rusbyana Fruit a dry, dehiscent capsule, thin-walled (≤ 0.2 mm thick), often pubescent both inside and out ........................................................................ 7 Fruit a berry, a fleshy dehiscent or indehiscent capsule, or a dry indehiscent or tardily indehiscent capsule, thick-walled (≥ 0.5 mm thick), never pubescent inside .......................................................................... 9 Fruit 3–6 mm long ........................................................................ C. sylvestris

Casearia 443

7. 8(7). 8.

9(6). 9. 10(9). 10. 11(10). 11. 12(11).

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

15(13). 15. 16(15). 16. 17(15). 17. 18(17). 18. 19(18). 19. 20(19). 20.

Fruit 8–15 mm long .................................................................................... 8 Capsule with reddish pubescence inside and out; leaves often drying brownish; sepals usually reflexed below the fruit .................. C. javitensis Capsule usually not pubescent on the exterior though occasionally with some appressed grayish hairs, the interior often with conspicuous golden pubescence; leaves often drying greenish; sepals usually remaining erect below the fruit ........................................ C. commersoniana Fruits usually ≤ 10 mm long .................................................................... 10 Fruits usually > 10 mm long .................................................................... 13 Fruit subsessile or on a pedicel to 2 mm in length; sepals persistent under the fruit ................................................................................. C. zizyphoides Fruit on a pedicel 2–6 mm long; sepals generally not persistent ........... 11 Leaves 7–15(–19) cm long, tending to be obovate ..................... C. guianensis Leaves 3–14 cm long, most of them usually ≤ 7 cm long, variable in shape but generally not obovate .................................................................... 12 Leaves usually drying light to olive green; trunk or branches often bearing spines; plants deciduous or semideciduous; fruits often at leafless nodes ........................................................................................... C. aculeata Leaves usually drying brownish; plants unarmed; leaves generally persistent; fruits generally at leafy nodes ......................................... C. sylvestris Fruits fleshy-dehiscent or indehiscent capsules or berries, with thin skin (exocarp) which often becomes wrinkled when dried ......................... 14 Fruits dry, spongy, or pulpy inside, but with hard exocarp that does not wrinkle on drying, indehiscent or tardily dehiscent .......................... 15 Fruits 15–30 mm long, borne directly in the leaf axils; pedicels the same color below and above the articulation .................................. C. spinescens Fruits ca. 3 cm diameter, borne on a 4–9 mm long branchlet the same color as the other twigs; pedicels often darker than the branchlet bearing the fruit, in dried specimens usually blackish (in some specimens, the articulation of the pedicel can be difficult to see, but the color difference between the pedicel and the branchlet is usually evident) ..................................................................................................... C. tremula Lower surface of leaves (or at least the midrib) densely pubescent....... 16 Lower surface of leaves generally glabrous ............................................ 17 Fruits glabrous ................................................................................. C. hirsuta Fruits pubescent ................................................................................ C. mollis Fruits 5–7 cm diameter when ripe; leaves usually entire ......... C. singularis Fruits 0.8–3 cm diameter when ripe; leaves subentire to serrate or crenate .................................................................................................. 18 Fruits tomentellous, 1–2 cm diameter ................................. C. mariquitensis Fruits glabrous or else pubescent only toward the apex, 0.8–3 cm diameter ....................................................................................................... 19 Lateral leaf veins 3 or 4 per side, leaves often drying discolorous, with lower surface greenish and upper surface brownish ............. C. prunifolia Lateral leaf veins 4–8 per side, leaves often drying discolorous, but lower surface greenish and upper surface often almost black ..................... 20 Fruit 20–30 mm diameter when ripe ............................................ C. pitumba Fruit 8–10 mm diameter when ripe ............................................ C. decandra

444

F LACOURTIACEAE

Casearia aculeata Jacq., Enum. Syst. Pl. 21. 1760. —Espuelo de gallo, Limoncillo. Shrub or tree 3–10 m tall, reportedly with spines on trunk and branches; leaves 3–14 × 1.5–5 cm, oblong to elliptic or obovate, usually glabrous but sometimes pubescent on lower surface, entire to serrulate or crenulate; petioles 3–8 mm long; flowers usually 3–4 mm long; stamens 8, disk lobes interstaminal; style entire; fruit a purpleblack, fleshy-walled capsule 5–10 mm diameter. Semideciduous to riparian forests, 0– 200 m; Delta Amacuro (Cerro Sacupana), northern Bolívar, Amazonas (San Juan de Manapiare). Anzoátegui, Apure, Aragua, Barinas, Distrito Federal, Falcón, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Trujillo, Yaracuy, Zulia; widespread in the Neotropics from Mexico to Paraguay. ŠFig. 379.

Shrub to tree to 15 m tall; leaves 4–25 × 2–9 cm, usually glabrous, drying greenish, crenate, serrate, or subentire; petioles 5–10 mm long; flowers 3–8 mm long; stamens 10– 20, disk lobes intrastaminal; style 3-parted, sepals usually persistent under the fruit; fruit a capsule 8–15 mm long, usually glabrous outside or sometimes with appressed gray pubescence. Shrub savannas, disturbed forests, forest borders, 100–900 m; Delta Amacuro (Cerro La Paloma near Río Cuyubini), Bolívar (Canaima, Gran Sabana, Río Paragua), Amazonas (El Pozo southeast of San Fernando de Atabapo, around Puerto Ayacucho, Río Cataniapo, Río Yaciba, San Carlos de Río Negro). Mérida, Zulia; southern Mexico to Panama, Guyana, Suriname, French Guiana, Brazil (south to Mato Grosso and Rio de Janeiro).

Casearia arborea (Rich.) Urb., Symb. Antill. 4: 421. 1910. —Samyda arborea Rich., Actes Soc. Hist. Nat. Paris 1: 109. 1792. —Caicareño, Palo de cují, Palo de escoba, Pasita. Shrub or small tree 2–12 m tall; leaves 5– 12 × 2–4.5 cm, usually narrowly oblong, usually bicolorous, the lower surface densely covered with very fine gray pubescence that is sparser on the veins, margins serrulate; petioles 2–8 mm long; flowers 3.5–4.5 mm long, on pedicels 2–4 mm long; stamens 10, disk lobes interstaminal; style entire; fruit a dry capsule 4–5 mm long, enclosed more than halfway by the sepals. Disturbed forests, savannas, evergreen lowland to lower montane forests, 100–900 m; Bolívar (Gran Sabana, Isla Anacoco, La Vergareña, Maripa, Río Aro), Amazonas (upper Río Cuao, San Carlos de Río Negro, Santa Bárbara del Orinoco, base of Sierra de la Neblina, Tamatama). Anzoátegui, Aragua, Barinas, Cojedes, Guárico, Lara, Mérida, Miranda, Táchira, Trujillo, Zulia; Guatemala to Panama, Greater Antilles, Colombia, Guyana, Suriname, French Guiana, eastern Ecuador, eastern Peru, Brazil, Bolivia.

Casearia decandra Jacq., Enum. Syst. Pl. 21. 1760. —Dujo (Yekwana). Samyda parviflora Loefl., Iter Hispan. 260. 1758. Small tree 8–15 m tall; leaves 2–6(–10) × 1.5–5 cm, elliptic, ovate, or lanceolate, drying dark brown or blackish, finely serrate, usually glabrous; petioles 2–7(–10) mm long; flowers 4–6 mm long, on pedicels 5–10 mm long; stamens 10, disk lobes interstaminal; style entire; fruit a tough capsule 8–10 mm diameter, red-orange, wall 1–2 mm thick, usually glabrous. Semideciduous forests, lower montane forests, riparian forests, 50– 800 m; Delta Amacuro (Río Acure), Bolívar (Río Botanamo, Río Caura, Río Suapure, Salto Pará), Amazonas (Simarowochi). Aragua, Carabobo, Falcón, Guárico, Lara, Miranda, Nueva Esparta, Sucre; Honduras to Panama, Antilles, south to Brazil (Rio Grande do Sul), Bolivia, Paraguay, northern Argentina, and northern Uruguay. ŠFig. 382.

Casearia commersoniana Cambess. in A. St.-Hil. et al., Fl. Bras. Merid. 2: 235. 1829 [1830]. Casearia densiflora Benth., J. Bot. (Hooker) 4: 113. 1841. Casearia laurifolia Benth., J. Bot. (Hooker) 4: 113. 1841.

Casearia grandiflora Cambess. in A. StHil. et al., Fl. Bras. Merid. 2: 168, t. 126. 1829 [1830]. —Caicareno, Fruta de paloma, Majagua sabanero, Nigüito, Peinesito, Uruate (Piaroa), Vara blanca. Shrub or tree 1.5–20 m tall; leaves 4–13 × 1.5–5 cm, narrowly oblong, markedly bicolorous with very dense tan pubescence on lower surface, minutely serrate; flowers 4–6 mm long, sessile; stamens 10, disk lobes interstaminal; style entire; fruit a dry capsule 7–8 mm long, enclosed more than half-

Casearia

way by the sepals. Secondary forests, savannas, deciduous to evergreen forests, forestsavanna borders, gallery forests, 100–1500 m; widespread in Bolívar and Amazonas. Anzoátegui, Barinas, Mérida, Sucre, Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Brazil. ŠFig. 383. A collection of this species from the slopes of Uaipán-tepui, Koyama & Agostini 7327 (VEN), is a host of the Rafflesiaceae parasite, Apodanthes caseariae Poit.

445

mm long, pedicels 4–6 mm long; stamens 8, disk lobes interstaminal; style entire; fruit a dark purple berry 15–20 mm diameter with walls ca. 1 mm thick. Semideciduous and evergreen forests, 100–500 m; Bolívar (Altiplanicie de Nuria, Caño Aliviadero near Tumeremo, 85 km southeast of Entrerios, Lago Guri area, Río Acanán, Río Suapure), Amazonas (slopes of Sierra de la Neblina). Barinas, Guárico, Portuguesa, Zulia; Costa Rica, Cuba, Jamaica, Colombia, northeastern Brazil. ŠFig. 384.

Casearia guianensis (Aubl.) Urb., Symb. Antill. 3: 322. 1902. —Iroucana guianensis Aubl., Hist. Pl. Guiane 329 (guianensis), t. 127 (guyannensis). 1775. —Athenaea guianensis (Aubl.) J.F. Gmel., Syst. Nat. 2: 629. 1791. —Cuspa, Rastrojero, Tapaculo. Casearia arguta auct. non H.B.K. 1821 [1823]: sensu Steyerm., Fieldiana, Bot. 28: 992. 1957. Casearia spinescens auct. non (Sw.) Griesb. 1863: sensu Pittier et al., Cat. Fl. Venez. 2: 170. 1947. Shrub to tree 4–15 m tall; leaves 7–19 × 3–7 cm, usually obovate, laxly pubescent on lower surface, at least on the major veins, coarsely serrate-crenate to subentire, usually drying brown or reddish or greenish brown; petioles 5–10 mm long; flowers 4–5 mm long, on pedicels 4–6 mm long; stamens 8, disk lobes interstaminal; style entire; fruit a fleshy capsule 6–10 mm diameter. Evergreen or deciduous to riparian forests, 100– 300 m; Delta Amacuro (Serranía de Imataca), northeastern Bolívar. Apure, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Sucre, Trujillo, Zulia; Panama, Antilles, Colombia, Guyana, Suriname, French Guiana, Brazil. ŠFig. 381.

Casearia javitensis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 366, t. 479. 1821 [1823]. —Chaetocrater javitensis (H.B.K.) Raf., Sylva Tellur. 149. 1838. —Apoykapundek (Pemón), Kawadi jodedi (Yekwana), Palo de ratón. Shrub or tree to 17 m tall; leaves 6–32 × 2–11 cm, oblong-ovate, usually glabrous and somewhat glossy, drying matte brownish or green, coarsely serrate to crenate or subentire, with 5–7 secondary veins per side; petioles 3–10 mm long; flowers 3–5 mm long, pedicels 3–10 mm long; stamens 10, drying blackish, disk lobes intrastaminal; styles 3branched; ovary and fruit with reddish pubescence inside and out; sepals usually reflexed under the fruit; fruit capsular, 8–15 mm long, brown to red when ripe; seed light brown with white aril. Secondary forests, evergreen lowland to montane forests, 100– 1200 m; Bolívar (Cerro Marutaní, 40–60 km south of El Dorado, Gran Sabana, 45 km east of Los Pijiguaos, Río Cucurital, Río Paragua, Río Paramichí, Río Paujil, Río Venamo), Amazonas (widespread). Guárico, Mérida; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 387.

Casearia hirsuta Sw., Fl. Ind. Occid. 755. 1798. —Samyda hirsuta (Sw.) Poir. in Lam., Encycl. 6: 493. 1804 [1805]. —Guidonia hirsuta (Sw.) M. Gómez, Dicc. Bot. Nom. Vulg. 94. 1889. Casearia mollis auct. non H.B.K. 1821 [1823]: sensu Steyerm., Acta Bot. Venez. 3: 141. 1968. Shrub to 4 m tall; leaves 4–18 × 2.5–8 cm, elliptic, usually with erect, dense, golden pubescence on lower surface and on veins above, minutely to coarsely serrate to subentire; petioles 3–8 mm long; flowers 5–7

Casearia mariquitensis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 363. 1821 [1823]. —Nigua, Tapaculo. Shrub to 5 m tall; leaves 5–18 × 2.5–4 cm, oblong to lanceolate or elliptic, usually glabrous or with sparse trichomes on veins, finely serrate; petioles mostly 2–5 mm long; flowers 4–6 mm long, on slender pedicels 3–8 mm long; stamens 10, disk lobes interstaminal; style entire; fruit a tough capsule, globose, pubescent, 10–20 mm diameter, with fine yellowish or reddish pubescence; seeds with orange aril. Semideciduous for-

446

F LACOURTIACEAE

ests, savannas, riparian forests, 100–800 m; scattered through much of Bolívar and Amazonas. Anzoátegui, Apure, Barinas, Cojedes, Guárico, Lara, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (south to Paraná). ŠFig. 389. Casearia mariquitensis, along with C. decandra, C. arguta H.B.K., C. mollis, and C. pitumba, form a poorly understood complex. Casearia mollis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 365, t. 480. 1821 [1823]. —Tapaculo. Casearia arguta, auct. non H.B.K. 1821 [1823]: sensu Pittier et al., Cat. Fl. Venez. 2: 169. 1947, pro parte. Casearia lasiophylla auct. non Eichler 1871: sensu Pittier et al., Cat. Fl. Venez. 2: 170. 1947. Shrub to small tree; leaves 2–5(–20?) × 1.5–3(–8?) cm, ovate to elliptic-oblong, covered with a thick golden tomentum all over the lower surface, irregularly serrate; petioles 2–5 mm long; flowers 2–5 mm long, pedicels 2–4 mm long; stamens 10, disk lobes interstaminal; style entire; fruit a tough capsule 10–20 mm diameter, finely pubescent. Deciduous forests, 50–200 m; Bolívar (Caicara–Puerto Ayacucho road). Widespread elsewhere in Venezuela; Cuba, Colombia. ŠFig. 391. Casearia neblinae Sleumer, Fl. Neotrop. Monogr. 22: 322. 1980. Tree to 15 m tall; leaves 7–12 × 4–5.5 cm; petioles ca. 8 mm long; flowers ca. 4 mm long on pedicels 3–4 mm long; immature capsule subglobose, tuberculate, ca. 4 mm diameter. Montane forests, ca. 1200–1300 m; Amazonas (expected on Sierra de la Neblina). Brazil (Amazonas: Serra Pirapucú). Casearia neblinae is known only from the type collection from Brazil, very close to the Venezuelan border. Casearia pitumba Sleumer, Blumea 24: 118. 1978. —Pitumba guianensis Aubl., Hist. Pl. Guiane 2(suppl.): 29, t. 385. 1775, non Casearia guianensis (Aubl.) Urb. 1902. —Samyda pitumba Poir. in Lam., Encycl. 6: 492. 1804 [1805]. —Dujo (Yekwana), Nigua. Casearia macrophylla Vahl, Eclog. Amer. 2: 32. 1798.

Shrub or tree 3–10 m tall; leaves 4–20 × 2–10 cm, elliptic-oblong, glabrous or with sparse trichomes on main veins of lower surface, serrate, crenate, or subentire; petioles 3–7 mm long; flowers 4–6 mm long, pedicels 2–6 mm long; stamens 10, the anthers with an apical gland covered with stiff trichomes, the disk lobes interstaminal; style entire; fruit a globose, woody capsule 20–30 mm diameter, mostly glabrous, yellow, the fruit wall 1–3 mm thick. Evergreen lowland to montane forests, riparian forests, 100–1300 m; Delta Amacuro (Río Toro), Bolívar (Cerro Venamo, Río Caura, Santa Elena de Uairén). Mérida, Táchira; Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil. ŠFig. 392. Correlated flowering and fruiting material, preferably from the same plants at different times of the year, is needed for this species, as well as others in this complex, especially Casearia decandra and C. mariquitensis. Casearia prunifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 364. 1821 [1823]. —Palo de conejo, Palo de gallineta. Shrub or tree to 20 m tall; leaves 5–15 × 2.5–4.5 cm, elliptic or oblong, glabrous, drying olive green, serrulate or crenulate to subentire; petioles 3–6 mm long; flowers 2.5– 3.5 mm long, pedicels 3–6 mm long; stamens 10, disk lobes interstaminal; style entire; fruit glabrous, 10–20 mm diameter. Evergreen lowland to lower montane forests, 100–600 m; Amazonas (upper Río Cuao, San Carlos de Río Negro, Serranía Batata 55 km southeast of Puerto Ayacucho). Colombia, Ecuador, Peru. Casearia rusbyana Briq., Annuaire Conserv. Jard. Bot. Genève 2: 73. 1898. —Palmito. Tree to 30 m tall; leaves ca. 15 × 5 cm, narrowly oblong, subcoriaceous, usually drying dark brown, serrate, lower surface covered with a fine pubescence; petioles 3–5 mm long; flowers 6–8 mm long, sessile; fruit a capsule ca. 7 mm long included more than halfway within the sepals. Evergreen lowland to lower montane forests, 50–700 m; Delta Amacuro (Santa Catalina, Serranía de Imataca), Bolívar (Altiplanicie de Nuria). Guyana, Suriname, French Guiana, Brazil (Amapá, Rondônia).

Casearia

Casearia singularis Eichl. in Mart., Fl. Bras. 13(1): 473, t. 95, fig. 2. 1871. —Maspara. Tree 10–20 m tall; leaves 8–25 × 4–10 cm, ovate to obovate, usually glabrous, margins entire or subsinuate; petioles 3–10 mm long; flowers 4–5 mm long, on pedicels 3–6 mm long; stamens 10, disk lobes interstaminal, glabrous; style entire; fruit a tough-skinned berry 5–7 cm diameter, edible, fruit wall to 15 mm thick. Evergreen lowland forests, 100–200 m; Delta Amacuro (Serranía de Imataca), Amazonas (Río Mawarinuma, upper Río Yatúa). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 388. Collections of this species from Venezuela were previously identified as Casearia combaymensis Tul., following the monograph of H. O. Sleumer (Fl. Neotrop. Monogr. 22: 305. 1971). That species, however, is characterized by much larger stipules and moretoothed leaves; it is restricted to Colombia, Ecuador, and Peru. Casearia spinescens (Sw.) Griseb., Cat. Pl. Cub. 10. 1866. —Samyda spinescens Sw., Prodr. 68. 1788. —Guidonia spinescens (Sw.) Griseb., Fl. Brit. W. I. 24. 1859. —Cordilla, Pardillo, Punteral. Casearia aculeata auct. non Jacq. 1760: sensu Pittier et al., Cat. Fl. Venez. 2: 169. 1947. Deciduous shrub or tree to 5 m tall, generally pyramidally shaped, shoots often vining and divaricating; leaves often clustered at the branch tips, 2–10 × 1.5–5 cm, elliptic to obovate, glabrous or with scattered appressed trichomes, entire to serrulate or crenate; petioles 2–8 mm long; flowers ca. 5 mm long, pedicels 2–5 mm long; stamens 10, the disk lobes interstaminal and adnate to the filaments, forming a high staminal tube; style entire; fruit a fleshy capsule 15–30 mm long, purple when ripe. Deciduous forests, spiny shrublands, riparian forests, 0–400 m; Delta Amacuro (near Sacupana), common in dry areas of northern Bolívar. Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Miranda, Monagas, Sucre, Zulia. Panama, Bahamas, Cuba, Hispaniola, Trinidad, Tobago, Guyana, Brazil (Roraima). ŠFig. 394. Casearia spruceana Benth. ex Eichler in Mart., Fl. Bras. 13(1): 486. 1871. —Ojito de pescado.

447

Shrub to small tree 2–5 m tall; leaves 6– 25 × 2.5–9.5 cm, narrowly oblong-lanceolate, glabrous or subglabrous, drying brownish, serrulate to subentire; petioles 5–8 mm long; flowers 6–15 mm long, pedicels 2–5 mm long; stamens 20–25, drying blackish, prominent against pale intrastaminal disk lobes; style divided into 3 branches toward the apex; fruit a dry capsule 10–15 mm long, included within the persistent sepals that are accrescent to 10–14 mm long. Seasonally flooded riparian forests, 50–200 m; Amazonas (Río Orinoco below mouth of Río Atabapo, mouth of Río Ventuari, San Carlos de Río Negro). Brazil. ŠFig. 393. Casearia sylvestris Sw., Fl. Ind. Occid. 2: 752. 1798. —Samyda silvestris (Sw.) Poir. in Lam., Encycl. 6: 492. 1804 [1805]. Casearia sylvestris var. carpinifolia (Benth.) Briq., Annuaire Conserv. Jard. Bot. Genève 2: 74. 1898. Casearia inaequilatera auct. non Cambess. 1829 [1830]: sensu Steyerm., Fieldiana, Bot. 28: 990. 1957. Shrub or tree to 15 m tall; leaves 3–14 × 2–7 cm, oblong to ovate or oblong-ovate, usually glabrous, serrulate or crenulate to entire; petioles 4–10 mm long; flowers 1.5–2 mm long, pedicels 1–5 mm long; stamens 10, disk lobes interstaminal; style 3-branched; fruit a globose capsule 3–6 mm long, splitting open widely at maturity, remnants of the 3-parted style often visible; seeds with an orange aril. Mexico to Argentina; 2 varieties, both in the flora area. This species might be confused with C. zizyphoides, but it is the only species in the flora area with a 3-parted style and intrastaminal disk lobes. Key to the Varieties of C. sylvestris 1. Leaves ± ovate to oblong-ovate, reticulations of veins and veinlets distinctly raised on both surfaces and often discolorous ........................ var. lingua 1. Leaves generally oblong, glabrous or laxly to densely gray-puberulous, reticulations of veins and veinlets rather obscure ............................. var. sylvestris C. sylvestris var. lingua (Cambess.) Eichler in Mart., Fl. Bras. 13(1): 482. 1871.

448

F LACOURTIACEAE

—Casearia lingua Cambess. in A. St.Hil. et al., Fl. Bras. Merid. 2: 171. 1829 [1830]. —Pan de acure, Tortolito. Shrub 1–4 m tall; trunk with corky layer resistant to fire. Savannas, 100–1100 m; Bolívar (widespread), Amazonas (Canaripó, Cerro Parú, Puerto Ayacucho). Widespread in northern Venezuela; Colombia, Guyana, Suriname, Brazil, Bolivia. ŠFig. 385. C. sylvestris var. sylvestris. —Tortolito. Shrub 0.5 m tall or tree to 15 m tall. Deciduous to moist forests, gallery forests, savanna edges, 50–1200 m; Delta Amacuro (10 km east-southeast of Castillos de Guayana, 13 km southeast of Piacoa), Bolívar (widespread in northern part, base of Ptari-tepui), Amazonas (Salto Salas on upper Río Orinoco, Santa Bárbara del Orinoco). Widespread in northern Venezuela; other distribution as in species. Casearia tremula (Griseb.) Griseb. ex Wright in Sauvalle, Anales Acad. Ci. Méd. Habana 5: 201. 1868. —Zuelania tremula Griseb., Cat. Pl. Cub. 9. 1866. —Guidonia tremula (Griseb.) Maza, Anales Soc. Esp. Hist. Nat. 23: 56. 1894. —Guinda. Shrub or tree to 10 m tall; leaves 3–16 × 2–8 cm, ovate-oblong to elliptic, with 8–12 lateral veins per side, glabrous or with trichomes on lower surface midrib, serrulate to crenulate or subentire; petioles 5–25 mm long; flowers 4–8 mm long, pedicels 5–15 mm long; stamens 15–25, disk lobes intrastaminal; style entire; fruit a fleshy capsule, large, tough, orbicular, ca. 3 cm diameter, borne on a 4–9 mm long branchlet the same color as the other twigs; pedicels often darker than the branchlet bearing the fruit; pulp reddish. Deciduous forests, 0–100 m; Bolívar (Puerto Ordaz, San Félix). Anzoátegui, Distrito Federal, Falcón, Dependencias Federales, Lara, Miranda, Monagas, Nueva Esparta, Sucre, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia. ŠFig. 380. Casearia ulmifolia Vahl ex Vent., Mém. Math. Phys. Inst. Nat. France 2: 150. 1808; Choix Pl. 46. 1808. —Guidonia ulmifolia (Vahl ex Vent.) Baill., Traité Bot. Méd. Phan. 2: 827. 1884. —Juasadi (Yekwana), Nirgua, Vara blanca.

Casearia celtidifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 363. 1821 [1823]. —Casearia celtidifolia (H.B.K.) Poepp. ex Eichler in Mart., Fl. Bras. 13(1): 477. 1871. Shrub or tree 2–30 m tall; buds pubescent; leaves 8–14 × 3–5.5 cm, oblong to elliptic, glabrous on lower surface or with some pubescence on larger veins, serrate; stipules linear, deciduous, 6–9 mm long; petioles 5–8 mm long; flowers 1.5–2.5 m long, sessile or pedicels to 3 mm long; stamens 10, filaments glabrous with a small apical gland, disk lobes interstaminal; style entire; fruit a dry capsule 3–4 mm long mostly covered by the persistent sepals. Deciduous to lower montane forests, edges of savannas, 50–700 m; Bolívar (Altiplanicie de Nuria, southwest of Caicara, near Maripa, Lago Guri, Río Pao, Río Paragua, Río Tabaro near Río Nichare, San Pedro de las Dos Bocas), Amazonas (Puerto Ayacucho and area to the north). Widespread in northern Venezuela; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 386. Casearia zizyphoides H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 362. 1821 [1823]. —Tortolito. Casearia celastroides Klotzsch in M.R. Schomb., Reis. Br.-Guiana 3: 1167. 1848 [1849], nom. nud. Casearia laevigata Willd. ex Klotzsch in M.R. Schomb., Reis. Br.-Guiana 3: 1167. 1848 [1849], nom. nud. Shrub or tree 1–8 m tall; leaves 3–6 × 2–3 cm, generally elliptic to oblong, glabrous, with a blunt narrowed tip, often shiny, usually entire and drying greenish; petioles 2–3 mm long; flowers 1.5–2 mm long, pedicels 1–2 mm long; stamens 10, filaments hairy, anthers without an apical gland, disk lobes interstaminal; style entire; fruits fleshy, 4–5 mm long, orange. Semideciduous forests, seasonally flooded savannas, 100–500 m; widespread in northeastern Bolívar, Amazonas (Puerto Ayacucho, Río Ocamo near Raudal Arata, San Juan de Manapiare). Apure, Distrito Federal, Falcón, Lara, Miranda, Monagas, Nueva Esparta, Sucre, Yaracuy, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ŠFig. 390.

Casearia

Fig. 379. Casearia aculeata

Fig. 380. Casearia tremula

449

450

F LACOURTIACEAE

Fig. 381. Casearia guianensis Fig. 382. Casearia decandra

Fig. 383. Casearia grandiflora

Casearia

451

Fig. 384. Casearia hirsuta

Fig. 385. Casearia sylvestris var. lingua

Fig. 386. Casearia ulmifolia

452

F LACOURTIACEAE

Fig. 387. Casearia javitensis

Fig. 388. Casearia singularis

Fig. 389. Casearia mariquitensis

Casearia

Fig. 390. Casearia zizyphoides

Fig. 391. Casearia mollis Fig. 392. Casearia pitumba

453

454

F LACOURTIACEAE

Fig. 393. Casearia spruceana

Fig. 394. Casearia spinescens

Euceraea

455

4. EUCERAEA Mart., Nov. Gen. Sp. Pl. 3: 90, t. 238. 1829 [1831]. Shrubs or small trees, the foliage grouped at the ends of the branchlets. Leaves spirally arranged, chartaceous to coriaceous, glandular-serrulate, pellucid-punctate and -lineate, pinnately veined, markedly reticulate, petiolate; stipules ovate-lanceolate, enveloping the terminal bud, but then soon deciduous. Inflorescences a simple spike or panicle of spikes, axillary to subterminal. Flowers staminate (with small abortive ovary) or bisexual (though some bisexual flowers may have abortive anthers), small, almost immersed in the rachis with their base, sessile, the subtending bract and 2 bracteoles subpersistent (plants either androdioecious or functionally dioecious). Sepals 4, slightly connate at base, white, imbricate; petals absent. Stamens 8, the 4 longer ones opposite the sepals, the 4 shorter ones alternating with them; filaments filiform; anthers small, bilocular, subglobose, introrse, basifixed, longitudinally dehiscent. Disk lobes 8, alternating with the stamens and connate with them at base, spatulate, barbate. Ovary superior, 1-locular, with 2 parietal placentas, each of them bearing a single ovule; style 0; stigma sessile, with 4–6 radiating lobes. Fruit a berry. Seeds 1 or 2, each with a short lacerate aril. Colombia, Venezuela, Guyana, Suriname, Brazil; 3 species, all in the flora area.

Fig. 395. Euceraea rheophytica

456

F LACOURTIACEAE

Fig. 396. Euceraea nitida

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

2.

Leaves narrowly lanceolate, 4–8 times longer than wide, 4–8 × 0.4–1.5 mm; flowers in a simple spike 2–4 cm long ................................. E. rheophytica Leaves narrowly to broadly elliptic, < 4 times as long as wide, much larger than above; flowers in a branched panicle > 4 cm long ........................ 2 Leaves narrowly elliptic, lateral veins 12–40 on either side of the midvein, margin crenulate throughout, the lower order reticulation prominent on both surfaces ............................................................................. E. nitida Leaves broadly elliptic, lateral veins 8–12 on either side of the midvein, margin serrate in distal half, entire and slightly revolute in lower half, the lower order reticulation often obscure on lower surface .................. ............................................................................................. E. sleumeriana

Euceraea nitida Mart., Nov. Gen. Sp. Pl. 3: 90, t. 238. 1829 [1831]. —Jabón. Shrub or small tree to 10 m tall. Gallery forests, streamsides, forest-savanna borders, evergreen lowland to montane forests, 100– 1700 m; Bolívar (Cerro Guaiquinima, Cerro Marujano in upper Río Caura basin, wide-

spread in the Gran Sabana, Macizo del Chimantá, headwaters of Río Túriba 45 km east of Los Pijiguaos), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Yutajé). Amazonian Colombia, Guyana, Suriname, Brazil. ŠFig. 396.

Hecatostemon 457

Euceraea rheophytica P.E. Berry & M. Olson, Brittonia 50: 493. 1998. Rheophytic shrub 1–2 m tall; leaves finely serrate, willow-like, grouped toward the branch tips. Rocky, stream banks in canyons, 300–800 m; Amazonas (Cañon Grande of upper Río Baría or Río Mawarinuma in the Sierra de la Neblina). Endemic. ŠFig. 395.

Euceraea sleumeriana Steyerm. & Maguire, Mem. New York Bot. Gard. 23: 870. 1972. Shrub or small tree to 3 m tall. Tepui shrublands and low forests, 1800–2100 m; Bolívar (Cerro Sarisariñama). Endemic.

5. HECATOSTEMON S.F. Blake, Contr. Gray Herb. 53: 42. 1918. Shrub or tree. Leaves alternate, deciduous, subserrate, subcrenate, or subentire, pinnately veined, pellucid-punctate and -lineate, petiolate, with deciduous stipules. Inflorescences axillary cymes, contracted, short-pedunculate, and rather few-flowered. Flowers bisexual, rather small, precocious, receptacle very shallow. Calyx lobes 5, imbricate, persistent; petals absent. Stamens 80–100, free,

Fig. 397. Hecatostemon completus

458

F LACOURTIACEAE

arranged in 3 series, inserted at the base of the calyx outside the disk, which consists of ca. 15 uniseriate, free, short-clavate, and flattened, distally hairy lobes or staminodes; filaments filiform; anthers oblong-elliptic, almost basifixed, dehiscing lengthwise, connective small. Ovary sessile, 1-locular, hairy, with 3 multiovulate placentas; style attenuate to rather short and glabrous; stigma small, capitate, faintly 3lobed. Fruit capsular, subglobose at maturity, dehiscent by 3 valves; pericarp coriaceous. Seeds numerous. Colombia, Venezuela, Guyana, Brazil (Roraima); 1 species. Hecatostemon completus (Jacq.) Sleumer, Blumea 24: 118. 1978. —Laetia completa Jacq., Enum. Syst. Pl. 24. 1760. Laetia guazumaefolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 356. 1821 [1823]. —Hecatostemon guazumaefolius (H.B.K.) Sleumer, Notizbl. Bot. Gart. BerlinDahlem 12: 55. 1934. Laetia hirtella H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 355. 1821 [1823]. Hecatostemon dasygynus S.F. Blake,

Contr. Gray Herb. ser. 2, 53: 43. 1918. Deciduous shrub or small tree to 10 m tall; lower surface of leaves softly puberulent. Deciduous forests, shrub savannas, disturbed areas, 50–200 m; Bolívar (Angosturita, Caicara, Ciudad Bolívar, Las Bonitas, across the Río Orinoco from Puerto Páez). Apure, Anzoátegui, Barinas, Cojedes, Falcón, Guárico, Miranda, Portuguesa, Zulia; other countries as in genus. ŠFig. 397.

6. HOMALIUM Jacq., Enum. Syst. Pl. 5, 24. 1760. Shrubs or trees, occasionally tall and with buttresses. Leaves (in American species) alternate, pinnately veined, usually serrate, sometimes repand-crenate, or subentire (the teeth glandular beneath), petiolate; stipules minute, deciduous or rarely absent. Flowers bisexual, in axillary and/or (sub)terminal, many-flowered racemes or panicles, solitary or fascicled along the rachis, sessile or pedicellate, the articulate pedicel subtended by an unusually small, deciduous or persistent bract. Calyx tube obconical or turbinate (may be pediform at the base), connate with the inferior part of the ovary, calyx lobes or sepals 5–8(–12), usually narrow, in American species hardly accrescent; petals inserted in the throat of the calyx, as many as and similar to, though usually larger than the sepals and alternating with them. Stamens epipetalous, in the American species in fascicles of (2)3–7; filaments filiform; anthers small, extrorse, dorsifixed. Disk represented by a (mostly pubescent) gland opposite each sepal. Ovary free in its upper part, conical, apex extenuate to the styles (as many in number as the placentas), in all American species forming a column, 1locular, placentas 2–6(–8), each with 1–few ovules; stigma simple or capitellate. Fruit capsular, half-inferior, rather coriaceous, indehiscent or 2–8-valved from the apex. Seeds solitary or few, small, angular, often hairy; testa crustaceous; endosperm copious. Pantropics; ca. 200 species (only 3 of these in the Americas), 2 in Venezuela, both in the flora area. Key to the Species of Homalium 1. 1.

Flowers sessile or subsessile; petals deltate-ovate, 2.5–3 mm long ................................................................................................. H. guianense Flowers distinctly pedicellate; petals ovate, 3.5–6 mm long .................. .............................................................................................. H. racemosum

Laetia

459

Fig. 398. Homalium guianense

Fig. 399. Homalium racemosum

Homalium guianense (Aubl.) Oken, Allg. Naturgesch. 3: 810. 1841. —Racoubea guyannensis Aubl., Hist. Pl. Guiane 590, t. 236. 1775. —Homalium racoubea Sw., Prodr. 86. 1788. —Dujo (Yekwana). Napimoga guyannensis Aubl., Hist. Pl. Guiane 590, t. 237. 1775. —Homalium napimoga Spreng., Syst. Veg. 4(cur. post.): 210. 1827. Homalium puberulum Klotzsch in M.R. Schomb., Reis. Br.-Guiana 985. 1848, nom. nud. Homalium densiflorum Spruce ex Benth., J. Proc. Linn. Soc., Bot. 4: 36. 1860. Shrub or tree to 25 m tall. Deciduous to evergreen lowland forests, riparian forests, usually in the understory, 50–400 m; Delta Amacuro (Río Amacuro between Amacuro and Agua Muerta), Bolívar (Isla Anacoco, upper Río Caroní, headwaters of Río Caura, Río Chiguao, mouth of Río Nichare, Río Paragua, Río Tabaro, Salto Pará on Río Caura). Apure; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 398.

Homalium racemosum Jacq., Enum. Syst. Pl. 24. 1760. —Caramacate blanco, Cedrito, Lute (Arekuna), Nigua, Palo gallineta. Homalium pedicellatum Spruce ex Benth., J. Proc. Linn. Soc., Bot. 4: 36. 1860. Homalium eleuthrostylum S.F. Blake, Contr. U.S. Natl. Herb. 20: 232. 1919. Homalium eurypetalum S.F. Blake, Contr. U.S. Natl. Herb. 20: 234. 1919. Homalium pittieri S.F. Blake, Contr. U.S. Natl. Herb. 20: 230. 1919. Homalium anzoateguiense Steyerm., Fieldiana, Bot. 28: 410. 1952. Homalium mituense Cuatrec., Trop. Woods 101: 19. 1955. Shrub or tree to 25 m tall. Seasonally flooded river banks and riparian forests, 50– 1400 m; widely scattered throughout Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Barinas, Carabobo, Distrito Federal, Falcón, Mérida, Miranda, Portuguesa, Zulia; Guatemala to Panama, West Indies, Colombia, Guyana, Suriname, French Guiana, eastern Peru, Brazil. ŠFig. 399.

460

F LACOURTIACEAE

7. LAETIA Loefl. ex L., Syst. Nat. ed. 10, 2: 1068, nom. cons. Shrubs or trees. Leaves alternate, distichous, entire or glandular-crenate or -serrate, sometimes pellucid-punctate, pinnately veined, petiolate; stipules deciduous. Inflorescences axillary, slightly supra-axillary, or subterminal, fasciculate or cymose. Flowers bisexual; pedicels articulate at or a little above the base; bracts and bracteoles free or sometimes united into a cupule. Sepals 4 or 5, imbricate, free or shortly connate at the base, often reflexed in anthesis; petals absent. Stamens 10– many, inserted on a small disk; filaments free, often of varying lengths; anthers elliptic to almost linear, introrse, basifixed or almost so, longitudinally dehiscent. Ovary free, 1-locular, with 3(–6) parietal, few- to many-ovulate placentas; style short, simple or shortly 3-lobed; stigmas capitate or slightly lobed. Fruit a berry-like capsule, tardily valvately dehiscent, sometimes resinous within; pericarp coriaceous. Seeds few to rather numerous, arillate; testa leathery; endosperm copious; embryo straight; cotyledons broad and foliaceous. Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 species, 5 species in Venezuela, all in the flora area. Key to the Species of Laetia 1. 1. 2(1).

2.

3(1).

3. 4(3).

4.

Bracts united into a cup that persists under the fruit; leaves (5–)7–18(–25) cm long, with 3–5(6) lateral veins; flowers in fascicles ......................... 2 Bracts free; leaves 3–16(–21) cm long, with 8–12(–14) lateral veins; flowers in fascicles or cymes ............................................................................... 3 Tips of youngest branches with minute pubescence; cups of bracts only slightly incised or lobed; leaves not shiny on upper surface when dry; flowering pedicels ± 5 mm long ................................................. L. cupulata Tips of youngest branches glabrous or subglabrous; cups of bracts deeply 2- or 3-lobed; leaves shiny on upper surface when dry; flowers sessile ..................................................................................................... L. coriacea Flowers in pedunculate dichasia, often with missing flowers or branches, flowers and fruits always borne directly from the leaf axils; leaves 3–7(–10) cm long ...................................................................... L. americana Flowers pedicellate but borne in sessile fascicles in or slightly above the leaf axils; leaves (6–)7–10(–21) cm long ................................................. 4 Flowers and fruits emerging ca. 2 mm above the leaf axils (even when not in flower or fruit it is usually possible to see cushions of bracts or scars above the leaf axils or leaf scars); stamens (12–)15–20; leaf base truncate ...................................................................................................... L. procera Flowers and fruits emerging directly from the leaf axils; stamens 10–12; leaf base cuneate ........................................................ L. suaveolens

Laetia americana L., Syst. Nat. ed. 10, 2: 1074. 1759. Laetia apetala Jacq., Enum. Syst. Pl. 24. 1760. —Guidonia apetala (Jacq.) Kuntze, Revis. Gen. Pl. 1: 44. 1891.

Azara umbellata C. Presl, Reliq. Haenk. 2: 92. 1835. Laetia acuminata Bonpl. ex Triana & Planch., Ann. Sci. Nat. Bot. sér. 6, 17: 104. 1862.

Laetia 461

Shrub or small tree to 10 m tall. Deciduous to semideciduous forests, 50–100 m; Bolívar (middle Río Orinoco). Barinas; Colombia, Ecuador, eastern Peru, Brazil, Bolivia, Paraguay. Laetia coriacea Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 84. 1861. —Guidonia coriacea (Spruce ex Benth.) Kuntze, Revis. Gen. Pl. 1: 44. 1891. —Kodojijoto grande (Yekwana). Shrub or small tree 1–6 m tall. Whitesand savannas, white-sand scrub (bana), Rio Negro caatinga, 100–600 m; Bolívar (El Cácaro between Rio Caura and Rio Paragua), Amazonas (slopes of Cerro Aracamuni, Cerro Vinilla, southwest of Cerro Yapacana, 10 km west of La Esmeralda, near Maroa, Río Atabapo basin, Río Autana, Río Guayapo, near San Carlos de Río Negro, base of Sierra de la Neblina). Northern Brazil. ŠFig. 400. Laetia cupulata Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 84. 1861. —Guidonia cupulata (Spruce ex Benth.) Kuntze, Revis. Gen. Pl. 1: 44. 1891. Shrub or tree 1–17 m tall. Evergreen lowland to lower montane forests, 100–600 m, Amazonas (Caño Negro east of Río Cunucunuma, La Esmeralda, Maroa, Río Sipapo, San Carlos de Río Negro, slopes of Sierra de la Neblina). Colombia (Vaupés), Guyana, Peru, Brazil. ŠFig. 402. Laetia procera (Poepp.) Eichler in Mart., Fl. Bras. 13(1): 453. 1871. —Samyda procera Poepp., Nov. Gen. Sp. Pl. 3: 67. 1845. —Casinga procera (Poepp.) Griseb., Fl. Brit. W. I. 710. 1864. —Guidonia procera (Poepp.) Kuntze, Revis. Gen. Pl. 1: 44. 1891. —Caimito cimarrón, Charo macho, Jobo macho, Pericoca, Piricoca, Sajokonudu (Yekwana). Laetia casearioides Sagot ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 85. 1861. Tree 8–40 m tall. Evergreen lowland to lower montane forests, riparian forests, 50– 600 m; Delta Amacuro (4–15 km east-southeast of Los Castillos de Guayana), Bolívar (slopes of Amaruay-tepui, Cerro Guaiquinima, near El Paraíso on San Félix to Upata road, La Escalera, Río Botanamo, Río Canaracuni, Río Supamo, Río Tabaro, Río Toro),

Amazonas (Río Cataniapo 48 km southeast of Puerto Ayacucho). Barinas, Mérida, Miranda, Monagas, Táchira, Zulia; Guatemala to Panama, Hispaniola, Puerto Rico, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 401. Laetia suaveolens (Poepp.) Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 85. 1861. —Samyda suaveolens Poepp., Nov. Gen. Sp. Pl. 3: 66, t. 274. 1845. —Casinga suaveolens (Poepp.) Griseb. ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 85. 1861. —Guidonia suaveolens (Poepp.) Kuntze, Revis. Gen. Pl. 1: 44. 1891. —Cazabe de tara. Laetia calophylla Eichler in Mart., Fl. Bras. 13(1): 454, t. 91. 1871. —Guidonia calophylla (Eichler) Kuntze, Revis. Gen. Pl. 1: 44. 1891. Samyda petiolaris Spruce ex Eichler in Mart., Fl. Bras. 13(1): 454. 1871. Casearia negrensis auct. non Eichler 1871: sensu Steyerm., Fieldiana, Bot. 28: 990. 1957. Shrub or small tree 3–5(–15) m tall. Seasonally flooded black- and clear-water riparian forests, 50–200 m; Amazonas (widespread). Apure, Guárico; Amazonian Colombia, Guyana, Peru, Brazil, Bolivia. ŠFig. 403.

Fig. 400. Laetia coriacea

462

F LACOURTIACEAE

Fig. 401. Laetia procera

Fig. 402. Laetia cupulata

Fig. 403. Laetia suaveolens

Lindackeria 463

8. LINDACKERIA C. Presl, Reliq. Haenk. 2: 89, t. 65. 1835. Shrubs or trees. Leaves alternate, persistent or deciduous, entire, undulate, or dentate, pinnately veined, often with fine dark hairs laxly scattered on lower surface; petioles often elongate, apically pulvinate; stipules subulate, usually deciduous. Inflorescences axillary or terminal racemes or panicles, few- to many-flowered, rarely reduced to a single flower. Flowers bisexual, with some staminate flowers often occurring in the same inflorescence, pedicellate; bracts ± deciduous. Sepals 3, imbricate; petals 6–12, imbricate, thinner than the sepals. Stamens 20–40; filaments linear to filiform, free or sometimes joined with sticky resin into a tube; anthers oblonglinear, basifixed, longitudinally dehiscent, often longer than the filaments. Ovary on a short stipe or sessile, smooth, muricate, or echinate, 1-locular; style simple, filiform, shortly 3- or 4-lobed. Fruit capsular, subglobose, often tuberculate or echinate, tardily dehiscent into 3 or 4 valves; pericarp coriaceous to woody. Seeds usually few, ovoid-triangular, with a rather fleshy reddish aril; embryo straight, large; cotyledons large, cordate, foliaceous. Mexico, Central America, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia, tropical Africa; 13 species (7 in Africa, 6 in the New World), 3 in Venezuela, 2 of these in flora area. The third species in Venezuela, Lindackeria laurina C. Presl, occurs in the western and Andean part of the country. It has capsules with spines intermediate in length between the two species below and much longer inflorescences (10–22 cm versus 1–6 cm long) with more numerous flowers. Key to the Species of Lindackeria 1.

1.

Mature capsule densely covered with obtuse conical, furrowed, or rugose tubercles 1–2 mm long, nearly glabrous; capsule 1.5–2(–3) cm diameter including the tubercles; flower buds obovoid-ellipsoid, not glossy; pedicels 3–15 mm long; petals 6, rarely more, 6–8 × 2 mm; stamens 20– 40; filaments glabrous or nearly so; anthers ca. 2 mm long; ovary completely covered with curved prickles to 1 mm long, glandular-pubescent, style not resinous; leaves often covered with tiny glands but usually not resinous and glossy ................................................................... L. paludosa Mature capsule rather sparsely covered with prickles 7–18 mm long, minutely glandular-puberulous, sometimes with scattered stellate hairs, not furrowed or rugose; with prickles, capsule measures (2–)3–6 cm long, (1.5–)2.5–5 cm broad; flower buds usually obovoid to clavate, frequently resinous and glossy; pedicels 15–25 mm long; petals ca. 8, 8– 10 × 5–6 mm; stamens ca. 50; filaments glandular-puberulent; anthers ca. 3 mm long; ovary with curved spines, but tomentose surface of ovary visible between them; leaves with tiny glands, often resinous-glutinous and glossy on both surfaces ........................................................ L. latifolia

Lindackeria latifolia Benth., Hooker’s J. Bot. Kew Gard. Misc. 3: 118. 1851. —Mayna latifolia (Benth.) Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 80. 1861. —Oncoba latifolia (Benth.) Eichler in Mart., Fl. Bras. 13(1): 440, t. 89, fig. l. 1871. —Badunuwa (Bale).

Shrub or small tree. Evergreen lowland forests, disturbed forests, riparian forests, 100– 200 m; Amazonas (Mamurividi on middle Río Pasimoni, Río Casiquiare between Guachapita and El Porvenir, Río Yatúa, Salto Yureba in lower Río Ventuari basin, Solano). Suriname, eastern Amazonian Brazil. ŠFig. 405.

464

F LACOURTIACEAE

Fig. 404. Lindackeria paludosa

Fig. 405. Lindackeria latifolia

Mayna 465

Lindackeria paludosa (Benth.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 404. 1925. —Mayna paludosa Benth., J. Bot. (Hooker) 4: 114. 1842. —Carpotroche paludosa (Benth.) Walp., Repert. Bot. Syst. 1: 203. 1842. —Kasakideimo, Raya, Trompilla. Mayna laxiflora Benth., J. Bot. (Hooker) 4: 114. 1842. Carpotroche laxiflora (Benth.) Walp., Repert. Bot. Syst. 1: 203. 1842. —Oncoba maynensis var. laxiflora (Benth.) Eichler in Mart., Fl. Bras. 13(1): 441. 1871. —Lindackeria maynensis var. laxiflora (Benth.) Mart. ex Pittier et al.,

Cat. Fl. Venez. 2: 166. 1947. Lindackeria latifolia auct. non Benth. 1851: sensu Pittier et al., Cat. Fl. Venez. 2: 166. 1947. Shrub or tree 4–15 m tall, trunk to 20 cm diameter. Flooded and nonflooded, riparian, and seasonally flooded evergreen lowland forests, also in savanna thickets, 100–500 m; widespread in Bolívar and Amazonas. Amazonian Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 404. The sap of Lindackeria paludosa is used as a beverage.

9. LUNANIA Hook., London J. Bot. 3: 317. 1844, nom. cons. Shrubs or trees; branchlets slender and ± zigzag. Leaves alternate, usually distichous, with 3(5) palmate veins and few shorter, upper lateral ones from the midrib, usually laxly pellucid-punctate, margin entire or subdenticulate, petiolate, exstipulate. Inflorescence axillary or subterminal, elongate, slender racemes or basally fewbranched panicles or spike-like racemes. Flowers bisexual, pedicellate, the pedicels articulate at the base, subtending bract and 2 bracteoles minute, deciduous. Calyx subglobose in bud, valvately dehiscing into 2, 3(–5) short, ± reflexed lobes; petals 0; disk hypogynous, cupular, 6–12-dentate, bearing the stamens at the margins and alternating with them. Stamens 6–12; filaments subulate; anthers oblongoid-ovoid, basifixed, 2-thecate, longitudinally dehiscent. Ovary ovoid, constricted and almost stipitate at the base, 1-locular with 3 multiovulate placentas; style thick, very short, with 3 stigmatic branches, each of these with a subcapitate inconspicuously 2-lobed stigma. Capsule 3-valved; pericarp coriaceous. Seeds few to many, small, rounded, usually finely foveolate, shiny, arillate; endosperm present. Mexico, Central America, Antilles (especially Cuba and Hispaniola), Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; 14 species, 1 in Venezuela. Lunania parviflora Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 90. 1861. Shrub or tree 3–15 m tall; leaves elliptic, the apex caudate-acuminate for 2–3 cm, glands present at the junction of midvein and

lateral veins; calyx splitting into 2 or 3 lobes 1.5–2 mm long. Evergreen lowland forests, 100–200 m; Amazonas (Salto Yureba in lower Río Ventuari basin). Táchira; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. ŠFig. 406.

10. MAYNA Aubl., Hist. Pl. Guiane 921, t. 352. 1775. Dendrostylis H. Karst. and Triana in Triana, Nuev. Jen. Esp. 26. 1854 [1855]. Dendrostigma Gleason, Phytologia 1: 29. 1933. Shrubs or small to medium-sized trees. Leaves alternate, pellucid-punctate or not, petiolate; stipules subulate, deciduous; petiole with apical pulvinus and often flexed; leaf margin entire or toothed; venation pinnate, the secondary veins closely approaching the margin of the leaf without forming loops with the adjacent secondary veins. Staminate flowers in few-flowered axillary fascicles or less commonly solitary; pistillate flowers axillary, solitary. Flowers scented, unisexual (plants dioecious). Sepals normally 3, imbricate, often persisting with the fruit; petals 6–8, rarely more, imbricate, exceeding the sepals in size. Stamens 20–50; filaments pubescent, free or

466

F LACOURTIACEAE

Fig. 406. Lunania parviflora

Mayna 467

slightly united at the base; anthers linear, basifixed, longitudinally dehiscent. Ovary with many tiny prickles, often coarsely pubescent, with 3 multiovulate placentas; styles (2)3 or 4(5), distally 2-lobed, the lobes often highly dissected. Fruit a dry berry, globose, covered with bristles, indehiscent or tardily opening at the end. Seeds numerous; testa fleshy, red to orange; endosperm copious; embryo straight; cotyledons cordate. Honduras to Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 6 species, 2 in Venezuela, 1 of these in the flora area. Mayna grandifolia (H. Karst.) Warb. in Engl. & Prantl, Nat. Pflanzenfam. 3(6a): 19. 1893. —Dendrostylis grandifolia H. Karst., Linnaea 28: 432. 1857. —Palo de cachicamo, Yaraiyara. Carpotroche zuliana Pittier, Arb. Arbust. Venez. 2/3 [reprinted from Bol. Comerc. Industr. Venez. no. 34]: 38. 1923. —Mayna zuliana (Pittier) A. Robyns, Ann. Missouri Bot. Gard. 55: 78. 1968. Carpotroche amazonica auct. non Mart. 1871: sensu Pittier et al., Cat. Fl. Venez. 2: 166. 1945.

Shrub or small tree. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro, Tamatama). Zulia; Panama, Colombia, Ecuador, Peru, Brazil. ŠFig. 407. Mayna grandifolia is vegetatively very similar to Carpotroche but often can be distinguished by leaf venation; in Mayna, the secondary veins frequently extend toward the leaf margin and run closer and closer to it, only contacting the other secondary veins through thin tertiary veins, or only after running parallel to the leaf margin for so long that the diameter of the vein has diminished

Fig. 407. Mayna grandifolia

468

FLACOURTIACEAE

to resemble that of one of the tertiary veins. In Carpotroche, the secondary veins often form distinct loops toward the leaf margins. The name Palo de cachicamo derives from the practice of using the bark to poison arma-

dillos. A bundle is made of scraped bark and leaves and then is used to block the entrance of a burrow. The animal is supposedly poisoned as it struggles to get through the bundle.

11. RYANIA Vahl, Eclog. Amer. 1: 51, t. 9. 1796 [1797], nom. cons. Shrubs or slender trees to 15 m tall, with simple and/or stellate hairs on branchlets, leaves, inflorescences, and fruits. Leaves alternate, distichous, epunctate, occasionally somewhat asymmetrical, membranous to coriaceous, pinnately veined, entire to denticulate or serrate, petiolate; stipules 2, acicular to elongate-lanceolate, often deciduous. Inflorescences axillary, 1–4-flowered, fasciculate or on a very short rachis. Flowers bisexual, often showy, white, yellowish-greenish, pink, or red, fragrant. Pedicels articulate at or above base, bracteolate at base; bracts small, deciduous. Sepals usually showy and petaloid, nearly free (usually with a very short calyx tube), quincuncial, spreading or erect, sometimes subpersistent under the fruit; petals absent. Stamens 30–70, free or almost so, inserted in 2 or 3 series at apex of calyx tube near base of sepals; filaments filiform, subequal in length, tapering to a fine point distally; anthers oblong to linear, subsagittate, attached near base, introrse, apex often conspicuously mucronate, opening their full length by 2 longitudinal slits. Disk coroniform or urceolate, villose outside, glabrous within, unequally toothed (to 1 mm). Ovary superior, sessile to stipitate, 1-locular; placentas 3–9, multiovulate in many ranks; style terminal, short to elongate, entire, or at apex 3–9fid (to 3 mm long); stigmas capitellate. Fruit capsular, indehiscent or ultimately valvately dehiscent, globular to pyriform, 1–6 cm long, slightly lobed, suberose, or with spongeous emergences, stellate-tomentose or subhirsute; exocarp to 8 mm thick; endocarp crustaceous or bony, to 6 mm thick. Seeds 25–150 per fruit, globular to bluntly angular, 3–5 mm diameter, hispidulous with scattered stellate, rarely also simple hairs; testa very finely pitted; aril at base of seed membranaceous, brown, resinous-punctate; endosperm copious; embryo straight; cotyledons flat and thin. Nicaragua to Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 8 species, 4 in Venezuela, all in the flora area. Key to the Species of Ryania 1. 1. 2(1). 2. 3(2). 3. 4(3).

Lower surface of leaves markedly canescent throughout .......................... ................................................................................. R. speciosa var. bicolor Lower surface of leaves glabrous to rusty-tomentose (but not canescent) ................................................................................................................ 2 Ovary stipitate; sepals deciduous; disk forming a collar 5 mm high .......... ................................................................................................. R. spruceana Ovary sessile or subsessile; sepals persistent; disk 1–2 mm high ............ 3 Petiole densely rusty-tomentulose; pedicels 3–10(–15) mm long at anthesis .................................................................................................... R. speciosa Petiole grayish-pubescent or glabrous; pedicels 10–27 mm long at anthesis ................................................................................................................ 4 Anthers (4–)5–8 mm long, distinctly mucronate; branchlets and petioles

Ryania 469

4.

glabrous or very sparsely pubescent; leaves narrowly lanceolate to broadly elliptic, narrowed at the base, the upper surface shiny, with prominent reticulation ......................................................... R. angustifolia Anthers 3.8–4.5 mm long, short-mucronate; branchlets and petioles variously pubescent; leaves ovate to ovate-lanceolate or elliptic, rounded or subcordate at the base, the upper surface dull, hirtellous on the midrib, reticulation hardly prominent .................................................... R. dentata

Ryania angustifolia (Turcz.) Monach., Lloydia 12: 21. 1949. —Tetracoryne angustifolia Turcz., Cat. Pl. Herb. Charcov. 1(app.): 14. 1857. Ryania acuminata Spruce ex Eichler in Mart., Fl. Bras. 13(1): 492. 1871. —Patrisia acuminata (Spruce ex Eichler) Kuntze, Revis. Gen. Pl. 1: 106. 1891. Shrub or tree to 8 m tall; flowers yellow. Forest edges, seasonally flooded riparian forests, 50–300 m; Bolívar (Río Parguaza, Río Parhueña, Río Sipapo), Amazonas (Caño San Miguel, Río Casiquiare, Río Cataniapo, San Carlos de Río Negro). Amazonian Colombia, Peru, and Brazil. ŠFig. 409. Ryania dentata (H.B.K.) Miq., Ann. Mag. Nat. Hist. 11: 16. 1843. —Patrisia dentata H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 357. 1821 [1823]. Patrisia affinis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 357. 1821 [1823]. Ryania kunthii Miq., Ann. Mag. Nat. Hist. 11: 16. 1843. Ryania sagotina auct. non Eichler 1871: sensu Pitter et al., Cat. Fl. Venez. 2: 168. 1947. Shrub or tree to 10 m tall. Colombia, Venezuela; 2 varieties, 1 in Venezuela. The second variety, var. toxica Dugand, occurs in eastern Colombia. R. dentata var. dentata Gallery forests, rocky or sandy places, secondary forests, savannas, 50–100 m; Amazonas (Caño Pavón on Río Atabapo, Puerto Ayacucho, Raudal de Atures, Río Cataniapo, mouth of Río Guayapo, Río Sipapo). Apure. ŠFig. 410. Ryania speciosa Vahl, Eclog. Amer. 1: 51, t. 9. 1796 [1797]. Shrub or small tree to 12 m tall. Nicaragua, Costa Rica, Panama, Colombia, Venezu-

ela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 varieties, 6 in Venezuela, 5 of these in the flora area. Key to the Varieties of R. speciosa 1. Leaves finely white- or grayish-canescent all over the lower surface, strongly bicolorous .......................... var. bicolor 1. Leaves glabrous to rusty-tomentose on the lower surface, not strongly bicolorous ............................................................... 2 2. Branchlets and lower surface of leaves softly tomentulose or tomentose ............ ..................................... var. tomentosa 2. Branchlets and lower surface of leaves glabrous or rusty-pubescent, but not softly toment(ul)ose ........................................ 3 3. Stipules persistent ........ var. stipularis 3. Stipules deciduous .................................. 4 4. Anthers 2.5–5 mm long; sepals 12–18 mm long ..................................... var. minor 4. Anthers 5–7 mm long; sepals 18–30(–35) mm long ..................... var. subuliflora R. speciosa var. bicolor (A. DC.) Monach., Lloydia 12: 19. 1949. —Patrisia bicolor A. DC. in DC., Prodr. 1: 256. 1825. —Ryania bicolor (A. DC.) Eichler in Mart., Fl. Bras. 13(1): 491, in obs. 1871. Ryania candollei Miq., Ann. Mag. Nat. Hist. 11: 16. 1843. Shrub 4–7 m tall. Gallery forests, flooded forests, 50–400 m; Bolívar (Caño Villacoa on Caicara to El Burro road, Río Parguaza, Serranía San Borja along lower Río Suapure, slopes of Sierra de Maigualida), Amazonas (Río Ugueto). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. R. speciosa var. minor Monach., Lloydia 12: 15. 1949. Shrub 1–4 m tall. Deciduous to evergreen

470

FLACOURTIACEAE

lowland forests, 50–800 m; Bolívar (22 km east of Túriba), Amazonas (Culebra, Río Mawarinuma, slopes of Sierra de la Neblina). Amazonian Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. R. speciosa var. stipularis (Linden & Planch.) Monach., Lloydia 12: 17. 1949. —Ryania stipularis Linden & Planch., Pl. Columb. 1: 22. 1863 [1874-1875]. —Fruto de burro. Shrub to small tree. Evergreen lowland to lower montane forests, 200–600 m; Bolívar (Cerro Camarón, Cerro Guaiquinima, upper Río Caura). Anzoátegui, Apure, Aragua, Carabobo, Cojedes, Distrito Federal, Mérida, Miranda, Monagas, Sucre, Yaracuy. R. speciosa var. subuliflora (Sandwith) Monach., Lloydia 12: 14. 1949. —Ryania pyrifera var. subuliflora Sandwith, J. Arnold Arb. 24: 219. 1943. —Borrachero, Sina-widiqui (Yekwana). Ryania pyrifera auct. non (Rich.) Uitten & Sleumer 1935: sensu J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4.1): 35. 1941, pro parte. Shrub 1–5 m tall. Savannas, evergreen lowland to lower montane forests, 50–1000

m; Bolívar (Caicara to El Burro road, Macizo del Chimantá [slopes of Chimantá-tepui], Río Villacoa), Amazonas (Capibara, slopes of Cerro Aracamuni and Cerro Huachamacari, Culebra, between Puerto Ayachucho and Samariapo, Río Cunucunuma, Río Pasimoni). Apure; Colombia, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil. R. speciosa var. tomentosa (Miq.) Monach., Lloydia 12: 16. 1949. —Ryania tomentosa Miq., Ann. Mag. Nat. Hist. 11: 15. 1843. —Patrisia tomentosa (Miq.) M. Roem., Fam. Nat. Syn. Monogr. 2: 136. 1846. —Ryania pyrifera var. tomentosa (Miq.) Sleumer in Pulle, Fl. Suriname 3: 286. 1935. Patrisia parviflora A. DC. in DC., Prodr. 1: 256. 1824. —Ryania parviflora (A. DC.) Griseb., Fl. Brit. W. I. 296. 1860. Ryania casiquiarensis Steyerm., Fieldiana, Bot. 28: 411. 1952. Ryania pyrifera auct. non (Rich.) Uittien & Sleumer 1935: sensu Ll. Williams, Field Mus. Nat. Hist., Bot. Ser. 15: 362. 1936. Ryania mansoana auct. non Eichler 1871: sensu Pittier et al., Cat. Fl. Venez. 2: 168. 1947.

Fig. 408. Ryania spruceana

Ryania 471

Fig. 409. Ryania angustifolia

Fig. 410. Ryania dentata var. dentata

472

FLACOURTIACEAE

Shrub 2–6 m tall. Evergreen lowland forests, 100–500 m; Bolívar (Las Claritas, Río Icabarú, Amazonas (Capibara). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Ryania spruceana Monach., Lloydia 12: 24. 1949. —Patrisia spruceana (Monach.) Lemée, Fl. Guyane Fr. 3: 68. 1953. —Bejuco de rana, Hoja salado.

Ryania stipularis auct. non Linden & Planch. 1863: sensu Pittier et al., Cat. Fl. Venez. 2: 168. 1947. Shrub or small tree 3–12 m tall. Riparian forests, lower montane forests, white-sand scrub (bana), 50–500 m; Amazonas (upper Río Cataniapo, upper Río Cuao, Río Guainía, Río Guayapo, Río Sipapo, San Carlos de Río Negro, San Fernando de Atabapo, San Pedro). Colombia, Peru, Brazil. ŠFig. 408.

12. XYLOSMA G. Forst., Fl. Ins. Austr. 72. 1786. Shrubs or trees; trunk, branches, and branchlets often armed with simple or branched axillary spines. Leaves alternate, persistent or rarely (semi)deciduous, pinnately veined, usually glandular-crenate or -serrate, rarely entire, petiolate, exstipulate. Flowers generally unisexual (plants dioecious or rarely andromonoecious), very rarely bisexual or seemingly so (the stamens then ± abortive), small, in short racemes often reduced to fascicles, very rarely solitary in foliate or defoliate leaf axils, rarely precocious from shoots of the past year; pedicels articulate at base or to the middle; bracts small. Calyx lobes or sepals 4 or 5, less often 6, very rarely 7, slightly united at base, imbricate, subpersistent; petals absent. Disk present, extrastaminal, consisting of few to several fleshy lobes or glands, these sometimes (mainly in the pistillate flowers) ± connate to an annulus. Staminate flowers with stamens 8 or more; filaments filiform, usually exceeding the sepals in length; anthers small, versatile, elliptic-globose, basifixed, 2-celled, longitudinally dehiscent; rudiment of ovary lacking. Pistillate flowers with stamens rarely present, generally few and in the form of staminodes without pollen; ovary superior, 1-celled, with 2 or 3(–6) few-ovulate placentas; style (sometimes short or practically 0) simple or short-branched distally, bearing 2(3 or 4) dilated stigmas, the latter sometimes flattened and very shortly lobed. Fruit a berry; pericarp thin-coriaceous, rather dry, rarely somewhat fleshy. Seeds generally few; testa smooth; endosperm copious; embryo large, with broad cotyledons. Central America, West Indies, South America, southeastern Asia and the Pacific; ca. 95 species (about half of these neotropical), 7 or 8 in Venezuela, 1 of these in the flora area. Xylosma tessmannii Sleumer was reported from Amazonas state in Sleumer (Flacourtiaceae. Fl. Neotrop. Monogr. 22: 181. 1980), based on Ernst 490 (BM), but this is apparently an error because Ernst did not collect in the flora area. A second collection, Ll. Williams 14668 (MO), from San Carlos de Río Negro, is sterile but might be Xylosma tessmannii. Liesner 24313 (MO), is a fruiting collection from San Ignacio de Yuruaní, Bolívar state, at 850 m elevation. The stems and foliage are densely erect-pubescent, which is uncharacteristic of Xylosma benthamii. Additional flowering material is needed to determine to what species this belongs. Xylosma benthamii (Tul.) Triana & Planch., Ann. Sci. Nat. Bot. sér. 6, 17: 1862. —Flacourtia benthamii Tul., Ann. Sci. Nat. Bot. sér. 3, 7: 291. 1847. —Hisingera benthamii (Tul.) Clos, Ann. Sci. Nat. Bot. sér. 6, 8: 225. 1857. —Myroxylon benthamii

(Tul.) Kuntze, Revis. Gen. Pl. 1: 44. 1891. Flacourtia nitida Benth., Hooker’s J. Bot. Kew Gard. Misc. 3: 119. 1851. Xylosma pallidifolium Sleumer ex Steyerm. and O. Huber, Fl. Avila 425. 1978, as synonym.

Zuelania 473

Fig. 411. Xylosma benthamii

Spiny-stemmed shrub or tree 2–8 m tall. Savanna edges, semideciduous forests, thorn scrub, 50–300 m; Bolívar (between Guasipati and El Mantecal, La Paragua). Apure, Dis-

trito Federal, Falcón, Guárico, Lara, Miranda, Monagas, Portuguesa; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 411.

13. ZUELANIA A. Rich. in Sagra, Hist. Phys. Cuba, Bot. Pl. Vasc. 10: 88. 1845. Shrubs or trees, often with sticky transparent sap. Leaves periodically deciduous, alternate, entire to subcrenate-serrate, pellucid-punctate, pubescent on the lower surface, pinnately veined, petiolate, with deciduous stipules. Flowers bisexual, often emerging before the leaves, in sparsely distributed fascicles from leafless axils or at branch tips, (sub)pedicellate, with small bracts at base. Sepals 4 or 5, barely connate at the base, imbricate, persistent; petals absent. Stamens 20–40, subperigynous. Disk appendages or staminodes 15–20, clavate, ± alternating with the stamens and united basally with them; filaments filiform; anthers small, almost basifixed, introrse, longitudinally dehiscent. Ovary free, 1-locular, with 3–many-ovulate placentas; stigma (sub)sessile, thick, peltate, the pistil having the appearance of a tiny urn. Fruit a fleshy capsule, globose, yellow-green to dark green, 3-sulcate, eventually dehiscing by 3 valves. Seeds numerous, arillate; endosperm fleshy. Mexico, Central America, West Indies, northern South America; 1 species. Zuelania guidonia (Sw.) Britton & Millsp., Bahama Fl. 285. 1920. —Laetia guidonia Sw., Prodr. 83. 1788. —Casearia guidonia (Sw.) Lundell, Wrightia 5: 41. 1974, non Benth. 1861. —Cabo de vela, Lechozo, Palo de camarón, Vela de barco, Yara yara. Deciduous shrub or tree 6–35 m tall. De-

ciduous to semideciduous forests, 100–300 m; Bolívar (Cerro Curichapo 25 km northeast of La Paragua, 15 km southeast of El Callao, Cerro San Mateo near Arekuna, 27 km south of El Dorado, La Yagua on San Félix to Upata road, 40 km south of Tumeremo), Amazonas (San Carlos de Río Negro). Zulia; other distribution as in the genus. ŠFig. 412.

474

FLACOURTIACEAE

Fig. 412. Zuelania guidonia

GENTIANACEAE by Lena Struwe, Paul J. M. Maas, Oliver Pihlar, and Victor A. Albert Trees, shrubs, vines, and perennial or annual herbs (saprophytic and without chlorophyll in Cotylanthera, Voyria, and Voyriella, rarely epiphytic, as in some Macrocarpaea and Voyria), generally glabrous, without latex but sometimes with resin at nodes (Potalia and Tachia). Stems and branches terete or often quadrangular and winged. Leaves opposite (rarely alternate or verticillate), simple, sometimes reduced to scales (Cotylanthera, Voyria, and Voyriella) or saccate (Saccifolium), sessile or petiolate, the blade often decurrent; exstipulate, with an interpetiolar line or ocrea, usually with colleters (small finger-shaped secretory glands) on the adaxial surfaces of

GENTIANACEAE 475

the leaf and the calyx bases; margins entire, rarely dentate, flat or recurved; venation brochidodromous (secondary veins terminating in a series of prominent arches forming a distinct intramarginal vein) or camptodromous (secondary veins curved upward and gradually diminishing distally within the margin, without forming conspicuous marginal loops), usually either pinnately veined with several pairs of straight secondary veins or with few, arcuate secondary veins diverging from the main vein close to the base of the blade. Inflorescences terminal or axillary, usually cymose, less often racemose, capitate, clustered, spicate, or flowers solitary; bracts and floral bracts leaf- or scale-like. Flowers bisexual, hypogynous, 4- or 5-merous, or rarely 6–16-merous. Calyx 4- or 5-merous (rarely lobes 2 or 6–8), connate at the base or rarely with free lobes, imbricate (decussate in Anthocleista, Chorisepalum, and Potalia), each lobe often with a dorsal wing, keel, or glandular area that can be decurrent on the calyx tube; corolla sympetalous, actinomorphic or rarely slightly zygomorphic, tubular, campanulate, salver- or funnelform, sometimes with nectaries on the inside, the lobes contorted in bud (basally valvate in Aripuana, imbricate in Bartonia, Obolaria, and Saccifolium). Stamens as many as the corolla lobes (reduced in number, e.g., Hoppea), inserted in the corolla tube or in the sinuses of the corolla lobes, actinomorphic or zygomorphic (e.g., Chelonanthus and Symbolanthus); filaments free (fused in Anthocleista and Potalia); anthers free or rarely connate, oblong, elliptic, to linear, often sagittate, dorsi- or basifixed, tetrasporangiate, sometimes with sterile apical appendages, longitudinally dehiscent (porate in Cotylanthera and some Exacum), usually introrse, after anthesis erect, curved, recurved, or helically twisted. Gynoecium syncarpous, bicarpellate, sessile or stipitate; ovary unilocular and/or bilocular, often with a glandular disk or glands at the base, with axile and/or parietal placentation, often with deeply protruding and curved placentas; ovules few to many; style one, terminal; stigmas capitate, bilamellate, peltate, or simple. Fruit usually dry, sometimes fleshy or leathery (Anthocleista, Chironia species, Fagraea, Potalia, and Symbolanthus), thin, woody, or fibrous, medially or apically dehiscent, seldom indehiscent (most species of the fleshy-fruited genera and Voyria species). Seeds usually small, rounded or angular, winged or not, nonarillate, the testa cells of variable morphology, often reticulate. Cosmopolitan except the Antarctic continent; ca. 86 genera and ca. 1550 species, 23 genera and 83 species in the flora area. Literature Cited in Gentianaceae Treatment Grothe, E., and P. J. M. Maas. 1984. A scanning electron microscopy study of the seed coat structure of Curtia Chamisso and Schlechtendahl and Hockinia Gardner (Gentianaceae). Proceedings Koninklijke Nederlandse Akademie van Wetenschappen. Series C. 87: 33–42. Kuntze, C. E. O. 1891. Revisio Generum Plantarum par. 2. Leipzig. Leeuwenberg, A. J. M., and P. W. Leenhouts. 1980. Taxonomy. Pages 8–96 in: Engler and Prantl’s Die Natürlichen Pflanzenfamilien, Angiospermae: Ordnung Gentianales, Fam. Loganiaceae. Vol. 28b (1). Leeuwenberg, A. J. M., ed. Duncker and Humblot, Berlin. Maas, P. J. M. 1985. Nomenclatural notes on neotropical Lisyantheae (Gentianaceae). Proceedings Koninklijke Nederlandse Akademie van Wetenschappen. Series C. 88: 405–412.

476

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Machado, I. C. S., I. Sazima, and M. Sazima. 1998. Bat pollination of the terrestrial herb Irlbachia alata (Gentianaceae) in northeastern Brazil. Plant Systematics and Evolution 209: 231–237. Maguire, B. 1981. Gentianaceae. In: The Botany of the Guayana Highland - Part XI. Maguire, B., and collaborators, eds. Memoirs of the New York Botanical Garden 32: 330–388. Maguire, B., and B. M. Boom. 1989. Gentianaceae (part 3). In: The Botany of the Guayana Highland - Part XIII. Maguire, B., and collaborators, eds. Memoirs of the New York Botanical Garden 51: 2–56. Pringle, J. S. 1995. Gentianaceae. Pages 1–131 in: Flora of Ecuador. Vol. 53. G. Harling and L. Andersson, eds. Department of Systematic Botany, Göteborg University, Göteborg. Struwe, L., and V. A. Albert. 1998. Lisianthius (Gentianaceae), its probable homonym Lisyanthus, and the priority of Helia over Irlbachia as its substitute. Harvard Papers in Botany 3: 63–71. Struwe, L., V. A. Albert, and B. Bremer. 1994 [1995]. Cladistics and family level classification of the Gentianales. Cladistics 10: 175–206. Struwe, L., P. J. M. Maas, and V. A. Albert. 1997. Aripuana cullmaniorum, a new genus and species of Gentianaceae from white-sands of southeastern Amazonas, Brazil. Harvard Papers of Botany 2: 235–253. Struwe, L., M. Thiv, J. W. Kadereit, A. S-R. Pepper, T. J. Motley, P. J. White, J. H. E. Rova, K. Potgieter, and V. A. Albert. 1998. Saccifolium (Saccifoliaceae), an endemic of Sierra de la Neblina on the Brazilian-Venezuelan frontier, is related to a temperate-alpine lineage of Gentianaceae. Harvard Papers in Botany 3: 199–214. The monotypic family Saccifoliaceae as well as the tribe Potalieae (comprised of Anthocleista, Fagraea, and Potalia, formerly in Loganiaceae) are here treated as members of Gentianaceae following recent phylogenetic findings (Struwe et al. 1994 [1995]; Struwe et al. 1997; Struwe et al. 1998; also compare Leeuwenberg and Leenhouts 1980). Gentians of the Venezuelan Guayana are distributed among several evolutionary lineages in the family, the largest group of genera belonging to the Irlbachia complex (Adenolisianthus, Chelonanthus, Irlbachia, Macrocarpaea, Rogersonanthus, Symbolanthus, Tachia, Tetrapollinia, Wurdackanthus, and possibly additional genera in the flora area [Struwe et al. 1997, Struwe et al. 1998]). Gentians (e.g., Chelonanthus, Gentiana, and Potalia) have pharmacological uses worldwide in traditional medicine, especially against inflammations, fevers, and fungal diseases. Clinical studies have shown that their seco-iridoids and xanthones have stimulating as well as depressing effects on the central nervous system and as well as other inhibiting features. Key to the Genera of Gentianaceae 1. 1. 2(1).

Plants without chlorophyll; leaves scale-like, yellow, pink, or purple ..... 2 Plants with chlorophyll; leaves not scale-like, green ................................ 3 Calyx distinctly connate basally, tubular to campanulate; corollas much longer than the calyx, persistent in fruit; plants often pink, yellow, bluish, or purplish, sometimes white ............................................... 21. Voyria

GENTIANACEAE

2.

477

Calyx with almost free lobes, rotate; corollas barely longer than the calyx, deciduous in fruit; plants always white .................................. 22. Voyriella 3(1). Leaves saccate, alternate, very tightly crowded at branch apices; bark with corky ridges ................................................................ 14. Saccifolium 3. Leaves flat, not saccate, opposite or verticillate, not very tightly crowded at branch apices; bark not prominently developed, or smooth without corky ridges ............................................................................................ 4 4(3). Flowers axillary (a terminal flower is present in Neurotheca), sessile, solitary or in clusters in the leaf axils ........................................................ 5 4. Flowers terminal, pedicellate or subsessile (sessile in Tapeinostemon sessiliforum), solitary or in cymes, spikes, capitula, or racemes, always terminal (sometimes with additional axillary cymose inflorescences) ..... 7 5(4). Plants woody, > 50 cm tall; corollas > 14 mm long; leaves > (6–)10 cm long; calyx woody or coriaceous ........................................................... 18. Tachia 5. Plants herbaceous, < 50 cm tall; corollas 4–8 mm long; leaves < 6(–10) cm long; calyx thin and membranaceous .................................................... 6 6(5). Leaves lanceolate, > 15 mm long; flowers in clusters of 2–8 in the leaf axils, terminal flower absent; corollas white; robust perennial herbs ............................................................................................... 7. Enicostema 6. Leaves linear, < 10 mm long; flowers 1 or 2 in each of the upper leaf axils, terminal flower present; corollas blue (rarely white); small annual herbs ..................................................................................... 11. Neurotheca 7(4). Corolla lobes 6–10; sepals free, decussate, 4 (2 outer and 2 inner) .......... 8 7. Corolla lobes 4 or 5; sepals united at least at the base (1/10–9/10 of the length), imbricate, 5 (4 in Celiantha chimantensis, Coutoubea, and Schultesia) .............................................................................................. 9 8(7). Fruit a dry capsule, dehiscing from the base in 4 valves; corolla lobes 6; corolla bud apex tapering; inflorescences 1–9-flowered; leaves < 15 cm long .................................................................................... 4. Chorisepalum 8. Fruit a fleshy berry, indehiscent; corolla lobes 8–10; corolla bud apex rounded; inflorescences 9–40-flowered; leaves > 20 cm long ... 12. Potalia 9(7). Stigmas capitate, sometimes slightly bilobed, but lobes never broad and flattened ............................................................................................... 10 9. Stigmas bilamellate, with flattened or linear lobes ................................ 11 10(9). Mature leaves > 20 mm long; suffrutescent herbs or shrubs (annual herbs in Tapeinostemon sessiliflorum, then with a capitate inflorescence) ........................................................................................ 19. Tapeinostemon 10. Mature leaves 1–20 mm long; small annual herbs ...........................9. Curtia 11(9). Inflorescence spicate or racemose, flowers sessile to shortly pedicellate (pedicel < 4 mm long); stamens inserted close to the corolla lobe sinuses ................................................................................................. 5. Coutoubea 11. Inflorescence cymose or flowers solitary, flowers distinctly pedicellate; stamens inserted in the middle or lower in the corolla tube .............. 12 12(11). Calyx with a prominent dorsal keel or wing, without glandular areas; ovaries without a basal disk or glandular areas; anthers erect after anthesis ........................................................................................... 15. Schultesia 12. Calyx with dorsal glandular areas, with keels or low ridges but not with broad distinct wings; ovaries with a basal glandular disk or area (ab-

478

G ENTIANACEAE

13(12).

13.

14(13). 14.

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

17.

18(17).

18.

19(15). 19. 20(19). 20. 21(20). 21. 22(21). 22.

23(21).

sent in Irlbachia pratensis, Rogersonanthus coccineus, and Tetrapollinia); anthers recurved after anthesis (rarely erect) ................... 13 Corolla tubes with a corona inside at the insertion point of the stamens; calyx divided ca. 7/8–9/10 of the length, (12–)20–45 mm long ................... ......................................................................................... 17. Symbolanthus Corollas without a corona (present in Wurdackanthus, then with leaves silvery below); calyx divided 1/4–3/4 of the length, 3–12(–20) mm long .............................................................................................................. 14 Corollas > 8 cm long; leaves very thick and coriaceous, broadly obovate ........................................................................................... 16. Sipapoantha Corollas < 8 cm long; leaves slightly coriaceous to chartaceous, linear, elliptic, ovate to lanceolate (obovate in Adenolisianthus and leaves then not thick and coriaceous) ..................................................................... 15 Calyx lobes triangular to lanceolate, acute to acutish; corolla lobes triangular; corolla bud apex tapering ......................................................... 16 Calyx lobes ovate to oblong, obtuse; corolla lobes ovate to elliptic; corolla bud apex rounded ................................................................................ 19 Flowers solitary, appearing axillary (but are not) ................ 10. Neblinantha Flowers in 2–many-flowered, terminal inflorescences (flowers rarely solitary and then with blue-purple or white corollas) .............................. 17 Leaves ovate to broadly lanceolate, > 3 mm wide; subligneous perennial herbs or subshrubs; corolla tubes conspicuously constricted below the insertion point of the stamens ................................................. 2. Celiantha Leaves linear to narrowly lanceolate, < 3 mm wide; annual herbs; corolla tubes not conspicuously constricted below the level of the insertion point of the stamens ............................................................................ 18 Leaves in > 6 pairs, most of them positioned at the base of the stem; corollas (18–)20–36 mm long; styles 15–19 mm long; pollen in polyads ................................................................................. 8. Irlbachia (pratensis) Leaves as many as 6 pairs, not most abundant close to the base of the stem; corollas 6–20(–25) mm long; styles 1–12 mm long; pollen in spinose tetrads ....................................................................... 20. Tetrapollinia Shrubs or trees, or subligneous perennials (distinctly woody at base), to 5 m tall ................................................................................................. 20 Annuals or short-lived perennial herbs, sometimes slightly woody at base, < 1(–2.5) m tall ..................................................................................... 24 Lower surface of leaves silvery; corollas red or pink ...... 23. Wurdackanthus Lower surface of leaves not silvery; corollas white, green, or yellow .... 21 Inflorescences with monochasial branches ............................................. 22 Inflorescences with dichotomous branching or a 1–3-flowered cyme .... 23 Leaves widest above middle, obovate; internodes between pedicels in inflorescence ≤ 10 mm long ...............................................1. Adenolisianthus Leaves widest below middle or at middle, linear, lanceolate, ovate, to elliptic; internodes between pedicels in inflorescence usually > 20 mm long .................................................................................... 3. Chelonanthus Bracts leaf-like; leaves chartaceous or slightly coriaceous, the lower surface with visible (but not always prominent) secondary veins; pollen in

Adenolisianthus 479

monads; inflorescences usually with > 3 flowers ............. 9. Macrocarpaea Bracts not leaf-like; leaves thick and coriaceous, the lower surface with invisible secondary veins; pollen in tetrads; inflorescences 1–3-flowered ............................................................................... 13. Rogersonanthus 24(19). Corollas red to orange; perennial herbs with vegetative shoots from the base of the plants ......................................................... 13. Rogersonanthus 24. Corollas white, yellow, green, brownish, pink, lavender, purple, or blue; annual herbs, sometimes very large, without vegetative shoots from the base of the plants ........................................................................... 25 25(24). Corolla tubes > 2 times as long as corolla lobes; corolla lobe apex obtuse; pollen in tetrads with smooth or reticulate exine, often also with globules, or in polyads with reticulate exine with loop-like structures, exine never spinose..................................................................... 3. Chelonanthus 25. Corolla tubes < 2 times as long as corolla lobes (except I. tatei, then with pink flowers); corolla lobe apex acutish; pollen in polyads with perforate exine with spines or globules ............................................ 8. Irlbachia 23.

1. ADENOLISIANTHUS Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. by Lena Struwe Branched shrubs or subshrubs. Branches terete to quadrangular, with 4 low decurrent ridges. Leaves sessile, broadly ovate to obovate, widest at or above the middle, coriaceous, thin and chartaceous when dry, the base attenuate and slightly auriculate, the apex acute to acuminate; with a low interpetiolar ocrea; margins not recurved; venation with 2 prominent, basally divergent and arcuate secondary veins. Inflorescence terminal, long-stalked, dichotomously branched into 1 or 2 (rarely 3) monochasial branches with flowers usually arranged along one side only, (5–)15–43-flowered, the internodes between the pedicels usually < 10 mm long; bracts and floral bracts scale-like. Flowers pedicellate, slightly zygomorphic, 5merous, erect or slightly nodding. Calyx campanulate, divided 1/2–2/3 of the length, thick and leathery, persistent in fruit (sometimes with spreading lobes and rupturing), the lobes elliptic, with a thickened dorsal ridge, hyaline-margined, the apex obtuse; corolla broadly funnelform with a basal narrow tube, pale green to greenish yellow, rather thick and leathery, deciduous in fruit but sometimes remaining at the apex of the fruit, the lobes ovate, obtuse, spreading, the corolla bud apex rounded. Stamens inserted close to the base in the corolla tube; filaments of unequal length, sometimes sharply bent at apex; anthers oblong, sagittate, slightly recurved after anthesis, versatile, with a small sterile apical appendage; pollen in tetrads, the exine reticulate. Ovary with a glandular disk; style long, slender, persistent in fruit; stigma bilamellate. Capsules ellipsoid to globose, dehiscing medially, erect or nodding. Seeds angular, not winged, the testa with dome-like and concave cells with thickened bands. Endemic to the Guayana Shield on white-sand savannas in southeastern Colombia and southern Venezuela; 1 species. This monotypic genus is part of the Irlbachia complex (Struwe et al. 1997; Struwe and Albert 1998). Adenolisianthus was included in Chelonanthus by Maguire and Boom (1989) and in Irlbachia by Maas (1985).

480

G ENTIANACEAE

Fig. 413. Adenolisianthus arboreus

Celiantha 481

Adenolisianthus arboreus (Spruce ex Progel) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. —Lisianthus arboreus Spruce ex Progel in Mart., Fl. Bras. 6(1): 240, fig. 64. 1865. —Helia arborea (Spruce ex Progel) Kuntze, Revis. Gen. Pl. 2: 428. 1891. —Irlbachia alata subsp. arborea (Spruce ex Progel) J.G.M. Pers. & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 410. 1985. —Chelonanthus fruticosus Ma-

guire & B.M. Boom, Mem. New York Bot. Gard. 51: 42. 1989. Shrub or subshrub to 3 m tall; leaves 30– 110 × 16–60 mm; calyx 5–7 mm long; corollas 12–17 mm long; capsules 8–9 mm long. White-sand savannas, 100–200 m; Amazonas (Maroa, Río Sipapo, San Fernando de Atabapo and vicinity, road between Yavita and Pimichín). Colombia (Amazonas: Araracuara, Vaupés: Cerro Chiribiquete, Mitú). ŠFig. 413.

2. CELIANTHA Maguire, Mem. New York Bot. Gard. 32: 382. 1981. by Lena Struwe Erect herbs, sometimes suffrutescent. Stems terete to slightly quadrangular, with or without decurrent wings. Leaves sessile to petiolate, elliptic, obovate, lanceolate, or ovate, (sub-)coriaceous, the base attenuate to obtuse, the apex acute to acuminate or obtuse; interpetiolar line prominent; petiole 0–22 mm long; margins recurved or not; venation with 1 pair of very prominent, basally divergent and arcuate lateral veins. Inflorescence to 60-flowered, a terminal diffuse panicle or irregular dichasium with side branches in the upper leaf axils (C. bella) or a few-flowered terminal cyme (C. chimantensis and C. imthurniana); bracts leaf-like, ovate, 6–30 mm long; floral bracts leaf-like or scale-like. Flowers pedicellate, actinomorphic, 4- (C. chimantensis) or 5-merous (C. bella and C. imthurniana), nodding. Calyx campanulate, divided ca. 1/3–1/2 of the length, thick, persistent in fruit, the lobes elliptic (C. bella) or triangular (C. chimantensis and C. imthurniana), abruptly acute to acuminate at apex, with a low dorsal keel, erect, usually spreading in fruit; corolla narrowly funnelform, purple, magenta, lilac, or pink (C. bella and C. imthurniana), or yellow (C. chimantensis), thin, persistent in fruit, the basal part of tube constricted, very narrow, 1–2 mm wide, abruptly widening at the insertion point of the stamens, the lobes ovate, lanceolate, triangular, acuminate, the corolla bud apex tapering. Stamens inserted in the lower half of the corolla tube; filaments of slightly unequal length; anthers linear to oblong, sagittate, recurved after anthesis; pollen in polyads, the exine with small globules. Ovary without a disk at the base of the ovary (at least in C. bella); style long, slender, deciduous or persistent in fruit; stigma bilamellate with ovate lobes. Capsule elliptic, medially and apically dehiscent, nodding. Seeds irregular to rounded, sometimes flattened, not winged, the testa cells dome-like or concave (C. imthurniana). Endemic to the Guayana Shield on tepui summits in southern Venezuela, Guyana, and northern Brazil; 3 species, all in the flora area. Key to the Species of Celiantha 1. 1. 2(1).

Flowers 4-merous; corolla yellow .......................................... C. chimantensis Flowers 5-merous; corolla pink, magenta, lilac, purple, or red, often with white stripes or spots on the inside ....................................................... 2 Lower leaves lanceolate, 55–170 mm long; calyx lobes broadly ovate and abruptly acute, ca. 1 mm long, < 1/4 of the length of the calyx tube, calyx lobe < 3 mm long; leaf apex acute to acuminate ............................. C. bella

482

G ENTIANACEAE

Fig. 414. Celiantha bella

Fig. 415. Celiantha imthurniana

Fig. 416. Celiantha chimantensis

Chelonanthus 483

2.

Lower leaves broadly elliptic, 20–40 mm long; calyx lobes triangular, narrowly acuminate, 3–4 mm long, ± equaling the length of the calyx tube; leaf apex obtuse to bluntly acute ........................................ C. imthurniana

Celiantha bella Maguire & Steyerm., Mem. New York Bot. Gard. 32: 383, figs. 88, 89. 1981. Suffrutescent, sparsely branched herb to 3 m tall, sometimes purplish; calyx 3–4 mm long, purple to dark green; corolla 30–65 mm long, magenta, lilac, to pink, with white lines inside, the lobes 8–22 mm long; capsule 10– 14 mm long. Tepui meadows and low forests on tepui summits, 1500–2200 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). ŠFig. 414. Celiantha chimantensis (Steyerm. & Maguire) Maguire, Mem. New York Bot. Gard. 32: 387. 1981. —Lisianthus chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17: 461. 1967. Branched herb to 1.5 m tall; leaves 8–90 × 5–38 mm, the base attenuate to nearly obtuse; leaf margins flat or slightly recurved; calyx 5–6 mm long, the lobes ca. 3 mm long; corolla 36–51 mm long, yellow, the lobes 13– 18 mm long; capsule ca. 6 mm long. Open

rocky areas on tepui summits and escarpments, 1800–2200 m; Bolívar (Angasimatepui, Macizo del Chimantá [Acopán-tepui]). Endemic. ŠFig. 416. Celiantha imthurniana (Oliv.) Maguire, Mem. New York Bot. Gard. 32: 387. 1981. —Lisianthus imthurnianus Oliv., Trans. Linn. Soc. London, Bot. 2: 279. 1887. —Calolisianthus imthurnianus (Oliv.) Gleason, Bull. Torrey Bot. Club 56: 402. 1929. Suffrutescent, branched herb to 1 m tall; leaves sessile, 8–38 × 9–14 mm, the base attenuate; leaf margins slightly recurved; calyx 5–6 mm long, the lobes ca. 3 mm long; corolla 33–50 mm long, purple, red, pink, or lavender, with white spots inside, the lobes 12–22 mm long; capsule 11–13 mm long. Montane forests and open areas on tepui summits and upper tepui slopes, 2000–2700 m; Bolívar (Aprada-tepui, Auyán-tepui, Macizo del Chimantá, Ilú-tepui, Kukenán-tepui, Roraima-tepui). Also on Mount Roraima in Guyana and Brazil. ŠFig. 415.

3. CHELONANTHUS Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. —Lisyanthus sect. Chelonanthus Griseb., Gen. Sp. Gent. 180. 1839 [1838]. Lisyanthus Aubl., Hist. Pl. Guiane 201. 1775., non Lisianthius P. Browne 1756. by Lena Struwe Annual or perennial herbs, sometimes woody at the base, to 3.5 m tall. Stems terete or quadrangular, sometimes winged. Leaves sessile or petiolate, ovate, lanceolate, or linear, chartaceous to seldom subcoriaceous, the base obtuse to acute, the apex obtuse, acute, to acuminate; with interpetiolar line; margins flat; venation with few, basally divergent and arcuate secondary veins. Inflorescence terminal, cymose, usually many-flowered with monochasial branches or reduced to a few flowers; bracts and floral bracts scale-like. Flowers pedicellate to subsessile, actinomorphic to slightly zygomorphic (with a slightly bent corolla tube with anthers clustered in upper or lower part of corolla mouth and style bent toward the lower side of the corolla mouth), 5-merous, horizontal or often nodding. Calyx campanulate, divided at least 1/2 of its length, thick and coriaceous, persistent in fruit, the lobes elliptic with a dorsal glandular ridge, hyaline-margined, the lobe apex obtuse; corolla campanulate to funnelform, blue, purple, green, or white, the tubes often bent, thin or thick and fleshy, deciduous in fruit but dry remnants often attached to apices of developing fruits, the lobes ovate to elliptic, the lobe apex obtuse, the corolla bud apex rounded. Stamens inserted close to the base of the corolla tube, often slightly exserted from

484

G ENTIANACEAE

the corolla mouth; filaments of subequal to unequal length, sometimes strongly bent close to apex; anthers oblong to linear, sagittate, erect or recurved (to 360°) after anthesis, with sterile apical appendages; pollen in tetrads or polyads (C. purpurascens), the exine unevenly reticulate, often with additional loops and globose structures (C. purpurascens). Ovary with a glandular disk at base of ovary; style long, slender, persistent in fruit; stigma bilamellate, the lobes ovate-elliptic. Capsule elliptic, dehiscing medially, often nodding. Seeds angular, not winged, the testa with dome-like and concave cells with band-like thickenings. Mexico, Belize, Honduras, Nicaragua, Guatemala, Costa Rica, Panama, Grenada, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, with the highest species diversity occurring on the Guayana Shield; ca. 8 species, 6 in Venezuela, all of these in the flora area. Chelonanthus has been included in Helia and Irlbachia and is a part of the Irlbachia complex (Struwe and Albert 1998; compare to Maas 1985). Bat pollination has been reported in Chelonanthus (Machado et al. 1998). Chelonanthus alatus is commonly used in the Neotropics as a remedy against many illnesses (fevers, fungal infections, etc.), and chemical isolates have shown promising results in clinical studies. Key to the Species of Chelonanthus 1. 1. 2(1).

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

4. 5(3).

5.

Corollas blue or purple, sometimes with a white throat (rarely completely white) .................................................................................. C. purpurascens Corollas white, yellow, to green, often with a green spot or stripe on each lobe ......................................................................................................... 2 Leaves linear-lanceolate or linear-elliptic, 4–12 times longer than wide, rarely narrowly lanceolate and then 2.5–4 times longer than wide ............................................................................................ C. angustifolius Leaves ovate, elliptic, to lanceolate, 1–3 times longer than wide ............ 3 Corollas green, yellow, sometimes greenish white (very rarely white); calyx 4–13 mm long; calyx lobes 3–7 mm long ......................................... 4 Corollas white, rarely pale yellowish green to greenish white; calyx 3– 6 mm long; calyx lobes 2–4 mm long ..................................................... 5 Leaves ovate, rarely elliptic, acute to acuminate at apex (rarely obtuse), thin; inflorescences usually with several inflorescence branches ....................................................................................................... C. alatus Leaves elliptic to oblong, obtuse at apex, coriaceous and slightly rugose; inflorescences usually with only one branch ................... C. schomburgkii Stems terete or only the upper part quadrangular, flowering branches to 4 mm diameter; style deciduous or less often partially or completely persistent until the dehiscence of the fruit; fruit nodding but style not distinctly stout and persistent and not pointing downward ......... C. albus Stems quadrangular, often distinctly winged, flowering branches 4–10 mm diameter; style and stigma completely persistent until the dehiscence of the fruit; fruit nodding with style and stigma distinctly persistent and stout, pointing downward ....................................... C. longistylus

Chelonanthus 485

Chelonanthus alatus (Aubl.) Pulle, Enum. Vasc. Pl. Surinam 376. 1906. —Lisyanthus alatus Aubl., Hist. Pl. Guiane 204, t. 80. 1775. —Helia alata (Aubl.) Kuntze, Revis. Gen. Pl. 2: 427. 1891. —Irlbachia alata (Aubl.) Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 408. 1985. —Sasafrás de loma, Tabaco de morrocoy, Tabaquilla. Lisianthus chelonoides L. f., Suppl. Pl. 134. 1781 [1782]. —Helia chelonoides (L. f.) Kuntze, Revis. Gen. Pl. 2: 427. 1891. —Chelonanthus chelonoides (L. f.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. Lisianthus acutangulus Ruiz & Pav., Fl. Peruv. 2: 14. 1799. —Helia acutangula (Ruiz & Pav.) Kuntze, Revis. Gen. Pl. 2: 428. 1891. —Chelonanthus acutangulus (Ruiz & Pav.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. Lisianthus trifidus H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 182. 1818 [1819]. —Lisyanthus trifidus (H.B.K.) Griseb., Gen. Sp. Gent. 185. 1839 [1838]. —Helia trifidus (H.B.K.) Kuntze, Revis. Gen. Pl. 2: 427. 1891. Lisianthus virgatus Progel in Mart., Fl. Bras. 6(1): 239. 1865. —Adenolisianthus virgatus (Progel) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. Branched or unbranched slender to robust annual herb to 3.5 m tall; leaves sessile or less often petiolate (lower leaves especially), ovate, elliptic, or rarely circular or obovate, widest below or at middle or a little bit above, (20–)80–280 × (8–)40–190 mm, the base long-attenuate to cordate, the apex usually acute or acuminate, less often obtuse; calyx 4–13 mm long, the lobes 3–7 mm long; corolla 15–45 mm long, the lobes 3–11 mm long; capsule elliptic, 9–23 mm long. Common and often weedy in lowland savannas, roadsides, and riversides or other secondary vegetation, 0–1100 m; Delta Amacuro (Serranía de Imataca), Bolívar (widespread except in the Gran Sabana), Amazonas (widespread). Widespread elsewhere in Venezuela; Mexico, Belize, Honduras, Nicaragua, Guatemala, Costa Rica, Panama, Grenada, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

Chelonanthus alatus is the most widespread and common of the tropical American gentians. It is also the most morphologically variable of all species of Chelonanthus and the majority of the species described in Chelonanthus are now considered synonyms of C. alatus (See Pringle 1995). Chelonanthus chelonoides is tentatively included here in C. alatus pending further studies of the phylogeny of this genus. Chelonanthus albus (Spruce ex Progel) Badillo in Pittier et al., Cat. Fl. Venez. 2: 293. 1947. —Lisianthus albus Spruce ex Progel in Mart., Fl. Bras. 6(1): 237. 1865. —Helia alba (Spruce ex Progel) Kuntze, Revis. Gen. Pl. 2: 428. 1891. —Irlbachia alata subsp. alba (Spruce ex Progel) J.G.M. Pers. & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 409. 1985. —Sarusa, Tabaco de venado, Tabaco picure. Slender and weak, annual or perennial herb to 2 m tall; leaves sessile, rarely petiolate, ovate to elliptic, widest below middle, 25–170 × 8–65 mm; calyx campanulate, 3–6 mm long, divided 2/3–4/5 of the length, the lobes 2–4 mm long; corolla 22–45 mm long, the lobes 6–8 mm long; capsule 9–17 mm long. Lowland moist savannas and open areas, riversides, forests (often on white sand or associated with granitic outcrops), 50–200 m; Amazonas (Río Baría, Río Casiquiare, Río Mawarinuma, Río Negro). Brazil (Amazonas: around Manaus and Rio Negro). ŠFig. 417. Chelonanthus albus can be distinguished from a rare white-flowered form of C. purpurascens by having erect or only slightly recurved anthers after anthesis (as opposed to the strongly recurved anthers of C. purpurascens) as well as thinner and weaker inflorescence branches. Chelonanthus angustifolius (H.B.K.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. —Lisianthus angustifolius H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 181. 1818 [1819], non Mart. 1826 [1827]. —Lisyanthus angustifolius (H.B.K.) Griseb., Gen. Sp. Gent. 186. 1839 [1838]. —Helia angustifolia (H.B.K.) Kuntze, Revis. Gen. Pl. 2: 428.

486

G ENTIANACEAE

1891. —Irlbachia alata subsp. angustifolia (H.B.K.) J.G.M. Pers. & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 409. 1985. —Sasafrás de loma. Lisianthus bifidus H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 181. 1818 [1819]. —Lisyanthus bifidus (H.B.K.) Griseb., Gen. Sp. Gent. 186. 1839 [1838]. —Helia bifida (H.B.K.) Kuntze, Revis. Gen. Pl. 2: 428. 1891. —Chelonanthus bifidus (H.B.K.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. Chelonanthus pyriformis Gleason, Bull. Torrey Bot. Club 58: 450. 1931. —Lisianthus pyriformis (Gleason) Steyerm., Fieldiana, Bot. 28: 498. 1953. Slender to robust, unbranched or sparsely branched, erect annual herb to 1.5 m tall; leaves sessile or rarely subsessile, linear to very narrowly ovate, elliptic, or obovate, widest at middle, (12–)20–150(–210) × 3–18(– 22) mm; calyx 4–7 mm long, divided 2/3–3/4 of the length, the lobes 3–6 mm long; corolla (19–)27–50 mm long, the lobes 4–9 mm long; capsule 5–10(–14) mm long. Savannas and secondary vegetation (often on white sand and rocky areas), 50–1600 m; Bolívar (Gran Sabana, Río Orinoco and tributaries), Amazonas (Cerro Duida, Cerro Moriche, La Esmeralda, Puerto Ayacucho, Sierra Parima). Guárico, Sucre(?); Colombia (Amazonas, Cundinamarca, Meta, Santander, Tolima, Vaupés), Guyana (Roraima), Peru, Brazil (northern Amazonas, Pará, Roraima). ŠFig. 418. Chelonanthus longistylus (J.G.M. Pers. & Maas) L. Struwe & V.A. Albert, Harvard Pap. Bot. 3: 70. 1998. —Irlbachia alata subsp. longistyla J.G.M. Pers. & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 410. 1985. Branched robust herb to 2.5 m tall; leaves sessile or petiolate, ovate, elliptic, or obovate, sometimes narrowly so, widest below middle, 25–250 × 10–120 mm, the apex acute; calyx 3–8 mm long, divided 2/3 of the length, the lobes 3–6 mm long; corolla 22–50 mm long, the lobes 3–11 mm long; capsule 11–16 mm long. Savannas, swamps, roadsides, riversides, secondary forests, open areas in forests; 300–1800 m; Bolívar (Cerro Altamira, Cerro Bolívar, Gran Sabana, Sierra de Lema). Trinidad, Guyana, Suriname,

French Guiana, Brazil (Roraima: Serra dos Surucucus). Chelonanthus purpurascens (Aubl.) L. Struwe, S. Nilsson & V.A. Albert, Harvard Pap. Bot. 3: 70. 1998. —Lisyanthus purpurascens Aubl., Hist. Pl. Guiane 201, t. 79. 1775. —Helia purpurascens (Aubl.) Kuntze, Revis. Gen. Pl. 2: 428. 1891. —Irlbachia purpurascens (Aubl.) Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 408. 1985. —Wakauyek (Arekuna). Lisyanthus uliginosus Griseb., Gen. Sp. Gent. 1812. 1839 [1838]. —Helia uliginosa (Griseb.) Kuntze, Revis. Gen. Pl. 2: 428. 1891. —Chelonanthus uliginosus (Griseb.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. Slender, branched, sometimes slightly suffrutescent herb or small shrub to 2 m tall; leaves sessile or shortly petiolate, ovate or elliptic, 13–92(–142) × 6–70 mm, the base attenuate, acute to obtuse, the apex acute to acuminate; calyx 5–10 mm long, divided 2/5– 1/2 of the length, the lobes 2–5 mm long; corolla 20–68 mm long, the lobes 5–14 mm long; capsule 7–17 mm long. Savannas, roadsides, riversides, secondary vegetation, forest edges, lowland and montane forests, most common in the highlands, 50–2000 m; widespread in Bolívar and Amazonas. Mérida; widespread in Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, and Brazil. Chelonanthus purpurascens is very variable morphologically, especially in leaf and corolla size. The individuals with the largest corollas seem to occur mainly in the easternmost area of distribution, from Bahia in Brazil to Guyana. Specimens with smaller corollas and smaller leaves are more common on the tepui summits and white-sand areas in southern Venezuela and Colombia. Chelonanthus schomburgkii (Griseb.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. —Lisianthus schomburgkii Griseb. in A. DC., Prodr. 9: 75. 1845. —Helia schomburgkii (Griseb.) Kuntze, Revis. Gen. Pl. 2: 428. 1891. Unbranched slender to robust herb to 3 m tall, sometimes suffrutescent; leaves elliptic, circular, or obovate, usually widest below

Chelonanthus 487

middle, 45–70 × 24–55 mm, the base obtuse to cordate, the apex obtuse; calyx 5–7 mm long, the lobes 3–5 mm long; corolla 29–38 mm long, the lobes 6–8 mm long; capsule elliptic, 9–13 mm long. White-sand savannas, 50–200 m; Amazonas (Maroa, Puerto Ayacucho). Colombia, Guyana. The status of this species is uncertain and it might later be shown to be conspecific with Chelonanthus alatus. Chelonanthus schomburgkii is here tentatively accepted as a distinct species pending further investigations in the genus.

Fig. 417. Chelonanthus albus

Fig. 418. Chelonanthus angustifolius

488

G ENTIANACEAE

4. CHORISEPALUM Gleason & Wodehouse, Bull. Torrey Bot. Club 58: 451. 1931. by Lena Struwe and Victor A. Albert Small trees and erect or scandent shrubs. Stems terete or quadrangular. Leaves petiolate, ovate to elliptic, glossy or rugose, often very coriaceous; the base obtuse to acute; the apex truncate, obtuse to acute; interpetiolar lines present; margins flat or recurved; venation pinnate with many pairs of straight secondary veins, inconspicuous or conspicuous, veins usually raised on the lower surface. Inflorescence terminal, dichasial, 1–9-flowered; bracts leaf-like. Flowers pedicellate, actinomorphic, erect. Calyx 4-merous, not fused at base, the free lobes decussate, 15–50 mm long (outer pair slightly larger than inner pair), lanceolate, not keeled or the outer pair with 2 dorsal keels (C. sipapoanum), hyaline-margined, thick and coriaceous, deciduous in fruit; corolla 6-merous, 25–70 mm long, tubular to salver- or funnelform, green, sometimes with purple margins, rather thin, deciduous in fruit, the lobes ovate to elliptic, sometimes aristate or acuminate, the corolla bud apex tapering. Stamens attached near the base of the corolla tube; filaments of equal length; anthers basifixed, linear, erect after anthesis; pollen in monads, the exine reticulate. Ovary with a disk around the base; style long, slender, deciduous in fruit; stigma bilamellate with elliptic lobes. Capsule elliptic, erect, with pericarp splitting from the base into 4 separate, deciduous parts and leaving fibrous traces of ovarian vasculature at the base. Seeds elliptic, flattened, winged, the testa cells papillose to rounded and smooth (C. psychotrioides). Endemic to the Guayana Shield in southern Venezuela, Guyana (Pakaraima Mountains), and Suriname (Tafelberg); 5 species, all in the flora area. Key to the Species of Chorisepalum 1.

1.

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

Upper surface of leaves dull, with prominent venation; secondary veins impressed, the lower surface with secondary venation raised; leaves usually brown when dry ........................................................................ 2 Upper surface of leaves glossy and smooth, with inconspicuous venation; secondary veins not conspicuously impressed, the lower surface with secondary venation not raised; leaves usually green when dry ........... 3 Upper surface of leaves punctate; leaves round-orbicular with truncate or obtuse apex; outer sepals 15–25 mm long ....................... C. rotundifolium Upper surface of leaves not punctate; leaves broadly ovate with acute or acuminate apex; sepals 25–33 mm long .......................... C. psychotrioides Leaves very thick with revolute margin; secondary veins invisible .......... .................................................................................................. C. carnosum Leaves thinner with flat margin; secondary veins usually visible ........... 4 Outer sepals each with 2 dorsal keels (1–2 mm tall) ............. C. sipapoanum Outer sepals not keeled .................................................................. C. ovatum

Chorisepalum carnosum Ewan, J. Wash. Acad. Sci. 37: 394. 1947. Climbing or scandent shrub, sometimes epiphytic, to 3 m tall; branches terete; leaves very broadly elliptic to broadly ovate, subsucculent, with obtuse or slightly acute api-

ces, 31–105 × 33–58 mm, the margin revolute; flowers solitary; outer sepals 36–50 mm long, not keeled; corolla 55–65 mm long with spreading, acute lobes; capsule ca. 25 mm long. Montane forests on tepui slopes and summits, 700–1600 m; Bolívar (Carrao-

Chorisepalum 489

tepui, Cerro Uroi, Cerro Venamo, Macizo del Chimantá [Abácapa-tepui]). Guyana (Pakaraima Mountains). ŠFig. 422. Chorisepalum ovatum Gleason, Bull. Torrey Bot. Club 58: 452. 1931. Chorisepalum breweri Steyerm. & Maguire, Bol. Soc. Venez. Ci. Nat. 32: 392. 1976. Shrub to 3 m tall; branches often slightly quadrangular; leaves elliptic, acute or acuminate apex, 30–110 × 20–55 mm; flowers solitary; outer sepals 35–45 mm long; corolla 50–70 mm long with circular and aristate lobes; capsule 16–30 mm long. Montane forests on tepui slopes and summits, 1000–2000 m; Bolívar (Cerro Guanay, Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Yutajé, Sierra de Maigualida). Endemic.

Fig. 419. Chorisepalum rotundifolium

Chorisepalum psychotrioides Ewan, J. Wash. Acad. Sci. 37: 394. 1947. Chorisepalum acuminatum Steyerm., Bol. Soc. Venez. Ci. Nat. 23: 75. 1962. —Chorisepalum psychotrioides var. acuminatum (Steyerm.) Maguire, Mem. New York Bot. Gard. 32: 349. 1981. Sprawling vine, upright shrub, or small tree to 8 m tall; young branches quadrangular, older terete; leaves 40–132 × 21–82 mm, the margin not recurved; inflorescence 2–9flowered; outer sepals 25–33 mm; corolla 30– 40 mm long, the lobes acute and sometimes aristate, spreading; capsule 13–15 mm long. Montane forests on tepui slopes and summits, 800–2000 m; Bolívar (Cerro Venamo, Ilú-tepui, Kukenán-tepui, Ptari-tepui, Roraima-tepui). Guyana (Pakaraima Mountains). ŠFig. 421.

Fig. 420. Chorisepalum sipapoanum

490

G ENTIANACEAE

Fig. 421. Chorisepalum psychotrioides Fig. 422. Chorisepalum carnosum

Two varieties, var. acuminatum (characterized by a scandent shrubby habit) and var. psychotrioides (characterized by an upright shrubby or arborescent habit), have been accepted earlier (Maguire 1981). However, the differences do not seem to be taxonomically consistent, but merely a natural variation in growth form, and therefore, these two varieties are not recognized here.

Chorisepalum rotundifolium Ewan, J. Wash. Acad. Sci. 37: 396. 1947. Low shrub to small tree to 2(–)4 m tall; branches terete or slightly quadrangular; leaves circular to obovate, 18–72(–142) × 12– 53(–110) mm, about as wide as long, with recurved margin, secondary venation prominent, impressed on the upper surface and raised on the lower surface; inflorescence 1–

Coutoubea 491

8-flowered; outer sepals 15–23 mm long; corolla 25–30 mm long, with acute, spreading lobes; anthers ca. 2 mm long, oblong; capsule 12–20 mm long. Montane forests and scrub on tepui slopes and summits, 1800–2400 m; Bolívar (Camarcaibarai-tepui, Ilú-tepui, Ptari-tepui, Sororopán-tepui, Tereké-Yuréntepui). Guyana (Mount Roraima). ŠFig. 419. The only known collection from Ilú-tepui (Maguire 33456, NY, VEN) is a tree to 4 m tall, with very large (to 14 cm long) leaves and might represent a different taxon. Chorisepalum sipapoanum (Maguire) L. Struwe & V.A. Albert, comb. nov. —Chorisepalum ovatum var. sipapoanum Maguire, Mem. New York Bot. Gard. 32: 344. 1981.

Shrub to 5 m tall; young branches quadrangular; leaves lanceolate to ovate, acute or acuminate tip, 40–70(–110) × 10–35(–55) mm, secondary venation nearly invisible; inflorescence 1(–3)-flowered; outer sepals 32– 40 mm long; corolla 45–58 mm long, the lobes aristate; capsule 15–21 mm long. Montane forests, savannas, and along streams on tepui slopes and summits, 1400–1700 m; Amazonas (Cerro Sipapo). Suriname (Tafelberg). ŠFig. 420. Maguire (1981) included Chorisepalum sipapoanum as a variety under C. ovatum. However, he must have misinterpreted the calyx lobes of the type collection of C. ovatum as being keeled as C. sipapoanum is the only species with keeled calyx lobes.

5. COUTOUBEA Aubl., Hist. Pl. Guiane 72, t. 27, 28. 1775. by Lena Struwe Annual or perennial herbs 5–120 cm tall, erect or rarely creeping. Stems terete to quadrangular. Leaves sessile or slightly petiolate, linear, lanceolate, obovate, to ovate, sometimes amplexicaul, chartaceous, the base obtuse to acute, the apex obtuse, acute, to acuminate; interpetiolar line present; venation pinnate, sometimes with arcuate secondary veins. Inflorescence terminal, a raceme or congested spike, sometimes also with axillary branches; bracts leaf-like. Flowers sessile or very shortly pedicellate, 4(5)-merous, often arranged horizontally or nearly so in whorls of 2–4. Calyx campanulate or tubular, divided nearly to the base, thin, persistent in fruit, the lobes narrowly triangular, acute, hyaline-margined, with or without low dorsal keels; corolla salverform with tube about as long as lobes, white, yellow, or greenish, often with blue, brown, green, or purple markings, thin, persistent in fruit, the corolla tube with conspicuous fibers (seen as white lines in old, dry corollas), the lobes lanceolate to ovate, acute, erect, spreading, or reflexed after anthesis. Stamens inserted in the upper part of the corolla tube, with a flat, hood-like structure at the insertion point; filaments equal in length, bent outward after anthesis; anthers oblong-linear, sagittate, erect after anthesis, soon deciduous; pollen in tetrads, the exine reticulate and microreticulate. Ovary without a basal disk; style slender, deciduous; stigma bilamellate, the lobes oblong. Capsule elliptic, enclosed by the persistent calyx and corolla, erect to horizontal. Seeds angular or globose, not winged, the testa cell morphology unknown. Mexico, Guatemala, Belize, Costa Rica, Panama, St. Vincent, Guadeloupe, Colombia, Venezuela, Trinidad, Tobago, Guyana, Suriname, French Guiana, Peru, Brazil; 5 species, 4 in Venezuela, all in the flora area. The only other species in the genus, Coutoubea humilis Sandwith, is a highland endemic of the Pakaraima Plateau (Guyana). Coutoubea is closely related to Schultesia, with which it shares persistent corollas in fruit and very similar pollen in tetrads (Struwe et al. 1998). Coutoubea spicata has been reported to be poisonous.

492

G ENTIANACEAE

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

3.

Small herbs 25 cm tall or less; leaves 6–20 × 1–5 mm ..................... C. minor Larger plants, 30–80 cm tall; leaves usually > 30 × 10 mm ..................... 2 Flowers tightly congested in spikes at the end of the usually unbranched stem, 3 or 4 in each whorl ............................................................ C. spicata Flowers loosely arranged in racemes in the upper part of the usually branched stem, 2 opposite each other in each whorl ............................ 3 Flowers shortly pedicellate (pedicel 1–3 mm long); corolla lobes erect or slightly spreading after anthesis; leaves spread out along stem, usually thin and membranaceous, lanceolate to ovate, rarely linear .... C. ramosa Flowers sessile; corolla lobes reflexed after anthesis; leaves aggregated close to the base of the stem, usually thick and coriaceous, oblanceolate to lanceolate-linear ....................................................................... C. reflexa

Coutoubea minor H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 179. 1818 [1819]. Small branched herb 5–25 cm tall; leaves narrowly ovate-elliptic, 6–20 × 1–5 mm, the base obtuse to cordate, the apex acute; inflorescence congested; flowers in whorls of 2(3); calyx 4–5 mm long; corolla 7–10 mm long, white or yellow, salverform, the lobes 5–6 mm long, elliptic, not reflexed after anthesis; capsule 3–4 mm long. Lowland savannas on sand, usually associated with granitic outcrops, often abundant, 50–200 m; Bolívar (Cerro San Borja), Amazonas (along Río Orinoco from Puerto Ayacucho to Río Atabapo and the mouth of Río Ventuari). Endemic. ŠFig. 424. Coutoubea ramosa Aubl., Hist. Pl. Guiane 74, t. 28. 1775. —Fregosillo, Lengua piapoco. Coutoubea racemosa G. Mey., Prim. Fl. Esseq. 86. 1818. —Exacum racemosum (G. Mey.) Roem. & Schult. in Schult. & Schult. f., Mantissa 3: 99. 1827. —Coutoubea ramosa var. racemosa (G. Mey.) Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 198. 1854. —Coutoubea ramosa f. racemosa (G. Mey.) Jonker in Pulle, Fl. Suriname 4(1)F: 407. 1936. Branched herb to 1 m tall; leaves narrowly lanceolate to linear, 24–116 × 4–24 mm, the base acute, the apex acute; inflorescences not congested, internodes over (5–)8 mm long between whorls; flowers 2 in each whorl; calyx 4–7 mm long; corolla 11–13 mm long, white, brownish, lavender, or pink, the lobes ovate, 4–5 mm long; capsule 5–6 mm long. Lowland savannas (sometimes on white sand), forests, riversides, secondary vegetation, 0–200(–500) m; widespread in

Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Guárico, Zulia; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 423. Some authors distinguish among 3 varieties of this species, Coutoubea ramosa var. longifolia Benth., var. racemosa (G. Mey.) Benth., and var. ramosa, but these subspecific taxa are not recognized here. Coutoubea ramosa is similar to C. reflexa, but differs in its shortly pedicellate flowers and leafier stems, especially immediately below the inflorescences. Coutoubea reflexa Benth., Ann. Nat. Hist. 2: 442. 1839. —Cadena. Suffrutescent herb to 10–30(–100) cm tall; leaves often aggregated close to the base, elliptic to lanceolate-linear, 20–100(–180) × 9– 21 mm, the base acute, the apex obtuse, acute, or rarely acuminate; inflorescences not congested, internodes > 10 mm long between flowers; flowers 2 or 3(4) in each whorl; calyx 6–9 mm long; corolla 14–19 mm long, white, yellowish, pinkish, or greenish, sometimes brown, purplish, or green in throat, the lobes 6–8 mm long, ovate, acute, or acuminate, reflexed after anthesis; capsule 7–9 mm long. Wet savannas associated with white sand, sandstone, or granite, 50– 1000 m; Bolívar (Gran Sabana, upper Río Caroní, Río Cuyuní), Amazonas (Río Atabapo, along Río Orinoco in the vicinity of Puerto Ayacucho), Guyana (Pakaraima and Rupununi savannas, Río Caroní), Brazil (Roraima). Coutoubea reflexa is probably closely related to C. ramosa (see note under that species).

Coutoubea 493

Fig. 423. Coutoubea ramosa

Fig. 424. Coutoubea minor

Fig. 425. Coutoubea spicata

494

G ENTIANACEAE

Coutoubea spicata Aubl., Hist. Pl. Guiane 72, t. 27. 1775. —Exacum spicatum (Aubl.) Vahl, Symb. Bot. 3: 17. 1794. Simple or sparsely branched herb, often suffrutescent, 25–120 cm tall; leaves 22–110 × 10–35 mm, narrowly ovate-triangular to narrowly oblong-lanceolate (basal leaves sometimes elliptic to subobovate and obtuse at apex), the base obtuse to cordate, amplexicaul or nearly so, the apex acute to acuminate; inflorescences elongated, congested, 15–230 mm long, internode between each whorl of flowers < 10 mm long; flowers

in whorls of 3 or 4; calyx 6–7 mm long, the lobes 3–5 mm long; corolla 10–14 mm long, white or greenish white, often brown-colored in the mouth, the lobes 4–7 mm long, ovate, reflexed after anthesis; capsule 7–8 mm long. Savannas (often white-sand), forest edges, roadsides, 50–900 m; widespread in Bolívar, Amazonas (Puerto Ayacucho, Río Atabapo, Río Ventuari basin). Nueva Esparta, Zulia; widespread from Mexico, Guatemala, Belize, Costa Rica, Panama, St. Vincent, Guadeloupe, Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 425.

6. CURTIA Cham. & Schltdl., Linnaea 1: 209, t. 4. 1826. Schuebleria Mart., Nov. Gen. Sp. Pl. 2: 113. 1826 [1827], “Schübleria.” Apophragma Griseb., Gen. Sp. Gent. 163. 1839 [1838]. by Lena Struwe Single-stemmed or sparsely branched annual herbs. Stems angled, with low decurrent ridges. Leaves opposite or verticillate (2–8 in each whorl), sessile to subsessile, linear to lanceolate or obovate, ovate, to rotund, the base obtuse to attenuate, the apex obtuse, acute, or acuminate; interpetiolar line present, inconspicuous; petiole 0–2 mm long; margins flat; venation inconspicuous. Inflorescences terminal, sometimes also with axillary branches, paniculate with small cymes, 1–100flowered; bracts and floral bracts leaf-like and scale-like. Flowers sessile or pedicellate, 4- or 5-merous, actinomorphic, sometimes heterostylous, erect. Calyx tubular, divided nearly to the base, thin, persistent in fruit, the lobes ovate-triangular to linear, keeled, the apex acute; corolla funnelform or tubular, often constricted at the throat, white, pink, yellow, purplish, or lavender, the inside of the tube often hairy, thin, deciduous in fruit, the lobes obovate, ovate, circular, or elliptic, with obtuse, acute, or acuminate apices, the corolla bud apex slightly tapering. Stamens inserted in the corolla tube at varying levels, usually included in corolla; filaments of equal length within each flower (differing between flowers due to heterostyly); anthers sagittate or not, erect after anthesis, with or without a very small sterile apical appendage, thecae rarely with basal hairy appendages, the connective inconspicuous or flattened and inconspicuous; pollen in monads, the exine reticulate, microreticulate or with a perforate tectum. Ovary without a disk at the base; style absent or short (to 5 mm), persistent; stigma capitate to clavate. Capsule ovoid, thinwalled, dehiscing apically, erect. Seeds rounded to angular, not winged, the testa cells reticulate and smooth. Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Uruguay, Argentina; ca. 6 species, 3 in Venezuela, all in the flora area. Key to the Species of Curtia 1. 1. 2(1). 2.

Leaves verticillate, (3)4 in each whorl ........................................... C. conferta Leaves opposite, 2 in each whorl (rarely 3 in each whorl) ....................... 2 Leaves broadly ovate to broadly elliptic, 3–10 mm wide; corollas 2–4 mm long ......................................................................................... C. obtusifolia Leaves linear, narrowly ovate, to narrowly spatulate, 0.5–3 mm wide; corollas 4–16 mm long ................................................................. C. tenuifolia

Curtia 495

Fig. 426. Curtia obtusifolia

Fig. 427. Curtia conferta

Fig. 428. Curtia tenuifolia

496

G ENTIANACEAE

Curtia conferta (Mart.) Knobl., Bot. Centralbl. 60: 357. 1894. —Schuebleria conferta Mart., Nov. Gen. Sp. Pl. 2: 115. 1826 [1827], “Schübleria conferta.” —Ají de morocoy, Garrapato. Curtia quadrifolia Maguire, Mem. New York Bot. Gard. 32: 352, 355, figs. 72h–q, 75. 1981. Branched herb to 35 cm tall; leaves broadly ovate, 1–8 × 1–6 mm; calyx lobes triangular, 1–2 mm long; corolla white, 2–3 mm long; capsule 2–3 mm long. White-sand or sandstone savannas and forest edges, 50–200 m; Amazonas (vicinity of Cerro Yapacana, Río Autana, Río Negro, Río Siapa). Colombia (Amazonas: Araracuara, Vaupés: Mitú, Río Caqueta, Río Kubiyú), Brazil (Amazonas: Rio Negro). ŠFig. 427. The name Curtia conferta was misused earlier for a southeast Brazilian taxon (renamed C. confusa by Grothe and Maas 1984). Curtia obtusifolia (Benth.) Knobl., Bot. Centralbl. 60: 357. 1894. —Schuebleria obtusifolia Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 194. 1854. Branched herb to 40 cm tall; leaves 1–5 (–14) × 1–3 mm; calyx lobes narrowly triangular, 1–3 mm long; corolla white, 5–7 mm long; capsule ca. 8 mm long. White-sand savannas, 50–200 m; Amazonas (Caño Pimichín, vicinity of Cerro Yapacana, Río Negro, Río Sipapo). Brazil (Amazonas: upper Rio Negro, Rio Uneiuxi). ŠFig. 426.

Curtia tenuifolia (Aubl.) Knobl., Bot. Centralbl. 60: 357. 1894. —Exacum tenuifolium Aubl., Hist. Pl. Guiane 70, t. 26, fig. 2. 1775. —Apophragma tenuifolium (Aubl.) Griseb., Gen. Sp. Gent. 163. 1839 [1838]. —Schuebleria tenuifolia (Aubl.) G. Don, Gen. Hist. 4: 202. 1838. Schuebleria tenella Mart., Nov. Gen. Sp. Pl. 2: 114. 1826 [1827]. —Curtia tenella (Mart.) Cham., Linnaea, 8: 13. 1894. —Curtia tenuifolia subsp. tenella (Mart.) Grothe & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 87: 40. 1984. Usually branched herb to 60 cm tall; leaves 1–20 × 0.5–3 mm; calyx 3–7 mm long, the lobes linear to narrowly triangular; corolla white, lavender, purple, or pink, 5–16 mm long; capsule 2–7 mm long. Savannas and open places, 50–1500 m; Bolívar (Cerro Médano, Gran Sabana), Amazonas (base of Cerro Camani, La Esmeralda, near Puerto Ayacucho, vicinity of Santa Bárbara). Apure, Guárico, Mérida, Portuguesa, Sucre, Yaracuy; Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Uruguay, Argentina. ŠFig. 428. Two subspecies, Curtia tenuifolia subsp. tenella and subsp. tenuifolia, have been recognized by some authors but the morphological distinction between these two taxa is very weak and they are not accepted here.

7. ENICOSTEMA Blume, Bijdr. 848. 1826 [1825]. by Lena Struwe Annual or perennial herbs, often suffrutescent. Stems terete to angular, often winged. Leaves sessile or subsessile, linear, narrowly ovate, to lanceolate, with obtuse or acute apex; venation prominent with 1 basal pair of arcuate secondary veins. Inflorescences axillary, dense clusters, 1–20-flowered; bracts leaf-like, small. Flowers sessile, 5-merous (rarely 3-, 4-, or 6-merous), actinomorphic, horizontal. Calyx narrowly campanulate, divided 2/3 or more of the length, thin, persistent in fruit, the lobes narrowly triangular (E. verticillatum) or elliptic-ovate, with white, hyaline margins; corolla tubular to narrowly funnelform, white, thin, persistent in fruit, the lobes triangular, ovate to oblong, the apex acute, the corolla bud apex tapering. Stamens inserted in the middle of the corolla tube, with a double hood-shaped appendix at the base of each filament; filaments of equal length; anthers linear to oblong, erect after anthesis, with a sterile apical appendage; pollen in monads, the exine reticulate. Ovary without a disk at the base; style shorter than the ovary, deciduous in

Enicostema 497

Fig. 429. Enicostema verticillatum

498

G ENTIANACEAE

fruit; stigma capitate, slightly bilobed. Capsule obovoid, dehiscing apically, erect to horizontal. Seeds rounded, not winged; testa reticulate. Costa Rica, Panama, Lesser Antilles, Venezuela, Trinidad, Tobago, Guyana, tropical Africa, Madagascar, India, Southeast Asia, Malesia; 3 species, 1 in Venezuela. Enicostema, which also has one African-Asian species and one species endemic to Madagascar, is closely related to American-African Neurotheca and the African genera Faroa, Pycnosphaera, and Urogentias (Struwe et al. 1998). Enicostema verticillatum (L.) Engl. ex Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 67. —Gentiana verticillata L., Syst. Nat. ed. 10, 2: 952. 1759. Perennial herb to 1 m tall; leaves 35–79 × 5–23 mm, lanceolate; calyx 4–6 mm long, the lobes 2–4 mm long; corolla 6–8 mm long, the lobes 1–2 mm long; capsule 3–5 mm long.

Lowland and middle elevation savannas and disturbed areas, 50–400 m; Bolívar (road between Tumeremo and Anacoco, vicinity of Upata). Lara, Sucre; Panama, Costa Rica, Antigua, Grenada, Dominica, Guadeloupe, Martinique, Nevis, St. Kitts, St. Lucia, Trinidad, Tobago, Guyana. ŠFig. 429.

8. IRLBACHIA Mart., Nov. Gen. Sp. Pl. 2: 101. 1826 [1827]. Pagaea Griseb. in DC., Prodr. 9: 70. 1845. Lisianthus sect. Brachycodon Benth., Hooker’s J. Bot.Kew Gard. Misc. 6: 200. 1854. —Brachycodon (Benth.) Progel in Mart., Fl. Bras. 6(1): 229. 1865. by Lena Struwe and Oliver Pihlar Annual herbs, sometimes woody at base. Stems terete, quadrangular, or 4winged. Leaves cauline, sometimes in a basal rosette and/or often most abundant close to the base, petiolate to sessile, elliptic, ovate, lanceolate, or linear, chartaceous to nearly coriaceous, the base obtuse, acute, or attenuate, the apex obtuse, acute, or acuminate; interpetiolar line present; venation conspicuous or inconspicuous with 0–2 pairs of basally divergent arcuate secondary veins. Inflorescence a terminal cyme usually with monochasial branches and with flowers often arranged along one side or alternating on opposite sides, 1–50-flowered; bracts and floral bracts scalelike (leaf-like in I. pumila and I. poeppigii). Flowers pedicellate, 5-merous, actinomorphic (sometimes with zygomorphic stamens), often becoming nodding with age. Calyx campanulate, divided 1/2–4/5 of the length, thick, persistent in fruit, the lobes elliptic with a dorsal glandular area or ridge, erect, the apex obtuse (acute in I. pratensis), hyaline-margined; corolla salverform, campanulate, or narrowly funnelform, white, pink, rose, lavender, blue, or purple, thin, deciduous in fruit, rarely persistent, the lobes 1–2 times as long as the corolla tube (3–6 times as long in I. tatei), broadly ovate to triangular, erect to recurved, the apex obtuse to acute, the corolla bud apex rounded (tapering in I. pratensis). Stamens inserted in the lower part of the corolla tube; filaments of equal or unequal length; anthers erect to recurved after anthesis, slightly sagittate, with sterile apical appendage; pollen in polyads, the exine with spines or globules. Ovary with a glandular disk at the base of the ovary (possibly absent in I. pratensis); style slender, persistent in fruit; stigma bilamellate, the lobes elliptic. Capsule ovoid to ellipsoid, dehiscing medially, often nodding. Seeds angular to globose, not winged; testa cells dome-like to concave with band-like thickenings. Southern Venezuela, Guyana, northern Brazil; 9 species, 8 in Venezuela, all in the flora area.

Irlbachia 499

This genus is endemic to the Guayana Shield, and most species are endemic to lowland white-sand areas (in Amazonas, Venezuela) or tepui summits (Bolívar and Amazonas). Irlbachia has also been used in a wider circumscription, including species from the genera Adenolisianthus, Calolisyanthus, Chelonanthus, Helia, Rogersonanthus, Tetrapollinia, and Wurdackanthus (Maas 1985). However, since the name Helia (a Brazilian endemic genus) currently has priority over Irlbachia, Irlbachia cannot be used for such a wide circumscription (Struwe and Albert 1998). Here Irlbachia is used in a narrow sense using the generic circumscription of Maguire (1981), Maguire & Boom (1989), and Struwe et al. (1997). Key to the Species of Irlbachia 1. 1. 2(1).

2.

3(1).

3.

4(3).

4.

5(3). 5. 6(5).

6. 7(6). 7.

Leaves linear or rarely narrowly lanceolate, 1–3(–5) mm wide, appearing 1-veined .................................................................................................. 2 Leaves lanceolate, ovate, or elliptic, > 5 mm wide, with 1–3 pairs of secondary veins ........................................................................................... 3 Small herb to 20 cm tall; leaves membranaceous; corollas 4–6 mm long, white; calyx 2–4 mm long; styles 1–2 mm long; capsules 4–7 mm long ....................................................................................................... I. pumila More robust herb, to 70 cm tall; leaves thick and coriaceous; corollas 18– 36 mm long; blue, blue-purple, or lavender, rarely white; calyx 4–7 mm long; styles 15–19 mm long; fruits over 7 mm long .................. I. pratensis Inflorescences long-pedunculate, peduncles (50–)100–300 mm long (as measured from the uppermost pair of leaves to the first flower in the inflorescence); corollas 10–33 mm long; leaves usually rather thick and subcoriaceous, with inconspicuous venation ........................................ 4 Inflorescences not long-pedunculate, peduncles 10–60 mm long; corollas 3–10(–12) mm long; leaves usually thin and membranacous, with 1 or 2 pairs of prominent secondary veins ................................................... 5 Corollas purple or blue (rarely white or lavender), 10–19 mm long, broadly funnelform with spreading lobes; corolla tubes 1–2 times as long as corolla lobes ................................................................... I. cardonae Corollas rose to pink, (17–)20–33 mm long, tubular or narrowly funnelform with erect or slightly spreading lobes; corolla tubes 3–6 times as long as corolla lobes ........................................................................... I. tatei Leaves in a basal rosette (or in a tight whorl at most a few cm up on the stem); plants to 15 cm tall ............................................... I. plantaginifolia Leaves cauline, not in a basal rosette; plants 15–50 cm tall, rarely smaller ................................................................................................................ 6 Inflorescences terminal and also with axillary inflorescence branches or solitary flowers present in the upper leaf axils; corollas ca. 12 mm long ................................................................................................. I. phelpsiana Inflorescences terminal, inflorescence branch in upper leaf axils not present; corollas 3–8 mm long ............................................................... 7 Bracts leaf-like ............................................................................... I. poeppigii Bracts scale-like ............................................................................ I. nemorosa

500

G ENTIANACEAE

Irlbachia cardonae (Gleason) Maguire, Mem. New York Bot. Gard. 32: 381, figs. 85AA–MM, 87A. 1981. —Chelonanthus cardonae Gleason, Brittonia 3: 189. 1939. —Lisianthus cardonae (Gleason) Steyerm., Fieldiana, Bot. 28: 498. 1953. Lisianthus acutilobus Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 439, fig. 9. 1966. Irlbachia paruana Maguire, Mem. New York Bot. Gard. 32: 380, figs. 82AA–MM, 87B–D. 1980. Sparsely branched herb to 1.5 m tall; leaves subsessile to shortly petiolate, 11–80 × 4–26 mm, narrowly elliptic to lanceolate, the margin entire; calyx 3–5 mm long; corolla 10–19 mm long, the lobes 5–6 mm long, spreading; style 4–12 mm long; capsule 6–9 mm long. Savannas and montane forests on tepui summits, 500–2300 m; Bolívar (Amaruay-tepui, Auyán-tepui, Camarcaibaraitepui, Cerro Camarón, Cerro El Venado, Cerro Guaiquinima, Cerro Jaua, Cerro Pauo, Cerro Venamo, Gran Sabana, Ilú-tepui, Macizo del Chimantá [Abácapa-tepui, Apacará-tepui, Toronó-tepui], Murisipántepui, Ptari-tepui, Serranía Marutaní, Sierra Pakaraima, Tereké-yurén-tepui), Amazonas (Cerro Parú, Cerro Huachamacari). Guyana (Pakaraima Mountains, Karowtipu). ŠFig. 433. Irlbachia nemorosa (Willd. ex Roem. & Schult.) Merrill, Proc. Amer. Philos. Soc. 86: 88. 1943. —Claytonia nemorosa Willd. ex Roem. & Schult., Syst. Veg. 5: 436. 1819. —Ditomaga nemorosa (Willd. ex Roem. & Schult.) Raf., Actes Soc. Linn. Bordeaux 6: 267. 1834. —Irlbachia bonplandiana Fenzl in Endl. & Fenzl, Nov. Stirp. 12. Dec. 1839, nom superfl. —Helia nemorosa (Willd. ex Roem. & Schult.) Kuntze, Revis. Gen. Pl. 428. 1891. —Pagaea nemorosa (Willd. ex Roem. & Schult.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 102. 1895. —Ají de morrocoy, Barui de venado, Tabaco de morrocoy, Tabaco de venado, Tabaquilla. Lisianthus recurvus Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 201. 1854. —Irlbachia recurva (Benth.) Progel in Mart., Fl. Bras. 6(1): 229. 1865. —Pagaea recurva (Benth.) Benth. & Hook. f., Gen. Pl. 2: 814. 1876.

Lisianthus subcordatus Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 200. 1854. —Irlbachia subcordata (Benth.) Progel in Mart., Fl. Bras. 6(1): 229. 1865. —Pagaea subcordata (Benth.) Benth. & Hook. f., Gen. Pl. 2(2): 814. 1876. Sparsely branched herb to 50 cm tall, sometimes epiphytic; leaves petiolate, 20– 115 × 11–63 mm, ovate to broadly elliptic, the margin entire; calyx 2–3 mm long; corolla 4–8 mm long, white or pale lilac, the lobes 2–3 mm long; style 1–5 mm long; capsule 5–10 mm long. White-sand savannas, along rivers, flooded riparian forests, sometimes on rocks and logs, 50–500 m; Bolívar (slopes of Macizo del Chimantá), Amazonas (Río Mawarinuma, Río Negro, Río Orinoco). Amazonian Colombia, Brazil (northern Amazonas). ŠFig. 430. Irlbachia nemorosa is quite variable in leaf and inflorescence morphology and is sometimes difficult to distinguish from I. elegans, a Brazilian species, which has narrowly lanceolate leaves with attenuate bases. Irlbachia phelpsiana Maguire, Mem. New York Bot. Gard. 32: 371. 1980. Herb to 30 cm tall; leaves petiolate, 30–45 × 10–15 mm, ovate-lanceolate, the margin crenulate or entire; calyx 2–3 mm long; corolla 12–15 mm long, white or bluish, the lobes 5–6 mm long; style 4–5 mm long; capsule 4–5 mm long. Montane forests, ca. 1800 m; Amazonas (Cerro Parú). Endemic. Irlbachia plantaginifolia Maguire, Mem. New York Bot. Gard. 32: 372, 377, fig. 84. 1980. —Verdolaga. Low herb to 10 cm tall; leaves appressed to the ground, rosulate, nearly succulent, petiolate to sessile, ovate to broadly ovate, 25–98(–105) × 12–65 mm; calyx 3–5 mm long; corolla white, 5–9 mm long, the lobes broadly ovate, 1–4 mm long; style ca. 1 mm long; capsule 7–10 mm long. White-sand savannas, caatinga forests, rocks along streams, 100–200 m; Amazonas (Río Casiquiare, Río Mawarinuma, Río Negro, Río Yatúa). Endemic. ŠFig. 432. This is the only species of Irlbachia having leaves in a tight basal rosette similarly to Plantago major L. (Plantaginaceae).

Irlbachia 501

Irlbachia poeppigii (Griseb.) L. Cobb & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 86: 131. 1983. —Pagaea poeppigii Griseb. in DC., Prodr. 9: 71. 1845. Lisianthus ramosissimus Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 200. 1854. —Brachycodon ramosissimus (Benth.) Progel in Mart., Fl. Bras. 6(1): 230. 1865. —Helia ramosissima (Benth.) Kuntze, Revis. Gen. Pl. 428. 1891. —Pagaea ramosissima (Benth.) Ewan, Proc. Biol. Soc. Wash. 63: 165. 1950. —Irlbachia ramosissima (Benth.) Maguire, Mem. New York Bot. Gard. 32: 370. 1980. Branched herb to 30 cm tall; leaves 16–50 × 5–12 mm, basal leaves smaller, lanceolateovate, the leaf margin nearly always crenulate; calyx 2–3 mm long, the lobes 1–3 mm long, nearly free; corolla 3–6 mm long, white, the lobes 1–3 mm long; style 1–2 mm long; capsule 4–8 mm long. Flooded forests, 300– 400 m; Bolívar (upper Río Paragua). Brazil (Amazonas: Rio Negro, Roraima, Pará). Irlbachia poeppigii is known from Venezuela by a single, somewhat atypical collection (Fernández 4522, MO) from the upper Río Paragua (Bolívar state), but is likely to occur also along the Río Negro (Amazonas state). Irlbachia pratensis (H.B.K.) L. Cobb & Maas, Proc. Kon. Ned. Akad. Wetensch., C. 86: 132. 1983. —Lisianthus pratensis H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 180. 1818 [1819]. —Helia pratensis (H.B.K.) Kuntze, Revis. Gen. Pl. 428. 1891. Lisianthus campanuloides Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 201. 1854. —Helia campanuloides (Spruce ex Benth.) Kuntze, Revis. Gen. Pl. 428. 1891. —Chelonanthus campanuloides (Spruce ex Benth.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 98. 1895. Basally branched or unbranched herb to 70 cm tall; leaves sessile, 12–40 × 1–3 mm, spread along stem or often grouped near the base; calyx 4–7 mm long, the lobes 2–4 mm long, often somewhat spreading; corolla 18– 36 mm long, broadly funnelform, very narrow at base, blue, blue-purple, or lavender, rarely white, the lobes 6–13 mm long; capsule 7–9 mm long. White-sand savannas, 100–300 m; Amazonas (Río Casiquiare, Río Negro, Río Orinoco, Río Pasimoni-Yatúa, Río Siapa). Brazil (Roraima). ŠFig. 434.

Irlbachia pratensis is distinguished from Tetrapollinia caerulescens by its larger, more abundant, and thicker leaves and more broadly funnelform flowers with ovate corolla lobes. Irlbachia pumila (Benth.) Maguire, Mem. New York Bot. Gard. 32: 372. 1980. —Lisianthus pumilus Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 200. 1854. —Brachycodon pumilus (Benth.) Progel in Mart., Fl. Bras. 6(1): 230. 1865. —Helia pumila (Benth.) Kuntze, Revis. Gen. Pl. 428. 1891. —Pagaea pumila (Benth.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 102, fig. 45. 1895. —Sardina, Yuwiji. Branched weedy herb to 20 cm tall; leaves sessile, 10–65 × 1–3 mm, linear (basal leaves smaller and linear-obovate); calyx 2–4 mm long, the lobes 2–3 mm long; corolla 4–6 mm long, broadly campanulate to broadly funnelform, white, the lobes broadly ovate, 1–3 mm long; capsule 4–7 mm long. On rocks near rivers, white-sand savannas, seasonally flooded forests (growing on logs, moss, or humus), 100–300 m; Amazonas (Caño Caname, Cerro Yapacana and vicinity, Río Casiquiare, Río Negro). Brazil (Amazonas: Rio Negro, Rio Madeira). ŠFig. 431. Irlbachia tatei (Gleason) Maguire, Mem. New York Bot. Gard. 32: 377. 1980. —Calolisianthus tatei Gleason, Bull. Torrey Bot. Club 58: 449. 1931. —Lisianthus tatei (Gleason) Steyerm., Fieldiana, Bot. 28: 498. 1953. Lisianthus scabridulus Steyerm., Fieldiana, Bot. 28: 496. 1953. Lisianthus jauaensis Steyerm. & Maguire, Mem. New York Bot. Gard. 23: 879, fig. 13. 1972. Unbranched or sparsely branched annual herb to 1 m tall; leaves petiolate, 6–78 × 3– 25(–36) mm, narrowly elliptic, lanceolate, or broadly ovate; calyx 3–6 mm long, the lobes 1–3 mm long; corolla lobes 4–8 mm long, broadly ovate or elliptic; style 11–26 mm long; capsule 6–11 mm long. Tepui meadows, (600–)1300–2200 m; Bolívar (Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Parú). Guyana(?). ŠFig. 435.

502

G ENTIANACEAE

Fig. 430. Irlbachia nemorosa

Fig. 431. Irlbachia pumila

Fig. 432. Irlbachia plantaginifolia

Irlbachia 503

Fig. 433. Irlbachia cardonae

Fig. 434. Irlbachia pratensis

Fig. 435. Irlbachia tatei

504

G ENTIANACEAE

9. MACROCARPAEA (Griseb.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 94. 1895. —Lisyanthus sect. Macrocarpaea Griseb., Gen. Sp. Gent. 173. 1839 [1838]. —Lisianthus sect. Megacarpaea Benth. in Benth. & Hook. f., Gen. Pl. 2: 814. 1876, orthographic error. Rusbyanthus Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 95. 1895. by Lena Struwe Branched shrubs and small trees (rarely epiphytic) or suffrutescent, perennial herbs (e.g., M. rubra). Stems terete or quadrangular. Leaves petiolate or sessile, elliptic, ovate, or obovate, the base cordate to acute, the apex obtuse to acute, chartaceous to coriaceous, smooth to rugose; interpetiolar lines or ocreae present, often conspicuous; venation often prominent with raised veins below and sunken veins above, pinnate, with arcuate secondary veins. Inflorescence terminal, a lax, many-flowered dichasium; bracts usually leaf-like. Flowers pedicellate, actinomorphic to slightly zygomorphic with anthers aggregated in the upper part of the corolla mouth and the style positioned below, 5-merous, erect. Calyx campanulate, divided 1/4–4/5 of the length, thick and woody or coriaceous, persistent in fruit, the lobes elliptic to ovate (rarely leaf-like M. bracteata), with a glandular dorsal keel on each lobe, hyaline-margined, the apex obtuse; corolla funnelform, often broadly so, green, yellow, or white (tubular and red in M. rubra), thick and fleshy, deciduous in fruit, the lobes oblong to ovate, the lobe apex obtuse, the corolla bud apex rounded. Stamens inserted in the middle of the corolla tube; filaments of unequal or subequal length; anthers linear to oblong, sagittate, recurved after anthesis, usually with a small sterile apical appendage; pollen in monads, the exine reticulate or verrucose. Ovary with a disk around the base; style long, slender, persistent in fruit; stigma bilamellate with broad lobes. Capsule ovoid to elliptic, woody, dehiscing medially, erect or nodding, Seeds angular, sometimes flattened, not winged, the testa cells rectangular, dome-like, or concave, sometimes with band-like thickenings. Costa Rica, Panama, Jamaica, Cuba, Dominican Republic, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; ca. 50 species, at least 6 species in Venezuela, 5 of these in the flora area. Macrocarpaea belongs to the Irlbachia complex (Struwe et al. 1997). All species of Macrocarpaea in the Venezuelan Guayana are endemic to tepuis of this region and show allopatric distributions; M. piresii and M. neblina occur in different altitudinal zones on Sierra de la Neblina. Key to the Species of Macrocarpaea 1. 1. 2(1). 2. 3(1). 3.

Leaves with veins not conspicuously impressed on upper surface, not rugose (papillose in M. autanae) ................................................................. 2 Leaves with veins strongly impressed on upper surface, rugose (at least lower leaves) .......................................................................................... 3 Leaf surface smooth; calyx 9–13 mm long; calyx lobes 6–8 mm long ................................................................................................... M. neblinae Leaf surface papillose; calyx 7–9 mm long; calyx lobes 3–4 mm long .................................................................................................... M. autanae Calyx, pedicels, and leaves glabrous; calyx 16–19 mm long ............ M. piresii Calyx, pedicels, and leaves puberulent; calyx 10–16 mm long ................ 4

Macrocarpaea 505

5 mm 3 cm

1 cm

Fig. 436. Macrocarpaea marahuacae

506

G ENTIANACEAE

Fig. 437. Macrocarpaea neblinae

Fig. 438. Macrocarpaea rugosa

Neblinantha

4(3). 4.

507

Leaf base cordate; calyx lobes triangular-ovate, keeled; petioles not winged .......................................................................................... M. rugosa Leaf base attenuate to obtuse; calyx lobes oblong, not keeled; petioles slightly winged ................................................................... M. marahuacae

Macrocarpaea autanae Weaver, Mem. New York Bot. Gard. 32: 336. 1981. Shrub to 4 m tall, glabrous; leaves petiolate, 190–250 mm × 120–145 mm, ellipticoblong to oblong, papillose and coriaceous, the base obtuse, the apex obtuse; calyx 7–9 mm long, the lobes 3–4 mm long, triangularovate, slightly keeled; corollas dull green, ca. 40 mm long, the lobes ca. 10 mm long; fruits unknown. Upland forests on tepui summits, ca. 1300 m; Amazonas (Cerro Autana). Endemic. Macrocarpaea marahuacae L. Struwe & V.A. Albert, Harvard Pap. Bot. 3: 182, fig. 1. 1998. Suffrutescent herb or shrub to 1 m tall, puberulent; leaves petiolate, 120–250 × 68– 135 mm, obovate-elliptic, rugose and coriaceous, the base obtuse to attenuate, the apex obtuse; calyx 12–16 mm long, the lobes 6–8 mm long, not keeled; corolla yellow-green, 35–45 mm long, the lobes 8–9 mm long; capsule 17–23 mm long. Rocky open areas on tepui summits, 2500–2700 m; Amazonas (Cerro Marahuaka). Endemic. ŠFig. 436. Macrocarpaea marahuacae is most similar to M. rugosa, an endemic of the Macizo del Chimantá (Bolívar state). Macrocarpaea neblinae Maguire & Steyerm., Mem. New York Bot. Gard. 32: 332, 335, figs. 62A–K, 63, 64. 1981. Suffrutescent herb, shrub, or treelet to 4 m tall, glabrous; leaves petiolate, 45–215 × 18–78 mm, elliptic-ovate, the base attenuate to obtuse, the apex usually acute, sometimes

obtuse; calyx 9–13 mm long, usually minutely verrucose, the lobes 6–8 mm long, not keeled; corolla 25–34 mm long, greenish white, the lobes 7–11 mm long; capsule oblong, 10–18 mm long. Forests on tepui summits, 1700–2000 m; Amazonas (Sierra de la Neblina). Endemic. ŠFig. 437. Macrocarpaea piresii Maguire, Mem. New York Bot. Gard. 32: 338, figs. 62L–M, 66, 67. 1981. Shrub or small tree to 5 m tall, glabrous; leaves petiolate, 90–120 × 60–90 mm, circular-elliptic to broadly ovate, coriaceous and rugose, the base obtuse, the apex obtuse; calyx 16–19 mm long, the lobes ca. 11 mm long, not keeled; corolla ca. 40 mm long, green; fruit unknown. Montane forests, 2400–2600 m; Amazonas (expected). Brazil (Amazonas: Serra da Neblina). Macrocarpaea piresii is known from a single collection in Brazil just across the border from Amazonas state, Venezuela. Macrocarpaea rugosa Steyerm., Bol. Soc. Venez. Ci. Nat. 25: 83, 85, fig. 4. 1963. Branched shrub or tree to 5 m tall, puberulent (very small hairs); leaves petiolate, 60–180 × 22–81 mm, oblong-elliptic, coriaceous and rugose, the base cordate, less often obtuse, the apex obtuse to slightly acute; calyx 10–14 mm long, the lobes 3–7 mm long, (slightly) keeled; corolla 24–33 mm long, dull green, the lobes 5–7 × 4–6 mm, elliptic, the apex obtuse; capsule ca. 16 mm long. Tepuisummit forests, 2100–2400 m; Bolívar (Macizo del Chimantá). Endemic. ŠFig. 438.

10. NEBLINANTHA Maguire, Phytologia 57: 311. 1985. by Lena Struwe Suffrutescent herbs or shrubs. Stems (sub)quadrangular, winged. Leaves sessile or subsessile (petiole to 3 mm long), ovate-lanceolate, coriaceous, the base attenuate-acute or obtuse, the apex acute; margin flat or recurved; interpetiolar line present (sometimes inconspicuous); venation with 1(2) pairs of basally divergent, arcuate secondary veins (invisible in N. parvifolia). Flowers solitary, appearing axillary (probably a single terminal flower with 1 or 2 branches developing in the axils of

508

G ENTIANACEAE

Fig. 439. Neblinantha neblinae

Neurotheca 509

the uppermost leaf pair, i.e., the bracts of the flower; N. neblinae), or terminal on short, leafy shoots (N. parvifolia). Flowers pedicellate, 5-merous, actinomorphic, erect or nodding. Calyx tubular, divided 4/5–7/8 its length, thin, persistent in fruit, the lobes narrowly triangular, long-acuminate, hyaline-margined, with a low dorsal keel; corolla salverform or narrowly funnelform, deep rose-pink or coral, thin, deciduous in fruit, the lobes circular or triangular, slightly spreading, the apex obtuse to acuminate, the corolla bud apex (slightly) tapering. Stamens inserted in the corolla tube; filaments of slightly unequal length, sometimes bent 180 degrees at apex; anthers ovate-oblong, sagittate, erect(?) after anthesis, with a small sterile apical appendage; pollen in tetrads, the exine reticulate. Ovary without(?) a disk at the base; style long, slender, persistent in fruit; stigma bilamellate, the lobes narrow. Capsule elliptic, dehiscing apically, erect or nodding. Seeds angular, not winged, the testa morphology unknown. Endemic to the Guayana Shield on the Venezuelan-Brazilian frontier (Sierra de la Neblina); 2 species, both in the flora area. Neblinantha is related to the genera of the Irlbachia complex. Key to the Species of Neblinantha 1.

1.

Flowers nodding, appearing axillary in the upper leaf axils; leaves 20– 36 mm long, with acute bases, venation prominent with 1 pair of arcuate secondary veins ................................................................... N. neblinae Flowers erect (rarely nodding), terminal on short leafy shoots; leaves 8– 15 mm long, with obtuse bases, venation invisible ............... N. parvifolia

Neblinantha neblinae Maguire, Phytologia 57: 311. 1985. Suffrutescent herb or subligneous shrub to 1.5 cm tall; leaves 20–36 × 10–15 mm, the base attenuate-acute; calyx tubular, 12–19 mm long, the lobes 8–13 mm long; corolla 35– 40 mm long, salverform, deep rose-pink, the lobes 7–10 mm long, circular, the apex obtuse to slightly mucronate; capsule ca. 10 mm long. Tepui meadows and upper slope forests, 2200– 2300 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). ŠFig. 439.

Neblinantha parvifolia Maguire, Phytologia 57: 311. 1985. Branched, suffrutescent herb to 60 cm tall; leaves 7–15 × 3–6 mm, the base obtuse, nearly amplexicaul; calyx tubular, 8–11 mm long, the lobes ca. 7 mm long; corolla 29–32 mm long, narrowly funnelform, coral, the lobes 6–12 mm long, triangular, the apex acuminate; capsule 6–10 mm long. Upper tepui-slope forests, 2200–2300 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina).

11. NEUROTHECA Salisb. ex Benth. & Hook. f., Gen. Pl. 2: 812. 1876. Octopleura Spruce ex Progel in Mart., Fl. Bras. 6(1): 212. 1865, non Octopleura Griseb. 1860. by Lena Struwe Simple or branched annual herbs. Stems quadrangular, with decurrent ridges or wings. Leaves sessile, linear to narrowly lanceolate, the basal leaves elliptic to spatulate, chartaceous, the base attenuate, the apex acute; interpetiolar line present, inconspicuous; the margins flat; venation with midvein only, inconspicuous. Inflorescence terminal, a lax or congested raceme (usually with a terminal flower) or rarely flowers appearing 1–3 in the leaf axils along the stem; bracts leaf-like, small. Flowers sessile, actinomorphic, 4-merous, horizontal. Calyx tubular, divided ca. 1/3 of

510

G ENTIANACEAE

its length, with 8(–12) distinct longitudinal veins plus several transverse veins on the lobes, thin, persistent in fruit, the lobes triangular, the apex acute; corolla tubular to narrowly funnelform, thin, deciduous, the lobes triangular to ovate, the apex acute, the corolla bud apex tapering. Stamens inserted below the middle of the corolla tube; filaments of subequal length; anthers oblong-ellipsoid, straight after anthesis, without sterile apical appendages; pollen in monads, the exine morphology unknown. Ovary without a disk around the base; style slender, deciduous in fruit; stigma bilobed, lobes small. Capsule oblong-ovoid, thin-walled, dehiscing apically, enclosed by the persistent calyx, horizontal to erect; seeds globose, the testa cells reticulate, smooth. Venezuela, Guyana, Suriname, French Guiana, Brazil, tropical Africa (Mozambique, Zaire, Angola, South Africa), western Madagascar; 3 species, 1 in Venezuela. Neurotheca is closely related to Enicostema, and to the African genera Djaloniella, Faroa, and Pycnosphaera (Struwe et al. 1998). Neurotheca loeselioides (Spruce ex Progel) Baill., Hist. Pl. 10: 138. 1888. —Octopleura loeselioides Spruce ex Progel in Mart., Fl. Bras. 6(1): 212, t. 58. 1865. Weak annual herb to 30 cm tall; leaves sessile, 3–18 × 1–8 mm; calyx 5–7 mm long, the lobes 1–2 mm long; corolla 3–7 mm long, blue (rarely pink or white), the lobes 1–2 mm long; capsule 2–3 mm long. Wet savannas (often sandy), on rocks in open places, 100– 1400 m; Amazonas (base of Cerro Yutajé). Guyana (Pakaraima Mountains), Suriname (Tafelberg), French Guiana, Brazil (Amapá, Amazonas, Mato Grosso, Pará, Rondônia), tropical Africa, western Madagascar. ŠFig. 440. Fig. 440. Neurotheca loeselioides

12. POTALIA Aubl., Hist. Pl. Guiane 394, t. 151. 1775. Nicandra Schreb., Gen. Pl. 1: 283. 1789, non Adans. 1763. by Lena Struwe and Victor A. Albert Monopodial or sparsely branched shrubs or trees. Stems terete with leaves crowded at branch apices. Leaves petiolate, narrowly elliptic to obovate, widest at middle or above, often very large, the base long-attenuate, the apex obtuse to acute (rarely truncate and sometimes with a short acumen); interpetiolar ocreae present; margins flat or slightly recurved; venation pinnate with many straight or slightly arcuate secondary veins. Inflorescence terminal, a dichasial cyme, to 50-flowered, the branches bright yellow or orange; bracts and floral bracts scale-like. Flowers pedicellate, actinomorphic, erect. Calyx 4-merous, divided to the base, yellow, whit-

Potalia 511

ish, or orange, coriaceous, persistent in fruit, the lobes decussate, convex, often ruptured by the fruit, usually hyaline-margined; corolla 8–10-merous, tubular with a narrow lower part and a wider upper part, white, yellow, or green, thick and fleshy, deciduous in fruit, the lobes lanceolate, sometimes with green margins, erect or slightly spreading. Stamens 8–10, inserted in the middle of the corolla tube, included in corolla; filaments very short, of equal length, completely fused into a staminal tube; anthers linear, sagittate, often with a small sterile apical appendage; pollen in monads (tetrads in P. maguireorum), the exine smooth, porate. Ovary with a glandular disk around the base, upper part sterile; style short; stigma capitate. Berry fleshy, globose, turbinate, ovoid, to obovoid, green, the apex acute, truncate to obtuse, largely consisting of sterile tissue (best seen in longitudinal sections), the fruit wall often with a thickened horizontal ring around the middle of the fruit (e.g., P. resinifera). Seeds elliptic, flattened, not winged, the testa cells reticulate. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 7 species, 4 in Venezuela, 3 of these in the flora area. Potalia was earlier considered to be a monotypic genus, only containing P. amara, but new revisionary studies have shown that several species exist and that P. amara is restricted to Guyana, Suriname, French Guiana, and Brazil (Amapá). Potalia is most closely related to the African-Malagasy genus Anthocleista and the Asian-Australian-Pacific genus Fagraea. Potalia is used widely in South America against snake bites, fevers, and inflammations. Flowers of P. elegans have been reported to be visited by hummingbirds. Key to the Species of Potalia 1. 1. 2(1).

2.

Leaves elliptic, broadest in the middle; corolla dark green ..... P. maguireorum Leaves obovate-spatulate, broadest toward the apex; corolla white, green, green with yellow margins, or yellowish orange .................................. 2 Inflorescence > 15 cm long, with conspicuously elongated basal branches; pericarp smooth, without a thickened ring; fruit in cross section without a prominent sterile apex; leaves usually very thick and coriaceous ...................................................................................................... P. elegans Inflorescence < 15 cm long, the basal branches not conspicuously elongated; pericarp with a thickened ring around the middle; fruit in cross section with a prominent sterile apex; leaves usually thin and chartaceous ............................................................................... P. resinifera

Potalia elegans L. Struwe & V.A. Albert, Harvard Pap. Bot. 3: 185, fig. 2. 1998. —Hoja de gavilán, Palo de lagarto mato, Palo de mato, Palo de rana, Palo de temblador, Temblador. Shrub or tree to 12 m tall; leaves 60– 100 × 10.5–21 cm, oblanceolate to obovate, widest above middle, coriaceous, the base attenuate, the apex acute, the margin thickened, sometimes slightly revolute; inflorescence (145–)170–290 mm long, longstalked, basal branches > (55–)100 mm

long; calyx lobes 7–9 mm long, yellow to white; corolla 11–15 mm, yellow and green, the lobes 6–8 mm long; berry 10–16 mm long, slightly longer than calyx, the apex acute. White-sand savannas, forests, 50– 200 m; Amazonas (Caño Caname, Capibara, Cerro Yapacana, La Esmeralda, Maroa, lower Río Baría, Río Casiquiare and tributaries, Río Cunucunuma, Río Pasimoni, Río Sipapo, San Carlos de Río Negro, Yavita). Colombia (Amazonas, Vaupés). ŠFig. 441.

512

G ENTIANACEAE

5 mm

3 mm

3 cm

1 cm

3 mm

Fig. 441. Potalia elegans

3 cm

Potalia 513

1 cm

3 cm

5 mm

2 mm

1 cm

2 mm

Fig. 442. Potalia maguireorum

514

G ENTIANACEAE

5 mm

3 cm

3 mm

1 cm

3 mm

1 cm

3 mm

Fig. 443. Potalia resinifera

Rogersonanthus 515

Potalia maguireorum L. Struwe & V.A. Albert, Harvard Pap. Bot. 3: 189, fig. 3. 1998. Shrub or treelet to 3 m tall; leaves 18–36 × 2.8–7.9 cm, narrowly elliptic to oblanceolate, widest at or slightly above middle, coriaceous, the base attenuate, the apex bluntly acute. the margins slightly revolute; inflorescence 32–135 mm long; calyx lobes 4– 5 mm long, yellow; corolla 7–8 mm long, dark green, the lobes 2–4 mm long; berry 6–10 mm long, at least twice as long as calyx, the pericarp without a thickened ring, apex acute. Savannas and forests on white-sand, 100–200 m; Amazonas (Río Casiquiare and tributaries toward Sierra de la Neblina). Brazil (Amazonas: upper Rio Negro). ŠFig. 442.

Potalia resinifera Mart., Nov. Gen. Sp. Pl. 2: 90, t. 170. 1826 [1827]. —Curarina. Shrub or treelet to 2 m tall; leaves 27–80 × 8.5–19 cm, broadly oblanceolate to obovate, widest above middle, chartaceous (rarely coriaceous), the base long-attenuate, the apex acute, the margins flat to slightly revolute; inflorescence 40–55 mm long; calyx lobes 4–7 mm long, yellow, usually recurved in fruit; corolla 13–20 mm long, yellow and green, the lobes ca. 5–10 mm long; berry 8–15 mm long, twice as long as calyx, the pericarp with a thickened ring, apex blunt. Savannas, forests, 100–200 m; Amazonas (Río Cataniapo, Río Negro, Río Yureba, between San Fernando de Atabapo and Santa Bárbara). Widespread in Colombia, Venezuela, Brazil, Peru, Ecuador, Bolivia. ŠFig. 443.

13. ROGERSONANTHUS Maguire & B.M. Boom, Mem. New York. Bot. Gard. 51: 3. 1989. by Lena Struwe Small perennial herbs, shrubs, or trees to 5 m tall. Stems quadrangular or terete, sometimes narrowly winged. Leaves short-petiolate or sessile, elliptic, lanceolate, ovate, to obovate, thick, subsucculent, coriaceous, leathery, the base attenuate, the apex acute to obtuse; interpetiolar line present; margins flat or recurved; venation pinnate, inconspicuous, with 1–several pairs of basally divergent arcuate secondary veins. Inflorescence terminal, a small dichasium, 1–12-flowered; bracts scale-like. Flowers pedicellate, actinomorphic, 5-merous, erect, horizontal, or nodding. Calyx campanulate, divided ca. 3/4 its length, thick and leathery (thinner in R. coccineus), persistent in fruit, the lobes elliptic, with a dorsal keel, the apex obtuse to slightly acute; corolla broadly funnelform, campanulate, or narrowly salverform, green, yellow, red, or orange, thick and leathery (thinner in R. coccineus), deciduous in fruit, the lobes circular to oblong, the apex obtuse, the corolla bud apex rounded. Stamens inserted in the lower half of the corolla tube, included in corolla; filaments of unequal length; anthers lanceolate, sagittate, recurved after anthesis; pollen in tetrads with reticulate exine. Ovary sessile, bilocular, with glandular area at the base of the ovary; style long, slender, basal part persistent in fruit; stigma bilamellate. Capsule elliptic, leathery, dehiscing medially or apically, sometimes nodding. Seeds angular, not winged, the testa cells concave with band-like thickenings (R. quelchii). Southern Venezuela, Trinidad, Guyana, Brazil(?) (restricted to tepuis except for Trinidad); 3 species, all in the flora area. Rogersonanthus belongs to the Irlbachia complex and has formerly been included in Macrocarpaea, but differs mainly in having pollen in tetrads (Struwe et al. 1997). It also shows morphological similaritites with the tetrad-bearing genus Chelonanthus.

516

G ENTIANACEAE

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

2.

Corolla red to orange, salverform; small herb to 60 cm tall ....... R. coccineus Corolla green, funnelform to campanulate; shrub or tree to 5 m tall ...... 2 Leaves broadest in the middle, (2–)3–5 times longer than wide, elliptic, the blade (39–)50–120(–143) mm long, the apex long-acute or acuminate ............................................................................................ R. arboreus Leaves broadest above middle, 1–2.5 times longer than wide, broadly elliptic-oblanceolate to obovate, 19–50(–70) mm long, the apex obtuse to bluntly acute ............................................................................... R. quelchii

Rogersonanthus arboreus (Britton) Maguire & B.M. Boom, Mem. New York. Bot. Gard. 51: 8. 1989. —Chelonanthus arboreus Britton, Bull. Dept. Agric. Trinidad & Tobago 19: 230. 1922. —Macrocarpaea arborea (Britton) Ewan, Contr. U.S. Natl. Herb. 29: 221. 1948. Calolisianthus tepuiensis Gleason, Brittonia 3: 188. 1939. —Macrocarpaea tepuiensis (Gleason) Steyerm., Fieldiana, Bot. 28: 498. 1953. —Rogersonanthus tepuiensis (Gleason) Maguire & B.M. Boom, Mem. New York. Bot. Gard. 51: 4. 1989. Macrocarpaea cerronis Ewan, Contr. U.S. Natl. Herb. 29: 223, pl. 2. 1948. Macrocarpaea salicifolia Ewan, Contr. U.S. Natl. Herb. 29: 224, pl. 3. 1948. Shrub or tree to 5 m tall; leaves petiolate, (39–)50–120(–186) × (16–)20–45(– 69) mm, elliptic; calyx 6–10 mm long, the lobes 4–8 mm long; corolla 26–45 mm long, green to yellowish green or greenish white, the lobes 9–12 mm long; capsule 14–19 mm long. Savannas and rocky areas on tepui summits and escarpments, 1200– 2600 m; Bolívar (Amaruay-tepui, Auyántepui, Cerro Guaiquinima, Cerro Upuigma, Gran Sabana, Kamarkawarai-tepui, Macizo del Chimantá [Apacará-tepui, Toronó-tepui], Murisipán-tepui, Ptari-tepui, Sororopán-tepui), Amazonas (Cerro Marahuaka). Trinidad. ŠFig. 445. Rogersonanthus arboreus and R. tepuiensis were considered two separate species by Maguire & Boom (1989). However, there are no consistent morphological differences between these two disjunct taxa and they are united here. A small-leaved form of Rogerso-

nanthus arboreus similar to R. quelchii is present on Auyán-tepui. Rogersonanthus coccineus Steyerm. ex L. Struwe, S. Nilsson & V.A. Albert, Harvard Pap. Bot. 3: 191, figs. 4, 5. 1998. Perennial herb 20–60 cm tall, sometimes with basal vegetative shoots; leaves sessile to subsessile, 16–28 × 6–9 mm, elliptic; calyx 6–10 mm long, the lobes 5–6 mm long; corolla 24–27 mm long, red to orange, salverform, the lobes 6–8 mm long; capsule 11–12 mm long. Meadows and bogs on tepui summits, 1900–2300 m; Amazonas (Sierra de la Neblina). Endemic. ŠFig. 446. Rogersonanthus quelchii (N.E. Br.) Maguire & B.M. Boom, Mem. New York. Bot. Gard. 51: 4. 1989. —Lisianthus quelchii N.E. Br., Trans. Linn. Soc. London, Bot. 6: 50, pl. 9, figs. 6–9. 1901. —Symbolanthus quelchii (N.E. Br.) Gleason, Bull. Torrey Bot. Club 56: 402. 1929. —Macrocarpaea quelchii (N.E. Br.) Ewan, Contr. U.S. Natl. Herb. 29: 233. 1948. —Irlbachia quelchii (N.E. Br.) Maas, Proc. Kon. Ned. Akad. Wetensch., C. 88: 410. 1985. Small tree or shrub 0.5–5 m tall; leaves shortly petiolate, 19–70 × 14–40 mm, broadly elliptic-oblanceolate to obovate; calyx 6–11 mm long, the lobes 4–8 mm long, corolla 21–44 mm long, green to yellowish green, funnelform, the lobes 7–11 mm long; capsule 11–15 mm long. Tepui meadows or forests on upper talus slopes, 1900–2800 m; Bolívar (Ilú-tepui, Kukenán-tepui, Roraimatepui, Yuruaní-tepui). Guyana (Cerro Upuima). ŠFig. 444.

Rogersonanthus 517

Fig. 444. Rogersonanthus quelchii

5 mm

2 mm

3 cm 1 cm

Fig. 445. Rogersonanthus arboreus

Fig. 446. Rogersonanthus coccineus

518

G ENTIANACEAE

Flower bud

Sepal

Cross section, dorsal, and frontal view of leaves

Fig. 447. Saccifolium bandeirae

Schultesia 519

14. SACCIFOLIUM Maguire & Pires, Mem. New York Bot. Gard. 29: 242. 1978. by Lena Struwe and Victor A. Albert Small branched shrubs. Branches with thick, corky, and ridged bark, with the leaves very tightly crowded at apices. Leaves possibly alternate or opposite and compressed, sessile, obovate, saccate, coriaceous to subsucculent, the base attenuate, the apex broadly obtuse; interpetiolar lines or ocreae absent; venation prominent with 3 parallel basal veins and secondary and tertiary reticulate venation. Flowers solitary in the leaf axils, sessile, (4)5-merous, actinomorphic, erect(?). Calyx divided nearly to the base, the tube about 1 mm long, the lobes lanceolate, with prominent glandular areas at the inner bases of the sepals; corolla tubular, constrained at the apex, white, thin, deciduous, the lobes ovate, imbricate in bud. Stamens inserted close to the base of the corolla tube; filaments of equal length; anthers linear, sagittate, latrorse at base and introrse near apex, probably erect after anthesis, with a prominent connective, with sterile apical appendages; pollen in monads with porate exine. Ovary without a glandular disk at the base; style slender, persistent in fruit; stigma small, simple, slightly bilobed. Fruit globose, probably a dry capsule (no mature fruits known). Seeds (immature) angular, the testa cells reticulate and smooth. Endemic to the Guayana Shield on the Venezuelan-Brazilian frontier (Sierra de la Neblina); 1 species. Saccifolium was formerly the only member of Saccifoliaceae, but recent phylogenetic studies have shown that it is nested within Gentianaceae (Struwe et al. 1998). Saccifolium bandeirae Maguire & Pires, Mem. New York Bot. Gard. 29: 242, 244, figs. 111–120. 1978. Branched subshrub to 60 cm tall; leaves 11–22 × 4–6 mm; calyx lobes 5–6 × 1–2 mm;

corolla 12–14 mm long, the lobes 1–2 mm long. Rock crevices and cliffs, 2700–3000 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Fig. 447.

15. SCHULTESIA Mart., Nov. Gen. Sp. Pl. 2: 103. 1826 [1827], nom. cons., non Schultesia Spreng. 1815, non Schultesia Schrad. 1821, non Schultesia Roth 1827. Reichertia H. Karst., Bot. Zeitung (Berlin) 5: 397. 1848. by Lena Struwe Annual herbs to 60 cm tall. Stems usually terete basally, becoming quadrangular and winged higher up. Leaves sessile, linear, lanceolate, ovate, to obovate, chartaceous, the base obtuse to attenuate, sometimes amplexicaul, the apex acute to short-acuminate or rarely obtuse; interpetiolar line present, inconspicuous; margins flat; venation with inconspicuous secondary veins. Inflorescence terminal, a dichasial cyme, 1–ca. 25-flowered; bracts leaf-like. Flowers sessile or pedicellate, 4merous (5-merous in S. pachyphylla), actinomorphic, erect to horizontal. Calyx tubular or urceolate, thin, persistent in fruit, the lobes narrowly triangular to ovate, keeled or broadly winged; corolla salver- to funnelform, constricted at the throat, thin, persistent in fruit, the lobes usually ovate to triangular, the corolla bud apex tapering. Stamens inserted in the corolla tube; filaments of equal length, sometimes winged at the base; anthers oblong, erect after anthesis, with inconspicuous sterile

520

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Fig. 448. Schultesia benthamiana

Fig. 449. Schultesia brachyptera

Schultesia 521

apical appendages; pollen in tetrads, the exine reticulate and microreticulate. Ovary with or without a disk at base; style slender, long, deciduous in fruit; stigma bilamellate. Capsule ovoid, dehiscing apically, erect to horizontal. Seeds and testa cell morphology not investigated. Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama Hispaniola, Cuba, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil (most species in eastern Brazil), Paraguay, one species also in western tropical Africa (Senegal); ca. 8–10 species, 5 in Venezuela, all in the flora area. Schultesia is closely related to Coutoubea, with which it shares very similar pollen tetrads. Key to the Species of Schultesia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

Corollas (26–)30–50 mm long; calyx 20–29 mm long .............. S. brachyptera Corollas < 28 mm long; calyx < 20 mm long ............................................. 2 Corollas 8–10 mm long; calyx 5–7 mm long, the lobes 1 mm long ............. ................................................................................................ S. subcrenata Corollas > 10 mm long; calyx 8–20 mm long, the lobes 5–10 mm long ................................................................................................................ 3 Leaf bases obtuse (rarely acute in very narrow leaves); leaves ovate to lanceolate, 3–10 mm wide ..................................................... S. guianensis Leaf bases acute to attenuate; leaves linear to narrowly lanceolate; 1–3 mm wide .......................................................................................... 4 Leaf apex acute; inflorescences > 2-flowered ........................ S. benthamiana Leaf apex obtuse; inflorescences 1(2)-flowered ............................ S. pohliana

Schultesia benthamiana Klotzsch ex Griseb., Linnaea 22: 34. 1849. Schultesia schomburgkiana Progel in Mart., Fl. Bras. 6(1): 206. 1865. Sparsely branched herb 10–30 cm tall; leaves 5–33 × 1–3 mm, linear to narrowly lanceolate, the base narrow and attenuate, the apex acute; calyx 8–20 mm long, divided ca. 1/2 of the length, with dorsal wings to 1 mm tall, the lobes 8–10 mm long, linear; corolla 18–26 mm long, yellow, rose, pale purple, or brownish, the lobes 6–12 mm long; capsule 6–8 mm long. Low and middle elevation savannas, riverbanks, 100–800 m; Bolívar (upper Río Caroní, Río Parguaza and vicinity). Colombia (Vichada), Guyana, Brazil (Roraima). ŠFig. 448. Schultesia brachyptera Cham., Linnaea 8: 8. 1833. Schultesia heterophylla Miq., Linnaea, 19: 136. 1847. —Schultesia brachyptera f. heterophylla (Miq.) Jonker, Recueil Trav. Bot. Néerl. 32: 250. 1936. Single-stemmed or rarely branched herb 15–80 cm tall; leaves 11–60(–80) × 2–5(–8)

mm (basal leaves often smaller and spatulate-obovate), linear to narrowly lanceolate, the base attenuate, the apex acute; calyx 20– 29 mm long, divided 1/3–1/2 of the length, mostly with conspicuous dorsal wings to 3 mm tall, the lobes 8–17 mm long, linear; corolla 26–50 mm long, narrowly funnelform, light purple, pink, or rose, the lobes 8–12 mm long, broadly elliptic to oblong; capsule 12– 16 mm long. Moist savannas, 50–800 m; Bolívar (upper Río Caroní), Amazonas (vicinity of Puerto Ayacucho). Aragua, Guárico; Panama(?), Cuba(?), Hispaniola(?), Colombia, Guyana, Suriname, French Guiana, Brazil. ŠFig. 449. It is uncertain whether the Central American and West Indian populations belong to this species, and some authors have segregated Schultesia heterophylla from S. brachyptera. Schultesia guianensis (Aubl.) Malme, Ark. Bot. 3(12): 9. 1904. —Exacum guianense Aubl., Hist. Pl. Guiane 68. 1775. —Miniatuia. Branched herb 3–24 cm tall; leaves 6–30 ×

522

G ENTIANACEAE

3–10 mm, ovate (rarely elliptic), base obtuse, apex acute; calyx 8–17 mm long, divided ca. 1/3 of the length, with prominent dorsal wings to 2 mm tall, the lobes 4–7 mm long, linear; corolla 8–20 mm long, salverform, purple, pink, orange, to yellow or yellow-brown, the lobes 5–6 mm long, ovate; capsule 8–10 mm long. Savannas (often on sand), river banks, 0–300 m; Bolívar (lowlands in the vicinity of Río Orinoco). Apure, Guárico, Táchira; Guatemala, Belize, Honduras, Costa Rica, Panama, Cuba, Guyana, Suriname, French Guiana, Brazil. Schultesia guianensis is a variable species that is similar to S. benthamiana but with smaller flowers, usually pink corollas, small bracts directly below the flowers, and more broadly ovate leaves. Schultesia pohliana Progel in Mart., Fl. Bras. 6(1): 205. 1865. Simple-stemmed weak herb to 30 cm tall; leaves 5–20 × ca. 1–3 mm, linear to narrowly

elliptic, the base attenuate, the apex obtuse (rarely acutish); calyx 9–20 mm long, divided ca. 3/5 of the length, with narrow dorsal wings 1(–2) mm tall, the lobes linear; corolla 10–25 mm long, salver- to funnelform, yellow, orange, purplish, brownish, or yellowish pink; capsule unknown. Savannas around granitic outcrops, 100–400 m; Bolívar (near Ciudad Piar), Amazonas (vicinity of Puerto Ayacucho). Guárico; Panama, Brazil. Schultesia subcrenata Klotzsch ex Griseb., Linnaea 22: 34. 1849. Branched herb to 6 cm tall; leaves 6–9 × 1–2 mm, linear, the base attenuate, the apex acute; calyx 6–8 mm long, divided ca. 2/3 its length, with prominent dorsal wings to 2 mm tall, the lobes 4–6 mm long, narrowly triangular; corolla 8–10 mm long, tubular, yellow, the lobes 2–4 mm long, elliptic; capsule 6–7 mm long. Tepui meadows, ca. 2700 m; Bolívar (Roraima-tepui). Guyana (Mount Roraima, Rupunini Savannas), Brazil.

16. SIPAPOANTHA Maguire & B.M. Boom, Mem. New York Bot. Gard. 51: 23, 25, figs. 17, 18. 1989. by Lena Struwe Unbranched annual herbs. Stems quadrangular, often with 4 narrow wings. Leaves mainly on the lower half of the stem, smaller toward the inflorescence, sessile, obovate (lower leaves), broadly elliptic (upper leaves), very thick and coriaceous, the base narrowly attenuate, the apex obtuse; interpetiolar line present; margin strongly revolute; venation invisible except for midvein. Inflorescence terminal, cymose, with 1–3 monochasial branches, 1–7-flowered; bracts and floral bracts similar to small leaves or scale-like. Flowers pedicellate, 5-merous, slightly zygomorphic (anthers aggregated in upper part of corolla mouth and style bent), erect to nodding. Calyx campanulate, divided ca. 2/3 of the length, thick and coriaceous, persistent, the lobes oblong, obtuse, with a thickened dorsal ridge, obtuse; corolla showy and large, funnelform, dark blue to purple, thin, deciduous in fruit, the lobes elliptic to circular, obtuse, the corolla bud apex rounded. Stamens inserted very close to the base of the corolla tube; filaments of unequal length, widened at the base; anthers lanceolate, sagittate, straight after anthesis, versatile, with sterile apical appendages; pollen in tetrads, the exine reticulate with small globules. Ovary with a glandular disk below the ovary; style long, slender, deciduous in fruit; stigma bilamellate, the lobes broadly circular. Capsule woody, oblong, dehiscing medially, nodding. Seeds angular, not winged, the testa cells concave with band-like thickenings. Endemic to the Guayana Shield in Venezuela (Cerro Sipapo); 1 species. Sipapoantha ostrina Maguire & B.M. Boom, Mem. New York Bot. Gard. 51: 26, 27, figs. 17, 18. 1989. Herb to 1.5 m tall; leaves (12–)30–75 × (10–)20–40 mm; calyx 8–10 mm long, the

lobes 5–6 mm long; corolla 85–96 mm long, the lobes 25–32 mm long; anthers 7–8 mm long; capsule 17–18 mm long. Tepui meadows, ca. 1500 m; Amazonas (Cerro Sipapo). Endemic. ŠFig. 450.

Sipapoantha 523

Fig. 450. Sipapoantha ostrina

524

G ENTIANACEAE

17. SYMBOLANTHUS G. Don, Gen. Hist. 4: 175. 1837. —Lisianthus sect. Symbolanthus (G. Don) Benth. in Benth. & Hook. f., Gen. Pl. 2: 814. 1876. Leiothamnus Griseb., Gen. Sp. Gent. 205. 1839 [1838]. by Lena Struwe Branched shrubs to small trees (seldom perennial herbs) to 5 m tall. Stems quadrangular and/or winged. Leaves petiolate or sessile, ovate (often broadly so) to elliptic, rarely linear or narrowly ovate, coriaceous to chartaceous, the base usually acute to attenuate, the apex usually acute, but sometimes obtuse; interpetiolar line or ocrea present; margins flat or recurved; venation prominent with a few basally divergent arcuate secondary veins. Inflorescence terminal, rarely axillary, a fewflowered dichasium or flowers solitary; small bracts present at the base of the pedicels. Flowers pedicellate, usually slightly zygomorphic (stamens and style zygomorphic within the slightly bent corolla tube), 5-merous, erect or horizontal. Calyx campanulate, divided 7/8–9/10 (or seldom less) of its length, thick and coriaceous, persistent and slightly enlarged in fruit, the lobes lanceolate to elliptic, with a dorsal glandular area on each lobe, sometimes keeled, hyaline-margined, the lobe apex acute, acuminate, or obtuse; corolla broadly salver- or funnelform, usually pink, magenta, or red, rarely purple, white, yellow, or green, often with white or purple stripes or white areas on the inside of the corolla, the lobes erect, recurved, or reflexed, the apex usually acute to acuminate or obtuse, the corolla bud apex tapering. Stamens inserted in the lower half of corolla tube, with a thin corona (or seldom only a ridge) at the base of the filaments; filaments of unequal length, sometimes sharply bent at apex; anthers oblong, sagittate, recurved after anthesis, with sterile apical appendages; pollen in tetrads, the exine reticulate. Ovary with a glandular disk at the base; style long, slender, only the base persistent in fruit; stigma bilamellate, lobes oblong. Capsule globose to elliptic, leathery, often glossy, nearly berry-like, medially dehiscent or possibly indehiscent, usually nodding. Seeds angular, not winged, the testa cells rectangular, dome-like, sometimes with band-like thickenings. Costa Rica, Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; ca. 20 species, ca. 10 in Venezuela, 7 of these in flora area. Symbolanthus is closely related to Wurdackanthus and is a member of the Irlbachia complex. Species delimitations outside the Guayana Highlands are uncertain and need further study. Key to the Species of Symbolanthus 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(3). 5.

Leaves 4–6 times longer than wide ........................................................... 2 Leaves 1.5–3(–4) times longer than wide .................................................. 3 Leaves linear, with strongly revolute margins ................. S. rosmarinifolius Leaves ovate-lanceolate, margins not strongly revolute ................ S. sessilis Leaves 8–22(–31) mm wide, coriaceous; calyx 12–20(–26) mm long ........ 4 Leaves (20–)30–90 mm wide, chartaceous and thin; calyx (22–)25–40 mm long ......................................................................................................... 5 Leaves (52–)60–124 mm long; petiole 6–12 mm long .............. S. camanensis Leaves 20–60 mm long; petiole 0–5 mm long .............................. S. yaviensis Corollas yellow; leaves sessile (rarely subsessile) ............................ S. aureus Corollas pink, rose, or magenta; leaves petiolate or rarely subsessile .... 6

Symbolanthus 525

6(5). 6.

Calyx 23–28 mm long, the lobes 2/3–3/4 the length of the corolla tube; corolla 50–60 mm long ................................................ S. huachamacariensis Calyx (23–)30–45 mm long, the lobes 1/3–5/8 the length of the corolla tube; corolla 60–95 mm long ........................................................... S. elisabethae

Symbolanthus aureus L. Struwe & V.A. Albert, Harvard Pap. Bot. 3: 194, fig. 6. 1998. Shrub or treelet; leaves sessile or rarely subsessile, 58–102 × 20–38 mm, elliptic to nearly obovate, chartaceous, the base attenuate, the apex acute; petiole 0–5 mm long; calyx 20–28 mm long (enlarged in fruit), the lobes ovate, the apex obtuse; corolla funnelform, yellow, white within, ca. 48 mm long, the lobes oblong, ca. 10 mm long; capsule ca. 30 mm long. Montane forest on tepui slopes, ca. 2500 m; Bolívar (Ilú-tepui). Endemic. ŠFig. 452. This species is known from only two collections and is the only yellow-flowered species of Symbolanthus from the flora area. The yellow to green-flowered Symbolanthus vasculosus (Griseb.) Gilg from the Coastal and Andean Cordilleras of Venezuela and the mountains of Colombia differs in having a much larger corolla (over 60 mm long) and petiolate leaves. Symbolanthus camanensis Maguire & B.M. Boom, Mem. New York Bot. Gard. 51: 17, fig. 9. 1989. Suffrutescent herb or subshrub to 2 m tall; leaves petiolate, (52–)60–90 × 20–31 mm, elliptic, coriaceous, base attenuate with slightly decurrent blade, the apex obtuse; petiole 7–15 mm long; calyx 15–17 mm long, the lobes oblong, the apex acute to obtuse; corolla 60–75 mm long, broadly funnelform, rose-lavender, the lobes 15–23 mm long; capsule unknown. Tepui summit, ca. 2200 m; Amazonas (Cerro Camani). Endemic. This species is only known from two collections. It shows similarities with Symbolanthus yaviensis but differs in having larger leaves with longer petioles. Symbolanthus elisabethae (M.R. Schomb.) Gilg, Bot. Jahrb. Syst. 22: 340. 1896. —Leiothamnus elisabethae M.R. Schomb., Verh. Gartenbau-Vereins Erfurt 5: 18, pl. 155, t. 1. 1847. —Lisianthus elisabethae (M.R. Schomb.) Griseb., Linnaea 22: 40. 1849. —Helia elisabethae

(M.R. Schomb.) Kuntze, Revis. Gen. Pl. 428: 1891. Branched herb or shrub to 3 m tall; leaves petiolate, (35–)90–180(–245) × (16–)50–90 mm, elliptic, chartaceous, the base attenuate, the apex acuminate; petiole 8–27 mm long; calyx (23–)30–45 mm long, the lobes ovate, the apex acuminate, acute to seldom obtuse; corolla 60–90 mm long, red, magenta, or pink, often with white and/or purple stripes inside the corolla and often whitish or greenish basally, the lobes 15–27 mm long; capsule 30–42 mm long. Montane forests and rocky areas on tepui slopes and summits, 900–2800 m; Bolívar (Aparamántepui, Auyán-tepui, Kukenán-tepui, La Escalera, Macizo del Chimantá, Ptari-tepui, Roraima-tepui, Uei-tepui), Amazonas (Sierra Parima). Guyana (Mount Roraima, Merume Mountains, Mount Ayanganna), Brazil (Amazonas: Serra da Neblina). ŠFig. 451. The species most similar to Symbolanthus elisabethae is S. huachamacariensis, which was earlier considered to be a subspecies of this species. They differ in that Symbolanthus elisabethae has larger flowers and larger, more broadly elliptic leaves. Some plants, particularly from the high-altitude summit of Roraima-tepui, have smaller leaves similar to Symbolanthus huachamacariensis but usually larger flowers like S. elisabethae, and they are here tentatively included in S. elisabethae. Symbolanthus elisabethae (and S. huachamacariensis) differs from the red- to pink-flowered S. magnificus Gilg, which is distributed in the Venezuelan Coastal Cordilleras, by having a cylindric corolla tube that barely widens toward the corolla mouth during anthesis as opposed to a broadly widened corolla tube. However, the species shows similarities with S. tricolor Gilg, also red-flowered and from the eastern and coastal Cordilleras but with a straight, not widened corolla tube. Further studies might show that these two species are conspecific. Symbolanthus huachamacariensis Steyerm., Ann. Missouri Bot. Gard. 75: 1083. 1988.

526

G ENTIANACEAE

Fig. 451. Symbolanthus elisabethae

1 cm 3 mm

3 cm

3 cm

3 cm

1 mm Fig. 452. Symbolanthus aureus

Fig. 453. Symbolanthus rosmarinifolius

Tachia 527

Symbolanthus elisabethae subsp. occidentalis Maguire & B.M. Boom, Mem. New York Bot. Gard. 51: 17, figs. 7, 8. 1989. Shrub to 1 m tall; branches quadrangular, winged; leaves petiolate, 94–124 × 32–40 (–44) mm, elliptic, chartaceous, the base acute to attenuate, the apex acute; petiole 6– 12 mm long; calyx 23–28 mm long, the lobes narrowly ovate, the apex acute; corolla 50–60 mm long, magenta-red, throat white with purple stripes, the lobes 16–21 mm long; capsule 20–28 mm long. Tepui summits, 1200– 1700 m; Amazonas (Cerro Huachamacari). Endemic. Symbolanthus huachamacariensis is similar to S. elisabethae and S. tricolor Gilg (the latter from the Venezuelan Andes), but differs in having smaller flowers (see also discussion under S. elisabethae). Future phylogenetic studies in Symbolanthus might show that these three taxa (and perhaps others from outside the flora area) are conspecific. Symbolanthus rosmarinifolius L. Struwe & V.A. Albert, Harvard Pap. Bot. 3: 194, fig. 6. 1998. Subshrub to 1 m tall; leaves sessile, 30–48 × 3–7 mm, linear, coriaceous, the base obtuse, the apex obtuse; calyx ca. 22 mm long, the lobes ca. 20 mm long; corolla unknown; capsule unknown. Tepui summit, ca. 2200 m; Amazonas (Cerro Coro Coro). Endemic. ŠFig. 453. This species is known only from one collection (Huber 12292, NY) and has character-

istic linear and revolute leaves that are reminiscent of Rosmarinus L. (Lamiaceae). Symbolanthus sessilis Steyerm. & Maguire, Mem. New York Bot. Gard. 23: 879. 1972. Subshrub to 1 m tall; branches quadrangular, winged; leaves sessile, 48–62 × 12–18 mm, lanceolate, coriaceous, ascending, the base attenuate, the apex acute; calyx 20– 26(–30 in fruit) mm long, the lobes ovate, the apex obtuse to acute; corolla ca. 75 mm long, deep rose-red, the lobes 15–20 mm long; capsule 23–25 mm long. Tepui meadows, 1900– 2100 m; Bolívar (Cerro Jaua). Endemic. Symbolanthus yaviensis Steyerm., Ann. Missouri Bot. Gard. 75: 1084. 1988. Symbolanthus yutajensis Maguire & B.M. Boom, Mem. New York. Bot. Gard. 51: 17–18, fig. 9. 1989. Suffrutescent herb or shrub to 2 m tall; branches acutely quadrangular, winged; leaves subsessile, 20–60 × 8–22 mm, elliptic, the base acute to attenuate, the apex obtuse to slightly acute; petiole 0–5 mm; calyx 14– 26 mm long, the lobes lanceolate to ovate, the apex acute; corolla 51–65 mm long, pink to rose and sometimes with a white, yellowish, or greenish basal part and white stripes or pinkish inside, broadly to narrowly funnelform and narrow at base, curved, the lobes broadly ovate, 13–16 mm long; capsule 14–30 mm long. Tepui-summit forests, 1700– 2100 m; Bolívar (Cerro Guanay), Amazonas (Cerro Yaví, Cerro Yutajé). Endemic.

18. TACHIA Aubl., Hist. Pl. Guiane 75. 1775. by Lena Struwe Shrubs or small treelets, sometimes single-stemmed. Branches terete or quadrangular and winged, often hollow, green or conspicuously yellow. Leaves petiolate, elliptic to obovate, coriaceous or chartaceous, the base acute to attentuate, the apex often acuminate; interpetiolar ocreae present; margins flat or recurved; venation pinnate with many straight secondary veins or with 1–3 pairs of basally divergent arcuate secondary veins. Flowers axillary, solitary or rarely in pairs, (sub)sessile on a very short branch/cushion in the upper leaf axils, 5-merous, slightly zygomorphic (corolla tube, stamens, and style bent), erect to horizontal; bracts absent. Calyx tubular, yellow, divided 1/3–4/5 of its length, thick and coriaceous, persistent or deciduous in fruit, the lobes elliptic to ovate, with obtuse to acute apices, with a dorsal glandular area, keel, or wing, usually hyaline-margined; corolla tubular to salverform, slightly inflated below the mouth, yellow, cream, orange, white, or greenish,

528

G ENTIANACEAE

thin to slightly fleshy, deciduous or partly persistent, the lobes erect, spreading, or reflexed, the lobe apex acute or apiculate, the corolla bud apex tapering. Stamens inserted close to the base in the corolla tube; filaments of unequal length, sometimes sharply bent at apex; anthers linear-oblong, recurved after anthesis, with sterile apical appendages; pollen in monads, the exine reticulate or smooth, sometimes with globules. Ovary with a glandular disk at the base; style long, slender, deciduous; stigma bilamellate, the lobes broad. Capsule elliptic, woody, horizontal to erect, dehiscing apically, the valves deciduous. Seeds globose, not winged, the testa cells papilla-like. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 species, 3 in Venezuela, all in the flora area. Tachia belongs to the Irlbachia complex but differs from all other genera in that group by having sessile, axillary flowers. Ant nests are common in the hollow stems, and the ants probably feed on the abundant secretions from the colleters (glands in the leaf axils) and glands on the calyx keels. Key to the Species of Tachia 1. 1. 2(1).

2.

Leaves inconspicously pinnately veined with straight secondary veins, coriaceous; corolla 20–38 mm long ........................................... T. grandifolia Leaves conspicuously 3- or 5(7)-veined, the secondary veins basally divergent and arcuate, chartaceous; corolla 52–60 mm long ....................... 2 Calyx broadly winged (wings to 2 mm tall), persistent in fruit, 25–40 mm long, divided 1/3 of the length (not including wing extension which is longer than calyx); calyx lobes 10–20 mm long, very narrow and with an acuminate apex; corolla tube 6 times longer than corolla lobes ....................................................................................... T. schomburgkiana Calyx slightly keeled, not winged, deciduous in fruit, 13–17 mm long, divided 1/2–2/3 of the length; calyx lobes 7–9 mm long, lanceolate with an acute apex; corolla tube 4 times longer than corolla lobes ......... T. gracilis

Tachia gracilis Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 203. 1854. —Cumaroa. Branched shrub to 3 m with slender branches; leaves 65–128(–162) × 18–55 mm, ovate; calyx 13–17 mm long; corolla 52–60 mm long, yellow, the lobes 9–10 mm long; capsule 19–20 mm long. Montane forests, 1000–1500 m; Amazonas (Caño Piedra on Cerro Sipapo massif, Cerro Cuao, Cerro Duida, Cerro Marahuaka, upper Río Ventuari, Sierra Parima). Guyana(?). Tachia grandifolia Maguire & Weaver, J. Arnold Arbor. 56: 109, figs. 1J–Q, 2, 3. 1975. Tall herb, shrub, or treelet to 4 m tall; leaves 105–350 × 26–145 mm, lanceolate, broadly elliptic to obovate; calyx 11–15 mm

long, with dorsal keels, the lobes 7–11 mm long; corolla 20–38 mm long, green, pale green, white, or cream-colored, sometimes with green tips, the lobes 4–10 mm long; capsule 15–20 mm long. Nonflooded lowland forests and montane forests on tepui slopes, 100–1800 m. Venezuela, Brazil (Amazonas, Roraima); 2 varieties, both in the flora area. Key to the Varieties of T. grandifolia 1. Petioles thick, 20–45 mm long, narrowly winged; leaves (180–)205–350 × (72–) 108–145 mm; flowering branches (5–) 10–13 mm thick ........ var. grandifolia 1. Petioles slender, 10–20(–30) mm long, not winged; leaves 105–148(–240) × 26–65 (–78) mm; flowering branches 2–5 mm thick ............................. var. orientalis

Tachia 529

Fig. 454. Tachia schomburgkiana

530

G ENTIANACEAE

T. grandifolia var. grandifolia Tall herb, shrub, or treelet to 4 m tall; corolla lobes 4–6 mm long; stamens included in the corolla. Lowland and montane forests, 100–800 m; Bolívar (El Paují), Amazonas (Río Mawarinuma, Río Negro). Amazonian Brazil. Tachia grandifolia var. grandifolia has the thickest branches and largest leaves in the genus. T. grandifolia var. orientalis Maguire & Weaver, J. Arnold Arbor. 56: 110. 1975. —Cosoiba. Small treelet to 3 m tall; corolla lobes 8– 10 mm long; stamens exserted from corolla mouth. Montane and lower montane forests,

500–1800 m; Bolívar (slopes of Cerro Jaua and Cerro Sarisariñama, Macizo del Chimantá [Toronó-tepui], Río Paragua), Amazonas (slopes of Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, and Sierra Parima). Brazil (Roraima). Tachia schomburgkiana Benth., Hooker’s J. Bot. Kew Gard. Misc. 2: 204. 1854. Shrub to 3 m tall; leaves 95–185 × 30–62 mm, ovate; corolla 50–55 mm long, the tube yellow, the lobes 12–15 mm long, yellow to green; capsule 20–22 mm long. Montane forests, often on white sand or sandstone 400– 1200 m; Bolívar (Gran Sabana). Guyana (Mount Roraima, Pakaraima Mountains). ŠFig. 454.

19. TAPEINOSTEMON Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 194. 1854. Stahelia Jonker, Recueil Trav. Bot. Néerl. 34: 494. 1937. by Lena Struwe Annual or perennial herbs or shrubs. Stems terete, not winged. Leaves sessile or petiolate, chartaceous or coriaceous, ovate, lanceolate, to elliptic, the base obtuse to acute, the apex acute; interpetiolar line present; venation conspicuous with 2– many arcuate secondary veins. Inflorescence terminal, sometimes also with axillary branches, lax, paniculate with numerous dichasial cymes and often very large (capitate in T. sessiliflorum); bracts leaf-like to scale-like. Flowers pedicellate (sessile in T. sessiliflorum), actinomorphic, 5-merous, usually nodding with age (erect or horizontal in T. sessiliflorum). Calyx tubular to campanulate, divided ca. 1/2–2/3 of its length, thin, persistent in fruit, the lobes lanceolate to narrowly triangular, not keeled, narrowly hyaline-margined, the lobe apex acute; corolla funnelform, salverform, or tubular, white, green, yellow, or orange, thin, deciduous in fruit, the lobes triangular to ovate, erect or slightly spreading, the apex acute, the corolla bud apex tapering. Stamens inserted in the middle of the corolla tube; filaments short, of equal length; anthers free or coherent, straight after anthesis, the connective often flattened and prominent, without sterile apical appendages; pollen in monads, the exine reticulate or porate. Ovary with a glandular disk at the base; style slender, persistent or partly deciduous in fruit; stigmas bilobed, small. Capsule ovoid to elliptic, dehiscing apically, nodding (erect or horizontal in T. sessiliflorum). Seed and testa morphology not investigated. Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil; 7 species, 6 in Venezuela, all in the flora area. The only species not present in the flora area, Tapeinostemon zamoranum Steyerm., occurs on the Andean slopes in southern Ecuador and northern Peru. Key to the Species of Tapeinostemon 1. 1.

Inflorescences capitate; herbs < 30 cm tall; corollas erect or horizontal ............................................................................................. T. sessiliflorum Inflorescences diffuse, a lax panicle with cymes; herbs or shrubs > 40 cm tall; most corollas nodding at anthesis ................................................. 2

Tapeinostemon 531

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

Corollas 8–16 mm long; styles 8–10 mm long at anthesis (upper part deciduous in fruit) ...................................................................................... 3 Corollas < 8 mm long; styles to 3 mm long at anthesis ............................ 4 Leaves petiolate, ovate; corollas white ..................................... T. longiflorum Leaves sessile, elliptic-lanceolate; corollas yellow and/or orange, some times with white lobes ................................................................. T. breweri Leaves distinctly petiolate ...................................................... T. spenneroides Leaves sessile or nearly so ......................................................................... 5 Leaves 15–32 × 5–15 mm, smooth, ovate-elliptic, widest at the middle ................................................................................................... T. jauaensis Leaves 40–70 × 20–30 mm, rugose, lanceolate, widest at base .... T. rugosum

Tapeinostemon breweri Steyerm. & Maguire, Acta Bot. Venez. 14: 24. 1984. Suffrutescent herb to 1 m tall; leaves 40– 70 × 15–30 mm, elliptic-lanceolate, coriaceous, conspicuously reticulate and rugose; corolla 10–13 mm long, yellow and/or orange, the lobes sometimes white; style ca. 4 mm long; capsule 7–9 mm long. Open rocky places and tepui meadows, 1400–2700 m; Bolívar (Serranía Marutaní), Amazonas (Cerro Marahuaka). Endemic. ŠFig. 455. Tapeinostemon jauaensis Steyerm. & Maguire, Bol. Soc. Venez. Ci. Nat. 32: 394. 1976. Herb to 50 cm tall; leaves 15–32 × 5–15 mm, ovate-elliptic, subcoriaceous, smooth; corolla 4–5 mm long, yellow and white; style ca. 2 mm long; capsule 3–4 mm long. Tepui meadows, 1700–1800 m; Bolívar (Cerro Jaua). Endemic. Tapeinostemon longiflorum Maguire & Steyerm., Mem. New York Bot. Gard. 32: 362. 1981. Shrub to 2 m tall; leaves 28–135 × 12–53 mm, ovate, slightly coriaceous to chartaceous, smooth; corollas white. Tepui meadows, forests, and streambanks, 1600–2400 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Key to the Varieties of T. longiflorum 1. Corolla 8–10 mm long, the lobes not spreading ....................... var. australe 1. Corolla 12–16 mm long, the lobes often spreading ................. var. longiflorum T. longiflorum var. australe Maguire & Steyerm., Mem. New York Bot. Gard. 32: 362. 1981. 1600–2400 m; Amazonas (southern part

Fig. 455. Tapeinostemon breweri

532

G ENTIANACEAE

Fig. 456. Tapeinostemon longiflorum var. longiflorum

Fig. 457. Tapeinostemon spenneroides

Tetrapollinia 533

of Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). T. longiflorum var. longiflorum 1600–2200 m; Amazonas (northern part of Sierra de la Neblina). Endemic. ŠFig. 456. Tapeinostemon rugosum Maguire & Steyerm., Mem. New York Bot. Gard. 32: 366. 1981. Herb to 1 m tall; leaves 40–70 × 20–30 mm, lanceolate, subcoriaceous, rugose with veins prominently raised on the lower surface; corolla ca. 4 mm long, green at base, white above; style ca. 1 mm long; capsule 2–4 mm long. Tepui-slope forests, 1700–1800 m; Bolívar (Macizo del Chimantá [Toronótepui]). Endemic. Tapeinostemon sessiliflorum (Humb. & Bonpl. ex Schult.) Pruski & S.F. Sm., Brittonia 49: 347. 1997. —Psychotria sessiliflora Humb. & Bonpl. ex Schult. in Roem. & Schult., Syst. Veg. 5: 191. 1819. Tapeinostemon capitatum Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 195. 1854. Tapeinostemon borrerioides Benth. ex Knobl., Bot. Centralbl. 60: 355. 1894. nom. nud.

Herb to 30 cm tall; leaves 20–70 × 10–20 mm, elliptic, chartaceous, smooth; corolla 2– 4 mm long, white; style 1 mm long; capsules 1–2 mm long. Moist caatinga forests on white sand, 100–200 m; Amazonas (Caño Ucata southeast of Síquita, Maroa, Río Casiquiare and tributaries). Colombia (Vaupés), Brazil (Amazonas: upper Rio Negro and Rio Vaupés). Tapeinostemon spenneroides Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 194. 1854. —Stahelia spenneroides (Benth.) Jonker, Recueil Trav. Bot. Néerl. 41: 149. 1948. Stahelia surinamensis Jonker, Recueil Trav. Bot. Néerl. 34: 494. 1937. Tapeinostemon ptariense Steyerm., Lloydia 14: 61. 1951. Annual herb to 3 m tall; leaves 22–115 × 9–41 mm, ovate to lanceolate, chartaceous, smooth; corolla 3–6 mm long, white or green, style 2–3 mm long; capsule 2–5 mm long. Lowland and upland forests, along streams, scrub forests, savannas, 300–2000 m; widespread in Bolívar and Amazonas. Colombia, Guyana, Suriname, Brazil. ŠFig. 457.

20. TETRAPOLLINIA Maguire & B.M. Boom, Mem. New York Bot. Gard. 51: 31. 1989. by Lena Struwe Unbranched annual herbs. Stem terete to quadrangular, narrowly 4-winged. Leaves sessile, evenly distributed along the stem, not aggregated close to the base, linear, rarely narrowly ovate, the base attenuate, the apex acute to seldom obtuse; interpetiolar lines present; margins flat; venation inconspicuous, usually with midvein only. Inflorescence terminal, cymose, with monochasial branches, 1–20flowered; bracts and floral bracts scale-like to linear. Flowers pedicellate to subsessile, actinomorphic, 5-merous, erect to horizontal (rarely nodding). Calyx campanulate, divided ca. 1/2–2/3 its length, with dorsal keels, thin, persistent in fruit, the lobes narrowly triangular, often spreading, the apex acute; corolla funnelform, white, light to dark blue, and light rose to dark purple, the base whitish, the inside of blue corollas sometimes white or blue/white-striped, thin, initally persistent in fruit but falling off before dehiscence of the fruit, the lobes triangular to narrowly triangular, rarely ovate to narrowly ovate, the lobe apex acute to acuminate, the corolla bud apex tapering. Stamens inserted in the lower half of the corolla tube; filaments of unequal length; anthers oblong, erect(?) after anthesis, with sterile apical appendages; pollen in tetrads, the exine spinose. Ovary without a disk at the base; style slender, deciduous in fruit; stigma bilamellate, the lobes linear to narrowly ovate. Capsule ovoid, dehiscing apically, erect to horizontal. Seeds angular, not winged, the testa cells concave without band-like thickenings. Venezuela, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay; 1 species. This monotypic genus belongs to the Irlbachia complex and was segregated from Irlbachia because of its unique pollen exine morphology.

534

G ENTIANACEAE

Tetrapollinia caerulescens (Aubl.) Maguire & B.M. Boom, Mem. New York Bot. Gard. 51: 31, figs. 13A–K, 20, 21. 1989. —Lisyanthus caerulescens Aubl., Hist. Pl. Guiane 207, t. 82. 1775. —Irlbachia caerulescens (Aubl.) Griseb., Gen. Sp. Gent. 195. 1839 [1838]. —Helia caerulescens (Aubl.) Kuntze, Revis. Gen. Pl. 428: 1891. Weak annual herb to 60 cm tall; leaves 3– 20(–33) × 1–3 mm; calyx 2–6 mm long, the lobes 2–5 mm long; corolla 5–18(–25) mm long, the lobes 2–10 mm long; capsule 3–9 mm long. Moist white-sand or clay savannas or other open places in lowlands and highlands, 200–1500 m; widespread in Bolívar. Eastern Venezuela; other countries as in genus. ŠFig. 458. Tetrapollinia caerulescens is morphologically most similar to Irlbachia pratensis but differs in having a smaller corolla, less leafy stems, and leaves not particularly aggregated toward the base of the stem.

Fig. 458. Tetrapollinia caerulescens

21. VOYRIA Aubl., Hist. Pl. Guiane 208. 1775. Leiphaimos Schltdl. & Cham., Linnaea 6: 387: 1831. by Paul J. M. Maas Erect, saprophytic, perennial herbs. Stems usually simple, or less often branched, terete, solitary or a few together. Roots poorly known, sometimes corallike or of the bird’s nest type. Leaves somewhat connate at bases, small, scale-like, the lower ones sometimes alternate. Inflorescence a terminal, few–30-flowered cyme or reduced to a solitary, terminal flower; bracts and bracteoles usually present, very similar to the foliate leaves. Flowers variously colored, (4)5(–7)-merous, shortly or long-pedicellate. Calyx tubular to campanulate, (4)5(–7)-lobed, persistent, sometimes provided with discoid scales at the inner base; corolla actinomorphic, salverform to funnelform, variously colored, far exceeding the calyx, marcescent, apical part often papillate inside; tube elongate; lobes (4)5(–7), contorted, spreading to recurved, rarely erect. Stamens (4)5(–7), included, inserted at various levels in the corolla tube; filaments conspicuous or virtually absent. Ovary 1-locular, the parietal placentae protruding, base of ovary provided with 2 opposite glandular marks, sometimes with 2 distinct glands, or eglandular; style filiform, gradually widened towards ovary; stigma funnelform, rotate, or capitate, with undulate margin, or weakly 2-lobed, appendages sometimes present; ovules many, anatropous. Capsule fusiform to globose, septicidally dehiscent, or often indehiscent. Seeds many, globose to filiform, in some species with 2 hair-like projections. Neotropics (18 species), tropical west Africa (1 species); 13 species in Venezuela, all in the flora area.

Voyria 535

Key to the Species of Voyria 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(1). 7. 8(7). 8. 9(8). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(11). 13. 14(13). 14.

Corolla tube > 25 mm long ......................................................................... 2 Corolla tube < 25 mm long ......................................................................... 7 Corolla funnelform, tube pale orange-brown, widened in the upper half (to 20 mm diameter), lobes white, erect ...................................... V. clavata Corolla salverform, differently colored, tube not widened, lobes spreading ................................................................................................................ 3 Calyx tube enclosed by sheathing bracts, limb often irregularly lobed; corolla blue, often fading to white ................................................ V. caerulea Calyx not enclosed by sheathing bracts, limb regularly lobed; corolla never blue ............................................................................................... 4 Yellow herbs; flowers always solitary; seeds filiform ...................... V. aphylla Differently colored herbs; flowers solitary or not; seeds ± globose .......... 5 Calyx lobes oblong-elliptic, apex obtuse; corolla pink to red .............. V. rosea Calyx lobes (narrowly) triangular, apex acute; corolla white to pink ...... 6 Corolla pinkish to white, lobes almost linear, soon recurved, the tube inflated below the throat ............................................................ V. tenuiflora Corolla purely white, the lobes ovate to narrowly ovate, spreading ................................................................................................. V. acuminata Stems and flowers purplish; filaments hairy, 3–10 mm long .......... V. pittieri Stems and flowers differently colored; filaments glabrous, 0–3 mm long ................................................................................................................ 8 Thecae with a long hairy tail at the base; corolla yellow to orange, often 4-merous .................................................................................. V. spruceana Thecae without a hairy appendage at the base; corolla yellow, white, or blue, usually 5-merous ........................................................................... 9 Ovary provided with 2 distinct glands; flowers solitary ......................... 10 Ovary eglandular or with indistinct glandular marks; flowers in a manyflowered cyme or solitary ..................................................................... 11 Glands sessile; flowers yellow; seeds ellipsoid ............................ V. flavescens Glands stipitate; corolla lobes generally blue; seeds filiform .......... V. tenella Flowers arranged in a many-flowered cyme (rarely solitary) ................ 12 Flowers solitary ........................................................................................ 13 Corolla yellow to orange; calyx 5–9 mm long ............................ V. aurantiaca Corolla white (to pale pink); calyx 2.5–6 mm long ..................... V. corymbosa Corolla pinkish to white, lobes almost linear, soon recurved, the tube inflated below the throat ............................................................ V. tenuiflora Corolla yellow to white, lobes spreading, the tube not inflated ............. 14 Flowers yellow; seeds filiform .......................................................... V. aphylla Flowers white to yellowish; seeds ovoid .......................................... V. chionea

Voyria acuminata Benth., J. Bot. (Hooker) 2: 46. 1840. Small saprophyte to 15 cm tall; flowers solitary or in a 2–6-flowered cyme; corolla 60–70 mm long, pure white. Evergreen lowland forests, 50–1000 m; Amazonas (Cerro

Aratityope, Cerro Duida, Cerro Parú, Río Maturacá below Salto de Huá on Brazilian border, Río Mawarinuma, Río Siapa, Sierra Parima). Brazil (Amazonas, Roraima). ŠFig. 459.

536

G ENTIANACEAE

Voyria aphylla (Jacq.) Pers., Syn. Pl. 1: 284. 1805. —Gentiana aphylla Jacq., Select. Stirp. Amer. Hist. 87, t. 60, fig. 3. 1763. —Leiphaimos aphylla (Jacq.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 104, fig. 46A–D, J–K. 1895. —Vohiria. Voyria araguensis H. Karst., Linnaea 28: 415. 1857. Small saprophyte 15–30 cm tall; flowers solitary, corolla 25–65 mm long, yellow to orange. Semideciduous forests, evergreen lowland forests, montane forests, on granitic inselbergs, white-sand caatinga forests and savannas, 50–1800 m; Delta Amacuro (Caño Jotajana off Caño Guiniquina, Coboina near Misíon San Francisco de Guayo), Bolívar (Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá [Toronó-tepui], upper slopes of Ptari-tepui, Sierra de Lema at Cerro Uiri), Amazonas (slopes of Cerro Aracamuni, Cerro Duida, saddle between Cerro Duida and Cerro Marahuaka, Río Mawarinuma, San Carlos de Río Negro, Sierra Parima). Aragua, Distrito Federal, Falcón, Miranda, Nueva Esparta, Sucre, Yaracuy; throughout the Neotropics. ŠFig. 464. This is the most common species in the genus. Its widespread distribution is facilitated by the filiform, wind-dispersed seeds. It occurs in a wide variety of habitats. Voyria aurantiaca Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 135, t. 1, fig. 3. 1840. —Leiphaimos aurantiaca (Splitg.) Miq., Stirp. Surinam. Select. 149. 1850 [1851]. Small saprophyte 10–25 cm tall with coral-shaped roots; inflorescence a loose cyme with as many as 30 flowers in several dichasia or sometimes reduced to 1 or 2 flowers; flowers yellow to orange, 5–9 mm long. Evergreen lowland forests, 0–400 m; Delta Amacuro (Serranía de Imataca south of Cerro Muerto), Bolívar (Altiplanicie de Nuria, Río Cuyuní, Serranía de Imataca), Amazonas (20 km south of Puerto Ayacucho, near San Pedro de Cataniapo, Tobogán de la Selva south of Coromoto). Panama, Pacific coast of Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 461. Voyria caerulea Aubl., Hist. P1. Guiane 211, t. 83, fig. 2. 1775.

Small saprophyte to 20 cm tall; inflorescence a dense 2–6(–10)-flowered cyme or flowers sometimes solitary; calyx irregularly lobed and longitudinally split; corolla blue to purplish, 40–80 mm long. Evergreen lowland forests in leaf mold, 50–300 m; Bolívar (Altiplanicie de Nuria, Serranía de Imataca), Amazonas (Río Yatúa below Piedra Arauicaua, San Carlos de Río Negro). Guyana, Suriname, French Guiana, Brazil. ŠFig. 460. Voyria chionea Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 197. 1854. —Leiphaimos chionea (Benth.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 104. 1895. Small saprophyte to 15 cm tall; flowers white or less often yellow, solitary; corolla 20–25 mm long. Evergreen lowland forests, ca. 100 m; Amazonas (below Piedra Arauicaua on Río Yatúa). Amazonian Colombia and Brazil. Voyria clavata Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 133, t. 1, fig. 1. 1840. Small saprophyte to 20 cm tall; flowers solitary or rarely in a few-flowered cyme, corolla funnelform, 70–115 mm long, yellow to white. Submontane forests, 200–400 m; Amazonas (Río Cunucunuma to base of Cerro Huachamacari). Guyana, Suriname, French Guiana, Amazonian Brazil. Voyria corymbosa Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 136, t. 2, fig. 4. 1840. Small saprophyte to 20 cm tall; inflorescence a corymb of several dichasia, to 30flowered, rarely 1–few-flowered; flowers white to pink, corolla 10–16(–22) mm long. Central America, Colombia, southern Venezuela, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of V. corymbosa 1. Filaments 0–1 mm long, inserted ca. 2 mm below the throat; floral tube inside papillate ........................... subsp. alba 1. Filaments 1–5 mm long, inserted 2–4 mm below the throat; floral tube glabrous ................................ subsp. corymbosa

Voyria 537

V. corymbosa subsp. alba (Standl.) Ruyters & Maas, Acta Bot. Neerl. 30: 144. 1981. —Leiphaimos alba Standl., Contr. U.S. Natl. Herb. 20: 198. 1919, “albus.” Lower montane forests, 200–300 m; Bolívar (Río Toro northeast to Río La Reforma). Central America, Colombia, Guyana, Suriname, French Guiana, Peru. V. corymbosa subsp. corymbosa Lower montane forests, ca. 500 m; Bolívar (slopes of Cerro Guaiquinima). Suriname, French Guiana, Peru, northernmost Brazil. Voyria flavescens Griseb. in A. DC., Prodr. 9: 85. 1845. —Leiphaimos flavescens (Griseb.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 105, fig. 46G. 1895. Small saprophyte 3–15 cm tall; flowers solitary, corolla yellow to yellowish white, 11–20 mm long; ovary with 2 elliptic glands near its base. Evergreen lowland and lower montane forests, 100–300 m; Bolívar (Río Toro to Río La Reforma), Amazonas (upper Río Orinoco, San Carlos de Río Negro). Mexico, Central America, tropical South America. ŠFig. 462. Voyria pittieri (Standl.) L.O. Williams, Fieldiana, Bot. 31: 413. 1968. —Leiphaimos pittieri Standl., Contr. U.S. Natl. Herb. 20: 197. 1919. Small saprophyte 3–15 cm tall; inflorescence a loose, few-flowered cyme or flowers solitary; corolla purplish white, 14–30 mm long; ovary with 2 elliptic glands near its base. Riparian Mora forests, near seal level to 100 m; Delta Amacuro (La Paloma in Río Cuyubini basin). Mérida, Táchira, Zulia; Panama, Colombia, Guyana, Peru, Amazonian Brazil. ŠFig. 466. Voyria rosea Aubl., Hist. Pl. Guiane 209, t. 83, fig. 1. 1775. Small saprophyte with pinkish stems to 20 cm tall; inflorescence a loose to dense (2)3–6(–10)-flowered cyme, or flowers solitary; corolla pink to red, 45–100 mm long. Evergreen lowland to upper montane forests, sometimes growing on termite nests, 100– 1000 m; Amazonas (Cerro Aratitiyope, Río Mawarinuma). Guyana, Suriname, French Guiana.

Voyria spruceana Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 197. 1854. —Leiphaimos spruceana (Benth.) Gilg in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 105, fig. 46E. 1895. Small saprophyte 2–25 cm tall; flowers solitary, corolla 7–30 mm long, yellow, rarely whitish to flesh-colored but then with yellow throat. Evergreen lowland forests, Manicaria swamp forests, Rio Negro caatinga forests, sometimes growing on termite nests, 0– 200 m; Delta Amacuro (lower Río Cuyubini), Amazonas (below Piedra Arauicaua on Río Yatúa, Río Baría, Río Mawarinuma, San Carlos de Río Negro). Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ŠFig. 465. This is one of the few species in the genus in which the corolla is often 4-merous. Voyria tenella Hook., Bot. Misc. 1: 47, t. 25, fig. B. 1829. —Leiphaimos tenella (Hook.) Miq., Stirp. Surinam. Select. 149. 1850 [1851]. —Vohiria. Small saprophyte to 20 cm tall; flowers solitary, corolla 9–22 mm long, the tube white to orange, the lobes blue or purplish; ovary with 2 stalked glands at its base. Evergreen lowland to montane forests, in leaf mold or sometimes on decaying logs, 50– 1400 m; Bolívar (La Escalera, Río Tírica), Amazonas (slopes of Cerro Huachamacari, road from lower Río Cataniapo to Gavilán). Aragua, Miranda, Zulia; Mexico, Central America, Antilles, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 463. This species is wind-dispersed by its sawdust-like seeds, . Voyria tenuiflora Griseb. in A. DC., Prodr. 9: 85. 1845. —Leiphaimos tenuiflora (Griseb.) Miq., Stirp. Surinam. Select. 149. 1850 [1851]. Small saprophyte 5–25 cm tall; flowers solitary, corolla 15–70 mm long, the tube salmon, pink, or white, the lobes creamy to white, becoming strongly recurved. Lower montane forests, 200–800 m; Amazonas (base of Cerro Duida opposite Culebra, east slopes of Cerro Huachamacari). Guyana, Suriname, French Guiana, northwestern Brazil.

538

G ENTIANACEAE

Fig. 459. Voyria acuminata

Fig. 460. Voyria caerulea

Fig. 461. Voyria aurantiaca

Voyria 539

Fig. 462. Voyria flavescens

Fig. 464. Voyria aphylla

Fig. 465. Voyria spruceana

Fig. 463. Voyria tenella

Fig. 466. Voyria pittieri

540

G ENTIANACEAE

22. VOYRIELLA (Miq.) Miq., Stirp. Surinam. Select. 146. 1850 [1851]. —Voyria sect. Voyriella Miq., Tijdschr. Wis.- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2: 122. 1849. by Paul J. M. Maas Erect, saprophytic, perennial, white herbs. Stems branched or unbranched, fleshy, quadrangular to slightly winged. Roots filiform. Leaves scale-like, small. Inflorescence a terminal or rarely axillary, ± contracted, 1–many-flowered cyme; bracts and bracteoles present. Flowers white, erect, (4)5(6)-merous, shortly pedicellate. Sepals almost free, narrowly triangular, persistent, provided with discoid scales at the inner bases; corolla actinomorphic, salverform to tubular, hardly exceeding calyx, caducous, papillate within, the lobes contorted, small. Stamens (4)5(6), included, inserted at various levels in corolla tube; filaments long or short. Ovary 2-locular at base, 1-locular in upper part, eglandular, placentae parietal; style filiform; stigma 2-lobed, papillate; ovules many, strictly orthotropous. Capsule globose to ovoid, indehiscent, provided with the persistent style. Seeds many, subglobose, pitted. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 1 species. Voyriella parviflora (Miq.) Miq., Stirp. Surinam. Select. 147. 1850 [1851]. —Voyria parviflora Miq., Tijdschr. Wis.Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 2: 122. 1849. Voyriella oxycarpha Sandwith, Bull. Misc. Inform. 1931: 56. 1931. Small, white saprophyte to 15 cm tall; corolla 3.5–12 mm long, to 2 mm diameter, the lobes 0.5–3 mm long and 1–2 mm wide. Ever-

green lowland to lower montane forests, 50– 800 m; Bolívar (upper Río Icabarú), Amazonas (slopes of Cerro Duida, Piedra Arauicaua, Río Autana near Raudal Ceguera, Río Mawarinuma, Río Siapa near Raudal Gallineta, Tobogán de la Selva south of Coromoto). Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil. ŠFig. 467.

Fig. 467. Voyriella parviflora

Wurdackanthus 541

23. WURDACKANTHUS Maguire, Phytologia 57: 312. 1985. by Lena Struwe Herbs or subshrubs. Stems sharply quadrangular and winged. Leaves subsessile or shortly petiolate, elliptic, lanceolate to ovate, coriaceous (W. argyreus) or chartaceous (W. frigidus), the base attenuate, the apex acute to acuminate; margins flat to slightly recurved; secondary venation indistinct with 1 or 2 pairs of basally divergent arcuate secondary veins; interpetiolar line present. Inflorescence terminal, cymose, 1–9-flowered. Flowers pedicellate, actinomorphic to slightly zygomorphic, 5-merous, horizontal to nodding. Calyx campanulate, divided ca. 3/4 its length, the lobes ovate to oblong, hyaline-margined, with acute to obtuse apices, somewhat spreading in fruit; corolla funnel- to salverform, lavender-pink, pink, or whitish (W. argyreus), or yellow to green-yellow (W. frigidus), thin, deciduous in fruit, the lobes broadly ovate, spreading, the lobe apex obtuse, acute, or acuminate, the corolla bud apex tapering. Stamens inserted close to the base of the corolla tube, with a corona at the insertion point (W. frigidus); filaments slightly unequal, often sharply bent near apex; anthers lanceolate-oblong, sagittate, recurved after anthesis, with sterile apical appendages; pollen in tetrads, the exine coarsely reticulate.

Fig. 468. Wurdackanthus argyreus

542

G ENTIANACEAE

Ovary with a glandular base; style long, slender, persistent in fruit; stigma bilamellate, the lobes linear-oblong. Capsule oblong, woody, dehiscing medially, nodding. Seeds angular, not winged; testa cells dome-like or concave with band-like thickenings (W. frigidus). Guadeloupe, St. Vincent, Dominica, Venezuela, Brazil; 2 species, 1 in Venezuela. Wurdackanthus is a part of the Irlbachia complex and is closely related to Symbolanthus, with which it shares the presence of corolline coronas and similar pollen morphology. Further studies may show that Wurdackanthus could be better placed within Symbolanthus to preserve the monophyly of the latter genus. Wurdackanthus argyreus Maguire, Phytologia 57: 312. 1985. Symbolanthus aracamuniensis Steyerm., Ann. Missouri Bot. Gard. 76: 958, fig. 11. 1989. Simple-stemmed subshrub or herb to 1.5 m tall; leaves 29–72 × 11–28 mm, ellipticlanceolate to ovate, the upper surfaces green, the lower surface silvery; calyx 7–12 mm

long, the lobes 4–8 mm long, ovate to oblong; corolla 40–55 mm long, lavender-pink, pink, or whitish, the lobes 16–25 mm long; capsule 13–18 mm long. Dwarf forests on tepui slopes, savannas on tepui summits, 1300– 2200 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). ŠFig. 468.

GESNERIACEAE by Christian Feuillet and Julian A. Steyermark Plants epiphytic or terrestrial herbs, subshrubs, shrubs, small trees, or vines, sometimes arising from tubers, scaly rhizomes, or stolons. Stems herbaceous and fleshy, suffrutescent, or woody, erect, ascending, pendent, or scandent, simple or branched, usually pubescent at least toward the apex. Leaves opposite, rarely whorled or alternate; blades equal to strongly unequal, simple, entire to variously toothed, membranous, fleshy to coriaceous; petioles usually present. Inflorescences axillary or terminal, modified cymes, or racemes, occasionally fascicles or the flowers solitary. Flowers usually zygomorphic, rarely nearly actinomorphic. Calyx of 4 or 5 lobes, free or connate at the base, equal or unequal, green or a different color, entire or variously toothed. Corolla variously colored, with a short or long tube of 5 connate petals, often oblique in the calyx, often gibbous to saccate at base, cylindric, ventricose, or ampliate above, the limb of 4 or 5 equal or unequal lobes, sometimes bilabiate, erect or spreading to reflexed. Stamens 2–4(5); staminodes (0)1–3, included or exserted, usually didynamous; filaments adnate to the base of the corolla tube, usually coiling or retracting after pollen shedding; anthers variously shaped, coherent at first, dehiscing by pores or longitudinal slits. Disk absent, or if present, annular, or of 1–5 separate or connate nectariferous glands. Ovary superior to inferior, 1-celled, or very rarely 2-celled, the 2 placentae parietal; style usually elongating as stamens retract, simple; stigma usually 2-lobed and ± concave. Fruit a berry, or a dry or

G ESNERIACEAE 543

fleshy capsule, 2- or 4-valved. Seeds numerous, ± fusiform or oblong, very small, usually striate or otherwise variously marked. Pantropics, rarely reaching into warm-temperate regions; 125–130 genera and ca. 2500 species, 17 genera and 58 species in the flora area. Key to the Genera of Gesneriaceae 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(2). 6. 7(6). 7. 8(7). 8. 9(7). 9. 10(9). 10. 11(10). 11. 12(10). 12. 13(12). 13. 14(6). 14. 15(14). 15. 16(14).

16.

Lateral aerial branches much elongated at their bases in the form and function of stolons ...................................................................... 10. Episcia Lateral branches not as above ................................................................... 2 Plants with leaves in a rosette, or stemless .............................................. 3 Plants with obvious internodes ................................................................. 6 Inflorescences and leaves arising directly from a tuber .......... 15. Rhoogeton Inflorescences and leaves held by a stem .................................................. 4 Inflorescences terminal ............................................................. 11. Koellikeria Inflorescences axillary ............................................................................... 5 Flowers in dense clusters obscuring the peduncles and pedicels from above; corollas > 2.5 cm long ......................................... 14. Paradrymonia Flowers in loose clusters with obvious peduncles and pedicels when seen from above; corollas < 1.5 cm long ....................................... 17. Tylopsacas Plants terrestrial, erect or decumbent ...................................................... 7 Plants epiphytic, lianescent, or creeping ................................................ 14 Ovary half-inferior to almost completely inferior; plants rhizomatous ... 8 Ovary superior or almost completely superior (Sinningia), plants with tubers, or lacking underground perennial stems, not rhizomatous ........ 9 Corollas orange, red, or pink ....................................................... 12. Kohleria Corollas white or slightly tinted with purple ............................... 8. Diastema Inflorescences always with peduncles and pedicels, lacking bracts .......... ..................................................................................................... 2. Besleria Inflorescences with bracts, or lacking peduncles .................................... 10 Corollas > 3 times the length of the calyx ............................................... 11 Corollas ≤ 2 times the length of the calyx ............................................... 12 Ovary half-inferior; flowers in inflorescences or on the uppermost axils ................................................................................................ 16. Sinningia Ovary superior; flowers not on the uppermost axils ........... 13. Nautilocalyx Inflorescences sessile or of solitary flowers; plants without tubers ........... ................................................................................................ 1. Alloplectus Inflorescences with obvious peduncles; plants with or without tubers . 13 Plant from a tuber; fruit a capsule ........................................ 3. Chrysothemis Plant without tuber; fruit a berry ......................................... 7. Corytoplectus Anthers dehiscing by pores ...................................................................... 15 Anthers dehiscing by longitudinal slits ................................................... 16 Calyx lobes 2 or 5, < 10 mm long .............................................. 4. Codonanthe Calyx lobes 5, > 12 mm long ........................................................ 9. Drymonia Corollas white or whitish, rarely yellow or cream-colored in Paradrymonia maculata and then one lobe closing the corolla tube; leaves of a pair nearly equal .................................................................................. 17 Corollas red or yellow; leaves of a pair nearly equal or unequal ........... 18

544

G ESNERIACEAE

17(16). Largest leaf of a pair < 14 cm long, leaves strongly unequal ..................... ........................................................................................ 5. Codonanthopsis 17. Largest leaf of a pair > 16 cm long or leaves subequal (Paradrymonia maculata) ....................................................................... 14. Paradrymonia 18(16). Calyx lobes cordate at base, if corolla red, then lower lobe yellow and not reflexed; fruit a fleshy capsule .............................................. 1. Alloplectus 18. Calyx lobes narrowed at base, or if cordate, then all five corolla lobes red and lower lobe reflexed (Columnea oerstediana); fruit a berry ............. .................................................................................................. 6. Columnea 1. ALLOPLECTUS Mart., Nov. Gen. Sp. Pl. 3: 53. 1829. Shrubs or coarse herbs, terrestrial or epiphytic; roots fibrous; stems erect or clambering, quadrangular. Leaves opposite, usually equal in a pair, often long-petiolate; blades elliptic to ovate or oblanceolate; margins entire to serrate. Flowers 1– few in fascicles, axillary, bracteate, peduncles not evident. Calyx of 5 lobes nearly free, often broad, entire to strongly serrate. Corolla tubular, yellow or red, oblique in the calyx, constricted or not in the throat, or laterally compressed, gibbous at base with a spur, often ventricose below mouth, the 5 lobes of the limb usually small. Stamen filaments adnate to the base of the corolla tube; anthers dehiscing by longitudinal slits. Disk gland large, sometimes 2-lobed. Ovary superior; style included. Fruit a fleshy, bivalved capsule surrounded by the persistent calyx.

Fig. 469. Alloplectus savannarum

Besleria 545

Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; about 60 species, 12 in Venezuela, 1 of these in the flora area. The plants usually occur in rain or cloud forests as terrestrials or epiphytes, and appear quite commonly at forest margins. Alloplectus savannarum C.V. Morton, Bull. Torrey Bot. Club 75: 563. 1948. Columnea calotricha var. austroamericanarum C.V. Morton, Bol. Soc. Venez. Ci. Nat. 23: 78. 1962. Columnea steyermarkii C.V. Morton, Bol. Soc. Venez. Ci. Nat. 23: 76. 1962. Epiphytic or terrestrial, scandent sub-

shrub; leaves of a pair unequal, blade with white or yellowish pubescence, the lower surface green or reddish; calyx red; corolla whitish. Disturbed slope forests, 500–1700 m; Bolívar (Gran Sabana, Los Testigos, Macizo del Chimantá, Río Cuyuní to Sierra de Lema). Táchira; eastern Colombia, Guyana, Suriname, eastern Peru, Brazil (Roraima). ŠFig. 469.

2. BESLERIA L., Sp. Pl. 619. 1753. Terrestrial, perennial herbs or shrubs with fibrous roots. Stems terete or quadrangular, to 5 m tall, often unbranched. Leaves opposite, equal or unequal in a pair; blades elliptic, ovate, or lanceolate, membranous to coriaceous, the apex acute to acuminate, base cuneate or rounded; margins entire to coarsely dentate; stomata randomly scattered; petioles short or long. Inflorescences axillary, fasciculate, cymose, or subumbellate, rarely the flowers solitary; peduncles lacking to elongated; bracts absent. Floral tube very short; calyx lobes united at least at base, campanulate, urceolate, or cylindric, 5-toothed or -parted, imbricate in aestivation, rounded to acuminate, entire to serrulate, green or otherwise colored; corolla yellow, orange, red, or white, erect or oblique in the calyx, the tube usually cylindric, sometimes spurred or gibbous at the base, and sometimes ventricose and contracted at the throat, the limb regular or irregular, 5-lobed, entire. Fertile stamens 4, didynamous, included; filaments broad, adnate to the base of the corolla tube, flattened, usually glabrous; anthers coherent at first, becoming free, the locules confluent at apex; staminode occasionally developing; disk annular, usually glabrous, entire, sometimes colored. Ovary superior, unilocular with the two parietal placentae ovuliferous only on the inner surface or on both surfaces; stigma 2-lobed. Fruit a fleshy globose berry, red, orange, or white; exocarp thick, verrucose or smooth. Seeds minute, ellipsoid, numerous, reddish brown, striate. Mexico, Central America, West Indies, south to Brazil (from an apparent center in northwestern South America); ca. 150 species, 30–35 in Venezuela, 8 of these in the flora area. Key to the Species of Besleria 1. 1. 2(1). 2.

3(2).

Inflorescence with a peduncle 1–6 cm long ............................................... 2 Inflorescence without a peduncle, or the peduncle ≤ 3 mm .................... 4 Corolla white, ca. 8 mm long; pedicels 2–6 mm long; calyx 2–3 mm long, the lobes suborbicular or oval ...................................................... B. flavovirens Corolla yellow, orange, or red, 17–40 mm long; pedicels 10–35 mm long; calyx 4–15 mm long, the lobes lanceolate-ovate, oblong-lanceolate, or ovate-oblong ........................................................................................... 3 Corolla yellow, spurred at base, 35–40 mm long; calyx lobes 4–7 mm long; rounded or obtuse at the apex; leaf blades subentire or inconspicuously

546

3.

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

8.

G ESNERIACEAE

shallowly serrulate with < 20 serrations on each margin; lower surface of leaf blades and petioles sparsely to densely sericeous ..... B. penduliflora Corolla red or orange, not spurred at base, 17–22 mm long; calyx lobes 7– 15 mm long, slenderly acuminate at the apex; leaf blades usually obviously serrulate with 25–40 teeth on each margin; lower surface of leaf blades and petioles glabrous or sparsely puberulous, the midrib and lateral veins slightly pubescent ................................................. B. laxiflora Leaves of a pair subequal, stem internodes straight ................................ 5 Leaves of a pair strongly unequal, stem internodes at an angle with each other ....................................................................................................... 7 Stem and petioles densely long-pubescent; corolla > 10 mm long ..... B. sp. A Stem and petioles glabrescent or short-pubescent; corolla 6–9 mm long .... 6 Lateral veins 5 or 6 on each side of the midrib .......................... B. parviflora Lateral veins 10–15 on each side of the midrib ........................ B. flavovirens Calyx, pedicels, petioles, and stems densely villosulous with lax spreading hairs to 2 mm long ........................................................................... B. sp. B Calyx, pedicels, petioles, and stems either sparsely pilose with shorter hairs or the pubescence antrorsely appressed ...................................... 8 Stems, petioles, and pedicels antrorsely appressed pubescent; calyx lobes 6–10 mm long, narrowly lanceolate, minutely appressed-pubescent .......................................................................................................... B. sp. C Stems, petioles, and pedicels sparsely pilose with ascending hairs 1 mm or less long; calyx lobes 4–5 mm long, ovate or ovate-lanceolate, sparsely hirsutulous with lax hairs ............................................ B. gibbosa

Besleria flavovirens Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 11: 49. 1823. Suffrutescent herb 0.3–1.5 m tall; leaves of a pair equal; inflorescence condensed, subsessile or peduncle to 3 cm long; corolla 8 mm long, erect in the calyx, white, lobes with hair or papillae inside. Forests near base of cliff, montane forests, 500–1600 m; Bolívar (Amaruay-tepui), Amazonas (Río Yatúa, Sierra de la Neblina). Costa Rica, Colombia (Amazonas), Guyana, Suriname, French Guiana, Peru (Loreto), Brazil (Amazonas, Bahia, Roraima). Besleria gibbosa (Poepp.) Hanst. in Mart., Fl. Bras. 8: 421. 1864. —Hypocyrta gibbosa Poepp., Nov. Gen. Sp. Pl. 3: 3, pl. 202. 1845 [1840]. Suffrutescent herb 0.2–1 m tall; leaves of a pair mostly equal or subequal; inflorescence subsessile; corolla 15–20 mm long, strongly oblique in the calyx, white with yellow throat. Moist forests on lateritic soil, riparian forests, 100–300 m; Bolívar (6 km

southeast of Salto Pará), Amazonas (base of Piedra Cocuy, near San Carlos de Río Negro). Apure, Barinas, Mérida, Táchira, Zulia; Colombia (Meta), Brazil (Amazonas). Besleria laxiflora Benth., London J. Bot. 5: 361. 1846. Suffrutescent herb or shrub 0.5–4(–7) m tall; leaves of a pair equal or subequal; inflorescence peduncle 1.5–6.5 cm long; corolla 13–23 mm long, erect in the calyx, yellow to red. Primary and secondary wet forests, 100– 1700 m; Bolívar (Cerro Ichún, El Paují, Río Canarucuni, upper Río Caroní, Río Caura, middle and upper Río Paragua, Salto Kamá on Gran Sabana, near Santa Elena, Santa Maria de Erebato, Serranía Pia-Zoi), Amazonas (Cerro Aratitiyope, Cerro Duida, slope of Cerro Huachamacari, Cerro Yureba, Cerro Yutajé, Río Coro Coro, Río Cunucunuma, Río Ugueto, Sierra Parima). Barinas, Districto Federal, Lara, Mérida, Miranda, Portuguesa, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, Brazil (Bahia, Roraima). ŠFig. 471.

Besleria 547

Fig. 470. Besleria penduliflora

Fig. 471. Besleria laxiflora

548

G ESNERIACEAE

Besleria parviflora L.E. Skog & Steyerm., Novon 1: 211. 1991. Suffrutescent herb 0.7–2 m tall; leaves of a pair equal; inflorescence sessile; corolla 5– 7 mm long, erect in the calyx, white to yellow, lobes glabrous inside. Near streams in montane forests, 400–1300(–2300) m; Bolívar (Amaruay-tepui, El Paují, upper Río Caroní, Serranía Pia-Zoi, Sierra de Lema, Sororopán-tepui), Amazonas (upper Río Orinoco, Sierra Parima). Adjacent Guyana and Brazil. Besleria penduliflora Fritsch, Repert. Spec. Nov. Regni Veg. 18: 9. 1922. Erect or climbing shrub 0.5–4 m tall; leaves of a pair unequal, the smaller 1/2 the size of the larger, to subequal; inflorescence peduncle pendent, 2.5–4 cm long, usually only one pedicel 15–20 mm long; corolla 35– 40 mm long, strongly oblique in the calyx, bright yellow. Montane forests, 800–2500 m; Bolívar (between El Dorado and Santa Elena, northeast of Luepa, Macizo del Chimantá [Amurí-tepui, Sarvén-tepui, Toronó-tepui], Ptari-tepui), Amazonas (Sierra de la Nebli-

na). Western Guyana. ŠFig. 470. Besleria sp. A Suffrutescent herb to 0.5 m tall; leaves of a pair equal; inflorescence sessile; corolla erect in the calyx, white (in bud). Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma). Endemic. Besleria sp. B Suffrutescent herb 0.3–2 m tall; leaves of a pair unequal, the smaller about 1/3 the size of the larger; inflorescence sessile; corolla 17–35 mm long, strongly oblique in the calyx, white. Lowland flooded forests, 100–200 m; Amazonas (upper Río Baria, Río Mawarinuma, Sierra de la Neblina). Endemic. Besleria sp. C Suffrutescent herb to 1 m tall; leaves of a pair unequal, the smaller about 1/3 the size of the larger; inflorescence sessile; corolla 25 mm long, strongly oblique in the calyx, white. Black-water flooded forests, 100–200 m; Amazonas (uppermost Río Yatúa). Endemic.

3. CHRYSOTHEMIS Decne., Rev. Hort. sér. 3, 3: 242. 1849. Tussacia Benth., London J. Bot. 5: 360, 363. 1846, non Tussaca Raf. 1814. Terrestrial or epiphytic herbs from perennial tubers. Stems succulent, erect or sometimes decumbent, subquadrangular. Leaves opposite, equal or subequal in a pair; blades with the margin dentate or crenate, usually dark green and sparsely pubescent. Inflorescences numerous, pedunculate, axillary but appearing terminal by exceeding the stem apex, cymose or umbellate, 1/6–1/2 the length of the subtending leaf; bracts small, leafy. Flowers somewhat zygomorphic, 1–9 per peduncle. Calyx of 5 connate sepals; corolla of 5 connate petals erect in the calyx, tube cylindric, gibbous, internally with a ring of hairs at the stamen insertion and glandular hairs above. Stamens inserted on the corolla base. Nectary posterior, 2-lobed or rarely to four separate glands. Ovary superior; stigma 2-lobed, glandular. Fruit a fleshy capsule included in the persistent calyx, globose to ovoid, bivalved. Guatemala to Panama, West Indies, Colombia, Venezuela, Trinidad, Tobago, Guyana, Suriname, French Guiana, Brazil; 6 species, 2 in Venezuela, both in the flora area. Key to the Species of Chrysothemis 1.

1.

Leaf blades not decurrent on the petiole; inflorescence 3–6-flowered, umbellate, with the pedicels ± equally elongated and arising equally from the summit of the peduncle; calyx inflated, cordate or subcordate at the base, to 25 × 15 mm; corolla lobes ± glabrous without .............. C. dichroa Leaf blades decurrent on the petiole; inflorescence usually 5–10(–15)flowered, subracemose-corymbose, with pedicels of unequal length and

Chrysothemis 549

sometimes inserted at different levels on the peduncle; calyx not inflated, rounded or obtuse at the base, to 18 × 11 mm; corolla lobes densely villous without ............................................................. C. pulchella Chrysothemis dichroa Leeuwenb., Acta Bot. Neerl. 7: 331. 1958. Tuberous herb 10–30 cm tall; calyx red; leaf blade decurrent on the petiole, margin with small teeth; corolla orange-yellow, with reddish round spots at the throat. Saxicolous herb under forest, 100–300 m; Amazonas (Río Orinoco basin from Gavilán to south of Río Atabapo). Colombia (Guainía). ŠFig. 473. Chrysothemis pulchella (Donn ex Sims) Decne., Rev. Hort. 21: 242. 1849. —Besleria pulchella Donn ex Sims, Bot.

Mag. 28: t. 1146. 1808. —Tussacia pulchella (Donn ex Sims) Rchb. ex Walp., Repert. Bot. Syst. 6: 740. 1847. Tuberous herb to 1 m tall; leaf blade not decurrent on the petiole, margin with large teeth; calyx red-orange, persistent around the fruit; corolla orange-yellow, with reddish longitudinal lines on the lobes and at the throat. Usually in damp forests on moist rocks, 0–500 m; Delta Amacuro (Cerro La Paloma, Río Amacuro), Bolívar (Cerro Pitón, base of Cerro Uroi, Las Trincheras, Parguaza, Salto Pará, near Turumbán). Panama, West Indies, Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Brazil, cultivated throughout the tropics as an ornamental. ŠFig. 472.

Fig. 472. Chrysothemis pulchella

Fig. 473. Chrysothemis dichroa

550

G ESNERIACEAE

4. CODONANTHE (Mart.) Hanst., Linnaea 26: 199, 208. 1853 [1854], nom. cons. —Hypocyrta sect. Codonanthe Mart., Nov. Gen. Sp. Pl. 3: 50. 1829. Epiphytes, usually growing on ant nests. Stems pendent, repent, or erect, to 2 m tall, becoming woody. Leaves opposite, equal to rarely strongly unequal in a pair; blades fleshy, becoming coriaceous when dry, entire, sinuate, or serrulate toward the apex. Flowers with the calyx 5-lobed or bilabiate, dorsal lobe often recurved. Corolla oblique in the calyx, the tube funnelform to subcampanulate, base spurred or rounded. Stamens inserted at the base of the corolla tube; anthers dehiscing by pores. Disk a single dorsal nectary, usually large. Ovary superior; style included, but elongating after pollen shed; stigma 2-lobed to stomatomorphic. Fruit a fleshy berrylike capsule, tardily dehiscent or dehiscent by 2 valves. Mexico, Central America, Lesser Antilles, Colombia, Venezuela, Trinidad, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; about 15 species, 4 in Venezuela, 2 of these in the flora area. The plants are usually associated with ants in various ways. They seem to be epiphytic on ant nests, and often have extrafloral nectaries on the lower surfaces of the leaves, between the bases of the calyx lobes, or at the nodes. Key to the Species of Codonanthe 1.

1.

Calyx bilabiate, the lower (posterior or dorsal) lobe narrow and linear-lanceolate, the upper (anterior or ventral) lobe broad, oblong, rounded, and entire or shallowly 4-toothed at the apex; leaves entire or serrulate toward apex .............................................................................. C. calcarata Calyx deeply 5-lobed, the lobes all linear-lanceolate; leaves always entire ................................................................................................. C. crassifolia

Codonanthe calcarata (Miq.) Hanst., Linnaea 34: 416. 1865. —Nematanthus calcaratus Miq., Linnaea 22: 472. 1849. —Flor de conejo, Koma koma (Yanomami). Codonanthe bipartita L.B. Sm., Bull. Torrey Bot. Club 60: 657, fig. 1–6. 1932 [1933]. Epiphyte, mostly growing on stinging-ant nests. Corolla white, pinkish, or pale lavender with occasional stripes or spots. Evergreen lowland to montane forests, 50–1100 m; Bolívar (Gran Sabana west to the Río Paragua bordering the base of Cerro Guaiquinima), Amazonas (Río Casiquiare, Río Cunucunuma, Río Mavaca, Río Mawarinuma, upper Río Orinoco). Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ŠFig. 474. The leaves of the plant are sometimes employed to cure skin rashes caused by irritating plants.

Codonanthe crassifolia (Focke) C.V. Morton, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1159. 1938. —Hypocyrta crassifolia Focke, Tijdschr. Wis- Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 5: 199. 1852. —Jedieji-chu (Yekwana), Salvajito biuri. Codonanthe confusa Sandwith, Bull. Misc. Inform. Kew 1931: 492. 1931. Epiphyte mostly growing on stinging-ant nests; corolla white or sometimes flushed with pink on the outer surface of the lobes, yellow on the lower interior surface. Evergreen lowland to montane forests, gallery forests, forests bordering savannas, 100– 1200 m; throughout Delta Amacuro, Bolívar, and Amazonas. Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 475.

Codonanthe 551

Fig. 474. Codonanthe calcarata

Fig. 475. Codonanthe crassifolia

552

G ESNERIACEAE

5. CODONANTHOPSIS Mansfeld, Repert. Spec. Nov. Regni Veg. 36: 120. 1934. Epiphytic herbs or subshrubs. Stems terete, erect or stiffly ascending. Leaves shortly petiolate, opposite but strongly unequal on mature stems so as to appear alternate, the smaller one stipule-like, early deciduous, the larger one elliptic to obovate, acute at the apex, thick and somewhat fleshy. Inflorescence axillary, of 1–few flowers in a cluster. Calyx lobes usually unequal; corolla oblique in the calyx, the tube spurred and gradually ampliate above. Stamen filaments connate and adnate to the base of the corolla tube; anthers coherent, the connective narrow. Fruit a fleshy, oblong capsule. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 4 or 5 species, 2 in Venezuela, both in the flora area. Key to the Species of Codonanthopsis 1.

1.

Leaves 6–11 × 2–3.5 cm, narrowly and obliquely elliptic; upper part of stem and portions of the leaf blades and petioles pilose with soft white hairs; flowers 1 or 2 in leaf axils; corolla white with yellow on the throat; sepals green, broadly lanceolate, 3–4 mm wide ..... C. dissimulata Leaves 10–30 × 3–7 cm, narrrowly elliptic-oblanceolate or elliptic-obovate; leaves and stem glabrous; flowers several or many, fasciculate on short peduncles; coralla pinkish or white with purple markings; sepals reddish, linear-lanceolate, 1–2 mm wide .......................................... C. ulei

Codonanthopsis dissimulata (H.E. Moore) Wiehler, Selbyana 5: 61. 1978. —Codonanthe dissimulata H.E. Moore, Baileya 19: 25. 1973. —Miel de pajarito. Epiphyte, associated with nests of stinging ants; leaves 6–11 × 2–3.5 cm, thick, fleshy-coriaceous; sepals pale green; corolla white with yellow on the lower interior of the tube, sometimes very small cleistogamous corolla in-

Fig. 476. Codonanthopsis dissimulata

cluded in the calyx. Lowland to lower montane forests along streams, 50–900 m; Delta Amacuro (Serranía de Imataca), Bolívar (upper Río Paragua). Amazonas (Río Baría at base of Sierra de la Neblina, upper Río Casiquiare, Sierra Parima). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 476.

Columnea 553

Codonanthopis ulei Mansfeld, Repert. Spec. Nov. Regni Veg. 36: 120. 1934. —Codonanthe ulei (Mansfeld) H.E. Moore, Baileya 19: 28. 1973. —Cupi. Epiphytic shrub associated with nests of stinging ants; stems to 9 mm diameter; leaves mainly 10–30 × 3–7 cm, thick-fleshy, often with red margins; sepals often red-

dish; corolla white or flushed with yellow, or pinkish with lavender or purplish marking on throat and/or lobes. Evergreen lowland forests along or near streams, 100– 200 m; Amazonas (lower Río Baría at the base of Sierra de la Neblina, near San Carlos de Río Negro). Colombia, Venezuela, Peru, Brazil.

6. COLUMNEA L., Sp. Pl. 638. 1753. Dalbergaria Tussac, Fl. Antill. 1: 141, pl. 19. 1808 [1811–1813]. Columnea subg. Pentadenia Planch., Fl. Serres Jard. Eur. 6: 45. 1850. —Pentadenia (Planch.) Hanst., Linnaea 26: 210. 1853 [1854]. Perennial suffrutescent or succulent herbs or small shrubs, axes often having adventitious roots at the nodes, epiphytic or terrestrial, habit erect, spreading, prostrate or pendulous. Leaves opposite, ± petiolate, those of a pair equal to strongly unequal; blades various in shape, usually with a characteristic indumentum. Flowers axillary, sometimes showy, ± pedicellate, bracteate but bracts sometimes soon caducous. Calyx lobes 5, usually free nearly to the base, usually equal; margins entire, toothed, or pectinate. Petals 5, normally fused into a tubular or ventricose corolla tube with slightly unequal to strongly unequal (zygomorphic) free corolla lobes, the 2 posterior lobes connate and themselves fused to 2 lateral lobes, and a single anterior lobe being the least fused when flowers zygomorphic; corolla lobes erect, spreading or reflexed, the tube gibbous at base. Stamens 4; filaments briefly connate at base and adnate to base of corolla tube; anthers coherent in 2 pairs, exserted or included, the loculi dehiscent by longitudinal slits. Disk normally reduced to a single 2-lobed gland or rarely 5 separate glands, the gland appressed to posterior side of ovary. Ovary superior, unilocular, with 2 parietal placentas; style terminal, about as long as stamens; stigma 2-lobed or stomatomorphic, exserted or included. Fruit a white or colored berry, sticky inside. Seeds small and numerous (often > 100), spindle-shaped, pale brown when ripe, with darker tips; testa obliquely striate; embryo straight. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 200 species, 10 in Venezuela, 2 of these in the flora area. Key to the Species of Columnea 1. 1.

Larger leaf of a pair to 10 cm long, the apex round or obtuse .................... .............................................................................................. C. microsepala Larger leaf of a pair > 20 cm long, the apex acuminate ............ C. sanguinea

Columnea microsepala (C.V. Morton) Kvist & L.E. Skog, Allertonia 6: 391. 1993. —Alloplectus microsepalus C.V. Morton, Fieldiana, Bot. 28: 523. 1953. —Pentadenia microsepala (C.V. Morton) Wiehler, Phytologia 27: 315. 1973. Epiphyte with fleshy stem; leaves of a

pair strongly unequal, both obtuse or rounded at apex, the larger to 10 cm long; corolla pale yellow; berry white. Habitat not indicated, ca. 1300 m; Amazonas (Sierra Parima). Aragua, Falcón, Miranda, Monagas, Yaracuy; western Ecuador, extreme northwestern Peru. ŠFig. 478.

554

G ESNERIACEAE

Fig. 477. Columnea sanguinea

Fig. 478. Columnea microsepala

Corytoplectus 555

Columnea sanguinea (Pers.) Hanst., Linnaea 34: 384. 1865. —Besleria sanguinea Pers., Syn. Pl. 2: 165. 1807. —Dalbergaria sanguinea (Pers.) Steud., Nomencl. Bot., ed 2, 1: 479. 1840. Columnea aureonitens Hook., Bot. Mag. 73: pl. 4294. 1847. —Dalbergaria aureonitens (Hook.) Wiehler, Phytologia 27: 316. 1973. Columnea affinis C.V. Morton, Fieldiana, Bot. 28: 529. 1953. Herb with fleshy stem, growing on rocks or lower trunks; leaves of a pair strongly unequal, both acuminate, the larger usually > 20

cm long; corolla white, yellow or green to orange-red, often with red hairs; berry pale green. On rocks or lower tree trunks in forests, 100–1800 m; Bolívar (Altiplanicie de Nuria, Amaruay-tepui, Cerro Jaua, La Danta, near El Palmar, Ilú-tepui, Macizo del Chimantá, Quebrada O-paru-má near Kavanayén, Sierra Pakaraima, Urimán-tepui), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Río Coro Coro, Sierra de la Neblina). Widespread elsewhere in Venezuela in wet forests; West Indies, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador (Napo), northern Brazil. ŠFig. 477.

7. CORYTOPLECTUS Oerst., Centralamer. Gesner. 45. 1858. Terrestrial herbs. Stem succulent, fleshy or suffrutescent. Leaves petiolate, opposite-decussate; blades usually of equal size. Inflorescences axillary, pedunculate, umbellate-cymose, shorter than the leaves; peduncles and pedicels erect; bracts present or lacking. Calyx lobes of equal size or dorsal lobe narrower; corolla tubular, erect in the calyx, inflated ventrally, constricted at the throat, limb small, lobes ± equal. Fruit a shiny globose berry. Colombia, Venezuela, western Guyana, Ecuador, Peru, northern Brazil; 8 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Corytoplectus 1. 1.

Inflorescence 5–10-flowered; pedunculate .................................. C. congestus Inflorescence 1–3-flowered; without peduncle ........................... C. deltoideus

Fig. 479. Corytoplectus deltoideus

556

G ESNERIACEAE

Corytoplectus congestus (Linden ex Hanst.) Wiehler, Phytologia 27: 313. 1973. —Alloplectus congestus Linden ex Hanst., Linnaea 34: 371. 1865. Herbaceous, terrestrial perennial 0.2–1.5 m tall; lower surface of leaves green or purple; flowers erect; bracts orange-red; corolla orange-red, pilose to densely villous. Montane forests along streams, 1100–2200 m; Amazonas (Sierra de la Neblina). Barinas, Falcón, Lara, Mérida, Trujillo, Zulia; Colombia, Brazil (Roraima). Corytoplectus deltoideus (C.V. Morton)

Wiehler, Phytologia 27: 313. 1973. —Alloplectus deltoideus C.V. Morton, Fieldiana, Bot. 28: 521. 1953. Herbaceous, terrestrial or rarely epiphytic perennial 0.6–1.5 m tall; leaves pale gray-green below suffused with lavender on veins or completely lavender; flowers erect; calyx lobes red; corolla yellow, pilose to densely villous. Wet, tall, montane forests along shaded slopes of streams at the base of sandstone bluffs, 100–2200 m; Bolívar (south of El Dorado, basins of Río Aponguao, Río Cuyuní, and Río Venamo, Sororopán-tepui). Western Guyana. ŠFig. 479.

8. DIASTEMA Benth., Bot. Voy. Sulphur 132. 1845. Slender, perennial, terrestrial herbs from scaly rhizomes. Stems short, usually unbranched, hirsute or villous. Leaves opposite, nearly equal in a pair; petiole usually elongate; blades coarsely crenate, dentate, or serrate. Inflorescences axillary or terminal racemes, pedunculate, bracteate. Calyx 5-lobed, each lobe free, elongate, entire; corolla white or yellowish sometimes with purple markings, or red, tubular, funnel-form or cylindric, erect in the calyx, neither spurred nor ventricose, only slightly broader toward the limbs, the limb regular or slightly bilabiate. Stamen filaments adnate to the base of the corolla tube; anthers orbicular, initially coherent by their apices, later separating. Ovary half-inferior. Fruit a dry thin-walled capsule, convex at apex, 2-valved. Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Bolivia; ca. 25 species, 3 in Venezuela, 1 of these in the flora area. Diastema racemiferum Benth., Bot. Voy. Sulphur 132. 1845. Small herb 2–25 cm tall, internodes often reddish; leaves 1.5–12 × 0.8–5 cm; corolla tube cylindric, white or yellowish, sometimes

Fig. 480. Diastema racemiferum

with purple marking at the throat. Wet bluffs, 1500–1800 m; Amazonas (upper Río Coro Coro). Mexico, Central America, Colombia, Ecuador, Peru. ŠFig. 480.

Drymonia 557

9. DRYMONIA Mart., Nov. Gen. Sp. Pl. 3: 57. 1829. Terrestrial or epiphytic shrubs or lianas. Stems quadrangular or terete, to 5 m long when climbing, often with spreading adventitious roots at the internodes; branches present or lacking. Leaves opposite, equal or nearly equal in a pair; blades membranous or rarely coriaceous; petioles sometimes elongate. Inflorescences axillary, of single flowers or several in a cluster; bracts often large and leaf-like, or small and inconspicuous, but often caducous. Flowers often showy and bright-colored. Sepals 5, often large and colored or leaf-like, free or briefly connate at the base, unequal with the upper most shortest. Corolla oblique in the clayx, usually funnelform and broader toward the mouth, spurred or saccate at the base, throat broad, limb of 5 lobes, usually spreading, rounded, often fimbriate, 3 basal lobes usually exceeding the 2 upper lobes. Stamens 4, adnate to the base of the corolla tube, didynamous, included; filaments occasionally contorted; anthers sagittate, broadest at the tip, narrowed toward the base to 2 separate spurs, before anthesis coherent by sides and faces with the spurred bases up, dehiscing by 2–4 pores, after anthesis the filaments coil and the anthers separate. Disc reduced to a single posterior gland. Ovary superior; style included; stigma 2-lobed. Fruit a fleshy capsule, becoming coriaceous, dehiscing by 2 valves that recoil to reveal a red, orange, or purple interior and a mass of seeds in a pulpy funicle. Seeds ellipsoid or fusiform, obliquely striate. Mexico, Central America, Lesser Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 100 species, 6 in Venezuela, 2 of these in the flora area. The genus Drymonia is one of the largest genera of Gesneriaceae in the Neotropics. Plants of Drymonia are usually lianas, which distinguishes the genus from most other Gesneriaceae. Another distinguishing character is the specialized porate dehiscence of the stamens described by H. E. Moore, Jr. (Baileya 3: 109–112. 1955). Key to the Species of Drymonia 1. 1.

Leaf blades entire; inflorescence compact with large pink-red bracts; corolla lobes not fimbriate-erose. .............................................. D. coccinea Leaf blades serrulate; flowers axillary; at least one of the corolla lobes fimbriate-erose ......................................................................... D. serrulata

Drymonia coccinea (Aubl.) Wiehler, Phytologia 27: 324. 1973. —Besleria coccinea Aubl., Hist. Pl. Guian. 632, pl. 255. 1775. —Alloplectus coccineus (Aubl.) Mart., Nov. Gen. Sp. Pl. 3: 57, and index. p. 189. 1829. —Columnea coccinea (Aubl.) Kuntze, Revis. Gen. 2: 472. 1891. Alloplectus circinatus Mart., Nov. Gen. Sp. Pl. 3: 56. t. 233, fig. 2. 1829. Vining or shrubby semiligneous epiphyte or hemiepiphyte, sometimes climbing to 8 m high; leaves subcoriaceous, the upper surface dark green, the lower surface silvery green; bracts deep red or rarely salmon-rose to or-

ange-red, often with yellowish margins or green tips; calyx greenish, yellowish, or greenish white; corolla 5–6 cm long, yellow with purple-brown spotted lobes; fruit pale yellow. Along streams, forested talus slopes in evergreen lowland or montane forests, 100–1600 m; Bolívar (headwaters of Río Paramichi by Venezuelan-Brazilian border), Amazonas (near Mavaca, affluents of Río Baría and Río Yatúa, along Río Orinoco southeast of Puerto Ayacucho, Sierra de la Neblina). Carabobo, Yaracuy; Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil (Acre, Mato Grosso, Pará, Piaui).

558

G ESNERIACEAE

Fig. 481. Drymonia serrulata

Drymonia serrulata (Jacq.) Mart., Nov. Gen. Sp. Pl. 3: 59. 1829. —Besleria serrulata Jacq., Hort. Schoenbr. 3: 21, t. 290. 1798. Besleria spectabilis H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 400. 1817. Vining shrub with adhesive roots, stems slightly succulent; leaves coriaceous-succulent, pale green or sometimes suffused with dull lavender or maroon, the leaf margins often red-toothed; bracts green; calyx green, the upper surface often wine-red; corolla creamy yellow or greenish white with reddish purple spots on lobes, turning in color

during anthesis, 3–5 cm long; fruit globose, the valves reflexed, fleshy, purple, dark wine-red, or orange-red within; seeds dark brown. Forested slopes along streams, rocky slopes, bases of sandstone bluffs, evergreen lowland forests, montane and deciduous forests, 0–500 m; Delta Amacuro (Caño Araguao), Bolívar (Altiplanicie de Nuria, middle Río Caura, headwaters of Río Chicanán), Amazonas (Sierra Parima). Venezuelan Coastal Cordillera and Andes; Mexico, Central America, lesser Antilles, south to southern South America. ŠFig. 481.

10. EPISCIA Mart., Nov. Gen. Sp. Pl. 3: 39. 1829. Stoloniferous, low, terrestrial or epiphytic herbs, rarely suffrutescent. Stems decumbent, creeping, or sprawling on the ground or fallen trees, to 1 m or more long, rooting at the nodes, branched or not, pubescent to pilose. Leaves often crowded, opposite, usually nearly equal; blades ovate or elliptic to lanceolate, upper surface dark green or with various patterns of variegation, lower surface often color other than green, the apex acute to rounded, base acute to cordate; petioles short. Inflorescences often of single, or 2–6 axillary flowers on slender pilose peduncles, bibracteate. Flowers zygomorphic, showy, the floral tube very short. Calyx often irregular, with the posterior lobe forced back around the corolla spur, sepals 5, free or

Episcia 559

shortly connate at the base, linear, oblong to lanceolate, apex acute, acuminate, or erose, green or otherwise colored, pilose; corolla salverform to campanulate, conspicuously spurred, inserted horizontally in the calyx, tubular, white, yellow, blue, purple, or red, contracted above the spur and at the throat, rarely ventricose, limb oblique, 5-lobed, spreading, lobes rounded, entire or minutely toothed, or fimbriate. Stamens 4, didynamous, included; filaments nearly straight, inserted at the base of the corolla, after anthesis depressed or coiling; anthers orbicular or oblong, coherent in pairs in a square or arc, becoming free, dehiscing by a longitudinal slit. Ovary superior; style included; stigma stomatomorphic, 2-lobed, capitate; disc gland one, at the base of the ovary, large, dorsal; placentae ovuliferous on both surfaces or only on the inner surface. Fruit an ovoid bivalved fleshy capsule. Seeds ellipsoid, shiny, obliquely striate, brown. Mexico, Central America, Colombia, Venezuela, Guyana, Surinam, French Guiana, Ecuador, Peru, Brazil; 12 species, 6 in Venezuela, 5 of these in the flora area. Key to the Species of Episcia 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Flowers red ................................................................................................. 2 Flowers yellow or white ............................................................................. 3 Upper surface of leaves bronze or olive-green; corolla bright yellow ......... ...................................................................................................... E. xantha Upper surface of leaves pale green; corolla white ...................... E. fimbriata Leaf blades pubescent, bullate; petiole 30–50 mm long ................ E. reptans Leaf blades obviously densely hairy; petiole 7–25 mm long ..................... 4 Calyx lobes 2–3 mm long; corolla tube sparsely hirsute ..................... E. sp. A Calyx lobes 7–8 mm long; corolla tube densely appressed-pubescent ... E. sp. B

Episcia fimbriata Fritsch, Bot. Jahrb. Syst. 37: 484. 1906. Herb with stolons; upper surface of leaves pale green, lower surface green or purple; corolla white, often with purple markings. Terrestrial or on wet boulders, 50–300 m; Amazonas (near Cerro Duida and Marahuaka, 60 km southeast of Puerto Ayacucho, near San Juan de Manipiare). Peru, Brazil. ŠFig. 482. Episcia reptans Mart., Nov. Gen. Sp. Pl. 3: 41, pl. 217. 1829. Herb with stolons; leaf surface pebbled, often discolored; corolla bright red. Edge of waterfalls, 200–300 m; Amazonas (Río Yureba). Mérida; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 483. The collection from the flora area (Delascio & Guanchez 10871, MO, VEN) is sterile but seems to represent this species. Episcia xantha Leeuwenb., Misc. Pap. Landbouwhoogeschool 19: 241. 1980. Herb with long stolons; upper surface of leaves dark green or bronze, lower surface

green or purple; corolla bright yellow. Dense forests, 50–100 m; Amazonas (40 km southeast of Puerto Ayacucho). Guyana, Suriname, French Guiana, adjacent Brazil. The collection from the flora area (Guánchez 406, US) is sterile, but most likely represents this species. Episcia sp. A Herb with short stolons; leaves green(?), the upper surface pebbled, the lower surface with dense indumentum; corolla crimson, tube nearly glabrous. Shaded, moist canyons, ca. 200 m; Amazonas (Río Coro Coro, near Yutajé). Endemic. Episcia sp. B Mat-forming herb, with a few stolons. Leaves densely appressed-pubescent, hairs long, yellow; corolla crimson, tube densely appressed-pubescent outside, lobes glabrous. Base of waterfalls, wet crevices, 1500–2000 m; Amazonas (Cerro Duida, Cerro Parú). Endemic.

560

G ESNERIACEAE

Fig. 482. Episcia fimbriata

Fig. 483. Episcia reptans

11. KOELLIKERIA Regel, Ind. Sem. Hort. Turic. [4]. 1847. Terrestrial perennial herbs. Stems from scaly rhizomes, with short internodes, unbranched, pilose. Leaves opposite, subequal in a pair; petiole usually short; blades pilose, crenate to serrate, randomly silvery-maculate. Inflorescences terminal, elongate, indeterminate racemes; peduncles elongate; bracts alternate, minute. Flowers numerous, small. Calyx lobes free; corolla tubular, nearly erect in the calyx, not spurred, the limb 2-lipped, upper 2 lobes short, lower 3 lobes much longer and toothed. Stamen filaments adnate to the base of the corolla tube, anthers coherent. Ovary half-inferior. Fruit a dry capsule. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Argentina; 2 or 3 species, 1 in Venezuela. Koellikeria erinoides (DC.) Mansfeld, Repert. Spec. Nov. Regni Veg. 38: 28. 1935. —Achimenes erinoides DC., Prodr. 7: 536. 1839. Dwarf herbaceous plant forming rosettes; upper surface of leaves striped with brownpurple, the lower surface gray with lavender; corolla strongly zygomorphic, the lower 3 lobes white with a few brownish red lines near base, the upper 2 lobes magenta with white along border tinged yellow. Lithophytic in forested canyon on igneous rocks along stream bed, 50–100 m; Bolívar (Río Ore affluent of Río Parguaza). Northern and central Venezuela; other distribution as in genus. ŠFig. 484. Fig. 484. Koellikeria erinoides

Nautilocalyx 561

Fig. 485. Kohleria hirsuta

12. KOHLERIA Regel, Ind. Sem. Hort. Bot. Turic. [4]. 1847. Isoloma Benth. ex Decne., Rev. Hort. 20: 465. 1848, non Isoloma J. Sm. 1841. Brachyloma Hanst., Linnaea 26: 203. 1853 [1854], non Brachyloma Sonder 1845. Terrestrial herb or subshrub from scaly rhizome. Stem erect or decumbent. Leaves opposite or in whorls of 3–6, crenate or serrate. Inflorescences axillary fascicles or umbels, or the flowers solitary. Corolla mostly orange, red, or pink. Ovary > half-inferior. Fruit a 2-valved capsule. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru; 17 species, 3 in Venezuela, 1 of these in the flora area. Kohleria hirsuta (H.B.K.) Regel, Flora 31: 250. 1848. —Gesneria hirsuta H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 394, t. 189. 1818. Gesneria rubricaulis Kunth & Bouché, Index Seminum (Berlin) 1847: 12. 1847. —Kohleria rubricaulis (Kunth & Bouché) Hassk., Bonplandia 8: 97. 1860. Brachyloma rhodomallos Hanst., Linnaea 29: 526. 1859. —Kohleria rhodomallos (Hanst.) Hanst., Linnaea 34: 441. 1865. Kohleria longipedunculata Hanst., Linnaea 34: 441. 1865. Isoloma pycnosuzygium Donn. Sm., Bot. Gaz. 61: 382. 1916. —Kohleria pycnosu-

zygium (Donn. Sm.) Badillo in Pittier et al., Cat. Fl. Venez. 2: 397. 1947. Herbaceous perennial 0.6–1 m tall, the rhizomes often with stolons or bearing scaly fleshy propagules; leaves membranous, the lower surface sometimes with wine-purple veins. Usually lithophytic on moist sandstone cliff faces, rarely terrestrial, humid montane tall forests, 800–1300 m; Bolívar (Amaruaytepui, Cerro Guaiquinima, vicinity of Kavanayén), Amazonas (Cerro Huachamacari, Sierra Parima). Venezuelan Coastal Cordillera and Andes; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador. ŠFig. 485.

13. NAUTILOCALYX Lind. ex Hanst., Linnaea 26: 181, 206. 1853 [1854]. Terrestrial or rarely epiphytic perennial herbs, or low shrubs, occasionally producing tubers. Stems erect or decumbent, succulent, sometimes rooting at the nodes. Leaves opposite, elongate, equal or unequal in a pair. Inflorescences axillary cymes or fascicles, or a solitary flowers, subtended by 2 or more leafy bracts. Flowers with an irregular calyx, lobes unequal; corolla tubular, oblique in the calyx, with a short

562

G ESNERIACEAE

spur, limb 5-lobed. Stamens 4, included; filaments basally connate, and adnate to the base of the corolla tube; anthers apically joined in 2 pairs, dehiscing by a longitudinal slit; disc of a single posterior gland or of 2 opposite glands. Ovary superior. Fruit a bivalved capsule. Mexico, Central America Lesser Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil, Bolivia; ca. 60 species, 23 species in Venezuela, 19 of these in the flora area. Key to the Species of Nautilocalyx 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(1). 6. 7(6). 7. 8(7). 8. 9(7). 9. 10(9). 10. 11(10). 11. 12(6). 12. 13(12). 13. 14(12). 14. 15(14). 15. 16(14). 16. 17(16). 17. 18(16). 18.

Leaf blade ovate or obovate ....................................................................... 2 Leaf blade elliptic or oblanceolate ............................................................. 6 Leaf blade ovate; corolla white ............................................... N. adenosiphon Leaf blade obovate; corolla red or white ................................................... 3 Corolla red .............................................................................. N. chimantensis Corolla white, with or without purple markings ...................................... 4 Lower surface of leaves with tertiary veins hidden by long hairs .... N. sp. A Lower surface of leaves with tertiary veins obvious ................................. 5 Leaf blade narrowly rounded or emarginate at the base, petiole thin ......................................................................................................... N. sp. B Leaf blade widely rounded or cordate at the base, petiole fleshy .............. ................................................................................................... N. cordatus Leaf blade surface flat ................................................................................ 7 Leaf blade bullate ..................................................................................... 12 Leaf blade decurrent on the petiole ........................................................... 8 Leaf blade not decurrent on the petiole .................................................... 9 Leaves and bracts reddish .......................................................... N. punctatus Leaves and bracts green or whitish ............................................... N. pallidus Leaf blade subsessile or very short; calyx tube longer than the lobes ................................................................................................. N. arenarius Leaf blade with an obvious petiole; calyx tube shorter than the lobes ...... 10 Leaf blade < 5 cm long ........................................................................ N. sp. C Leaf blade > 8 cm long ............................................................................. 11 Calyx lobes ca. 10 times as long as wide ................................. N. fasciculatus Calyx lobes ca. 3–4 times as long as wide .......................................... N. sp. D Corolla white ............................................................................................ 13 Corolla red or bright pink ........................................................................ 14 Leaf blade oblanceolate, margin large-crenate ....................... N. pemphidius Leaf blade elliptic, margin short-crenate-serrate ........................ N. resioides Leaf blade oblanceolate, margin bicrenate ............................................. 15 Leaf blade elliptic ..................................................................................... 16 Corolla tube ≤ 2.5 cm long ............................................................N. maguirei Corolla tube > 2.5 cm long .................................................................. N. sp. E Petiole ≤ 1.5 cm long ................................................................................ 17 Petiole > 2 cm long ................................................................................... 18 Corolla tube ca. 1.5 cm long .................................................. N. cataractarum Corolla tube ca. 2 cm long ................................................................... N. sp. F Calyx lobes entire; corolla tube 2–3 cm long ...................................... N. sp. G Calyx lobes dentate; corolla tube 3–5 cm long ................. N. porphyrotrichus

Nautilocalyx 563

Nautilocalyx adenosiphon (Leeuwenb.) Wiehler, Selbyana 5: 29. 1978. —Episcia adenosiphon Leeuwenb., Acta Bot. Neerl. 18: 858. 1969. Terrestrial trailing herb; petiole about as long as leaf blade; leaf blade to 4 cm long, base mostly rounded, apex obtuse to acute, margin crenate; calyx lobes wide-elliptic; corolla white, the tube about 3.5 cm long. Dwarf forests, on igneous rocks, elevation not indicated; Bolívar (Sierra Imataca). French Guiana. ŠFig. 486.

base broadly rounded, cordate, apex broadly rounded, margin (crenate to) bicrenate; calyx lobes lanceolate; corolla white, lobes and throat often lavender, tube 3.5–5 cm long. Lowland and slope forests, 100–1500 m; Bolívar (Cerro Duida, Cerro Jaua, Cerro Maraveni near Maijía, upper Río Caura, Río Curumú in Río Ichún basin, headwaters of Río Parimichi, Serranía Marutaní), Amazonas (near base of Cerro Aracamuni, Cerro Aratitiyope, Cerro Huachamacari, Cerro Marahuaka, Río Siapa). Guyana. ŠFig. 487.

Nautilocalyx arenarius L.E. Skog & Steyerm., Novon 1: 215. 1991. Terrestrial herb; petiole nearly sessile to 5 mm long; leaf blade to 15 cm long, base rounded to decurrent, apex acute, margin minutely crenulate-serrulate; calyx lobes shorter than the tube; corolla white suffused with purple, tube to 4 cm long. Low forests on white sand, 100–200 m; Amazonas (near Maroa, near San Carlos de Río Negro). Endemic.

Nautilocalyx fasciculatus L.E. Skog & Steyerm., Novon 1: 219. 1991. Terrestrial suffrutescent herb; petiole 3–4 cm long; leaf blade 10–18 cm long, base acute, apex acute, margin minutely crenateserrulate; calyx lobes linear-lanceolate; corolla red, tube 3–5 cm long. Damp forests, 400–1000 m; Amazonas (base of Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina).

Nautilocalyx cataractarum Wiehler, Selbyana 1: 165. 1975. Terrestrial and saxicolous low herb; petiole 0.5–2 cm long; leaf blade to 15 cm long, base cuneate to attenuate, apex obtuse to acute, margin crenate; calyx lobes lanceolate; corolla bright pink, tube 1.0–1.5 cm long. Wet rocks or soil in lower montane forests, ca. 500 m; Bolívar (near Salto Angel). Endemic. ŠFig. 490. Nautilocalyx chimantensis L.E. Skog & Steyerm., Novon 1: 217. 1991. Terrestrial or saxicolous herb; petiole 2–7 cm long; leaf blade 7–15 cm long, base rounded to subcordate, apex broadly rounded or subobtuse, margin shallowly crenate-dentate; calyx lobes ligulate-oblong; corolla deep red, tube about 3 cm long. On exposed or shaded wet banks, 1400–1700 m; Bolívar (Macizo del Chimantá). Endemic. Nautilocalyx cordatus (Gleason) L.E. Skog, Novon 1: 217. 1991. —Episcia cordata Gleason, Bull. Torrey Bot. Club 58: 466. 1931. Centrosolenia hirsuta Benth., London J. Bot. 5: 362. 1846. —Episcia hirsuta (Benth.) Hanst., Linnaea 34: 350. 1865. Terrestrial or saxicolous trailing herb; petiole 1–7 cm long; leaf blade 5–16 cm long,

Nautilocalyx maguirei L.E. Skog & Steyerm., Novon 1: 219. 1991. Terrestrial or saxicolous herb; petiole 2.5– 8 cm long; leaf blade 12–26 cm long, base mostly oblique, one side acute to rounded, other side rounded to cordate, apex shortly acute or subacuminate, margin biserrate; calyx lobes narrowly lanceolate; corolla red, tube 2.3 cm long. Moist cliff faces, shaded banks, 1000–1200 m; Bolívar (Cerro Guaiquinima). Endemic. Nautilocalyx pallidus (Sprague) Sprague, Bull. Misc. Inform. 1912: 89. 1912. —Alloplectus pallidus Sprague, Bull. Misc. Inform. 1911: 346. 1911. Terrestrial or low-trunk epiphytic herb; petiole to 10 cm long; leaf blade 9–30 cm or more long, base decurrent, apex acuminate, margin serrate; calyx lobes foliaceous; corolla white, tube about 3 cm long. Whitesand lowland forests, 100–200 m; Amazonas (around San Carlos de Río Negro). French Guiana, Amazonian Peru and Brazil. ŠFig. 491. Nautilocalyx pemphidius L.E. Skog, Syst. Bot. 14: 281. 1989. Terrestrial or saxicolous herb; petiole nearly sessile to 3 cm long; leaf blade 5–20

564

G ESNERIACEAE

cm long, base attenuate or cuneate, apex acute to acuminate, margin scalloped to crenate; calyx lobes ligulate to subulate; corolla white, tube ca. 12 mm long. Wet forests near streams, often on rocks, 100–1300 m; Amazonas (Río Mawarinuma, Sierra de la Neblina). Endemic. Nautilocalyx porphyrotrichus (Leeuwenb.) Wiehler, Phytologia 27: 308. 1973. —Episcia porphyrotricha Leeuwenb., Acta Bot. Neerl. 7: 311. 1958. Terrestrial or saxicolous herb; petiole 0.5– 5 cm long; leaf blade 6–20 cm long, base cuneate to decurrent, apex acuminate, margin serrate to serrulate; calyx lobes linear; corolla red, tube 3.7–4.2 mm long. Wet forests, 400–1300 m; Bolívar (Amaruay-tepui, Cerro Venamo, south of El Paují, La Escalera, Macizo del Chimantá, upper Río Caroní, upper Río Cuyuní, Río Pauo affluent of Río Caura, Sierra de Lema, Sierra Ichún). Guyana. ŠFig. 488. Nautilocalyx punctatus Wiehler, Selbyana 5: 40. 1978. Terrestrial suffrutescent herb; petiole 1–2 cm long or less; leaf blade 10–30 cm long, base decurrent, apex acuminate, margin serrate; calyx lobes lanceolate; corolla creamwhite, spur yellow, tube 3–4 cm long. Wet forests near streams, 100–900 m; Amazonas (base of Cerro Yapacana, Río Asisa, Río Casiquiare, Río Cunucunuma, Río Mawarinuma, Río Yatúa, Sierra Parima). Endemic. Nautilocalyx resioides (Leeuwenb.) Wiehler, Selbyana 5: 42. 1978. —Episcia resioides Leeuwenb., Acta Bot. Neerl. 13: 59. 1964. Terrestrial or saxicolous herb; petiole 1–2 cm long; leaf blade 8–20 cm long, base decurrent, apex acute, margin minutely crenateserrate; calyx lobes lanceolate; corolla white with lilac on lobes and throat, tube 8 mm long. Lower montane forests, 500–900 m; Bolívar (Cerro Guaiquinima, La Escalera, Sierra de Lema). Endemic. ŠFig. 492. Nautilocalyx sp. A Saxicolous herb; petioles 1–2 cm long; leaf blades 3–7 cm long, base acute, apex rounded to obtuse, margin serrate-crenate; calyx lobes linear; corolla white in bud. Shaded bluffs of narrow rocky stream, 700–800 m; Bolívar (Cerro Guaiquinima). Endemic. ŠFig. 489.

Nautilocalyx sp. B Terrestrial(?) herb; petiole 4–6 cm long; leaf blade 7–9 cm long, base narrowly rounded to emarginate, apex rounded to broadly obtuse, margin obscurely crenate; calyx lobes narrowly lanceolate; corolla white, tube ca. 3 cm long. Slopes, elevation not indicated; Amazonas (Sierra Guaharibo on upper Río Orinoco). Endemic. Nautilocalyx sp. C Terrestrial herb; petiole 1 cm long; leaf blade 1.5–2.5 cm long, base acute to rounded, apex rounded, margin serrate; calyx lobes narrowly lanceolate; corolla unknown. In pockets of soil along escarpment edges, 1200–1300 m; Amazonas (Cerro Autana). Endemic. Nautilocalyx sp. D Terrestrial herb; petiole 2–5 cm long; leaf blade 7–13 cm long, base obtuse to acute, apex obtuse to subacuminate, margin biserrate; calyx lobes oblong to ovate; corolla red, tube 2–2.5 cm long. Roadside, 800–900 m; Bolívar (17 km east of El Paují). Endemic. Nautilocalyx sp. E Terrestrial or saxicolous herb; petiole nearly sessile to 3 cm long; leaf blade 6–17 cm long, base looks decurrent but mostly narrowly rounded, apex rounded to acute, margin wide-crenate to bicrenate; calyx lobes broadly oblanceolate; corolla red to orange-red, tube 2.5–3.5 cm long. Streambanks, mossy rocks, 300–1400 m; Amazonas (Cerro Sipapo, Cerro Yutajé, near Yutajé, Río Coro Coro). Endemic. Nautilocalyx sp. F Saxicolous herb; petiole nearly sessile to 0.5 cm; leaf blade 3–8 cm long, base attenuate to decurrent, apex obtuse to slightly acute, margin crenate to bicrenate; calyx lobes oblanceolate; corolla red, tube 2.3 cm long. Crevices on rocks, 800–1200 m; Amazonas (Cerro Yapacana). Endemic. Nautilocalyx sp. G Saxicolous herb; petiole nearly sessile to 3 cm long; leaf blade 6–15 cm long, base decurrent, apex acute to acuminate, margin minutely serrate to biserrate; calyx lobes oblanceolate; corolla red, tube 1.5–2 cm long. On boulders, in bluff crevices, 500–900 m; Bolívar (near El Paují, near Icabarú). Endemic.

Nautilocalyx 565

Fig. 486. Nautilocalyx adenosiphon Fig. 487. Nautilocalyx cordatus

Fig. 488. Nautilocalyx porphyrotrichus

566

G ESNERIACEAE

Fig. 489. Nautilocalyx sp. A

Fig. 490. Nautilocalyx cataractarum

Nautilocalyx 567

Fig. 491. Nautilocalyx pallidus

Fig. 492. Nautilocalyx resioides

568

G ESNERIACEAE

14. PARADRYMONIA Hanst., Linnaea 26: 180, 206. 1853 [1854]. Epiphytic or terrestrial herbs or subshrubs. Stems erect, ascending, or scrambling, often succulent, bearing adventitious roots. Leaves equal to strongly unequal in a pair, often appearing to be in a rosette, but usually large, elongated and surpassing the stem; blades usually lanceolate, membranous to fleshy, apex acuminate, base cuneate or long-decurrent into the petiole. Inflorescences of many flowers, usually congested in the axils of the leaves, but rarely exceeding the length of the petiole. Flowers often hidden by the leaves. Calyx lobes linear or lanceolate, free and usually long-attenuate at the apex; corolla white or yellow with red or purple spots or lines, funnelform or trumpet-shaped, spurred at the base. Stamens included; anthers oblong, coherent, bearded or not, ± dehiscing by a longitudinal slit; disc reduced to 1 or 2 glands. Ovary superior. Fruit a bivalved capsule. Central America, northern South America; ca. 30 species, 7 in Venezuela, 6 of these in the flora area. Key to the Species of Paradrymonia 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(3). 5.

Tertiary venation evident on lower leaf surface; leaf blades closely crenate-serrate ....................................................................................... 2 Tertiary venation not evident on lower leaf surface; leaf blades either entire or loosely serrate or toothed ........................................................... 3 Petiole 5–15 cm long, thin; leaf blade apex acute to acuminate, base cuneate .......................................................................... P. metamorphophylla Petiole to 3 cm long, thick; leaf blade apex blunt, base asymmetrically acute to obtuse ........................................................................................... P. sp. E Calyx lobes elliptic ..................................................................................... 4 Calyx lobes partly linear ............................................................................ 5 Flowers yellow ...................................................................................... P. sp. A Flowers cream or white ....................................................................... P. sp. D Leaf blade narrow, oblong, and long-decurrent; corolla white with white hairs .................................................................................................. P. sp. B Leaf blade ovate and short-decurrent; corolla white with magenta hairs .......................................................................................................... P. sp. C

Paradrymonia metamorphophylla (J.D. Sm.) Wiehler, Phytologia 27: 327. 1973. —Alloplectus metamorphophyllus J.D. Sm., Bot. Gaz. (Crawfordsville) 52: 52. 1911. Terrestrial or low epiphyte; stem less than 5 mm thick; leaf petiole (3–)8–15 cm long, villous; blade oblong to ovate-elliptic, base cuneate, apex acute, margin crenate to bicrenate, glabrous above; pedicels and calyx densely villous; sepals linear-lanceolate; corolla white, tube densely villous. Flooded to wet forests, slope forests, 100–200 m; Amazonas (Río Baría, Río Mawarinuma, Río Yatúa, near Sierra de la Neblina). Costa Rica, Panama, Colombia, Ecuador, Peru.

Paradrymonia sp. A Epiphyte on lower part of trunks, or saxicolous; stem > 5 mm thick; petiole 20–35 cm long, appressed-pubescent to glabrous; leaf blade obovate to ovate-elliptic, slightly decurrent on the upper part of the petiole at base, abruptly acuminate at apex, margin entire to repand, the upper surface glabrous; pedicels and calyx appressed-pubescent to glabrescent; sepals narrowly to broadly lanceolate; corolla not seen. Boulders in lowland and montane forests, 100–1300 m; Amazonas (Sierra de la Neblina). Endemic. Paradrymonia sp. B Terrestrial(?); stem ca. 5 mm thick; petiole

Paradrymonia 569

Fig. 493. Paradrymonia sp. D

8–11 cm long; leaf pubescent; blade broadly ovate, base cuneate, apex broadly acuminate, margin obscurely to clearly serrate, the upper surface minutely puberulous to glabrous; pedicels and calyx with long, erect, multicellular, red hairs; sepals narrowly triangular with a long, linear acumen; corolla white, tube with white hair. Creeping on rocks, often partly buried in duff, or climbing lower tree trunks, ca. 1200 m; Amazonas (Cerro Yapacana, Sierra de la Neblina). Endemic. Paradrymonia sp. C Terrestrial, saxicolous or low epiphyte; stem ca. 5 mm thick; petiole 8–15 cm long; leaf densely to sparsely appressed-pilose; blade broadly ovate to obovate, base cuneate, apex acuminate to caudate, margin glandular-dentate, the upper surface appressed-pilose to glabrous; pedicels and calyx pilose; sepals subulate; corolla white, tube with magenta hairs. Growing on rocks and lower tree trunks, 800–1200 m; Amazonas (Cerro Marahuaka, Cerro Yapacana). Endemic. Paradrymonia sp. D Saxicolous or terrestrial epiphyte; stem 1 cm thick; petiole 6–15 cm long, glabrous; leaf blade broadly obovate to elliptic, long-decurrent at base, acuminate at apex, margin entire, glabrous; pedicels and calyx sparsely pilose; sepals oblanceolate to lance-elliptic; corolla creamy white, upper 3/4 of the tube pilose. Tepui slopes and summits, 100–1400 m; Amazonas (Cerro Aracamuni, Piedra Arauicaua). Endemic. ŠFig. 493.

Paradrymonia sp. E Epiphyte; stem 5 mm thick or more; petiole ca. 2 cm long, thick, covered with dense, long, brown-red hairs; leaf blade elliptic, asymetrically acute to obtuse at base, widely rounded at apex, margin biserrate, the upper surface velutinous; pedicels and calyx with a dense, long, brown-red indumentum; sepals narrowly triangular; corolla with red hairs outside the lobes. Slope forests, ca. 1000 m; Amazonas (Cerro Marahuaka). Endemic.

570

G ESNERIACEAE

15. RHOOGETON Leeuwenb., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 146: 321. 1958. Small, acaulescent, terrestrial, herbaceous plants. Leaves basal, petiolate. Inflorescence axillary, umbellate, subcymose, or thyrsoid, long-pedunculate, 1–6-flowered. Peduncles longer than the leaves. Subtending bracts lanceolate. Calyx somewhat zygomorphic, deeply divided with 5 free lobes. Corolla red or orange, tubular or trumpet-shaped, nearly straight, slightly widened upward with 5 spreading, slightly unequal, rounded lobes, spurred at the base, the throat within bearing glandular hairs. Stamens 4, didynamous, included; filaments free, large, inserted near the base of the corolla. Disc gland 1, dorsal, emarginate, glabrous. Ovary superior. Stigma 2-lobed. Placentae ovuliferous only on the outer face. Endemic to the Guayana Shield in Venezuela and Guyana; 2 species, 1 in Venezuela. Rhoogeton viviparus Leeuwenb., Acta Bot. Neerl. 7: 323, 431, fig. 32. 1958. Rhoogeton leeuwenbergianus C.V. Morton,

Bol. Soc. Venez. Ci. Nat. 23: 80. 1962. Terrestrial, herbaceous plant 10–30 cm tall; leaves broadly elliptic, coarsely dentate,

Fig. 494. Rhoogeton viviparus

Sinningia 571

the lower surface silvery; petiole magentared; calyx dull orange-brown and green; corolla scarlet or red, 2.5–3 cm long. Base of wet sandstone bluffs, vertical faces of waterfalls, wet tall forests, 100–2000 m; Bolívar (Cerro Jaua, Cerro La Danta, Cerro Pauo,

Cerro Venamo, Macizo del Chimantá, Sierra de Lema), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Parú, Cerro Sipapo, Sierra de la Neblina, upper Río Siapa). Guyana. ŠFig. 494.

16. SINNINGIA Nees, Ann. Sci. Nat. (Paris), sér. 1, 6: 296. 1825. Gesnera Mart., Nova Gen. Sp. 3: 27. 1829. Gesneria sect. Corytholoma Benth., Pl. Hartw. 230. 1846. —Corytholoma (Benth.) Decne., Rev. Hort. 20: 466. 1848. Rechsteineria Regel, Ind. Sem. Hort. Bot. Turic. [4]. 1847. Terrestrial pubescent herbs or shrubs, perennial from woody tubers, or from rhizomes. Stems erect or decumbent, often unbranched. Leaves opposite or in whorls, or congested on short stems from the tuber, nearly equal in a pair or whorl; blades oblong, lanceolate, or orbicular; petioles of various length or lacking. Inflorescences axillary, with one or more flowers in umbels or cymes, or appearing terminal in racemes; peduncles and pedicels short or long, bracteate. Floral tube very short; calyx tube campanulate, lobes 5, triangular; corolla usually red or orange, tube campanulate to cylindric, erect, gibbous at the base, often ventricose above, limb 5-lobed, nearly regular, but more often bilabiate. Stamens 4, in 2 pairs; filaments adnate to the base of the corolla tube; anthers coherent, dehiscing by longitudinal slits; disc of 1–5 glands, sometimes with 2 larger and connate. Ovary half-inferior to superior; style slender; stigma stomatomorphic. Fruit an ovoid capsule dehiscing by 2 or 4 valves. Seeds small, numerous, fusiform, shiny, striate. Mexico, Central America, south to Bolivia, northern Argentina and Uruguay; ca. 75 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Sinningia 1.

1.

Leaves usually ternate, linear to narrowly ovate, ± sessile and straight or obtuse at base, pubescence long-erect; internode between leaves and flowers usually > 3 times as long as the upper leaves, flowers in racemes ....................................................................................................... S. elatior Leaves usually opposite, lanceolate, petiolate or at least acute-decurrent at base, pubescence short; progressive transition between leaves and bracts with first blooming leaves usually longer than the internode, first flowers axillary, the following gradually forming a raceme .................................................................................................. S. incarnata

Sinningia elatior (H.B.K.) Chautems, Candollea 45: 383. 1990. —Gesneria elatior H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 393. 1818. Herbaceous perennial 0.5–1 m tall; leaves usually in whorls of 3, rarely opposite; corolla tubular, scarlet or dark red. Wet savannas, cliffs, 50–500 m; Bolívar (road between Los Pijiguaos and Puerto Ayacucho, Serranía de los Pijiguaos), Amazonas (near Puerto

Ayacucho). Northern and western Venezuela; wide-ranging in Colombia, Peru, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 495. Sinningia incarnata (Aubl.) Denham, Baileya 19: 126. 1974. —Besleria incarnata Aubl., Hist. Pl. Guiane. 635, t. 256. 1775. —Rechsteineria incarnata (Aubl.) Leeuwenb., Acta Bot. Neerl. 7: 320. 1958.

572

G ESNERIACEAE

Fig. 495. Sinningia elatior

Gesneria caracasana Otto & Dietr., Allg. Gartenzeitung 6: 345. 1838. —Rechsteineria caracasana (Otto & Dietr.) Kuntze, Revis. Gen. Pl. 2: 474. 1891. Gesneria vargasii DC., Prodr. 7: 527. 1839. Gesneria lindeniana Brongn., Rev. Hort. sér. 3, 1: 363. 1847. Gesneria gollmeriana Hanst., Ind. Sem. Hort. Berol., App. 1861: 7. 1861. —Rechsteineria gollmeriana (Hanst.) Kuntze,

Revis. Gen. Pl. 2: 474. 1891. Herbaceous perennial 0.5–1 m tall; leaves opposite or in whorls of 3; corolla tubular, scarlet or dark red. Rocky savannas and cliffs, 100–1200 m; Bolívar (headwaters of Río Chicanán, near Santa Elena, Serranía de los Pijiguaos). Northern and western Venezuela; wide-ranging in Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil.

17. TYLOPSACAS Leeuwenb., Taxon 9: 220. 1960. Tylosperma Leeuwenb., Acta Bot. Neerl. 7: 323, 432, fig. 33. 1958, non Boschantzev 1952, nec Donk 1957. Terrestrial herbaceous plants. Leaves basal, oblanceolate. Inflorescence axillary, shorter than the leaves, paniculate, several- to many-flowered. Peduncles and pedicels slender, ebracteate. Flowers small. Sepals connate at the base, linear-lanceolate. Corolla white, tubular, gibbous at base, the lobes spreading. Stamens 4, included; filaments free, inserted near the base of the corolla; anthers reniform, dehiscing along longitudinal fissures. Ovary superior; stigma capitate. Nectary a narrow ring with 2 prominent dorsal lobes or teeth. Fruit a 2- or 4-valved capsule. Seeds subglobose, pustulate. Endemic to the Guayana Shield in southern Venezuela and Guyana; 1 species.

G NETACEAE 573

Fig. 496. Tylopsacas cuneatum

Tylopsacas cuneatum (Gleason) Leeuwenb., Taxon 9: 221. 1960. —Episcia cuneata Gleason, Bull. Torrey Bot. Club 58: 467. 1931. —Tylosperma cuneatum (Gleason) Leeuwenb., Acta Bot. Neerl. 7: 323, 432, fig. 33. 1958. Small terrestrial herb 10–30 cm tall; leaf blades with numerous sharp serrations, the lower surface silvery pale green; petioles often dull lavender or purplish; calyx green; corolla white. Frequent on moist, shaded

sandstone bluffs near water or by waterfalls, moss-covered rocks on humid forested slopes and well-drained sandy soils in forests, 100– 700 m; Bolívar (Cerro Ichún, Cerro Jaua, Cerro Pauo, Cerro Sarisariñama, Macizo del Chimantá, Sierra de Lema), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Parú, Cerro Sipapo, upper Río Siapa, Sierra de la Neblina). Guyana. ŠFig. 496.

GNETACEAE by Dennis W. Stevenson Woody vines (two arborescent Old World species). Wood with anomalous secondary growth, appearing as concentric rings of secondary vascular bundles. Leaves opposite, papyraceous to very coriaceous; venation reticulate. Plants dioecious or functionally so. Reproductive axes arranged in whorls of spikes or panicles, ultimate units also whorled, each whorl subtended by pair of fused bracts. Microsporangiate reproductive structures consisting of an obconical enclosing bract with an exserted slender stalk bearing 2 distal pollen sacs. Sterile ovulate structures often associated with the upper whorls of microsporangiate reproductive stuctures, consisting of 2 envelopes enclosing the nucellus: the inner one, the integument, and the outer one bract-like. Fertile ovulate structures enclosed by 3 layers, the innermost (the inner

574

G NETACEAE

integument) with a long tapering and exserted apex that exudes a small pollination droplet. Seeds yellow to red to deep purple when mature, appearing drupaceous with a thin coriaceous outer layer, a middle layer that has an outer fleshy part and an inner fibrous part, and an inner papyraceous layer, all surrounding the megagametophyte containing a single embryo with 2 cotyledons. Pantropics; 1 genus and ca. 35 species, 6 of these in the flora area. 1. GNETUM L., Mant. Pl. 1: 18. 1767. Characters and distribution the same as for the family. All species in Venezuela are known in the venacular as Towauri. The seeds of these species are eaten either boiled or roasted. Key to the Species of Gnetum 1. 1. 2(1).

2.

3(2).

3.

4(1).

4. 5(4).

5.

Leaves with dense subepidermal fibers so that the upper surface appears silky and the lower sculptured; mature leaves to 15 × 7 cm ................ 2 Leaves without subepidermal fibers, the surface not appearing silky or sculptured; mature leaves to 26 × 15 cm .............................................. 4 Bark winged; mature leaves oblong to oblong-elliptic; bract collars of ultimate reproductive units distant, in microsporangiate axes 10–15 mm, in ovulate axes 15–20 mm apart .................................... G. schwackeanum Bark smooth; mature leaves elliptic; bract collars of ultimate reproductive units close, in microsporangiate axes 1–2 mm, in ovulate axes < 10 mm apart ........................................................................................... 3 Mature leaves very coriaceous, to 6 × 4 cm, rounded to acute at apex, truncate at base, lower surface with broad prominent primary vein and secondary veins > 2 mm wide ................................................ G. camporum Mature leaves chartaceous to rarely subcoriaceous, 12(–15) × 6(–8) cm, acute to acuminate at apex, unequally obtuse at base, lower surface with very narrow, secondary veins < 1 mm wide ......................... G. urens Tertiary and higher order veins quite distinct as fine lines < 1 mm wide; adult leaves to 17 × 8 cm, leaves turning dark brown to black when dry ............................................................................................ G. paniculatum Tertiary and higher order veins indistinct and 1–2 mm wide when apparent; adult leaves to 26 × 15 cm, remaining green or tan when dry ..... 5 Nodes thickened and obviously articulate; adult leaves heavily coriaceous, to 20 × 15 cm, cordate to rounded at base; bract collars of ultimate reproductive units close, in microsporangiate axes 1–2 mm, in ovulate axes < 10 mm apart ..................................................... G. leyboldii Nodes unthickened and obscurely articulate; adult leaves coriaceous, to 26 × 9 cm, oblique at base; bract collars of ultimate reproductive units distant, in microsporangiate axes 10–20 mm, in ovulate axes 15– 20 mm apart .......................................................................... G. nodiflorum

Gnetum camporum (Markgr.) D.W. Stev. & T. Zanoni in Görts, Fl. Guianas, ser. A, fam. 7 (Gnetaceae) 14. 1991. —Gnetum urens var. camporum Markgr., Acta Bot. Venez. 6: 371. 1972 [1971]. Woody vine; twigs tan, smooth to slightly

striated, internodes often < 10 cm long. Along the margins of rocky streams, gallery forests, dwarf gallery forests bordering savannas, 500–1500 m; Bolívar (Gran Sabana). Endemic.

Gnetum 575

Fig. 497. Gnetum schwackeanum

576

G NETACEAE

Gnetum leyboldii Tul., Ann. Sci. Nat. Bot. sér. 4, 10: 117. 1858. Gnetum paraense Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr. 3: 403. 1902. Woody vine; branches articulate, bark of older stems often grooved. Along rivers, flooded forest margins, wet forests bordering savannas, 50–700(–1700) m; widespread in Bolívar and Amazonas. Colombia, eastern Ecuador, Peru, northern Brazil. Gnetum nodiflorum Brongn. in Duperrey, Voy. Monde (Phan.) 12. 1829. —Aarana, Lengua de tigre. Gnetum amazonicum Tul., Ann. Sci. Nat. Bot. sér. 4, 10: 116. 1858. Gnetum oblongifolium Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr. 3: 403. 1902. Gnetum cruzianum Gleason, Bull. Torrey Bot. Club 52: 196. 1925. Woody vine; twigs thick-barked with sparse but coarse lenticels. River and flooded forest margins, rocky slopes, sandstone bluffs of wet areas of tepuis, 100–1000(– 1800) m; widespread in Bolívar and Amazonas. Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. The resin of this species is collected and used as incense. Gnetum paniculatum Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 8: 357, t. 2, 3. 1856.

Gnetum microstachyum Benth. ex Tul., Ann. Sci. Nat. Bot. sér. 4, 10: 119. 1858. Robust woody vine; twigs with numerous large lenticels, dark brown to almost black. River margins, flooded forests, 200–1100 m; Bolívar (Amaruay-tepui), Amazonas (Río Siapa). Guyana, French Guiana, northern Brazil (Amazonas: upper Rio Negro). Gnetum schwackeanum Taub. ex Markgr., Bull. Jard. Bot. Buitenzorg sér. 3, 10: 450. 1930. Slender vine; bark with numerous broad, usually thin, wings of cork; nodes barely swollen. Mainly along rivers and on whitesand soils at base of rocky outcrops, 50–1000 m; southern Amazonas. Northwestern Brazil. ŠFig. 497. Gnetum urens (Aubl.) Blume, Tijdschr. Natuurl. Gesch. Physiol. 1: 162. 1834. —Thoa urens Aubl., Hist. Pl. Guiane 874, t. 336. 1775. Gnetum thoa Brongn. in Duperrey, Voy. Monde (Phan.) 2: 12. 1829, nom. illeg. Gnetum melinonii Benoist, Bull. Mus. Hist. Nat.(Paris) sér. 2, 17: 66. 1945. Gnetum leyboldii var. woodsonianum Markgr., Ann. Missouri Bot. Gard. 52: 385. 1965. Slender vine; twigs smooth, light gray to tan; leaves obviously and finely striate. River basins in flooded forests, black-water swamps, tepuis, 500–1700 m; widespread in Bolívar and Amazonas. Northern Brazil.

HAEMODORACEAE by Paul J. M. Maas and Hiltje Maas Perennial, rhizomatose, rarely cormiferous, sometimes stoloniferous herbs, the rhizomes often with red latex. Leaves parallel-veined, distichous, equitant, linear or

Pyrrorhiza 577

ensiform, mainly radical, sheathing at the base, longitudinally folded and connate towards the apex (apex unifacial or bifacial). Inflorescence a terminal, bracteate thyrse, sometimes corymbiform or capituliform, the partial florescences lax or dense cincinni; indument of inflorescence composed of simple hairs with or without swollen top cell. Flowers bisexual, actinomorphic to zygomorphic, shortly pedicellate. Tepals 6, arranged in 1 (not in the flora area) or 2 whorls, imbricate, free or basally connate, persistent, petaloid, subequal; tube when present (absent in the flora area) short to elongate, straight or curved. Stamens 6 in 2 whorls (not in the flora area), or 3 opposite the inner tepals thus representing the inner whorl (Xiphidium), or 3 combined with 2 staminodes representing the outer whorl (Schiekia), or only 1 combined with 2 staminodes representing the inner whorl (Pyrrorhiza); filaments free or adnate to the perianth tube; anthers basifixed or versatile (not in the flora area), 2locular, introrsely opening by longitudinal slits. Ovary superior (in the flora area) to more often inferior, 3-locular, ovules many, anatropous, placentation axile, septal nectaries present; style 1, filiform, persistent; stigma capitate to slightly 3-lobed. Fruit a 3-locular loculicidal capsule, seeds 2–many, per locule, tuberculate, hairy or glabrous; embryo small, embedded in abundant starchy endosperm. Neotropics, temperate North America, South Africa, Australia, New Guinea; 14 genera and ca. 80 species, 3 genera and 3 species in the flora area. The South African genus Wachendorfia is cultivated in Venezuela for its showy flowers. Key to the Genera of Haemodoraceae 1. 1. 2(1). 2.

Inflorescence a hairy thyrse composed of 2–4 cincinni; cincinni 5–7-flowered; stamen 1; cormiferous herbs ......................................... 1. Pyrrorhiza Inflorescence a glabrous thyrse composed of many cincinni; cincinni 3– 25-flowered; stamens 3; rhizomatose herbs .......................................... 2 Outer side of tepals hairy; staminodes 2; fruit with 2–4 seeds per locule .................................................................................................... 2. Schiekia Tepals glabrous; staminodes none; fruit with many seeds per locule ........ ................................................................................................. 3. Xiphidium

1. PYRRORHIZA Maguire & Wurdack, Mem. New York Bot. Gard. 9: 318. 1957. Cormiferous herbs with few radical leaves. Inflorescence a congested hairy thyrse, consisting of 2–4 cincinni; cincinni 5–7-flowered. Flowers actinomorphic, tepals free, equal. Stamen 1, staminodes 2. Capsule with 2 seeds per locule, seeds discoid with hairy margin. Endemic to the Guayana Shield in southern Venezuela; 1 species. Pyrrorhiza neblinae Maguire & Wurdack, Mem. New York Bot. Gard. 9: 318, fig. 63a–g. 1957. Herb 50–85 cm tall; corms bright red-orange inside; leaves 18–45 × 0.6–1.1 cm; inflorescence 4.5–7 × 3–5 cm, ferrugineous orange to red; flowers creamy, often with or-

ange-red apex or base; tepals 12–16 mm long; capsule shiny, maroon to brownish red, ellipsoid, 12–15 × 10–16 mm. Locally frequent or occasional in open tepui meadows, 1800– 2100 m; Amazonas (Sierra de la Neblina). Endemic. ŠFig. 498.

578

H AEMODORACEAE

Fig. 498. Pyrrorhiza neblinae

Fig. 499. Schiekia orinocensis subsp. orinocensis

Schiekia 579

2. SCHIEKIA Meisn., Pl. Vasc. Gen. Tab. Diagn. 397. 1842; Meisn., Pl. Vasc. Gen. Comm. 300. 1842. Rhizomatose herbs, rooting at the nodes. Leaves radical and scattered, margin apically serrulate. Inflorescence composed of many cincinni, glabrous; cincinni 2–25flowered. Flowers zygomorphic, tepals slightly unequal in size, the 3 abaxial ones basally connate as are the 3 adaxial ones, creating a ± bilabiate appearance. Stamens 3, the 2 adaxial ones smaller than the abaxial one, staminodes 2. Capsule with 2–4 seeds per locule, seeds subglobose, tuberculate. Southeastern Colombia (Boyacá, Guainía, Meta, Vaupés, Vichada), southern and eastern Venezuela, Guyana, Suriname, French Guiana, Brazil (Amazonas, Goiás, Maranhão, Mato Grosso, Pará, Rondônia, Roraima); 1 species. Schiekia orinocensis (H.B.K.) Meisn., Pl. Vasc. Gen., Tab. Diagn. 397. 1842; Meisn., Pl. Vasc. Gen. Comm. 300. 1842. —Wachendorfia orinocensis H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 319. 1815 [1816]. Herb 15–100 cm tall, sprouting from the lower leaves, rhizomes horizontally creeping, to 20 cm long, latex red; roots fibrous; leaves 10–60 × 0.2–2.5 cm; inflorescence 2–35 × 1–9 cm, orange to cream; flowers orange, yellowish, or white, often outer side of outer tepals green; tepals 5–12 mm long; capsule green, maturing orange to shiny dark purple, globose, 4–8 mm diameter. Distribution as in genus; 2 subspecies, both in the flora area. The species Schiekia orinocensis is subdivided here into two subspecies, one of them roughly representing the forest plants (Schiekia orinocensis subsp. silvestris), the typical subspecies comprising the savanna plants. Within these subspecies the plants are again rather variable. One might discern a form characterized by mainly basal, very narrow leaves from the Venezuelan savannas as Maguire & Wurdack (1971) did in their subspecies savannarum. Plants from Mato Grosso in Brazil are characterized by broad basal and cauline leaves and manyflowered rhipidia. However, in our opinion the rate of variation is too large to discern more than two subspecies within Schiekia orinocensis. Maguire & Wurdack (1971) state that the tepals of the extreme savanna form are basically white with orange stripes, whereas the extreme woodland form has solid orange tepals. The flowers of Schiekia have a definitely zygomorphic perianth, the three abaxial tepals and the three adaxial tepals being basally connate creating a slightly bilabiate appearance of the flower,

the lips being separated by basal slit-like pouches. Here again there is a difference between the savanna subspecies and the forest subspecies: the three abaxial tepals forming the “lower lip” of the flower are definitively reflexed in subsp. silvestris, but in subsp. orinocensis the three abaxial tepals together with the adaxial ones form a quite tubular flower. The reflexed tepals may function as a landing platform for pollinators in subsp. silvestris, the differently shaped flower of subsp. orinocensis may suggest a different pollinator. Key to the Subspecies of S. orinocensis 1. Leaves radical, 0.2–1.5 cm wide; inflorescence mostly exceeding the leaves (if not, then leaves < 0.3 cm wide); rhizome short ...................... subsp. orinocensis 1. Leaves scattered, 1.5–2.5 cm wide; leaves exceeding the inflorescence; rhizome long-creeping ........... subsp. silvestris S. orinocensis subsp. orinocensis Schiekia flavescens Maury, J. Bot. (Morot) 16: 269, 1889, fig. 12 annotated as “Schieckia congesta.” Schiekia orinocensis subsp. savannarum Maguire & Wurdack, Mem. New York Bot. Gard. 9: 320. 1957. Moist savannas, sometimes on granitic outcrops, 50–600 m; Bolívar (Río Ore affluent of Río Parguaza), Amazonas (Caño Caname, Caño San Miguel, base of Cerro Yapacana, Piedra Pintado, Puerto Ayacucho area, Río Atabapo). Guárico, Trujillo; Colombia (Boyacá, Guainía, Meta, Vichada), Guyana, Suriname, Brazil (Amazonas, Goiás, Maranhão, Mato Grosso, Pará, Rondônia, Roraima). ŠFig. 499.

580

H AEMODORACEAE

Fig. 500. Xiphidium caeruleum

H ALORAGACEAE 581

S. orinocensis subsp. silvestris Maas & Stoel, Fl. Neotrop. 61: 21. 1993. Forest edges, often on white sand or on granitic outcrops, sandy river banks, around small plantations, 50–200(–600 m); Delta Amacuro (San Víctor, Serranía de Imataca), Bolívar (southwest of Caicara), Amazonas (Caño Caname, Chapezón, Nericagua, Río

Atacavi, upper Río Autana, Río Guayapo, middle Río Pasimoni, lower Río Ventuari, abandoned road from San Antonio to San Fernando de Atabapo, San Pedro de Cataniapo, San Carlos de Río Negro). Amazonian Colombia (Vaupés), Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará, Roraima).

3. XIPHIDIUM Aubl., Hist. Pl. Guiane 33. 1775. Rhizomatose, stoloniferous herbs rooting at the nodes. Leaves mainly radical; margins serrulate. Inflorescence composed of many cincinni; cincinni 3–25-flowered, simple or branched. Flowers subactinomorphic, tepals slightly unequal in size, the adaxial outer petal connate with the 2 adaxial inner ones. Stamens 3, the abaxial stamen longer than the 2 adaxial ones. Capsule with many seeds per locule. Seeds subglobose, tuberculate. Neotropics; 2 species, 1 in Venezuela. Xiphidium caeruleum Aubl., Hist. Pl. Guiane 33, t. 11. 1775. Ixia xiphidium Loefl., Iter Hispan. 179. 1758. Xiphidium floribundum Sw., Prod. 17. 1788, Ixia xiphidium Loefl., as synonym. Xiphidium fockeanum Miq., Linnaea 17: 63. 1843. Xiphidium giganteum Lindl., Edward’s Bot. Reg. 32: t. 66. 1846. Herb 15–180 cm tall, rhizomes horizontally creeping, 4–20 cm long, latex red; stolons 5–55 cm long; leaves 20–65 × 1.5–6 cm; inflorescence 2–44 × 2–15 cm, the cincinni 5–

25-flowered, sometimes with young plantlets; flowers white to yellowish white, sometimes with yellow honey-guide; the 3 adaxial tepals basally adnate, 4–13 mm long; capsule green when young, maturing shiny orange, red, and finally black, (sub)globose, 5– 10 mm diameter. Sandstone or granitic substrate, open or shaded areas in forests, forest edges, disturbed forests, river banks, tops of exposed granite boulders, 0–1500 m; Delta Amacuro (scattered), northern Bolívar, Amazonas (along Río Orinoco). Widespread in Venezuela and the Neotropics, also cultivated in several neotropical countries. ŠFig. 500.

HALORAGACEAE by Luther J. Raechal Annual or perennial, aquatic or terrestrial herbs, sometimes with a persistent woody base. Leaves alternate, opposite, or whorled, diverse in form and size, often pinnately veined, exstipulate. Flowers solitary and axillary, or in spikes, racemes, or panicles, usually very small, unisexual or bisexual (plants polygamous or hermaphroditic), individually subtended by a pair of bracteoles. Sepals (3)4, valvate, persistent in fruit; petals usually 4, sometimes 3 or absent, commonly deciduous. Stamens usually 8, sometimes (3)4, in 1 or 2 whorls, the outer set opposite the sepals; filaments mostly short or very short; anthers large, opening by longitudinal slits. Ovary inferior, (2)3or 4-carpellate and -locular (partitions sometimes poorly developed); styles distinct, feathery. Fruit small, nut-like or drupaceous, sometimes separating into distinct mericarps. Endosperm present. Cosmopolitan; 8 genera and 100 species, 1 species in the flora area.

582

H ALORAGACEAE

1. LAUREMBERGIA P.J. Bergius, Descr. Pl. Cap. 350. 1767. Serpicula L., Mant. Pl. 3: 16. 1770. Single or branched, glabrous to pilose herbs, with woody rhizome. Leaves opposite, alternate, or subverticillate. Flowers miniscule, in congested inflorescences, staminate or bisexual ones pedicellate, pistillate ones sessile. Calyx 4-lobed with an ellipsoid or urceolate tube, 8-veined, generally with costas between the veins; petals 4 or rudimentary to abortive in pistillate flowers. Stamens 4, 8, or absent. Ovary initially 4-locular with 1 ovule per locule, becoming 1-locular; styles 4 or absent; stigma plumose or capitate and papillose. Fruit small, nut-like with woody endocarp, with 4–8 well-defined or inconspicuous costas, with persistent calyx lobes. Seeds with membranous testa. Pantropics, introduced in Australia and New Zealand; 4 species, 1 in Venezuela. Laurembergia tetrandra (Schott ex Spreng.) Kanitz in Mart., Fl. Bras. 13(2): 378. 1882. —Haloragis tetrandra Schott ex Spreng., Syst. Veg. 4(cur. post.): 405. 1827. Succulent herb with elongate, ascending stems and dull red flowers. Edges of rivers or swampy meadows, 800–2200 m; Bolívar (Gran Sabana, Macizo del Chimantá). Lara, Sucre; Peru, Brazil (Bahia to Santa Catarina). ŠFig. 501.

Fruit Staminate flower

Cross section of pistillate flower Fig. 501. Laurembergia tetrandra

Heliconia 583

HELICONIACEAE by Lennart L. Andersson Rhizomatic herbs, often very tall and with gigantic leaves; vegetative stems mostly very short, more rarely (in smaller species) elongated with leaves dispersed along the stem. Leaves alternate, distichous, differentiated into sheath, petiole (sometimes indistinct), and blade; sheaths tightly clasping, in acaulescent species forming a pseudostem, without a distinct ligule; blade entire (sometimes torn into narrow segments), with a coarse midrib and closely set, parallel, slightly sigmoid lateral veins fusing near margin, lateral veins interconnected by tertiary cross veins. Inflorescence terminal, structurally well defined, of cincinnate, much-condensed flower clusters in the axils of spathaceous bracts, flowers bracteolate, bracteoles keeled. Flowers zygomorphic, basically 3-merous and 5-cyclic. Perianth and androecium fused to form a closed tube at base, perianth fused above that to form an open tube with an abaxial slit. Androecium of 5 stamens and a ± scale-like staminode; filaments free above the closed tube; staminode inserted in the slit of the open tube. Ovary inferior, 3-locular; ovules anatropous, 1 in each locule, subbasal; style 1, filiform, stigma capitate. Fruit a drupe with 1–3 irregularly rugose, operculate pyrenes. Seed lacking aril. Tropical and subtropical America, Malesia, Melanesia, east to Samoa; 1 genus and 160 species, 14 species in the flora area. 1. HELICONIA L., Mant. Pl. 147. 1771. Characters and distribution as in the family; ca. 160 species, 18 in Venezuela, 14 of these in the flora area. Key to the Species of Heliconia 1. 1. 2(1). 2. 3(2).

3. 4(3). 4.

5(1).

5.

Inflorescence pendent ................................................................................ 2 Inflorescence erect ..................................................................................... 5 Spathes solid red; rachis with a brownish, floccose, cobweb-like indument; perianth white ............................................................................ H. revoluta Spathes red with yellow or green margin; rachis hirtellous to hirsute; perianth yellow or green ........................................................................... 3 Spathes waxy, pink to dull reddish with green margin; pedicels of laterformed flowers longer than those of earlier ones; perianth green ......... ................................................................................................. H. chartacea Spathes not waxy, bright red with yellow margin; pedicels of later-formed flowers shorter than those of earlier ones; perianth yellow ................. 4 Leaf blades subcoriaceous, stiffly erect, remaining ± intact throughout their life span; bracteoles stiff and persistent in fruit ......... H. marginata Leaf blades herbaceous, arching, old ones torn along lateral veins and ± conspicuously dilacerated; bracteoles membranous, dissolving at anthesis ............................................................................... H. platystachys Spathes shed with the mature fruit while top of inflorescence is still actively growing, the result being a small, compact inflorescence on top of a naked and conspicuously scarred rachis ........................... H. episcopalis Spathes persistent, the whole inflorescence wilting at the same time .... 6

584

H ELICONIACEAE

6(5). 6.

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

10(6).

10.

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

Flowers largely hidden within the spathes at anthesis; spathes usually ± deeply boat-shaped, > 2 cm deep when dry (ca. 2.5 cm when fresh) ... 7 Flowers wholly (including ovary) exposed at anthesis; spathes ± shallowly trough-shaped and usually < 2 cm deep when dry (ca. 2.5 cm when fresh) .................................................................................................... 10 Flowers resupinate, orange with blackish green “eye-spot” near apex ................................................................................................. H. densiflora Flowers not resupinate, yellow, white, or whitish and green ................... 8 Plants usually < 1 m tall; spathes greenish outside, dark purplish inside; perianth pure white ............................................................... H. lourteigiae Plants usually > 1 m tall; spathes bright red and/or yellow outside, dull yellowish inside; perianth yellow or greenish ...................................... 9 Spathes with ± sharply defined green border, tip straight; perianth white at base shading into pale green distally ........................................ H. bihai Spathes without sharply defined green margin (usually solid red in the flora area), tips circinately recurved from late bud onward; perianth yellow ............................................................................ H. spathocircinada Leaves all ± sessile, petiole, when distinguishable, < 1 cm long and all ± same length; perianth strongly flattened laterally, narrowly V-shaped in cross section ............................................................................ H. hirsuta Lower leaves clearly petiolate, petioles well over 1 cm long, those toward base of plant progressively longer; perianth ± isosceles-triangular (to subelliptic or subcircular) in cross section .......................................... 11 Perianth strongly sigmoid; inflorescence with bright red rachis and yellow to greenish yellow spathes ............................................ H. richardiana Perianth straight or almost so; inflorescence not with above combination of colors ................................................................................................ 12 Rachis ± hirsute or hirtellous, at least on inner flattened side; plants cannoid (Canna-like, leaves dispersed along stem) ................... H. julianii Rachis glabrous or with ± ephemeral cobweb-like indument; plants ± musoid (Musa-like, leaves clustered at the base) ............................... 13 Spathes diverging from rachis at an angle of < 40°; perianth obtusely angled in cross section; plant totally glabrous .................... H. psittacorum Spathes diverging from rachis at an angle of 55–100°; perianth sharply angled in cross section; plants usually with an inconspicuous cobweblike indument on rachis, pedicels, and midrib of lower surface of leaf ................................................................................................ H. acuminata

Heliconia acuminata Rich., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 15: 26, t. 11. 1831. Rather slender musoid plant, usually 1–2 m tall. Well-drained ground in forest understory and small light-gaps. Widespread in southern Venezuela, Guyana, Suriname, French Guiana, Amazonian Colombia, Peru, and Brazil; 4 subspecies, 2 in Venezuela, both in the flora area. The other two subspecies occur in the Trapecio Amazónico area of Colombia, Peru, and Brazil (subsp. psittacorastra L. Anders-

son) and in the Manú area of southeastern Peru (subsp. immaculata L. Andersson). Key to the Subspecies of H. acuminata 1. Perianth usually < 4.5 cm long, dull green with white or yellowish tip, or yellow to white with distinct “eye-spots” toward tip, spot < 5 mm from tip ........................ ............................... subsp. acuminata 1. Perianth usually > 4.5 cm long, yellowish or white with distinct “eye-spots” > 5 mm from tip ............... subsp. occidentalis

Heliconia 585

H. acuminata subsp. acuminata Evergreen lowland to montane forests and forest openings, 50–1000 m; widespread in Bolívar and Amazonas. Guyana, Suriname, French Guiana, northeastern Brazil. ŠFig. 505. H. acuminata subsp. occidentalis L. Andersson, Opera Bot. 82: 61. 1985. Herb to 2 m tall. Rock outcrops, 100 (–1000) m; Amazonas (basin of Río Negro and Río Orinoco). Brazil (Amazonas, Pará). Heliconia bihai (L.) L., Mant. Pl. 211. 1771. —Musa bihai L., Sp. Pl. 1043. 1753. Heliconia aurea G. Rodr., Bol. Soc. Venez. Ci. Nat. 15: 120. 1954. Heliconia schaeferiana G. Rodr., Acta Biol. Venez. 1: 210. 1954. Heliconia humilis auct. non Musa humilis Aubl.: sensu Jacq., Pl. Rar. Hort. Caes. Schoenbr. t. 48. 1797 (plant, not name; note that Heliconia humilis sensu Aristeg. in Gén. Heliconia Venez., is Heliconia stricta Huber). Heliconia caribaea auct. non Lam. 1783 [1785], nec Aristeg. 1961. Robust musoid plant 2–5 m tall. Along pool and river margins, on scree grounds, roadsides, sea level to 1200 m; widespread in Delta Amacuro and Bolívar. Venezuelan Coastal Cordillera; Guyana, Suriname, French Guiana, Brazil (southwestern Pará). ŠFig. 506. Heliconia chartacea Lane ex Barreiros, Revista Brasil. Biol. 32: 205. 1972. Robust cannoid plant 3–5 m tall. Young secondary growth in clearings and light-gaps in rain forests, on well-drained ground, 50– 600 m; scattered in Bolívar and Amazonas. Guyana, Suriname, French Guiana, central and western Amazon basin. ŠFig. 508. Heliconia densiflora B. Verl., Rev. Hort. 40: 112. 1869. Rather slender plant 1–3 m tall. Welldrained ground in forest understory and small light-gaps, 200–500 m; Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 2 subspecies, both in the flora area. Key to the Subspecies of H. densiflora 1. Petiole usually < 25% of blade length; blade width usually < 25% of length ............................. subsp. angustifolia

1. Petiole usually > 25% of blade length, blade width usually > 25% of length ................................ subsp. densiflora H.

densiflora subsp. angustifolia L. Andersson, Opera Bot. 82: 77. 1985. Southeastern Bolívar. Guyana, Brazil (Pará). H. densiflora subsp. densiflora Bolívar (Canaima, La Vergareña, San Juan de Manapiare). Southeastern Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Acre, Amapá, Amazonas, Pará), Bolivia. ŠFig. 502. Heliconia episcopalis Vell., Fl. Flumin. 107. 1825 [1829]. Musoid plant 1.5–4 m tall. Riversides, light swamp forests, secondary vegetation on moist to swampy ground, ca. 800 m; Amazonas (upper Río Orinoco). Maracaibo Basin, western Llanos; Colombia, Ecuador, Peru, Brazil, Bolivia. Heliconia hirsuta L. f., Suppl. Pl. 158. 1781 [1782]. Heliconia costanensis Aristeg., Bol. Soc. Venez. Ci. Nat. 25: 206. 1964. Slender plant, usually 1–3 m tall; leaves ± sessile, dispersed ± all along the stem. Semideciduous to pluvial forests, usually on well-drained ground in light-gaps, sea level to 1100 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Common throughout Venezuela except in arid regions; throughout tropical America. ŠFig. 504. Heliconia julianii Barreiros, Rodriguésia 41: 130, fig. 2. 1976. Slender cannoid plant 1–3 m tall. Edges of and light-gaps in swamp forests and varzea, 100–200 m; Amazonas (basins of Río Negro and Río Orinoco). Colombia, Ecuador, Peru, Brazil (Acre, Amazonas, Rondônia, Mato Grosso). ŠFig. 511. Heliconia lourteigiae Emygdio & Santos, Bol. Mus. Nac. Rio de Janeiro 43: 1. 1977. Slender musoid plant to 0.8 m tall. Rainforests, usually on poorly drained soils, ca. 500 m; Bolívar (Río Samay affluent of Río Icabarú). Guyana, Suriname, French Guiana, Brazil (Amazonas, Roraima).

586

H ELICONIACEAE

Heliconia marginata (Griggs) Pittier, Man. Pl. Usual. Venez. 299. 1926. —Bihai marginata Griggs, Bull. Torrey Bot. Club 42: 323. 1915. Heliconia marginata f. lutea Aristeg., Gén. Heliconia Venez. t. 19a. 1961. Robust musoid plant 1.5–5 m tall. Seasonal swamps, 0–100 m; Delta Amacuro (widespread). Maracaibo Basin, Llanos; Costa Rica, Panama, most of tropical South America. ŠFig. 509. Heliconia platystachys Baker, Ann. Bot. 7: 194. 1893. Robust cannoid plant 3–6 m tall. Edges and light-gaps of seasonal evergreen to semideciduous forests on well-drained grounds, ca. 200 m; Bolívar (widespread). Maracaibo Basin, northwestern Llanos; Panama, Colombia. ŠFig. 503.

Bolívar (southwest of El Manteco). Monagas; Guyana, Suriname, French Guiana, Brazil (southern Bahia, Espírito Santo, Maranhão, Pará). Heliconia spathocircinada Aristeg., Bol. Soc. Venez. Ci. Nat. 22: 18. 1961. Rather slender to rather robust musoid plant, 1–3 m tall. Well-drained ground in forest understory and light-gaps, 100–800 m; widespread in Delta Amacuro and Bolívar. Monagas, Sucre; Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil (Bahia). ŠFig. 510.

Heliconia psittacorum L. f., Suppl. Pl. 156. 1781 [1782]. Heliconia cannoidea Rich., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 15: 24, t. 9–10. 1831. Heliconia psittacorum var. rhizomatosa Aristeg., Bol. Soc. Venez. Ci. Nat. 22: 20. 1961. Slender musoid plant, usually 0.5–1.5 m tall. Savanna scrub, especially on seasonally flooded ground, 0–1200 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Wide distribution over most of tropical South America but absent from the Western Amazon Basin. ŠFig. 507. Heliconia revoluta (Griggs) Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 4. 1930. —Bihai revoluta Griggs, Bull. Torrey Bot. Club 42: 322. 1915. Robust musoid plant, usually 2–4 m tall. Not yet known from the flora area but may be expected at ca. 2000 m in the vicinity of Sierra de la Neblina. Coastal Cordillera; Colombia, Brazil (Amazonas: Serra da Neblina). Heliconia richardiana Miq., Linnaea 18: 70. 1844. Heliconia glauca Poit. ex B. Verl., Rev. Hort. 40: 112. 1869. Rather slender musoid plant 1–2 m tall. Moist to swampy ground in forest understory and small light-gaps, 100–200 m; Delta Amacuro (La Paloma in Río Cuyubini basin),

Fig. 502. Heliconia densiflora subsp. densiflora

Heliconia 587

Fig. 503. Heliconia platystachys

Fig. 505. Heliconia acuminata subsp. acuminata

Fig. 504. Heliconia hirsuta

588

H ELICONIACEAE

Fig. 506. Heliconia bihai Fig. 507. Heliconia psittacorum

Heliconia 589

Fig. 508. Heliconia chartacea

Fig. 509. Heliconia marginata

590

H ELICONIACEAE

Fig. 510. Heliconia spathocircinada Fig. 511. Heliconia julianii

Hernandia 591

Fig. 512. Hernandia guianensis

592

H ERNANDIACEAE

HERNANDIACEAE by James S. Miller and Paul E. Berry Trees, shrubs, or lianas. Leaves alternate, simple (sometimes lobed) or palmately compound, exstipulate. Inflorescences cymose, paniculate, thyrsiform, or corymbose, axillary. Flowers small, bisexual or unisexual. Perianth of 4–8 parts in a single whorl or 3–5 parts in 2 whorls. Stamens 3–5(–7), opposite the outer tepals when the perianth is double, alternate with the tepals when the perianth is single; filaments often with nectariferous appendages; anthers bithecal, dehiscing longitudinally by 2 valves. Ovary inferior, 1-locular with a single pendulous ovule. Fruit dry or drupaceous, 1-seeded, lightly ridged to prominently winged. Seeds lacking endosperm, cotyledons wrinkled or flat, large. Pantropics; 4 genera and ca. 60 species, 2 genera and 3 species in the flora area. Key to the Genera of Hernandiaceae 1. 1.

Trees; flowers unisexual, greenish yellow ................................. 1. Hernandia Lianas or small trees; flowers bisexual, grayish white .... 2. Sparattanthelium

1. HERNANDIA L., Sp. Pl. 981. 1753. Soft-wooded, monoecious, evergreen trees. Leaves alternate, simple, usually broad, entire, long petiolate. Inflorescences clustered near the ends of branches, paniculate or thyrsiform, long-pedunculate, bracteate, consisting of involucrate clusters of 2 pedicellate staminate flowers with 1 subsessile pistillate flower. Staminate flowers with 3 inner and 3 outer perianth lobes, these connate basally into an elongate tube, with 3 stamens and 3–6 glandular staminodia; pistillate flowers with 4 outer and 3 inner perianth lobes, these connate basally into a tube, the entire flower subtended by a globose cupule, ovary inferior, unilocular; stigma fleshy, surrounded by 4 glandular staminodia, these sessile in the mouth of the perianth tube. Fruits borne in the greatly enlarged, accrescent cupule, drupaceous, ovoid to ellipsoid. Pantropics (mostly tropical Asia through Oceania); 24 species, 1 in Venezuela. Hernandia guianensis Aubl., Hist. Pl. Guiane 848, t. 329. 1775. —Aguacatillo blanco, Cajoro, Cococoroli, Cocojoro, Coro-cororu. Hernandia sonora Kosterm., Med. Bot. Mus. Utrecht 25: 45. 1936, non L. 1763. Tree 5–10(–30) m tall. Riparian forests,

evergreen lowland to lower montane forests, 0–400 m; Delta Amacuro (Caño Araguabisi, between Caño Guayo and Caño Sacupana, Caño Güiniquina, Caño Jobure, Río Toro), Bolívar (Altiplanicie de Nuria). Monagas, Sucre; Trinidad, Guyana, Suriname, French Guiana, Amazonian Brazil. ŠFig. 512.

2. SPARATTANTHELIUM Mart., Flora 24(2) Beibl.: 40. 1841. Evergreen shrubs or lianas. Leaves alternate, simple, lanceolate to narrowly elliptic or ovate, trinerved or triplinerved, usually with distinctly percurrent tertiary veins, margin entire, short-petiolate. Inflorescence a large, lax axillary or terminal dichasium, ebracteate. Flowers bisexual; perianth 4- or 5(–7)-lobed, in a single whorl. Stamens 4 or 5(–7), alternate with the calyx lobes; filaments glabrous; anthers elongate, apiculate. Infructescence dichotomous, pendent, with nodular thickenings at points of ramification. Fruits drupaceous, ellipsoid to ovoid. Neotropics; 13 species, 2 in Venezuela, both in the flora area.

Sparattanthelium 593

Key to the Species of Sparattanthelium 1. 1.

Lower surface of mature leaves only slightly pubescent along the veins ........................................................................................ S. tupiniquinorum Lower surface of mature leaves densely gray-tomentulose ..... S. uncigerum

Sparattanthelium tupiniquinorum Mart., Flora 24(2) Beibl. 40. 1841. Liana or small tree to 5 m tall. Evergreen lowland to lower montane forests, 50–400 m; Delta Amacuro (Serranía de Imataca), Bolívar (Río Botanamo, Serranía de Parguaza), Amazonas (Puerto Ayacucho, San Carlos de Río Negro, Yavita, Yutajé). Guyana, Suriname, Brazil. ŠFig. 513.

Sparattanthelium uncigerum (Meisn.) Kubitzki, Bot. Jahrb. Syst. 89: 196. 1969. —Sparattanthelium botucudorum var. uncigerum Meisn. in Mart., Fl. Bras. 5(2): 293. 1866. Liana or small tree. Evergreen lowland and lower montane forests, 100–600 m; Bolívar (Río Botanamo, Serranía de los Pijiguaos). Guyana, Suriname.

Fig. 513. Sparattanthelium tupiniquinorum

594

H IPPOCRATEACEAE

HIPPOCRATEACEAE by Alberta M. W. Mennega and Jennifer P. Hedin Trees, shrubs, scandent shrubs, or lianas, generally glabrous, pilose or scabrous in a few genera. Branches mostly opposite. Leaves nearly always opposite or pseudo-opposite, simple, petiolate; stipules minute or absent; blades with margins entire, undulate, or crenate-serrate. Inflorescences axillary, cymose, corymbose, paniculate, thyrsoid-paniculate, or fasciculate. Flowers actinomorphic, usually very small (2–4 mm) to small (8–10 mm), rarely over 15 mm diameter (to 30 mm in some species of Salacia), bracteolate, pedicellate. Sepals 5, imbricate, basally united, often persisting; petals 5, free, inserted under the disk, imbricate. Disk always present, variable, an annular ring, or cushion-shaped, cup-shaped, pulvinate-fleshy, truncate, or with a flat basal part and a truncate central portion, continuous or consisting of 3(5) staminal pockets (Cheiloclinium). Stamens 3(5), inserted within the disk; filaments ligulate, often conspicuously widened towards the base, often recurved in the open flower; anthers basifixed, 2-locular, dehiscing by extrorse, horizontal or oblique apical clefts (vertical clefts in Peritassa). Ovary superior, often immersed in the disk or adnate to the disk by partitions, 3-locular; ovules 2–14 per locule, collateral, superposed, or 2-ranked; style short or absent; stigmas obscure or conspicuous, entire or bifid, free or ± adnate to summit of ovary, alternate or opposite the stamens. Fruits capsules or drupes, dehiscent or indehiscent; capsular fruits with 3 mericarps, free or laterally ± united; drupaceous fruits with an indehiscent pericarp, leathery, fleshy, or woody. Seeds in capsular fruits few to numerous, attached by a basal wing of various size; in the drupaceous fruits, seeds usually embedded in a mucilaginous pulp. Mainly tropics, also subtropics of both hemispheres; 24 genera and 356 species; 11 genera and 38 species in the flora area. The genera of Hippocrateaceae probably belong in Celastraceae as a subfamily, but do not form a monophyletic group. The genera with indehiscent fruit (four in the flora area) appear to be closely related, but the genera with dehiscent fruit probably are not. In this flora, the two families are treated separately as a matter of convenience. Vegetatively, Hippocrateaceae can often be recognized by the presence of scattered, few to numerous, small, circular black spots (corky warts around large stomata) on the lower surface of the leaves, and occasionally also on the upper surface. Yellowish exudations on branches, petioles, or inflorescences may be found in several genera, mostly in Cheiloclinium. Rubberoid threads often appear on breaking of petioles, pedicels, or other organs in several species of various genera, visible in both fresh and dried material. To distinguish between the genera with capsular versus drupaceous fruits, inspection of the cross section of a twig may be helpful. In the genera with capsular fruits (Anthodon, Cuervea, Elachyptera, Hippocratea, Hylenaea, Prionostemma, and Pristimera), the wood rays are wide and conspicuous, often appearing as lighter-colored spokes of a wheel, whereas in the genera with drupes (Cheiloclinium, Peritassa, Salacia, and Tontelea), the rays are narrow and not at all conspicuous. In mature wood of the latter genera, the structure of the wood is often anomalous by the occurrence of included phloem either as isolated strands or imbedded in concentric bands of sclerenchymatic tissue. Anomalous wood structure does not occur in

H IPPOCRATEACEAE 595

the group with capsular fruits (in Hippocratea, the bark forms indentations in the woody cylinder). Key to the Genera of Hippocrateaceae 1.

1.

2(1).

2. 3(2). 3.

4(3).

4.

5(3). 5. 6(5).

6.

Leaves scabrous; red latex present; flowers ca. 10 mm wide, greenish, the petals clawed at the base, the apex pectinate; disk flattened; fruit with 3 separately attached mericarps, each ca. 6–8 × 5 cm, the pericarp thick, woody, scabrous ...................................................... 8. Prionostemma Leaves smooth and glabrous; latex absent; flowers without above combination of characters, the petals not clawed, margins serrate, erose, or fimbriate; fruits a glabrous capsule with either free or connate mericarps or a drupe.............................................................................. 2 Disk consisting of 3(5) small staminiferous pockets; stigmas sessile, for the greater part adnate to ovary; anthers small; fruit a round to ovoid, small to medium-sized drupe, often with 3 longitudinal stripes at the base .................................................................................... 2. Cheiloclinium Disk various, but without staminiferous pockets; style present; anthers small to relatively large; fruit a capsule or a drupe ............................. 3 Disk fleshy, annular-pulvinate or conical, often the basal part considerably flattened or upturned; stigmas obscure ........................................ 4 Disk membranous to subfleshy, forming a flat inconspicuous ring, or cupshaped with a short to very short rim; stigmas conspicuous or obscure ................................................................................................................ 5 Inflorescence corymbose-paniculate, young parts densely brown-puberulent; leaves medium-sized, 3–14 cm long; secondary veins equidistant; flowers 4–8 mm wide, often laterally arranged along ultimate branchlets, cream to yellowish green, the petals transversely barbellate; disk conspicuous, fleshy, truncate-conical, puberulous; fruit a capsule with free, elliptic, thin-walled mericarps, the seeds affixed by a conspicuous basal papery wing ................................................................ 5. Hippocratea Inflorescence either fasciculate or branching, glabrous (in S. amplectens brown-tomentellous); leaves medium-sized or very large; in the middle part of the large leaves the secondary veins very wide apart, the tertiary venation parallel, often impressed or flat; flowers small to large, 2–30 mm wide, in the larger type of flowers the petals orbiculate and flabellately veined; disk variable, annular-pulvinate, the base flattened or not, the apex rounded or produced in thin lobes between the stamens; fruit a drupe, subglobose, ellipsoid, often large, to 10 cm diameter, the seeds embedded in a mucilaginous pulp .................... 10. Salacia Anther locules dehiscing by vertical clefts, the connective in some species conspicuously produced; fruit a drupe, often pruinose ........... 7. Peritassa Anther locules dehiscing by horizontal confluent clefts or by oblique clefts; fruit a capsule or a drupe ............................................................ 6 Petals conspicuously and regularly serrate; ovary with 8–14 ovules per locule; fruits flat, to 18 cm diameter, the 3 mericarps connate at the base for 2–4 cm, but completely surrounded by a 3-lobed thin-coriaceous, broad rim ....................................................................... 1. Anthodon Petals entire or ± erosulous, but not regularly serrate; ovules 2–6 per loc-

596

H IPPOCRATEACEAE

ule; fruit a capsule with free mericarps or a drupe .............................. 7 Flowers relatively large, 10–18 mm wide, white, fragrant; inflorescence fewflowered, 5–12 cm long; peduncle long, slender; leaves with 6–9 pairs of secondary veins; fruits capsular with 3 large, broadly subovate to orbicular convex mericarps; seeds with a short basal wing ......... 3. Cuervea 7. Without above combination of characters ................................................. 8 8(7). Flowers minute; inflorescence 3–20 cm long, the branchlets extremely thin, capillaceous; flowers very numerous, for the greater part abortive; bark purplish, often peeling; leaves with 8–12 pairs of secondary veins; fruits with 3 large oblong mericarps, ≤ 10 cm long; seeds with a conspicuous basal wing ............................................................. 6. Hylenaea 8. Without above combination of characters ................................................. 9 9(8). Leaves chartaceous, subcoriaceous to coriaceous (papery in Tontelea cylindrocarpa), margin usually entire; intersecondary veins (shorter secondary veins between and parallel to full-length secondary veins) often prominent; flowers generally ca. 4 mm wide; disk cup-shaped 0.2–0.4 mm tall; style with 3 entire or ± split stigmas, opposite or alternate with the stamens, seldom stigmas obscure (Tontelea cylindrocarpa); anthers usually rather large, butterfly-like; fruit a drupe, ellipsoid to globose or cylindric, in some species very large (16 cm long and ca. 4.5 cm diameter) .................................................................. 11. Tontelea 9. Leaves papery, chartaceous, or subcoriaceous, margin crenulate or undulate, intersecondary veins not obvious; flowers 2–4 mm wide; disk minute, flat or cup-shaped; style with obscure stigmas or ending in a minute triangular stigmatic field; anthers small, not butterfly-like; fruit a capsule; seed with a conspicuous basal wing (narrow ring in Pristimera tenuiflora) .......................................................................... 10 10(9). Inflorescence usually copiously branched, 2–11 cm long, axillary, often with accessory inflorescences in the leaf axils; pedicels 1–2 mm long; flowers minute, fragrant, white or creamish, the petals suberect, subfleshy; disk cup-shaped, the rim ca. 0.1 mm tall; upper surface of blades ± shining; tertiary venation flat on upper surface, prominulous on lower surface, not yellowish ........................................... 4. Elachyptera 10. Inflorescence compact or ca. 5 times branched, < 7 cm long, usually much shorter, solitary in the leaf axils; pedicels absent or < 1 mm long; flowers small, not fragrant, greenish or white, the petals spreading (suberect in Pristimera nervosa); disk either flat or with a short erect rim; upper surface of blades not shiny, tertiary venation prominulous on both sides, when dry the veins often yellowish ..................... 9. Pristimera 7(6).

1. ANTHODON Ruiz & Pav., Fl. Peruv. 1: 45. 1798. Lianas, occasionally trees, glabrous throughout. Leaves opposite, crenulate. Inflorescences axillary, cymose, 3–6 times dichotomously branched, pedunculate. Flowers 5–11 mm wide, white, yellowish, or green, subtended by 2 or 3 minute bracteoles. Pedicels slender, very short. Sepals orbiculate, the margins slightly serrate; petals spreading, oblong, the margins conspicuously serrate. Disk short-cylindric with a semi-erect short, undulate margin, subfleshy. Stamens 3, erect; filaments short, ligulate; anthers broadly reniform, dehiscing by elongate, horizontal, apical clefts. Ovary depressed-trigonous; ovules 8–14 per locule; style stout, short, crowned by a triangular stigmatic knob. Fruit large, capsular, flattened, the 3 mericarps con-

Cheiloclinium 597

Fig. 514. Anthodon decussatum

nate at basal margins for about half their length, dehiscing along a median suture, adjacent halves coherent; pericarp coriaceous, flabellate-costate without. Seeds 8– 14, erect from base, affixed by a basal wing, the portion containing the embryo apical, coriaceous, much shorter than the membranous wing. Panama, Colombia, Venezuela, French Guiana, Peru, Brazil, Bolivia, Paraguay; 2 species, 1 in Venezuela. Anthodon decussatum Ruiz & Pav., Fl. Peruv. 1: 45. 1798. —Curabayek (Arekuna), Mulato. Salacia brevistaminea Pittier, Bol. Soc. Venez. Ci. Nat. 5: 309. 1939. Liana. Disturbed evergreen lowland to lower montane forests, 100–900 m; Bolívar

(Caño Cañi 40 km northwest of Salto Pará, 20 km east of La Paragua, Río Karún), Amazonas (Isla Ratón, Río Cunucunuma, base of Sierra de la Neblina). Coastal Cordillera, Falcón, Mérida, Sucre; Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ŠFig. 514.

2. CHEILOCLINIUM Miers, Trans. Linn. Soc. London 28: 420. 1872. Glabrous shrubs or trees with scandent branches, or lianas; much of plant, especially the inflorescences, flowers, and fruits often covered by a bright yellow exudate. Leaves opposite, petiolate; margins entire, crenulate or serrulate; tertiary venation often parallel, subparallel, or orthogonal-reticulate; petiole often with threads. Inflorescences axillary, thyrsoid, corymbose or cymose, dichotomously branched; peduncle very short to relatively long, 2–11 cm long. Flowers small, 2–4 mm wide, sessile or pedicellate, often crowded, bracteoles very small. Sepals 5, imbricate, orbiculate or ovate, the margin often erosulous; petals 5, imbricate, oblong, thin-fleshy, often the lower part at the inner side thickened in the middle, threads sometimes in sepals and petals. Disk discontinuous, forming 3(5) saccate staminal pockets. Stamens 3 (5 in Cheiloclinium anomalum), inserted in the pockets of the

598

H IPPOCRATEACEAE

disk and enclosed by its lips; filaments thin, short-erect; anthers very small, 2-locular, transversely ellipsoid, opening by extrorse, confluent, horizontal clefts. Ovary trigonous (pentagonous in Cheiloclinium anomalum), subglobose, the apex truncate, locules with 2–4 ovules; style absent; stigmas 3(5) opposite the stamens, adnate to the apex of the ovary, linear or oblong-deltoid with ± free, entire or bilobed apices. Fruits drupaceous, mostly medium-sized, ovate to globose, orange-brown or green, 3(–5)-locular, the septa soon evanescent; pericarp coriaceous, often with 3 longitudinal lighter-colored stripes in the basal part. Seeds ovate-angular, embedded in a mucilaginous pulp. Neotropics from Central America south to Paraguay; 11 species, 8 in Venezuela, all in the flora area. Key to the Species of Cheiloclinium 1.

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

4.

5(4). 5.

6(3).

6.

7(6).

7.

Inflorescence thyrsoid with a straight rachis; flowers brownish red, orange, or dark yellow with a crimson margin on the petals; generally shrubs or trees to 20 m tall, seldom lianas; petiole with threads .................................................................................................. C. cognatum Inflorescence dichotomously branched; flowers white, yellowish, or greenish; generally lianas, occasionally trees; petiole without threads ....... 2 Stamens, stigmas, and locules 5; fruit ovate, ≤ 6 cm long and 3.5 cm diameter, greenish, drying purplish brown, pericarp smooth ...... C. anomalum Stamens, stigmas, and locules 3; fruit not as above ................................. 3 Inflorescence loosely branched, 4–11 cm long, the peduncle slender, 1–5 cm long ......................................................................................................... 4 Inflorescence compact or many times branched, subspherical, ca. 4 cm long, the peduncle absent or < 1 cm long and rather stout .................. 6 Leaf blades (when dry) dark green on upper surface, light green on lower surface; pedicels 3–4 mm long, slender; stigmas with an emarginate apex; fruits ovate, pericarp smooth, pale orange when mature ............ C. klugii Leaf blades similarly colored on both surfaces; pedicels 1–3 mm long; stigmas entire or deeply bifid; fruits roundish, yellow or with a pericarp with white lenticels ................................................................................ 5 Flowers yellowish, 2 mm wide; stigmas deeply bifid; tertiary venation of the blades hardly visible; fruit globose, yellow .................. C. habropodum Flowers white or creamish, 2–4 mm wide; stigmas linear, apex entire; tertiary venation of the blades ± raised; fruit globose to ovate, 4.5 × 3.5 cm, in clusters, the pericarp green to brown with white lenticels .... C. belizense Leaves thick-coriaceous, entire, broadly elliptic, the apex obtuse; flowers almost sessile; stigmas broad, adnate to the ovary, the apex truncate or emarginate ................................................................................. C. obtusum Leaves chartaceous, margins slightly to conspicuously crenate, the apex short-acuminate; flowers sessile or pedicellate; stigmas entire, bilobed, or emarginate ......................................................................................... 7 Leaves with tertiary venation slightly prominulous or obscure, not closely horizontally reticulate; stigma lobes bifid; fruit ellipsoid, yellow when ripe, ca. 25 × 40 mm, the pericarp smooth, with 6–10 faint ribs at the base ................................................................................. C. hippocrateoides Leaves with tertiary venation closely subhorizontally reticulate, ± prominulous; stigma lobes entire, narrow; fruit unknown .... C. diffusiflorum

Cheiloclinium 599

Cheiloclinium anomalum Miers, Trans. Linn. Soc. London 28: 420. 1872. —Salacia anomalum (Miers) Peyr. in Mart., Fl. Bras. 11(1): 144. 1878. Cheiloclinium glaziovii A.C. Sm., Brittonia 3: 549. 1940. Liana. Evergreen lowland to lower montane and riparian forests, 100–500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Cerro Ichún), Amazonas (Río Siapa near base of Cerro Aracamuni, Río Cunucunuma, Río Mawarinuma, Río Negro). Panama, Guyana, Suriname, Peru, Brazil, Bolivia. Cheiloclinium belizense (Standl.) A.C. Sm., Brittonia 3: 540. 1940. —Salacia belizensis Standl., Field Mus. Nat. Hist., Bot. Ser. 8: 19. 1930. Salacia podostemma Sandw., Bull. Misc. Inform. 1931: 187. 1931. —Cheiloclinium podostemmum (Sandw.) A.C. Sm., Brittonia 3: 542. 1940. Cheiloclinium jenmanii A.C. Sm., Brittonia 3: 542. 1940. Cheiloclinium meianthemum A.C. Sm., J. Arnold Arbor. 27: 88. 1946. Liana. Lower montane forests, 100–500 m; Bolívar (Cerro Ichún), Amazonas (Isla Ratón). Apure, Sucre, Táchira; Central America, Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ŠFig. 515. Cheiloclinium cognatum (Miers) A.C. Sm., Brittonia 3: 529. 1940. —Kippistia cognata Miers, Trans. Linn. Soc. London 28: 417. 1872. —Salacia cognata (Miers) Peyr. in Mart., Fl. Bras. 11(1): 144. 1878. —Cimbra pote, Gaspadillo marrón, Melocotoncito. Salacia sphaerocarpa Rusby, Descr. S. Amer. Pl. 52. 1920. Salacia lineolata A.C. Sm., Bull. Torrey Bot. Club 66: 234. 1939. —Cheiloclinium lineolatum (A.C. Sm.) A.C. Sm., Brittonia 3: 533. 1940. Shrub or slender tree 3–20 m, with scandent branches, seldom a liana. Primary to disturbed evergreen lowland to upper montane forests, 50–2000 m; Delta Amacuro (lower Río Orinoco, Río Toro), Bolívar (widespread, including Gran Sabana and tepui slopes), Amazonas (Río Casiquiare, Río Mawarinuma, Río Siapa, San Carlos de Río Negro). Miranda, Monagas, Táchira; Costa Rica, Panama, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 516.

Cheiloclinium diffusiflorum (Miers) A.C. Sm., Brittonia 3: 539. 1940. —Kippistia diffusiflora Miers, Trans. Linn. Soc. London 28: 418. 1872. —Salacia diffusiflora (Miers) Peyr. in Mart., Fl. Bras. 11(1): 143. 1878. Tontelea parviflora Miers, Trans. Linn. Soc. London 28: 386. 1872. —Cheiloclinium parviflorum (Miers) A.C. Sm., Brittonia 3: 537. 1940. Cheiloclinium lucidum A.C. Sm., Brittonia 3: 537. 1940. Liana or clambering shrub. Evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, Río Mawarinuma, upper Río Orinoco). Guyana, Suriname, French Guiana, Amazonian Brazil. Cheiloclinium habropodum A.C. Sm., Contr. U.S. Natl. Herb. 29: 332. 1950. —Gengibrillo. Liana. Evergreen lowland forests, 100– 200 m; Amazonas (Capibara). Endemic. Cheiloclinium hippocrateoides (Peyr.) A.C. Sm., Brittonia 3: 546. 1940. —Salacia hippocrateoides Peyr. in Mart., Fl. Bras. 11(1): 142. 1878. Salacia gleasoniana A.C. Sm., Lloydia 2: 191. 1939. —Cheiloclinium gleasonianum (A.C. Sm.) A.C. Sm., Brittonia 3: 546. 1940. Salacia krukovii A.C. Sm., Bull. Torrey Bot. Club 66: 237. 1939. —Cheiloclinium krukovii (A.C. Sm.) A.C. Sm., Brittonia 3: 544. 1940. Liana or low tree. Evergreen lowland to montane forests, 100–1100 m; Bolívar (Cerro Ichún, Kavanayén), Amazonas (upper Río Orinoco, Río Siapa, base of Sierra de la Neblina). Distrito Federal; Costa Rica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Cheiloclinium klugii A.C. Sm., Brittonia 3: 543. 1940. Liana. Riverine forests, white-sand savannas, scrub vegetation, 100–200 m; Bolívar (Río Caura), Amazonas (base of Cerro Sipapo). Zulia; Panama, Colombia, Peru, Brazil. Cheiloclinium obtusum A.C. Sm., Brittonia 3: 545. 1940. Liana or low tree. Montane forests on lower tepui slopes, ca. 1000 m; Bolívar (Río Tírica in the Macizo del Chimantá). Peru, Brazil.

600

H IPPOCRATEACEAE

Fig. 515. Cheiloclinium belizense

Fig. 516. Cheiloclinium cognatum

3. CUERVEA Triana ex Miers, Trans. Linn. Soc. London 28: 370. 1872. Lianas, occasionally shrubs. Leaves opposite, glabrous; stipules reduced; blades subcoriaceous, entire or crenulate. Inflorescence axillary, usually pedunculate, paniculate-corymbose. Flowers large, ca. 10 mm wide, loosely arranged, pedicellate. Sepals unequal, suborbicular, membranous; petals spreading, membranous, flabellately veined. Disk low, flat or suberect, membranous. Stamens 3, erect or reflexed; filaments ligulate, short; anthers transversely ellipsoid, dehiscing by an extrorse-apical horizontal cleft. Ovary trigonous, sulcate, gradually tapering into the short, truncate style; ovules 4–9 per locule; stigmas conspicuous or not, opposite the stamens. Fruit of 3 capsules, separately attached to a swollen receptacle, convex, rounded at the apex, dehiscing along an unobtrusive median suture; pericarp thincoriaceous. Seeds affixed by a short basal wing or without a wing. Neotropics, African tropics; 5 species, 1 in Venezuela. Cuervea kappleriana (Miq.) A.C. Sm., Brittonia 3: 396. 1940. —Hippocratea kappleriana Miq., Linnaea 26: 220. 1853. —Metsajabaja.

Hippocratea mitchellae I.M. Johnst., Contr. Gray Herb. 81: 93. 1928. —Cuervea mitchellae (I.M. Johnst.) A.C. Sm., Brittonia 3: 401. 1940.

Elachyptera 601

Seed

Fruit Fig. 517. Cuervea kappleriana

Liana or shrub, ca. 5 m tall. River banks, swampy places, near sea level to 300 m; Delta Amacuro (Río Acure, Río Cuyubini, Río Toro), Bolívar (Río Caura, Salto Pará, Serranía de Imataca), Amazonas (40 km southeast of Puerto Ayacucho). Mexico, Honduras,

Costa Rica, Panama, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay. ŠFig. 517. In the sterile state it is generally very difficult to distinguish Cuervea kappleriana from Hylenaea comosa.

4. ELACHYPTERA A.C. Sm., Brittonia 3: 383. 1940. Lianas or shrubs with scandent branches, glabrous except occasionally the inflorescence puberulent. Leaves opposite, slenderly petiolate. Inflorescences axillary or terminal, many-flowered, paniculate-corymbose, peduncle and branches quadrangular, often accessory inflorescences in a leaf axil; pedicels slender; bracteoles minute. Flowers very small, fragrant, white, purplish white, or cream. Sepals obtuse or acute at the apex; petals suberect, thin-fleshy, rounded at the apex. Disk erect, short-cylindric, inconspicuous. Stamens 3, semi-erect; filaments ligulate, short; anthers transversely ellipsoid, extrorsely nodding, dehiscing by extrorse-apical horizontal clefts. Ovary depressed-trigonous, 3-locular; ovules 2–4 per locule; style very short, with a deltoid stigmatic shield with 3 minute stigmatic knobs opposite the stamens. Fruit of 3 free mericarps, the mericarps thin, dehiscing by 2 caducous valves; pericarp thin-coriaceous, green. Seeds winged or not, the wing basal or lateral, the portion containing the embryo coriaceous. Neotropics, Africa, Asia; 7 species, 1 in Venezuela.

602

H IPPOCRATEACEAE

Elachyptera floribunda (Benth.) A.C. Sm., Brittonia 3: 383. 1940. —Hippocratea floribunda Benth., Bot. Voy. Sulphur 78. 1844. Liana, or slender tree to 15 m with scandent branches; dried leaves dark brownish, the upper surface glossy; dried leaves, flowers, or wood boiled in water yield purplish extract. Riverine, swamp, and savanna forests, 0–200 m; Delta Amacuro (Caño Arature north of Río Grande), Amazonas (Río Casiquiare, Río Cataniapo). Apure; Mexico, Guatemala, Honduras, Nicaragua, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 518.

Seed Fig. 518. Elachyptera floribunda

5. HIPPOCRATEA L., Sp. Pl. 1191. 1753. Coarse woody lianas, seldom shrubs; young parts densely and minutely brown tomentellous-papillate. Leaf blades broadly ovate to oblong-elliptic, the base rounded to acute, the margins generally conspicuously crenate or serrate, the apex rounded, obtusely short-cuspidate, or acuminate, threads in petioles. Inflorescences axillary, panicles pedunculate, the branches elongate, dichotomously or pseudodichotomously arranged, often with short lateral alternate branchlets. Flowers 4–6 mm diameter, congested toward apices of, or laterally arranged along ultimate branchlets, pedicellate, bracteolate. Sepals tomentellous, ovate to deltoid, irregularly ciliolate at margins; petals spreading, transversely barbellate within, white, cream, yellowish green, or green. Disk fleshy, annular-pulvinate or truncateconical, closely puberulent without. Stamens 3, suberect to reflexed; filaments widened toward the base, sometimes connate to upper margin of disk; anthers transversely ellipsoid, dehiscing by extrorse confluent clefts. Ovary completely immersed in the disk, 3-locular; ovules 4–8 per locule; style subulate; stigmas inconspicuous. Fruits few per infructescence; capsules 3, divergent, separately attached to a swollen receptacle, flattened, dehiscing along an inconspicuous median suture, pericarp thin. Seeds erect from the base, attached by a conspicuous basal wing, the portion containing the embryo apical, coriaceous. Tropics and subtropics; 3 species, 1 in Venezuela. Hippocratea volubilis L., Sp. Pl. 1194. 1753. —Bejuco de sapo. Liana, occasionally a shrub. Evergreen lowland to lower montane forests, 0–400

(–1000) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Throughout most of Venezuela; worldwide distribution as in genus. ŠFig. 519.

Hylenaea 603

Fig. 519. Hippocratea volubilis

6. HYLENAEA Miers, Trans. Linn. Soc. London 28: 366. 1872. Woody lianas or small trees, glabrous throughout. Leaves opposite; stipules minute, deciduous. Inflorescences axillary and/or terminal, often aggregate in the leaf axil, paniculate-corymbose, pedunculate (in H. comosa with numerous capillaceous branches and abortive flowers), to 20 cm long; pedicels slender, articulate, short. Flowers very small, ca. 1.5–3 mm wide, sometimes abortive, greenish or yellowish. Calyx lobes narrow or ovate; petals membranous, rounded at the apex. Disk erect, short-cylindric, thin. Stamens 3, erect to strongly reflexed; filaments ligulate and broadened towards the base; anthers very small, subglobose or transversely oblong, opening by extrorse-apical, horizontal, confluent clefts. Ovary subglobosetrigonous, 3-locular, ovules 4–8 per locule; style fleshy, subulate; stigmas obscure. Fruit with 3 free mericarps; mericarps flat to convex, large, obovate-oblong, ca. 9 × 6 cm, dehiscing along an obscure median suture; pericarp coriaceous or woody, flabellate-costate or ecostate and strongly scaly. Seeds erect from the base, affixed by a basal wing prolonged in a narrow dorsal wing enveloping wholly or to half way the portion containing the embryo, the embryoniferous part thick, coriaceous. Neotropics; 3 species, 1 in Venezuela.

604

H IPPOCRATEACEAE

Fig. 520. Hylenaea comosa

Hylenaea comosa (Sw.) Miers, Trans. Linn. Soc. London 28: 367. 1882. —Hippocratea comosa Sw., Prodr. 17. 1788. Hippocratea crinita Pittier, Bol. Soc. Venez. Ci. Nat. 6: 10. 1939. Liana, seldom a slender tree with scandent branches; young twigs with a reddish or purplish peeling bark. Riparian forests, 0–

300 m; Delta Amacuro (Caño Güiniquina, Río Amacuro, Río Toro), Bolívar (Río Nichare, Río Oris in middle Río Paragua basin, Salto Pará), Amazonas (Río Mawarinuma). Costa Rica, Haiti, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Brazil. ŠFig. 520. See remarks under Cuervea.

7. PERITASSA Miers, Trans. Linn. Soc. London 28: 402. 1872. Shrubs, small trees with scandent branches, or lianas, usually glabrous, but in some species the young branches and inflorescences pilose. Leaves opposite or alternate (outside the flora area); blade crenulate-serrate to ± entire, the base rounded, acute, or cuneate, the apex rounded, attenuate, acute, or cuspidate with an obtuse often glandular tip, often dark green on upper surface and lighter green on lower surface, when dried usually olivaceous to dark purplish brown on upper surface, lighter colored on lower surface; texture (lens needed) finely puncticulate below, ± wrinkled above. Inflorescences axillary, thyrsoid-paniculate, or corymbose, dichotomously divided from the base or with a short to long peduncle; bracts small, boatshaped, generally fimbriate. Flowers very small, 2–4 mm wide, cream, yellowish green, or green, almost sessile or with a short pedicel (≤ 3 mm). Sepals suborbicular, often forming a cup, erosulous; petals semi-erect to recurved, thin-fleshy, elliptic, rounded at apex, margins often erosulous. Disk short-tubular, free, the margin dentate or crenulate. Stamens 3, erect; filaments ligulate, in some species produced into an apiculum above the anthers; anthers dorsally adnate to the filaments, dehiscing vertically by clefts. Ovary with a short truncate style, 3-locular; ovules 2–4 per loc-

Peritassa 605

ule; stigmas inconspicuous. Fruits drupaceous, 2–6 cm long, depressed-globose or oblong-elliptic; pericarp leathery, rugulose to pitted, green or tan, turning orange when ripe, when dried often bluish green, pruinose. Seeds angled, ellipsoid, embedded in a mucilaginous pulp. New World tropics and subtropics; ca. 14 species, 5 in Venezuela, all in the flora area. Key to the Species of Peritassa 1. 1.

2(1).

2.

3(1).

3.

4(3). 4.

Stamens with a pronounced filament; leaf margin crenulate, undulate, or entire; fruit depressed globose, containing to 6 compartments ........... 2 Stamens without a pronounced filament; leaf margin entire; fruit oblongellipsoid, number of compartments difficult to assess, the septa soon evanescent .............................................................................................. 3 Leaf apex short-acuminate, margin slightly or conspicuously crenate or undulate, secondary veins 5–11 per side, prominulous beneath; inflorescence ≤ 8 cm long, lax, usually divided from the base; flowers pale yellow with orange disk; fruit ≤ 6 cm long and 10 cm diameter, orange, becoming bluish and pruinose when dry; seeds brown, covered by a juicy white aril .......................................................................... P. laevigata Leaf apex rounded, emarginate, or short-acuminate, margin entire, secondary veins 9–11 per side, usually immersed or at least inconspicuous; inflorescence 2–4 cm long on slender peduncle 7–20 mm long; flowers pale yellow to green; fruit ≤ 3 cm long and 4.5 cm diameter, bluish ....................................................................................................... P. compta Leaves thin-coriaceous, elliptic, 6–19 cm long, rounded to acute at base, short-acuminate at apex; inflorescence branches glabrous; ripe fruit orange ......................................................................................... P. pruinosa Leaves membranous, 7–18 cm long, cuneate or sometimes rounded at base, cuspidate to acute at apex; inflorescence branches glabrous or tomentellous; ripe fruit yellow ......................................................................... 4 Leaves attenuate in the lower part, the base cuneate, the apex acuminate; inflorescence and young branches glabrous ........................... P. peruviana Leaves obovate-elliptic, the base rounded to cuneate, the apex often cuspidate with a short, broad, obtuse acumen; inflorescence and young branches rufous-tomentellous ............................................... P. huanucana

Peritassa compta Miers, Trans. Linn. Soc. London 28: 405. 1872. —Ají de paloma. Peritassa laevigata subsp. obtusa Peyr. in Mart., Fl. Bras. 11(1): 153. 1878. Small tree, scandent shrub, or liana. Savannas, seasonally flooded riparian forests, 100–1500 m; Bolívar (near La Tigrera), Amazonas (Cerro Duida, Río Negro, 20 km southeast of San Fernando de Atabapo). Tobago, Colombia, Guyana, Suriname, Peru, northern Amazonian Brazil. Peritassa huanucana (Loes.) A.C. Sm., Brittonia 3: 521. 1940. —Hippocratea huanucana Loes., Repert. Spec. Nov.

Regni Veg. 1: 163. 1905. Liana or low shrub with scandent branches. Semideciduous to evergreen lowland or lower montane forests, 200–1000 m; Bolívar (scattered in Gran Sabana, Represa Guri), Amazonas (Cerro Huachamacari, Simarawochi). Costa Rica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 522. Peritassa laevigata (Hoffmanns.) A.C. Sm., Brittonia 3: 508. 1940. —Tonsella laevigata Hoffmanns. in Link, Jahrb. Gewächsk. 1(3): 68. 1820. —Cachete de vieja, Derello.

606

H IPPOCRATEACEAE

Fig. 521. Peritassa laevigata

Fig. 522. Peritassa huanucana

Prionostemma 607

Salacia granulata Urb., Repert. Spec. Nov. Regni Veg. 14: 33. 1916. —Peritassa granulata (Urb.) A.C. Sm., Brittonia 3: 512. 1940. Liana, seldom a shrub or small tree. Evergreen lowland to montane forests, 0–1800 m; Delta Amacuro (Sacupana), Bolívar (widely scattered), Amazonas (Cerro Sipapo, Puerto Ayacucho, Capibara, Río Manapiare, Río Negro). Apure, Falcón, Guárico, Lara, Miranda, Táchira; Panama, Colombia, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 521. Peritassa peruviana (Miers) A.C. Sm., Brittonia 3: 519. 1940. —Sicyomorpha peruviana Miers, Trans. Linn. Soc. London 28: 411. 1872. —Curawei-yek (Arekuna). Liana. Montane forests, 900–1100 m; Bolívar (Kavanayén), Amazonas (Sierra Pari-

ma). Ecuador, Peru, Brazil, Bolivia. Peritassa peruviana and P. huanucana are very similar morphologically, and difficult to separate when the pilose inflorescence or pilose young branches of P. huanucana are missing. Peritassa pruinosa (Seem.) A.C. Sm., Brittonia 3: 520. 1940. —Salacia pruinosa Seem., Bot. Voy. Herald 90. 1853. —Sicyomorpha pruinosa (Seem.) Miers, Trans. Linn. Soc. London 28: 411. 1872. —Kahka kaichipen mén (Panare). Liana, shrub, or tree. Evergreen lowland to lower montane forests, riparian forests, 100–600 m; Bolívar (Cerro Bolívar, northeast of Ciudad Piar, Corozal Village near Caicara, Isla Anacoco, Río Caura), Amazonas (Capibara, La Esmeralda). Falcón, Zulia; Costa Rica, Panama, Colombia.

8. PRIONOSTEMMA Miers, Trans. Linn. Soc. London 28: 354. 1872. Lianas or shrubs with scandent branches, scabrous in the American species, elsewhere glabrous. Leaves opposite, scabrous in the American species. Inflorescences paniculate, lax. Flowers medium-sized to large, 5–12 mm wide, pedicellate.

Fig. 523. Prionostemma aspera

608

H IPPOCRATEACEAE

Sepals unequal; petals orbicular with a narrow clawed base, thin, veins flabellate. Disk subfleshy, patelliform or a flat cushion, the middle part glabrous or pilose (in the American species), the border zone different. Stamens 3, erect in the young flower, afterwards strongly recurved; anthers small, dehiscing by horizontal or apical confluent clefts. Ovary pyramidal; style subulate with an orange stigmatic knob. Fruit 3-carpellate, dehiscing by a median suture; pericarp coriaceous, green. Seeds winged, the basal wing longer than the portion containing the embryo. Pantropics; 5 species, 1 in Venezuela. Prionostemma aspera (Lam.) Miers, Trans. Linn. Soc. London 28: 355. 1872, “asperum.” —Hippocratea aspera Lam., Tabl. Encycl. 1: 101. 1791. —Sangrito. Salacia catalinensis Rusby, Descr. S. Am. Pl. 53. 1920, “Salacea.” High-climbing liana, scandent shrub, or tree. Semideciduous to evergreen lowland,

riparian, and lower montane forests, gallery forests, secondary vegetation, 100–300(–800) m; common in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in lowland Venezuela; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia. ŠFig. 523.

9. PRISTIMERA Miers, Trans. Linn. Soc. London 28: 360. 1872. Lianas or scandent shrubs, glabrous. Leaves opposite; blades denticulate or undulate-crenulate, seldom entire; venation (including tertiary veins) raised and prominent on both sides. Inflorescences axillary, sometimes aggregated on short leafless branches, pedunculate, in few- to many-flowered dichotomously branched cymes. Flowers pedicellate or almost sessile, minute to very small, 2–5 mm wide, usually loosely arranged, green, yellowish green, or white. Sepals short, entire or denticulate; petals orbicular to ovate, thin-fleshy to membranous, entire or faintly erosulous, often glandular-punctate. Disk fleshy, flat or slightly cupular, short, inconspicuous. Stamens 3, suberect to spreading; filaments strongly widened towards the base; anthers small, transversely ellipsoid, dehiscing by horizontal confluent clefts. Ovary 3-lobed; ovules 2–10 per locule; style short, subulate, obtuse or terminated by a minute triangular stigmatic shield; stigmas alternate with stamens. Fruit with 3 free elliptic mericarps, each dehiscing by a median suture; pericarp thin-coriaceous or papery. Seeds with a conspicuous basal wing and 2 conspicuous longitudinal veins, the portion containing the embryo much smaller than the wing; in P. tenuiflora mericarps roundish, slightly convex, the seeds with a narrow wing surrounding the embryoniferous part. Pantropics; 24 species, 3 in Venezuela, all in the flora area. Key to the Species of Pristimera 1.

1.

Leaves with a long-acuminate apex, the tip acute; inflorescence slender; flowers almost sessile, white or greenish white; style thick, short, terminated by a minute triangular stigmatic shield or 3 minute stigmas; fruit with roundish mericarp; seeds with wing around the portion containing the embryo ................................................................... P. tenuiflora Leaves with an abruptly short-acuminate apex, the tip blunt, or the apex obtuse to acute and without an acumen; flowers short-pedicellate, cream, yellow, or yellowish green; style short, obtuse; stigmas obscure; fruit with elliptic mericarps; seeds with a basal wing .......................... 2

Pristimera 609

2(1).

2.

Leaves thin, ovate or broadly elliptic, the apex abruptly short-acuminate, drying greenish, ≤ 18 × 9 cm; inflorescence slender or not, ± compact, with short quadrangular branches; peduncle 1–2.5 cm; bracts and bracteoles conspicuously fimbriate ............................................. P. nervosa Leaves subcoriaceous, ovate, the apex obtuse, drying yellowish, ≤ 8 × 4.5 cm; inflorescence slender; peduncle 1–1.5 cm; bracts and bracteoles pointed, not fimbriate ............................................................... P. verrucosa

Fig. 524. Pristimera tenuiflora

Fig. 525. Pristimera nervosa

610

H IPPOCRATEACEAE

Pristimera nervosa (Miers) A.C. Sm., Brittonia 3: 370. 1940. —Sicyomorpha nervosa Miers, Trans. Linn. Soc. London 28: 412. 1872. Liana or small tree. Evergreen lowland forests, often in swampy areas, 0–500 m; Delta Amacuro (Río Cuyubini, near Santa Catalina), Bolívar (Río Paramichí), Amazonas (upper Río Baría, Salto Yureba in lower Río Ventuari basin). Apure; Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil, Bolivia. ŠFig. 525. Pristimera tenuiflora (Mart. ex Peyr.) A.C. Sm., Brittonia 3: 382. 1940. —Hippocratea tenuiflora Mart. ex Peyr. in Mart.,

Fl. Bras. 11(1): 132. 1878. Liana or small shrub. Evergreen lowland forests, 0–300 m; Delta Amacuro (Caño JotaSabuca near Caño Mariusa), Bolívar (Río Asa south of La Paragua, south of Tumeremo). Costa Rica, Nicaragua, West Indies, Colombia (Chocó), Suriname, French Guiana, Peru, Brazil. ŠFig. 524. Pristimera verrucosa (H.B.K.) Miers, Trans. Linn. Soc. London 28: 375. 1872. —Hippocratea verrucosa H.B.K., Nov. Gen. Sp. quarto ed. 5: 138. 1821 [1822]. Liana. Evergreen lowland and lower montane forests, 100–300 m; Bolívar (Altiplanicie de Nuria, 30 km north of El Dorado). Falcón, Zulia; Colombia (Atlantic region), Guyana.

10. SALACIA L., Mant. Pl. 159. 1767. Lianas, shrubs, or slender trees, glabrous (inflorescence tomentellous in Salacia amplectens), bark occasionally with orangish exudate. Branches often at right angles to main stem, the nodes swollen or flattened. Leaves usually opposite (always so in the flora area); petioles slender or stout; blades entire to crenulate, the base acute, rounded, or cordate, the apex rounded, emarginate, acute, acuminate, or cuspidate, concolorous or lighter green on lower surface, when dried light reddish brown to dark purplish brown and various shades of green, texture coriaceous to papery, in some species minutely foveolate on both sides, size very variable, 5–50 × 2–25 cm; secondary veins occasionally very widely spaced in the middle of the blades. Inflorescences axillary or arising from defoliate branchlets, fasciculate, crowded-racemose, thyrsoid-paniculate, or cymose; bracts small; pedicels slender, short to long (1–30 mm). Flowers loosely arranged or congested, usually small to medium-sized, in some species to 30 mm wide, greenish, greenish yellow, or reddish brown. Sepals usually broadly imbricate, often unequal, broadly ovate to deltoid, often scariose and erosulous at the margins; petals narrowly imbricate, mostly rotate at anthesis, rounded at apex, entire or erosulous. Disk fleshy, annular-pulvinate, truncate-conical, in some species the central part produced into 3 teeth, or with the base flattened, seldom ± cupular (not in the flora area). Stamens 3, usually sharply reflexed; filaments ligulate, slightly widened towards the base; anthers small, transversely ellipsoid, reniform, or obtrullate, the clefts extrorse or apical, horizontal or oblique, confluent. Ovary depressed-conical, often subtrigonous, generally deeply immersed in the disk and connate to it, 3-locular; ovules 2–8(–11) per locule, collateral or superposed; ovary tapering into a short subulate style; stigmas obscure. Fruits globose, subglobose, ellipsoid, or cucumber-like, 1–12 cm long, the septa evanescent at maturity, green, bluish green, often turning orange at maturity; pericarp coriaceous, glabrous, smooth, wrinkled, or with warts, protuberances, or thin scales, sometimes speckled or striped. Seeds several, embedded in mucilaginous pulp, angled, oblong. Pantropics, occasionally subtropics; 180–200 species, 10 in Venezuela, all in the flora area.

Salacia 611

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

2.

3(2).

3.

4(3).

4.

Inflorescence dichotomously branched or thyrsoid-paniculate; fruit globose or ellipsoid ...................................................................................... 2 Inflorescence fasciculate; fruit globose ...................................................... 5 Leaves generally coriaceous, 20–50 × 10–25 cm, texture on both sides finely pitted, base cordate, obtuse, or rounded, apex nearly always rounded; inflorescence glabrous, 4–10 cm long, many-times subdichotomously branched from the base or with a very short peduncle; flowers ca. 2.5 mm wide, subcampanulate, petals erect, white or reddish to orange-brown; disk with a low, flat basal part, the upper part with 3 thin, upright, deltoid extensions, alternating with the stamens; fruit globose, ca. 6 cm diameter, the pericarp thin, coriaceous, flaky, pale greenish brown or dull orange; seeds embedded in an orange pulp ..................................................................................................... S. cordata Leaves subcoriaceous or papery, oblong to obovate, generally < 20 × 10 cm, texture not pitted; base acute to obtuse, apex short-cuspidate, mucronate, or rounded; inflorescence glabrous or ± tomentellous, 3–5 cm long, dichotomously or paniculately branched; peduncle 2–30 mm long; flowers 4–11 mm wide, petals spreading, green to yellow-green; disk not as above; fruits globose or ellipsoid, pericarp smooth or flaky, bluegreen or orange, sometimes white-speckled ......................................... 3 Leaves elliptic, petiole 4–6 mm; inflorescence 4–11 cm long, axillary, glabrous, dichotomously branching with slender internal branches, peduncle 20–30 mm long; flowers 4–6 mm wide, disk flattened, 0.2–0.4 mm high; fruit globose, 2–3 cm diameter ....................................... S. opacifolia Leaves oblong to obovate, petiole 8–12 mm; inflorescence 2–3 cm long, axillary or on short lateral branches, tomentellous or mostly glabrous, dichotomously or paniculately branching with stout internal branches, peduncle 2–20 mm long; flowers 6–13 mm wide; disk not as above; fruit globose or ellipsoid ................................................................................. 4 Leaves oblong, the apex cuspidate with a blunt, not glandular tip; inflorescence brown-tomentellous, dichotomously branched; peduncle 1–2 cm long; flower ca. 4 mm wide; pedicels not articulate, 1–3 mm long; petals erect, fleshy, oblong, puberulent on both sides; disk with a thin upturned lower part, and a conical upper part, glabrous, concolorous; fruit solitary, globose to broadly ellipsoid, 5 cm diameter, glaucous when young, orange when ripe, the pericarp smooth and leathery; seeds 4–6, brown, embedded in creamy pulp ........................ S. amplectens Leaves obovate, the apex short, acuminate, sometimes mucronate, the midrib often terminated by a thickened dark gland; inflorescence almost glabrous, paniculate; peduncle absent or < 5 mm long; flower 8– 13 mm wide; pedicels articulate, 4–12 mm long; petals spreading, subfleshy, often slightly farinose-puberulent; disk conical-truncate, the basal half often dark brown and tomentellous, the upper part yellowish brown and glabrous; fruit solitary or in pairs, broadly ellipsoid, ≤ 7 × 5 cm, the pericarp grayish ferruginous, rugose, speckled when young, later smooth; seeds with an orange skin ........................................ S. multiflora

612

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

6.

7(5).

7.

8(7).

8.

9(7).

9.

H IPPOCRATEACEAE

Flowers < 12 mm wide; disk gradually flattened at the base or not flattened at all; pedicels 5–30 mm long ...................................................... 6 Flowers > 12 mm wide; disk conspicuously flattened basally; pedicels usually slender, 3–20 mm long .................................................................... 7 Leaves obovate, the base acute, the apex gradually or abruptly shortacuminate, when dried dark green on upper surface, lighter green on lower surface, 6–8 × 2.5–3 cm; flowers tan to salmon-orange, > 9 mm wide; pedicels 1.3–2 cm long; fruit globose, ca. 2 cm diameter, grayish green, the pericarp corky, slightly tuberculous; seeds orange ............... ............................................................................................. S. kanukuensis Leaves elliptic, the base obtuse, the apex rounded, seldom with an abrupt short tip, when dried concolorous, width and length variable, but generally 2–2.5 times longer than wide; flowers green, dull orange, or petals yellow with a dark orange base, 5–9 mm wide; pedicels slender, 5– 10 mm long (fruiting pedicels ca. 1.5–2 cm long); fruit 4 cm diameter, glaucous green with a grayish hue, turning orangish, the pericarp smooth; seeds unknown .............................................................. S. elliptica Leaves narrowly elliptic, ca. 3 times longer than wide, to 34(–60) cm long, the secondary veins more widely separated in the middle part of leaf; flowers yellowish green; pedicels short to moderately long (3–16 mm), thickened; pericarp rugulose or pitted .................................................. 8 Leaves ovate to elliptic, 15–33 cm long, 2–2.5 times longer than wide, the secondary veins evenly separated throughout the leaf; flowers yellowish green or petals brown at base; pedicels slender, ≤ 2 cm long; pericarp smooth and thin, wrinkled, or tuberculate ................................... 9 Low trees or lianas; leaves when dried liver-colored, the base acute; tertiary veins distinct, horizontally parallel; flowers 24–30 mm wide; pedicels 12–16 mm long; fruit globose, ca. 4 cm diameter, pale yellow to orange, pericarp tuberculate ................................................. S. macrantha Trees; leaves when dried clear brown or grayish brown-green, the base rounded to acute; tertiary veins obsolete; flowers ca. 15–18 mm wide; pedicels ca. 3 mm long; fruit globose, 4.5 cm wide, grayish green with white powder or white stripes, turning yellow when ripe, pericarp smooth ........................................................................................ S. gigantea Lianas, occasionally trees to 8 m; blades ovate-oblong, twice as long as wide, to 26 × 10 cm, usually smaller, the base rounded, the apex acuminate; venation impressed on the upper surface, flat to prominulous on lower surface; petals yellowish green to orangish, often reddish brown at base; fruit ≥ 3 cm diameter, blue-green when young, dull orange when mature, the pulp medium brown, the pericarp coriaceous, smooth or tuberculate ......................................................... S. impressifolia Mostly trees to 15 m, sometimes lianas; leaves elliptic, about 2 times as long as wide, 14–25 × 6–11 cm, the base rounded, the apex obtusely cuspidate or rounded; secondary veins prominulous on upper surface and sunken in a groove, prominulous on lower surface; petals yellowish green; fruit 4 × 4 cm, yellow when mature, pulp color unknown, the pericarp wrinkled or tuberculate ............................................... S. juruana

Salacia 613

Salacia amplectens A.C. Sm., Brittonia 3: 443. 1940. Liana. Evergreen lowland to montane forests, 100–1000 m; Bolívar (upper Río Paragua), Amazonas (Cerro Sipapo, Río Coro Coro). Guyana, Suriname, French Guiana, Brazil (Amazonas). Salacia cordata (Miers) Mennega, Novon 2: 232. 1992. —Thermophila cordata Miers, Trans. Linn. Soc. London 28: 401. 1872. Salacia megistophylla Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 4: 222. 1929. Salacia acreana A.C. Sm., Bull. Torrey Bot. Club 66: 243. 1939. Small tree or robust, high-climbing liana. Evergreen lowland to montane forests, 50– 800 m; Bolívar (Serranía de Imataca), Amazonas (Isla Ratón, Río Matacuni, Río Orinoco, Simarawochi). Yaracuy, Zulia; Mexico, Costa Rica, Panama, Colombia, Peru, northern Brazil, Bolivia. ŠFig. 528. This species is very variable in the size and outline of its leaves, but the texture and surface are consistent, as are the inflorescences, flowers, and fruits. Salacia elliptica (Mart.) G. Don, Gen. Syst. 1: 628. 1831. —Anthodus ellipticus Mart. in Schult., Mant. 1: 348. 1822. —Melocotoncito. Tree or liana. Gallery forests, black-water swamps, wooded slopes, 50–300 m; Bolívar (mouth of Río Caura, Río Paragua, near Upata), Amazonas (Lago Tití near San Fernando de Atabapo, Río Baría, Río Negro). Anzoátegui, Apure, Barinas, Táchira; Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. The present concept of Salacia elliptica is probably too broad, in view of the great variation in leaf size, form, and structure, as well as flower size and color, but constant differentiating characters were not found. Salacia gigantea Loes., Verh. Bot. Vereins Prov. Brandenburg 48: 182. 1907. Understory tree or liana, 3–15 m. Riparian forests, 100–200 m; Amazonas (Caño Yagua, El Merey on Río Casiquire, Río Casiquiare near mouth of Río Paciba,

Río Orinoco 5 km east of San Fernando de Atabapo). Panama, Colombia, Peru, Brazil. Salacia impressifolia (Miers) A.C. Sm., Bull. Torrey Bot. Club. 66: 247. 1939. —Raddia impressifolia Miers, Trans. Linn. Soc. London 28: 392. 1872. —Cachete de viejo. Salacia grandiflora Peyr. in Mart., Fl. Bras. 11(1): 157. 1878. High-climbing liana, occasionally shrub or tree to 12 m. Evergreen lowland to montane forests along rivers and creeks, 50–300 m; Bolívar (middle Río Paragua), Amazonas (La Esmeralda, Río Casiquiare near El Porvenir, Samariapo, San Carlos de Río Negro, San Fernando de Atabapo). Coastal Cordillera, Apure, Yaracuy; Mexico, Honduras, Nicaragua, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 526. Salacia juruana Loes., Verh. Bot. Vereins Prov. Brandenburg 48: 181. 1907. —Cachete de vieja, Pilón rosado. Small tree or liana; fruits mottled. Periodically flooded black-water river banks, lower montane forests, 50–300 m; Bolívar (Altiplanicie de Nuria, southeast of Villa Lola), Amazonas (Caño Minicia, Río Casiquiare, near San Fernando de Atabapo). Ecuador, Peru, Brazil, Bolivia. Salacia kanukuensis A.C. Sm., Lloydia 2: 192. 1939. Liana. Evergreen lowland forests, blackwater swamps, 100–300 m; Bolívar (Serranía de Imataca), Amazonas (Río Baría). Guyana, Suriname, French Guiana. Salacia macrantha A.C. Sm., Bull. Torrey Bot. Club 66: 245. 1939. Shrub or slender tree. Dense montane forests, ca. 1300 m; Bolívar (along Río Caruay at base of Ptari-tepui). Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. Salacia multiflora (Lam.) DC., Prodr. 1: 570. 1824. —Hippocratea multiflora Lam., Tabl. Encycl. 1: 101. 1791. Nicaragua, Costa Rica, Panama, Colom-

614

H IPPOCRATEACEAE

Fig. 526. Salacia impressifolia

Fig. 527. Salacia multiflora subsp. mucronata

Fig. 528. Salacia cordata

Tontelea 615

bia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, both in the flora area.

1. Inflorescences glabrous, strictly dichotomously and regulary branched; apex of leaf mucronate; fruit ellipsoid, often furrowed, pale orange, white-speckled ........ ............................... subsp. mucronata 1. Inflorescences occasionally reddish-tomentellous, not strictly dichotomously branched, primary branches often congested on the peduncle; apex of leaf generally not mucronate; fruit ovoid or ellipsoid, furrowed or not, the pericarp with fine scales or white-speckled .................. ................................ subsp. multiflora

S. multiflora subsp. multiflora. —Bejuco de pereza. Thermophila rugulosa Miers, Trans. Linn. Soc. London 28: 400. 1872. Salacia obovata Peyr. in Mart., Fl. Bras. 11(1): 154. 1878. Salacia pittieriana A.C. Sm., Brittonia 3: 440. 1940. Low- or high-climbing liana, occasionally a shrub. Riverbanks, evergreen lowland to montane forests, 100–1300 m; Bolívar (Gran Sabana, Río Caura, Río Nichare, Serranía de Maigualida), Amazonas (Caño Iguapo on upper Río Orinoco, Río Casiquiare, Río Mawarinuma, Río Yureba, San Carlos de Río Negro). Aragua, Miranda, Sucre, Táchira; Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

S. multiflora subsp. mucronata (Rusby) Mennega, Novon 2: 232. 1992. —Salacia mucronata Rusby, Descr. S. Amer. Pl. 53. 1920. Liana to 40 m long. Evergreen lowland to lower montane forests, 0–400 m; Delta Amacuro (lower Río Orinoco, Río Toro, Santa Catalina), Bolívar (Sierra de Maigualida). Suriname, French Guiana. ŠFig. 527.

Salacia opacifolia (J.F. Macbr.) A.C. Sm., Brittonia 4: 434. 1940. —Hippocratea opacifolia J.F. Macbr., Field Mus. Nat. Hist. Dept. Bot. Leafl. 8(2): 123. 1930. Salacia gracilis A.C. Sm., Bull. Torrey Bot. Club 66: 249. 1939. Liana. Riparian forests, 100–200 m; Amazonas (Caño Libreta 35 km southeast of La Esmeralda). Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil.

Key to the Subspecies of S. multiflora

11. TONTELEA Aubl., Hist. Pl. Guiane 31. 1775. Lianas, shrubs, or medium-sized trees, glabrous throughout. Branches and leaves opposite, occasionally alternate (not in the flora area). Leaves usually dark green on upper surface, sometimes glossy, lighter on lower surface, margin usually entire, sometimes slightly crenulate; presence of intersecondary veins is characteristic, tertiary venation mostly reticulate. Inflorescences axillary, thyrsoid-paniculate or pseudodichotomously branching; bracts very small. Flowers small to very small (ca. 2–4 mm wide, 5–7 mm in Tontelea cylindrocarpa), white, cream, greenish, or yellow, usually arranged in loose clusters at the end of the branchlets, subtended by minute bracteoles. Petals submembranous to thin-fleshy, often fimbriate or farinosepuberulent at the apex. Disk thin-fleshy, short-tubular, free from the ovary or connate to it by interstaminal septa. Stamens 3; filaments ligulate; anthers transversely ellipsoid, broader than high, giving the impression of a butterfly, the locules subglobose, dehiscing by horizontal extrorse clefts. Ovary depressed-conical, 3-locular; ovules 2(4–8) per locule; style short; stigmas 3, obvious, divaricate, entire, emarginate or bilobed or so deeply divided that a wheel with 6 equal spokes is formed, alternate or opposite the stamens, sometimes absent. Fruits stipitate, subglobose to ellipsoid, slightly angled (cylindric in T. cylindrocarpa), medium-sized to large (size of an infant’s head); pericarp coriaceous or woody. Seeds imbedded in a mucilaginous pulp.

616

H IPPOCRATEACEAE

Costa Rica, Honduras, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay; ca. 26 species, 8 in Venezuela, 6 of these in the flora area. Key to the Species of Tontelea 1.

1.

2(1).

2.

3(1).

3.

4(3).

4.

5(4).

5.

Leaves coriaceous, white-dotted on the lower surface when dried (lens needed); pericarp of the fruit slightly or strongly pitted; stigmas bilobed, alternate with the stamens, or obscure ...................................... 2 Leaves chartaceous to thin-coriaceous, not punctate on the lower surface when dried; pericarp smooth or slightly flaky; stigmas either entire and alternate with the stamens, or so deeply lobed that it is difficult to see whether alternate with or opposite to the stamens, or stigmas absent ......................................................................................................... 3 Leaves ovate to elliptic, not shining above, with conspicuous white dots on lower surface; fruit rounded or obovate, strongly pitted, blue-green, turning yellow ..................................................................... T. mauritioides Leaves obovate, shining above, with white dots not pronounced on lower surface; fruit 3-angular with a slightly pitted or wrinkled surface, olive-green, turning orange ........................................................ T. coriacea Lianas; leaves chartaceous, elliptic to oblong, drying light green, secondary veins ± 6 per side, arcuate-ascending; inflorescence solitary or aggregate in the axils, ca. 7–20 cm long; peduncle very slender, ≤ 16 cm long; pedicels 6–10 mm long; flowers ca. 6 mm wide, yellow; style subulate, stigmas absent; fruits cylindric, woody, ≤ 16 cm long .................... ........................................................................................... T. cylindrocarpa Shrubs, trees, or lianas; leaves chartaceous to sub-coriaceous, narrowly elliptic to ovate, drying dark green or brownish, secondary veins 6– 10 per side, arcuate-ascending; inflorescence axillary, mostly compact or loosely branched, not over 8 cm long; peduncle ca. 1 cm long or absent; pedicels 1–7 mm long; flowers 3–5 mm wide, white, light green, or yellowish orange; style terminated by 3 stigmas, lobed or not; fruits globose or ellipsoid, 3–11 cm long ......................................................... 4 Inflorescence loosely branched, ≤ 8 cm long; pedicels 3–5 mm long; flowers yellow or light green; stigmas deeply bilobed, forming a shield with 6 equal parts; fruit 3-angled, ellipsoid, ca. 2.5 cm long, the pericarp leathery, greenish, turning orange ............................................ T. laxiflora Inflorescence compact, semi-globular, ≤ 4(–5) cm long; pedicel < 1 mm long; flowers white to greenish; stigmas entire, alternate with the stamens; fruit 3-angled, globose or ellipsoid, 2.5–11 cm long, the pericarp leathery or woody, green or brown ........................................................ 5 Leaves chartaceous to subcoriaceous, narrowly elliptic, ca. 3–4 times as long as broad, base cuneate, apex acuminate, venation obscure above; flowers fragrant, white to greenish; fruit ± 3-angled, ca. 2.5 cm long, the pericarp leathery, dark grayish green, turning orange .... T. attenuata Leaves coriaceous, elliptic to somewhat (ob)ovate, ca. 2–2.5 times as long as broad, base rounded, apex rounded or obtuse, rarely somewhat cuspidate, venation flat above; flowers not fragrant, yellowish to greenish; fruit globose or elliptic, 6–12 cm diameter, the pericarp leathery to woody, green turning brown ..................................................... T. ovalifolia

Tontelea 617

Tontelea attenuata Miers, Trans. Linn. Soc. London 28: 385. 1872. —Salacia attenuata (Miers) Peyr. in Mart., Fl. Bras. 11(1): 149. 1878. —Cachete de vieja. Liana, shrub, or tree. Evergreen lowland and riparian forests, 50–300 m; Amazonas (Río Casiquiare, Río Guainía, Río Negro, Río Siapa). Anzoátegui; Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil. ŠFig. 529. Without flowers, it is difficult to distinguish Tontelea attenuata from T. laxiflora. Tontelea coriacea A.C. Sm., Brittonia 3: 480. 1940. Liana or small tree. Lower montane forests, 200–500 m; Bolívar (Caño Pablo 10 km southeast of Salto Pará, Serranía de Imataca). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. Tontelea cylindrocarpa (A.C. Sm.) A.C. Sm., Brittonia 3: 501. 1940. —Salacia cylindrocarpa A.C. Sm., Bull. Torrey Bot. Club 66: 240. 1939. Liana. Premontane to lower montane forests, 100–500 m; Amazonas (Sierra de la Neblina). Suriname, French Guiana, Peru, Brazil.

Salacia fluminensis Peyr. in Mart., Fl. Bras. 11(1): 149. 1878. —Tontelea fluminensis (Peyr.) A.C. Sm., Brittonia 3: 477. 1940. Large liana, occasionally shrub or tree. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, both in Venezuela, 1 in the flora area. This species shows great variation in leaf and fruit characters. The specimens with fruits with a leathery pericarp are considered to be the typical subspecies. The other subspecies, subsp. richardii Peyr. is known from Panama, Colombia, and French Guiana. T. ovalifolia subsp. ovalifolia High-climbing liana or tree. Gallery and evergreen lowland to montane forests, 0–800 m; Delta Amacuro (Caño Mariusa), Bolívar (near Salto Pará), Amazonas (Río Cataniapo, Sierra de la Neblina). Apure; other distribution as for the species.

Tontelea laxiflora (Benth.) A.C. Sm., Brittonia 3: 481. 1940. —Anthodon laxiflorus Benth., Hooker’s J. Bot. Kew Gard. Misc. 4: 10. 1852. —Salacia laxiflora (Benth.) Peyr. in Mart., Fl. Bras. 11(1): 145. 1878. —Hoja de venado. Liana or shrub. Evergreen lowland to montane forests, 100–1000 m; Bolívar (Icabarú, Río Caruay, Río Hacha), Amazonas (Río Atabapo, San Carlos de Río Negro, Simarawochi). Guyana, Suriname, French Guiana, Brazil. Tontelea mauritioides A.C. Sm., Brittonia 3: 481. 1940. Tree or liana. Lower montane forests, ca. 500 m; Bolívar (Cerro Ichún). Brazil. Tontelea ovalifolia (Miers) A.C. Sm., Brittonia 3: 476. 1940. —Cuervea ovalifolia Miers, Trans. Linn. Soc. London 28: 371. 1872.

Fig. 529. Tontelea attenuata

618

H UGONIACEAE

HUGONIACEAE by Nelson Ramírez, Paul E. Berry, and Antony Jardim Shrubs, trees, or woody vines. Leaves alternate (opposite in Ctenolophon), simple, margin entire or less often undulate- crenate or glandulose-subserrate; stipules deciduous. Inflorescences axillary to terminal racemes, spikes, or panicles, or else axillary fascicles. Flowers bisexual, commonly 5-merous; sepals imbricate, distinct or joined at the base, often ± dissimilar; petals distinct, convolute. Stamens (5)10(–15), mostly unequal; filaments connate for much of their length to form a tube; anthers dithecal, opening by longitudinal slits; nectary disk extrastaminal, usually represented by 2–5 glands. Ovary superior, compound, with 2–5 carpels and locules, with axile-apical placentation and without false partitions, sometimes 1-locular at the very top with the partitions not reaching the summit; ovules 1 or 2 per locule, pendulous; styles distinct or the single style evidently cleft. Fruit a drupe or a nut (Ctenolophon). Seeds with straight or slightly curved embryo; endosperm thick (Indorouchera) or more often scanty or lacking. Pantropics; 5 genera and ca. 60 species, 2 genera and 8 species in the flora area. This segregate family is often treated in the Linaceae; the delimitations of the species and genera are in need of greater scrutiny. Key to the Genera of Hugoniaceae 1.

1.

Secondary leaf veins arching upward near the margin and reticulateveined in between; petals villous on inner surface and clawed at the base; styles usually 5 .......................................................... 1. Hebepetalum Secondary leaf veins very fine and closely parallel, numerous, without reticulate veins in between; petals glabrous on inner surface, not clawed at the base; styles 3 ..................................................................2. Roucheria

1. HEBEPETALUM Benth. & Hook. f., Gen. Pl. 1: 244. 1862. Glabrous trees. Leaves alternate, coriaceous, entire or slightly undulatecrenate; stipules small, broadly triangular. Inflorescences cymules arranged in terminal panicles or in the upper leaf axils. Flowers small, short-pedicellate. Sepals 5, base subconnate; petals 5, imbricate or contorted, deciduous, briefly clawed at base, midvein with a prominent callus dorsally, densely villous on inner surface. Stamens (8) 10, with 5 extrastaminal nectary glands; filaments deltoid. Ovary globose, 4- or 5locular, with 1 or 2 ovules per locule; styles 4 or 5, distinct, with small, subcapitate stigmas. Fruit an ovoid-globose drupe, 1–5-locular, pulp sparse and black, endocarp hard. Seeds solitary or rarely paired, slightly compressed; embryo slightly curved. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, all in the flora area. Key to the Species of Hebepetalum 1. 1.

Ovary 4-locular, styles 4, stigmas villous; petals with short villous pubescence on inner surface; anthers villous apically .................. H. roraimense Ovary 5-locular, styles 5, stigmas glabrous; petals with long villous pubescence on inner surface; anthers glabrous apically ................................ 2

Hebepetalum 619

2(1). 2.

Leaf apex acute, margin entire to subcrenulate, never subrepand ........... .......................................................................................... H. humiriifolium Leaf apex emarginate, margin subrepand ......................................... H. sp. A

Hebepetalum humiriifolium (Planch.) Benth. in Benth. & Hook. f., Gen. Pl. 1: 244. 1862. —Roucheria humiriifolia Planch., London J. Bot. 6: 143. 1847. —Kerosenillo. Roucheria angulata Gleason, Bull. Torrey Bot. Club 58: 373. 1931. Small to large tree to 30 m tall with small butresses and reddish inner bark; sap sticky, translucid; petals yellow-orange with white apex. Nonflooded evergreen forests, edges of

Fig. 530. Hebepetalum humiriifolium

savannas, 100–600 m; border area of Delta Amacuro and Bolívar (Río Toro), Bolívar (Cerro Abismo), Amazonas (La Esmeralda, Río Puruname, Yavita to Maroa). Southeastern Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ŠFig. 530. Hebepetalum roraimense Secco & S.M.B. Silva, Bol. Mus. Paraense Emilio Goeldi, Sér. Bot. 8: 260. 1992.

620

H UGONIACEAE

Small tree 4–8 m tall. Low forests, on white sand or oxisol, ca. 100 m; Amazonas (Río Guayapo). Brazil (Pará, Roraima).

Hebepetalum sp. A Tree 10–15 m tall; lower trunk angular, with clear, sticky exudate. Lowland evergreen forests on white sand, 100–200 m; Amazonas (Río Mawarinuma). Endemic.

2. ROUCHERIA Planch., London J. Bot. 6: 141. 1847, spelling variant: Rouchera. Shrubs or trees. Leaves alternate, oblong, entire or glandulose-subserrate, coriaceous, secondary veins numerous, fine, and parallel, nearly perpendicular to midvein, the apex acuminate, the margins strongly involute in bud; stipules small. Inflorescence paniculate or fasciculate, terminal and/or axillary. Flowers small, yellowish, sessile or subsessile. Sepals 5, oblong, imbricate, basally connate; petals 5, plane, glabrous or pilose along margins, convolute, deciduous. Stamens 10; filaments flattened-subulate, of two heights, joined at the base in a staminal tube, with usually 10 extrastaminal nectary glands. Ovary ovoid, 1–5-locular, 1 or 2 subapical ovules per locule; styles (2)3(–5), filiform; stigmas complanate. Fruits barely fleshy, subglobose, endocarp hard and 3–5-angled. Seeds 1 or 2 per locule, flattened; embryo slightly curved. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 7 species, 5 in Venezuela, all in the flora area. In the Venezuelan Guayana, Roucheria laxiflora and R. punctata cannot always be clearly distinguished. Key to the Species of Roucheria 1. 1. 2(1). 2.

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

Inflorescences axillary, in fascicles or very tight panicles; leaf apex longacuminate ............................................................................................... 2 Inflorescences terminal and axillary, in large, well-branched panicles; leaf apex short-acuminate or obtuse ............................................................ 3 Leaves and young growth glabrous or nearly so; flowers in fasicles not exceeding the petioles in length, no rachis readily visible ........ R. calophylla Leaves pubescent on lower surface, young growth with dense brown or reddish brown pubescence; flowers in very tight panicles ca. 2–4 times as long as the petiole, the rachis easily visible ................ R. schomburgkii Leaves 4–8 × 1.5–3 cm, the blade slightly inrolled at margin, more so at base, the apex obtuse; secondary venation raised .......................... R. sp. A Leaves larger than above, the blade never inrolled, the apex acute; secondary venation sunken ........................................................................ 4 Flowers in lax, terminal and axillary panicles, sometimes with additional subapical flowering shoots ......................................................... R. laxiflora Flowers usually in dense, terminal inflorescences, occasionally axillary as well ....................................................................................... R. columbiana

Roucheria calophylla Planch., London J. Bot. 6: 142, t. 2. 1847. Hebepetalum parviflorum Ducke, Bull. Mus. Hist. Nat. (Paris) sér. 2, 4: 735. 1932. —Roucheria parviflora (Ducke) Ducke, Trop. Woods 43: 21. 1935. Shrub to small tree 1.5–10 m tall. Seasonally flooded savannas and riparian forests, 50–200 m; scattered in eastern and southern

Amazonas. Colombia, Suriname, Peru, Brazil. ŠFig. 532. Roucheria columbiana Hallier f., Beih. Bot. Centralbl. 39(2): 49. 1923, "Rouchera." —Simallo. Hebepetalum punctatum Ducke, Bull. Mus. Hist. Nat. (Paris), sér 2, 4: 735. 1932. —Roucheria punctata (Ducke)

Roucheria 621

Fig. 531. Roucheria laxiflora

Fig. 532. Roucheria calophylla

622

H UGONIACEAE

Fig. 533. Roucheria sp. A

H UMIRIACEAE 623

Ducke, Trop. Woods 43: 21. 1935. Small to large tree to 25 m tall. Evergreen forests, 100–1100 m; Amazonas (Cerro Yutajé, near San Carlos de Río Negro, Sierra de la Neblina, Yavita to Maroa). Nicaragua, Colombia, Ecuador, Peru, Brazil, Bolivia. Roucheria laxiflora H. Winkl., Repert. Spec. Nov. Regni Veg. 7: 109. 1909. Shrub to large tree to 30 m tall. Cloud forests, tepui-slope forests, upland riparian forests, 1000–1700 m; Bolívar (La Escalera, Macizo del Chimantá, Río Aponguao, upper Río Cuyuní), Amazonas (Cerro Duida, Cerro Marahuaka, Sierra de la Neblina). Western Bolivia. ŠFig. 531.

Roucheria schomburgkii Planch., London J. Bot. 7: 526. 1848. —Hebepetalum schomburgkii (Planch.) Ducke, Arch. Jard. Bot. Rio de Janeiro 6: 38. 1933. Tree 5–15(–20) m tall; fruits green, turning orange. Lowland and lower montane forests, 300–400 m; Bolívar (upper Río Caura, upper Río Paragua north of Cerro Ichún). Guyana, Peru, Brazil, Bolivia. Roucheria sp. A Shrub or small tree 3–10 m tall. Montane scrub and among rocks, 1500–2000 m; Amazonas (Cerro Sipapo). Endemic. ŠFig. 533.

HUMIRIACEAE by José Cuatrecasas and Otto Huber Subshrubs to large trees. Leaves alternate, simple, entire, crenate, or serratecrenate, pinnately veined, sessile or pseudopetiolate; stipules tiny and caducous or absent; petiole (or pseudopetiole) a gradual or abrupt narrowing of the proximal part of the blade, usually strongly canaliculate adaxially or else flat; margins frequently dotted with glands abaxially. Inflorescences axillary, pseudoterminal, or seldom terminal, paniculate or corymbiform, with dichasial branching and terminal 3-flowered cymes, or partially monochasial; bracts small, persistent or deciduous. Flowers bisexual, actinomorphic or nearly so, 5-merous. Sepals 5, with a cupular or tubular, connate and fleshy base, thinner toward the margins, the free part suborbicular or triangular, sometimes with marginal or dorsal glands; aestivation usually quincuncial, all sepals equal or the 2 outer ones smaller; petals 5, free, the aestivation contorted, cochlear or quincuncial, usually thick and fleshy, sometimes membranaceous, oblong, linear, or somewhat lanceolate, obtuse or acute, rarely with a gland at the top, white, greenish white, yellowish, or rarely red or purplish. Stamens monadelphous, numerous and fascicled in Vantanea, in the other genera 10–30 and biseriate or uniseriate; filaments united at base into a ring or a short tube, alternating in different lengths, usually 1-anthered, but sometimes the 5 alternate to the petals trifurcate and with 3 anthers; anthers dorsifixed or sub-basifixed; thecae 2, bilocular, laterally attached and each cell dehiscing by longitudinal slits, or else (2)4 unilocular, disjunct thecae dehiscing by detachment or sometimes by a slit; connective thick, fleshy, ovoid or lanceoloid, obtuse, acute, or long-acuminate at apex; some stamens occasionally staminodial. Nectary disk intrastaminal, surrounding the ovary, membranaceous or subcarnose, tubular or cupular and acutely dentate, or composed of 10–20 (or more) free or connate scales. Ovary syncarpous, superior, ovoid or ellipsoid, (4)5(6 or 7)-carpellate, usually 5-septate with axile placentation; ovules 1 or 2 per locule; style single, entire, columnar, as long as stamens or shorter, rarely longer; stigma narrowly or broadly 5-lobed-capitate, the rays usually glandular-glutinous. Fruit drupaceous, with a fleshy or fibrous, perishable exocarp and a

624

H UMIRIACEAE

woody endocarp, usually (4)5(6 or 7)-septate, the surface smooth or verrucose, slightly striate or strongly costate, germinal dehiscence by longitudinal valves; exocarp when dry often turning hard and coriaceous, but decaying spontaneously on the ground; endocarp woody, usually compact or with many round cavities, or spongy. Seeds 1 or 2, rarely more per fruit, oblong, bitegumented; cotyledons oblong or ovate; endosperm fleshy and oily. Costa Rica, Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, one species in coastal West Africa; 8 genera and ca. 60 species, 6 genera and 24 species in the flora area. Key to the Genera of Humiriaceae 1.

1.

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

3. 4(3).

4. 5(4).

5.

Stamens numerous (30–180) and pluriseriate; anthers with 2 bilocular sacs; endocarp with longitudinal, oblong valves, and no resinous cavities; carpels 2-ovulate ............................................................... 6. Vantanea Stamens 10–30; anthers with unilocular, free sacs, the filaments united into a short tube at base; endocarp various, with or without resinous cavities, or spongy; carpels 1-ovulate .................................................... 2 Anthers with 4 unilocular sacs; stamens 20–30; endocarp sharply costate and furrowed ......................................................................... 1. Endopleura Anthers with 2 unilocular sacs; stamens 10–20; endocarp various ......... 3 Anther sacs hirsute; filaments conspicuously glandular; locules of ovary with 2 superposed ovules; endocarp longitudinally striate, valvate, with 5 small holes at apex, not resinous-lacunose, the valves irregularly oblong or lingulate ............................................................ 2. Humiria Anther sacs glabrous; filaments smooth, glabrous; locules of ovary with 1 ovule; endocarp sometimes with resinous cavities or spongy ........... 4 Stamens 10; style usually twice as long or longer than the ovary (rarely shorter); endocarp resinous-lacunose, often bullate, the valves very broad, alternating with filiform, inconspicuous ribs, usually no small holes at apex .......................................................................... 4. Sacoglottis Stamens 20; style equaling the ovary or shorter; endocarp not as above ................................................................................................................ 5 Stamens all with a single anther, 10 longer ones alternating with 10 shorter ones; sacs of anthers basal; endocarp with 5 elliptic, short valves at the upper half alternating with 5 subapical small holes ...... 3. Humiriastrum Stamens more differentiated than above: the 5 opposite the sepals longest, distally trifurcate and triantheriferous, the 5 opposite the petals a little shorter, entire with a single anther, and the remaining 10 between the petals and sepals and shorter, also with single anthers; sacs of anthers laterally attached; endocarp resinous-lacunose, the surface rugulose or bullate, the valves broad, the separating ribs thin, inconspicuous, usually no small holes at the apex ................... 5. Schistostemon

1. ENDOPLEURA Cuatrec., Contr. U.S. Nat. Herb. 35: 80. 1961. Medium to large trees, branchlets and leaves glabrous. Leaves elliptic, apex narrowly acute or acuminate, base cuneate, margin serrate, veins reticulateprominulous. Inflorescences cymose-paniculate, axillary, much shorter than the leaves; peduncles 1–5 cm long; bracts persistent, ovate, minute. Sepals orbiculate, hirtellous, united at base; petals greenish, somewhat thick, linear-oblong, subacute

Endopleura 625

to obtuse. Stamens 22–30; filaments thick, united in lower 1/2–1/3, different lengths alternating; anthers with 4 globose-elliptic sacs, 2 basal and 2 apical or lateral. Scales of the disk 10, triangular, united at the base. Ovary glabrous, subglobose, carpels and locules 5, each locule with 1 ovule; style stout; stigma capitate, 5-lobed. Drupe oblong-ellipsoid, 4–6 × 2–4 cm when mature, rounded at both ends; exocarp coriaceous when dry, mesocarp fleshy; endocarp woody, deeply 5-grooved, with 5 projecting ribs divided halfway into two, giving a 10-radiate shaped cross section. Seeds usually 2 or 3 per fruit, oblong, ca. 30 × 7 mm. Southern Venezuela and Amazonian Brazil (Amazonas, Pará); 1 species. Endopleura uchi (Huber) Cuatrec., Contr. U.S. Nat. Herb. 35: 81. 1961. —Sacoglottis uchi Huber, Bol. Mus. Paraense 2: 489. 1898. —Pata de picure.

Tree 10–15 m tall; inflorescence greenish. Lower montane forests, 400–500 m; Amazonas (upper Río Matapire). Amazonian Brazil. ŠFig. 534.

Stamens

Fig. 534. Endopleura uchi

626

H UMIRIACEAE

2. HUMIRIA Aubl., Hist. Pl. Guiane 564. 1775, orth. cons., “Houmiri.” Low subshrubs or shrubs to large trees. Leaves usually glabrous and shiny, seldom puberulous or hirsute, often decurrent; margin entire or shallowly crenate, dentate or serrate. Inflorescences paniculate, usually corymbiform with dichasial or monochasial branching, axillary or subterminal; bracts persistent. Sepals united at base into a cupular calyx, the lobes imbricate, suborbicular or ovate; petals white, sometimes greenish, carnose, ovate-oblong to linear, subacute to obtuse. Stamens 20, uniseriate, 10 longer ones alternating with 10 shorter ones; filaments complanate, united into a tube about half way up from the base, the free part narrowing upward and densely papillose or muricate; anthers ovoid-lanceolate, dorsifixed above the base; thecae 2, unilocular, subglobose, hirsute, inserted adaxially-laterally at the base. Disk annular, surrounding the ovary, formed by 20 linear, ± united scales. Ovary ovoid or globose, carpels 5, locules (4)5, each locule with 2 anatropous, pendulous, superposed ovules; style straight, erect, as long or longer than the filaments, patent-hirsute except for the distal part; stigmas 5, stellate, each ray subglobose or club-shaped, glutinose. Drupe rather small, to 20 × 15 mm, ovoid, ellipsoid, or oblong; epicarp thin; mesocarp fleshy, usually sweet and fragrant, edible; endocarp woody, ellipsoid, ovoid, or oblong, 10-striate, rarely less, the stria equidistant, marking 5 longitudinal, narrow germinal valves, usually alternating with 5 small apical holes, in the process of maturation, each of the original 5 biovular cavities becomes 2 superposed monospermous cells. Seeds usually 1–4 per fruit, pyriform-ellipsoid or oblong, 3–5 mm long, the outer episperm scaly, the endosperm carnose. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 4 species, all in the flora area. Key to the Species of Humiria 1. 1.

2(1).

2.

3(1).

3.

Leaf blades either < 4 cm long or linear and to 10 cm long; sepals subacute or rounded .............................................................................................. 2 Leaf blades larger and broader, usually > 4 cm long, glabrous, or the midrib pubescent abaxially, rarely the surface sparsely hirtellous; sepals rounded .................................................................................................. 3 Leaf blades elliptic-oblong, sessile, small, 1.5–4 × 0.5–1.5 cm, minutely patent-pilose on both surfaces; young branchlets and inflorescences minutely hyaline-pilose; petals and calyx sparsely pilose; sepals subacute .......................................................................................... H. fruticosa Leaf blades linear, 2.5–10 × 0.3–0.8 cm, with a canaliculate pseudopetiole 0.5–2 cm long; plant glabrous; petals rarely sparsely and minutely pilose; sepals rounded ................................................................ H. wurdackii Leaves petiolate, thick-coriaceous, rigid, subrounded, broadly obovate, broadly elliptic, or oblong-obovate, 7–18 × 4–10 cm; petiole stout, winged, canaliculate, amplexicaul; drupe ellipsoid, 10–12 × 7–9 mm; plant glabrous; calyx glabrous except for the ciliate margin; petals hirtellous ..................................................................................... H. crassifolia Leaf blades sessile or ± pseudopetiolate, chartaceous or coriaceous, oblong-elliptic or oblong-obovate, elliptic or attenuate, 4–14 × 2–6 cm, sometimes smaller (1.5–5 × 0.8–2.5 cm), rarely larger, the pseudopetiole flat; drupe oblong or ellipsoid, 10–11.4 × 4–8 mm; plant glabrous or somewhat pubescent ........................................ H. balsamifera

Humiria 627

Humiria balsamifera Aubl., Hist. Pl. Guiane 564, pl. 225. 1775, “Houmiri.” —Humiria balsamifera J. St.-Hil., Expos. Fam. 2: 374. 1805, orthogr. var. —Humirium balsamiferum Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 102. 1853, orthogr. var. Myrodendrum balsamiferum Raeusch, Nom. Bot. 196. 1797., comb. illeg. Myrodendrum amplexicaule Willd., Sp. Pl. 2(2): 1171. 1800. Low shrub or medium to large tree to 25 m tall, very variable in form and size; flowers in small or large panicles, bright white or greenish white, usually showy. Evergreen lowland to upland forests, more frequent in scrub and open savannas or meadows, on rocks and on white-sand soils, usually in dry sites, but also in temporarily flooded sites, also in high-tepui scrub, 0–2100 m. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 14 varieties, 12 in Venezuela, all in the flora area. There are occasional transitional forms between the varieties, and the number of varieties is likely to be reduced considerably with further study. Key to the Varieties of H. balsamifera 1. Leaves sessile or subsessile, base of blades broad and amplectant or broadly attenuate ............................................... 2 1. Leaves petiolate or pseudopetiolate, base of blades ± attenuate ............................ 5 2. Leaf blades elliptic, rounded at both ends, amplectant at base, sessile, coriaceous, rigid, 2.5–5 × 1.7–3.2 cm; branchlets subterete ......... var. imbaimadaiensis 2. Leaf blades oblong or broader and slightly narrowing toward the base, sessile or subsessile, usually flexible or subrigid; young branchlets flat and ± winged or angular to terete ................................... 3 3. Leaf blades 6–14 × 2.5–6 cm, obovate, oblong-obovate, obovate-elliptic, rounded or obtuse at apex; young branchlets flat and ± winged ............ var. balsamifera 3. Leaf blades 3.5–10 × 1.5–4 cm, oblong, subelliptic-oblong, or sublanceolate-elliptic, slightly narrowing toward the apex, rather obtuse; branchlets angulate to terete, rarely narrowly winged ........ 4 4. Leaf blades 3.5–10 × 1.5–4 cm, sublanceolate-elliptic, attenuate to the obtuse apex, slightly narrowed, obtuse or

auriculate at base ...... var. subsessilis 4. Leaf blades 5–10 × 1.5–3.5 cm, oblong or subelliptic-oblong, attenuate, cuneate at base ........................ var. savannarum 5. Leaf blades abruptly contracted into a long winged pseudopetiole ................... 6 5. Leaf blades gradually attenuate to a cuneate base or suddenly contracted, with a 1–7 mm-long petiole .......................... 7 6. Leaf blades broadly obovate, oblong-obovate, suborbicular or elliptic, 3–12 × 2–6 cm; pseudopetiole 0.5–2.5 cm long ......... .................................... var. guianensis 6. Leaf blades narrower, elliptic-oblong 3–10 × 1.2–3 cm; pseudopetiole 0.5–1.5 cm long .................................. var. laurina 7. Leaf blades obovate or obovate-elliptic or rarely oblong-elliptic, long-attenuate, cuneate at base; young branchlets hirtellous or glabrous ................................ 8 7. Leaf blades usually elliptic or orbicular, shortly attenuate at base, the veins abaxially prominulous; young branchlets always hirtellous .......................... 11 8. Leaves medium-sized to large, 5–12 × 2–6 cm, subcoriaceous, the veins not prominulous abaxially .......... var. floribunda 8. Leaves smaller, 1.5–7 cm long, coriaceousrigid or subrigid, the veins prominulous abaxially ............................................... 9 9. Petals hirsute; branchlets glabrous .......... ........................................... var. iluana 9. Petals glabrous; young branchlets hirtellous or glabrous .................................. 10 10. Leaf blades rigidly coriaceous, (2.5–)4– 7 × 2.4–5.2 cm; petiole 2–6 mm long; young branchlets hirtellous or glabrous; endocarp ellipsoid, about 10 × 5 mm ...... ........................................ var. coriacea 10. Leaf blades thinly coriaceous, 3–5.5 × 2– 3.6 cm; petiole ca. 5–7 mm long; young branchlets glabrous; endocarp oblong, 11–13.5 × 3.8–4.8 mm ............................. .................................... var. stenocarpa 11. Leaf blades glabrous, 4–7 × 3–5.2 cm, orbicular, or suborbicular-elliptic, rounded at apex, cuneate at base into a thick short petiole (1–2 mm long), the margin entire, almost eglandular ....................... ............................ var. guaiquinimana 11. Leaf blades hirtellous, at least on the midrib abaxially, oblong-elliptic, narrowed and obtuse at apex, slightly narrowed at base into a 2–3-mm-long petiole, the margin usually slightly crenulate, with few glands ......... var. pilosa

628

H UMIRIACEAE

H. balsamifera var. balsamifera Well-developed forest tree to 23 m tall, more often a savanna shrub 0.5–2 m tall; ripe drupes black, juicy, edible. Lowland forests, forest edges, white-sand scrub, open savannas, 100–300 m; Bolívar (scattered), Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 536. H. balsamifera var. coriacea Cuatrec., Contr. U.S. Natl. Herb. 35: 110. 1961. Shrub or small tree (0.2–)1–5(–15) m tall, the young branches puberulous, hirsute, or glabrate; leaves small, (2.5–)4–7 × 2.4–5.2 cm, rigid-coriaceous, glabrous, obovate or obovate-elliptic, obtuse, rather rounded and often emarginate and mucronate at apex, shortly attenuate at base into a winged petiole 2–6 mm long; midrib slightly conspicuous or obsolete adaxially, abaxially prominent with secondary veins prominulous. Common in scrub and savannas of open rocky places, also in montane forests and along rapids of uplands and highlands, (400–)600–1700(– 2100) m; Bolívar (± all tepuis, Gran Sabana, upper Río Caroní, Río Paragua), Amazonas (± all tepuis, Cerro Mahedi). Monagas; Guyana, Suriname, Brazil (Amazonas: Serra Aracá, Roraima: Serra Tepequém). The rigid and sclerophyllous texture of the leaves and their size and shape is the distinctive feature of this variety. H. balsamifera var. floribunda (Mart.) Cuatrec., Contr. U.S. Natl. Herb. 35: 99. 1961. —Humirium floribundum Mart., Nov. Gen. Sp. Pl. 2: 143, pl 199. 1826 [1827]. —Humiria floribunda (Mart.) Urb. in Mart., Fl. Bras. 12(2): 437. 1877. Tree or shrub 0.5–5(–15) m tall; fruits juicy, black, edible. Evergreen lowland forests, mixed montane forests, savannas, white-sand meadows, other humid grounds, 50–1000(–1300) m; Bolívar (Canaima, Cerro Guaiquinima, Gran Sabana, Sierra de Lema), Amazonas (Cerro Moriche, base of Cerro Sipapo, San Carlos de Río Negro). Guyana, Suriname, French Guiana, Peru, Brazil (Amapá, Amazonas, Bahia, Ceará, Maranhão, Mato Grosso, Minas Gerais, Pará, Roraima). H. balsamifera var. guaiquinimana Cuatrec., Contr. U.S. Natl. Herb. 35: 113. 1961.

Shrub to low tree to 12 m tall; young branchlets hirtellous, soon becoming glabrous; leaves rigid, thick, suborbicular or suborbicular-elliptic, subsessile, rounded, or very obtuse, often retuse, emarginate, and mucronate at apex, suddenly attenuate to shortly cuneate-petiolate at base. Scrub and shrub savanna, 300–1600 m; Bolívar (Cerro Guaiquinima, Río Trueno). Endemic. H. balsamifera var. guianensis (Benth.) Cuatrec., Contr. U.S. Natl. Herb. 35: 103. 1961. —Humirium guianense Benth., London J. Bot. 2: 374. 1843. Humirium surinamense Miq., Stirp. Suriname Select 86. pl. 24. 1850 [1851]. Humiria floribunda var. guianensis Urb. in Mart., Fl. Bras. 12(2): 439. 1877. Humiria cassiquiari Suess. & Bergdolt, Repert. Spec. Nov. Regni Veg. 39: 16. 1935. Shrub or small tree; leaves obovate, ovate, or elliptic, suddenly contracted above the base into a long, narrow pseudopetiole (the most distinctive feature for this variety, but also found to intergrade with vars. floribunda, laurina, and others). Shrublands, caatingas, savannas, along forest margins, 50–200 m; Amazonas (from the base of Cerro Sipapo to San Carlos de Río Negro). Anzoátegui; eastern Colombia, Guyana, Suriname, Brazil (Amazonas, Pará, Roraima). H. balsamifera var. iluana Cuatrec., Contr. U.S. Natl. Herb. 35: 115. 1961. Bushy shrub 1–2 m tall; leaves small, rigid-coriaceous, obovate-elliptic, glabrous, ± attenuate and obtuse, emarginate or mucronate at apex, cuneate at base; similar to var. coriacea but differing mainly by its hirtellous petals. Savannas, open shrubland, 900–1600 m; Bolívar (Cerro Guaiquinima, Gran Sabana, base of Ilú-tepui). Endemic. H. balsamifera var. imbaimadaiensis Cuatrec., Contr. U.S. Natl. Herb. 35: 115. 1961. Low shrub with glabrous branchlets and thick, rigid, broadly sessile leaves; flowers greenish white; fruits oblong, ellipsoid, ca. 7 mm long. Shrublands, forest edges, 500– 1400 m; Bolívar (northern Gran Sabana). Guyana.

Humiria 629

H.

balsamifera var. laurina (Urb.) Cuatrec., Contr. U.S. Natl. Herb. 35: 107. 1961. —Humiria floribunda var. laurina Urb. in Mart., Fl. Bras. 12(2): 439. 1877. —Niña. Humiria floribunda var. spathulata Gleason, Bull. Torrey Bot. Club 58: 374. 1931. Low sprawling shrub 0.5–3 m tall, rarely a tree; leaf blades elliptic-oblong, contracted at base into a long petiole; branchlets usually glabrous; fruits purplish black and edible. White-sand savannas and meadows, shrublands, or a tree in caatinga or lower montane forests, 50–300 m; Bolívar (Sabana Cusimi), Amazonas (widespread). Southeastern Colombia, Guyana, Brazil (Amazonas, Pará, Roraima). H. balsamifera var. pilosa (Steyerm.) Cuatrec., Contr. U.S. Natl. Herb. 35: 116. 1961. —Humiria pilosa Steyerm., Fieldiana, Bot. 28: 270. 1952. —Uraurai-yek (Arekuna). Shrub or tree to 12 m tall; leaves rigid, coriaceous, elliptic or oblong-elliptic, hirtellous at the abaxial side, at least at the midrib; young branchlets and inflorescences hirtellous; flowers greenish white. Low montane forests and shrublands, 1100–1600 m; Bolívar (Ayavaparú near base of Ilú- and Wadakapiapué-tepui, Ptari-tepui). Endemic. H.

balsamifera var. savannarum (Gleason) Cuatrec., Contr. U.S. Natl. Herb. 35: 108. 1961. —Humiria savannarum Gleason, Bull. Torrey Club 58: 378. 1931. Low shrub 0.2–1(–2) m tall; leaves narrow, oblong, typically narrow-lanceolate with blunt apex and cuneate base, sessile or shortly (to 2 mm) petiolate. White-sand savannas, low granitic hills, 100–300 m; Bolívar (Cerro Marimarota along the middle Río Orinoco), Amazonas (Canaripó, Cerro Camani, Cerro Moriche, La Esmeralda, Santa Bárbara). Brazil (Amazonas: base of Serra Aracá). H. balsamifera var. stenocarpa Cuatrec., Contr. U.S. Natl. Herb. 35: 114. 1961. Shrub or low tree (1–)2–4(–8) m tall, similar to var. coriacea, but with thinner and longer attenuate leaves and narrower fruits

with a slender, oblong endocarp obtuse at both ends, 11–13.5 × 3.8–4.5 mm. Savannas and shrublands of lower mountains, 400– 800(–1200) m; Bolívar (Cerro Altamira east of Ciudad Piar, Río Asa, Río Hacha near Canaima). Brazil (Roraima: Serra Tepequém). H. balsamifera var. subsessilis (Urb.) Cuatrec., Contr. U.S. Natl. Herb. 35: 102. 1961. —Humiria floribunda var. subsessilis Urb. in Mart., Fl. Bras. 12(2): 439. 1877. —Midi, Niña. Small or medium-sized tree (5–20 m tall) or sprawling shrub (0.2–1.5 m). Lowland forests, savanna edges, white-sand savannas, 50–400 m; Bolívar (Túriba, Los Pijiguaos), Amazonas (widespread in lowlands). Southeastern Colombia, Brazil (Amazonas: Rio Uaupés basin). Humiria crassifolia Mart. ex Urb. in Mart., Fl. Bras. 12(2): 441. 1877. Shrub or treelet 1.5–5(–7) m tall, densely branched. Shrublands, open savannas, 300– 1600 m; Bolívar (Cerro Chiricayen, Río Uaiparú), Amazonas (Cerro Autana, Cerro Aracamuni, Cerro Cuao, Sierra de la Neblina). Colombia (Sierra de Isibukuri), Guyana (Kaieteur Plateau), to be expected on the Brazilian side of Serra da Neblina. ŠFig. 538. Humiria fruticosa Cuatrec., Contr. U.S. Natl. Herb. 35: 118. 1961. Subshrub or low shrub 0.2–1 m tall, with small, narrow, sessile leaves and with pubescence of minute, pointed hairs on leaves and branchlets; drupe globose to ovoid, 5–10 × 6– 8 mm. White-sand savannas and meadows, 100–200 m; Amazonas (near and around base of Cerro Yapacana). Endemic. ŠFig. 537. Humiria wurdackii Cuatrec., Contr. U.S. Natl. Herb. 35: 119. 1961. Slender shrub or treelet 0.5–8 m tall, the branches twisted; leaves narrow, glabrous; drupe ellipsoid, small. Thickets or low forests around savannas in seasonally flooded areas, 100–200 m; Amazonas (Caño Pimichín, Caño Yagua, Río Atabapo basin, Río Autana, Río Guainía, Río Guayapo). Adjacent Colombia. ŠFig. 535.

630

H UMIRIACEAE

Fig. 535. Humiria wurdackii

Fig. 537. Humiria fruticosa

Fig. 536. Humiria balsamifera var. balsamifera

Fig. 538. Humiria crassifolia

Humiriastrum 631

3. HUMIRIASTRUM (Urb.) Cuatrec., Contr. U.S. Natl. Herb. 35: 122. 1961. —Sacoglottis subg. Humiriastrum Urb. in Mart., Fl. Bras. 12(2): 443. 1877, "Saccoglottis." —Sacoglottis sect. Humiriastrum (Urb.) Reiche in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 37. 1890, "Saccoglottis." Small to large trees. Leaves evergreen, coriaceous or subcoriaceous, alternate, petiolate, usually crenate-dentate and acuminate, rarely entire or obtuse; stipules small and deciduous, or lacking. Inflorescences axillary or pseudoterminal, mostly dichotomously or trichotomously branched and paniculate; bracts deltoid, amplectant, persistent or deciduous. Sepals suborbicular, united at base; petals thick-membranaceous, oblong. Stamens 20, in 2 alternating lengths; filaments complanate, glabrous and united in a tube to half their length; anthers ovate or oblong, acuminate or obtuse, attached to filament near the base; thecae 2, unilocular, globose or ellipsoid, inserted at the base of the thick connective, usually lanceolate and acute. Disk cupular, encircling the ovary, finely dentate or deeply fissured. Ovary 5-locular, 5-ovulate, the ovules pendulous; style short; stigma capitate, lobed. Drupe small or medium-sized, ellipsoid or subglobose; exocarp smooth, carnose when fresh, coriaceous or subcoriaceous when dry; endocarp woody, usually without resinous cavities, 5 foramina (small holes) at apex, alternating with 5 oblong germinal opercula or valves on the upper part (rarely reaching near the base). Seeds 1, 2, or rarely 5, oblong. Costa Rica, Panama, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 16 species, 6 in Venezuela, all in the flora area. Key to the Species of Humiriastrum 1. 1. 2(1).

2.

3(2).

3.

Leaf blades obtuse or shortly obtusely apiculate, never subacute or cuspidate; terminal branches hirsute or minutely hirtellous ....................... 2 Leaf blades acutely acuminate or cuspidate, glabrous; branchlets glabrous ....................................................................................................... 4 Bracts deciduous; leaf blades obovate, rounded at apex, or obtusely rhomboid, the margin brevicrenate, the base cuneate, glabrous on the upper surface, pilose mainly at base when young on the lower surface, (2.2–) 3–4.8 × (1.5–)2.8–4.8 cm; petals with the midline dorsally hirsute pseudopetiole 1–2 mm long, the base thickened pulvinate and glandular; sepals copiously hirsute ....................................................... H. liesneri Bracts persistent; leaf blades broadly elliptic, obovate-elliptic, or oblongobovate, 2.5–9 × 1.4–4.8 cm; petals distally hirtellous; pseudopetiole 1– 3 mm long; sepals hirtellous to glabrous .............................................. 3 Leaf blades oblong-obovate, 2.5–9 × 1.4–4 cm, the apex rounded, truncate or obtuse, also retuse, the base cuneate, the margin revolute and entire, the upper surface glabrous except for the hirsute midrib, the lower surface hirsute all over; branchlets ceriferous, hirsute, the trichomes ca. 1 mm long, firm, patent; drupe 20 × 10 mm, smooth ....... H. obovatum Leaf blades broadly elliptic or obovate-elliptic, 4–7 × 2.5–4.8 cm, the apex abruptly contracted and shortly obtusely apiculate, the base shortly cuneate, the margin entire or slightly subcrenate, the apiculum 3.5 mm long, the midrib on young leaves puberulous, the surface on both sides glabrous; young branchlets minutely hirtellous, later glabrous and

632

4(1).

4.

5(4).

5.

H UMIRIACEAE

smooth, often ceriferous, the white wax becoming fibrous; drupe shortly ellipsoid, smooth, 30 × 25 mm (immature) ................ H. ottohuberi Bracts deciduous; leaf blades thin-coriaceous, elliptic or ovate-elliptic, 5– 8 × 3.5 cm, the tip abruptly narrowed and acuminate or cuspidate with 10–17 mm long cusp, the base abruptly cuneate, the margins obsoletely crenate; petiole 2–4 mm long; peduncles of inflorescences minutely hirsute; ovary hispidulous; drupe ellipsoid, rugose when dry, 20–23 × 12– 15 m ................................................................................... H. colombianum Bracts persistent; leaf blades 6–15 × 3–5.5 cm, ovate-oblong, elliptic-oblong, broadly ovate, elliptic, or obovate-elliptic, thick-coriaceous, the apex cuspidate or not, the base not abruptly cuneate, the margins conspicuously crenate; petiole longer than above; peduncles of inflorescences glabrous; drupe oblong or globose, smooth ................................ 5 Leaf blades 8.5–15 × 3.4–5.5 cm, ovate-oblong or elliptic-oblong, acuminate, the cusp included; pseudopetiole 8 mm long; sepals hirsute; ovary sparsely pilose; drupe oblong, 31–36 × 19–24 mm ......... H. piraparanense Leaf blades 6–10 × 3–5 cm, broadly ovate or elliptic or obovate-elliptic, the tip abruptly longly acuminate or cuspidate, the cusp 8–20 mm long; pseudopetiole 4–10 mm long; sepals dorsally glabrous; ovary glabrous; drupe globose, 10–20 mm diameter .................................... H. cuspidatum

Humiriastrum colombianum Cuatrec., Contr. U.S. Natl. Herb. 35: 134. 1961. —Sacoglottis excelsa var. colombiana Cuatrec., Brittonia 8: 196. 1956. Buttressed tree to 30 m tall, with an orange brownish trunk, inner bark and resinous exudate reddish. Evergreen lowland forests, often on sandy soil, 100–200 m; Amazonas (western base of Sierra de la Neblina, upper Río Negro and Río Guianía). Zulia; Colombia, Peru. Humiriastrum cuspidatum (Benth.) Cuatrec., Contr. U.S. Nat. Herb. 35: 130. 1961. —Humirium cuspidatum Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 101. 1853. —Sacoglottis cuspidata (Benth.) Urb. in Mart., Fl. Bras. 12(2): 444. 1877. Medium-sized tree; leaves coriaceous, glabrous. Seasonally flooded lowland forests, Mauritia palm swamps, 100–200 m; Bolívar (middle Río Orinoco at Piedra Marimare), Amazonas (Río Baría, San Carlos de Río Negro). Brazil. Humiriastrum liesneri Cuatrec., Phytologia 71: 165. 1991. Small tree; inflorescences usually shorter than the upper leaves; flowers and fruits unknown. Open tepui forests, ca. 1400 m; Amazonas (Cerro Aracamuni). Endemic.

Humiriastrum obovatum (Benth.) Cuatrec., Contr. U.S. Natl. Herb. 35: 125. 1961. —Humirium obovatum Benth., London J. Bot. 2: 737. 1843. —Sacoglottis obovata (Benth.) Urb. in Mart., Fl. Bras. 12(2): 443. 1877. Tree to 40 m tall; leaves typically coriaceous, rigid, with recurved margins. Lowland to upland forests, 400–900 m; Bolívar (western base of Amaruay-tepui, Río Caura basin, Río Icabarú). Guyana. Humiriastrum ottohuberi Cuatrec., Phytologia 68: 260. 1990. Small tree 4–10 m tall. Seasonally flooded riparian forests, 50–200 m; Amazonas (Caño San Miguel, Río Casiquiare, Río Pasimoni). Endemic. Humiriastrum piraparanense Cuatrec., Contr. U.S. Natl. Herb. 35: 127. 1961. Small to medium-sized tree to 20 m tall; fruits oblong, with a thick green exocarp, reddish when ripe. Riparian forests, 100–200 m; Amazonas (Caño Yagua, Río Sipapo, Río Guainía). Colombia (Vaupés), Brazil (Amazonas). ŠFig. 539. The fruits of Humiriastrum piraparanense are said to be edible.

Sacoglottis 633

Fig. 539. Humiriastrum piraparanense

4. SACOGLOTTIS Mart., Nov. Gen. Sp. Pl. 2: 146. 1827, spelling variant: Saccoglottis. Aubria Baill., Adansonia 2: 265. 1862. Sacoglottis subg. Eusaccoglottis Urb. in Mart., Fl. Bras. 12(2): 442, 448, pl. 94– I, 95. 1877, "Saccoglottis." —Sacoglottis sect. Eusaccoglottis (Urb.) Reiche in Engler & Prantl, Nat. Pflanzenfam. 3(4): 37, fig. 32. 1890, "Saccoglottis." —Sacoglottis sect. Eusaccoglottis (Urb.) Winkler in Engl. & Prantl, Nat. Pflanzenfam. 19a: 128, figs. 58, 59, 1931. Medium-sized or large evergreen trees to 40 m tall, or shrubs with densely branched and leafy crown. Leaves coriaceous or subcoriaceous, pseudopetiolate; blades entire or slightly dentate-crenate; venation prominent or immersed, often with glandular dots near the base. Flowers usually small, white or greenish, pentamerous. Calyx imbricate in aestivation, the sepals united at base, the lobes ovate or rounded; petals free, in aestivation cochlear or quincuncial, thick-membranaceous, oblong, narrowing distally, usually white or greenish white. Stamens 10; filaments complanate, glabrous, united at base into a ± extensive tube, lanceolate and acute distally, the 5 opposite the sepals longer than the alternate 5; anthers ovoid or oblong attached dorsally near the base; thecae 2, unilocular, ellipsoid, dehiscing longitudinally, the connective ovate-acuminate, rather thick at base. Intrastaminal

634

H UMIRIACEAE

disk cupular, dentate, encircling the ovary. Ovary globose or ovate, 5-locular, the cells uniovulate, the carpels opposite to sepals; ovules anatropous, pendulous, with ventral raphe; style as long as or exceeding the stamens, columnar; stigma capitate, 5-lobed, the rays glutinose. Drupe ellipsoid or spherical, large or small; exocarp carnose, subcoriaceous when dry; endocarp woody, ± bullate, usually marked with 5 ribs, filled with resinous, vacuous, round cavities and usually with 1 or 2 oblong seeds, dehiscent at germination by valves almost from apex to base. Costa Rica, Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, disjunct in West Africa [Sacoglottis gabonensis (Baill.) Urb.]; ca. 10 species, 6 in Venezuela, all in the flora area. Key to the Species of Sacoglottis 1. 1. 2(1). 2. 3(2).

3.

4(2).

4.

5(4).

5.

Leaf blades small, 3.5 × 3 cm, the apex obovate, obtuse, or retuse ............ ................................................................................................... S. maguirei Leaf blades 6–22 cm long, the apex attenuate or apiculate ..................... 2 Bracts deciduous; fruits 4–6 cm long ......................................................... 3 Bracts persistent; fruits < 3 cm long ...................................................... ...4 Drupe and endocarp oblong, acute or apiculate, usually pointed, 4–5 × 1.3–2 cm; inflorescences contracted, subglomerate; leaf blades elliptic or ovate, rigidly coriaceous, 8–22 × 3.5–10 cm, usually the crenate margins revolute .......................................................................... S. ceratocarpa Drupe and endocarp ellipsoid, obtuse at both ends, 4–6 × 2.6–3.5 cm; inflorescences short, lax; leaf blades oblong-elliptic, sublanceolate, acuminate or cuspidate, slightly crenate at the flat margin, thin, flexuous, usually prominously veined above, conspicuously reticulate beneath, 6–15 × 2.5–5.7 cm .................................................................. S. amazonica Drupe oblong, usually acute, 1.5–3 × 0.9–1.2 cm; style more than twice as long as the ovary (2–3 mm long); petals 3.5–4.5 mm long; leaves usually rigidly coriaceous, 5–15 × 3–6 cm, the lower surface ± conspicuously reticulate, elliptic or ovate-elliptic, ± apiculate at apex ....... S. guianensis Drupe globose or subglobose, 1.5–2.5 cm diameter; style shorter than to more than twice as long as the ovary; petals 2.5–4 mm long; leaves rigid-coriaceous to thin-coriaceous, 5–13 × 2–6.5 cm, prominently reticulate on one or both sides .................................................................. 5 Style equal or shorter than the ovary, (0.5–0.7 mm long); petals 2.5–3 mm long; drupe 1.5–2.0 cm diameter, with double exocarp 3–5 mm thick; leaf blades rigid-coriaceous, 6–13 × 2.5–6.5 cm, smooth and shining, the upper surface with inconspicuous venation, the lower surface prominently reticulate .......................................................... S. cydonioides Style more than twice as long as the ovary (2–3 mm long); petals 4 mm long; drupe 1.7–2.8 cm long, with simple exocarp 1–2 mm thick; leaf blades thin-coriaceous, 5–11 × 2–2.5 cm, prominuously or prominently reticulate on both surfaces ............................................ S. mattogrossensis

Sacoglottis amazonica Mart., Nov. Gen. Sp. Pl. 2: 146. 1826 [1827]. Tree to 20 m tall; fruits large, ellipsoid, the endocarp full of hollow or resinous cavities making them float easily in water. Sea-

sonally flooded riparian and nonflooded forests, 0–50 m; Delta Amacuro (Caño Araguabisi, Caño Joba-Suburu, Caño Jobure). Lesser Antilles (St. Vincent, Trinidad), Guyana, Ecuador, Peru, Brazil. ŠFig. 541.

Sacoglottis 635

Fig. 540. Sacoglottis maguirei

Fig. 541. Sacoglottis amazonica

Fig. 542. Sacoglottis guianensis

636

H UMIRIACEAE

The washed, naked, usually strongly bullate endocarps are often found among drift material stranded on tropical and even temperate or northern beaches. Sacoglottis ceratocarpa Ducke, Bol. Técn. Inst. Agron. N. 4: 13. 1945. Medium-sized to tall tree; fruits elongate, often pointedly apiculate; inflorescences very short, with glomerate flowers and deciduous bracts. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (Río Caura, Río Paragua), Amazonas (base and slopes of Sierra de la Neblina). Colombia, Peru, Brazil. Sacoglottis cydonioides Cuatrec., Contr. U.S. Natl. Herb. 35: 183. 1961. Medium-sized or large tree to 40 m tall. Evergreen lowland forests, 200–600 m; Delta Amacuro (Río Toro), Bolívar (Río Paragua, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil (Acre, Amapá, Amazonas). Sacoglottis guianensis Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 103. 1853. —Peru-yek (Arekuna), Ponsigué montañero. Sacoglottis amazonica Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 104. 1853. Sacoglottis guianensis f. dolichocarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 179. 1922. Sacoglottis guianensis var. major Ducke, Arq. Inst. Biol. Veg. 4: 27. 1938. Sacoglottis guianensis var. hispidula Cuatrec., Contr. U.S. Natl. Herb. 35(2): 180. 1961. Shrub or medium-sized tree (1.5–)3–10 (–30) m tall. Gallery forests, evergreen lowland forests, occasionally in savannas, 50–

1300 m; widely scattered in Bolívar and Amazonas. Colombia, Guyana, French Guiana, Suriname, Peru, Brazil. ŠFig. 542. This is an extremely polymorphic species with several described varieties and forms, usually difficult to determine because most of the collections are lacking some of the necessary parts. The variations are in the size of leaves, their margins, the size of the fruits, and the glabrous or hirsute condition of branchlets and petals and the length of petioles. Sacoglottis maguirei Cuatrec., Contr. U.S. Natl. Herb. 35: 163. 1961. Small tree with spreading branchlets. Low tepui-summit forests, ca. 1200 m; Amazonas (summit of Cerro Yapacana). Endemic. ŠFig. 540. Sacoglottis mattogrossensis Malme, Ark. Bot. 22(7): 9. 1929 [1928]. Sacoglottis guianensis f. sphaerocarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 178. 1922. Small to medium-sized tree to 15–25 m tall. Riparian, nonflooded, and secondary forests, 50–200 m; Bolívar (Lago Guri area, Río Caroní), Amazonas (Isla Ratón, Río Baría, Río Cunucunuma, Río Manapiare basin). Colombia, Guyana, Peru, Brazil. Sacoglottis mattogrossensis is polymorphic, presenting several forms based on the glabrous or hirsute nature of the branchlets and of the petals, characters mostly not present together in the specimens, for which reason many collections cannot be named safely to the variety. It is closely related to S. guianensis, from which it differs by its globose fruits and its usually thinner, more flexible, more conspicuously veined leaves.

5. SCHISTOSTEMON (Urb.) Cuatrec., Contr. U.S. Natl. Herb. 35: 146. 1961. —Sacoglottis subg. Schistostemon Urb. in Mart., Fl. Bras. 12(2): 443, 445. 1877, "Saccoglottis." —Sacoglottis sect. Schistostemon (Urb.) Reiche in Engl. & Prantl., Nat. Pflanzenfam. 3(4): 37, fig. 32. 1890, "Saccoglottis." Large to small evergreen trees. Leaves coriaceous or subcoriaceous, rigid, the margin ± crenate or entire, sessile or pseudopetiolate, often with glandular marks. Inflorescences axillary or terminal, cymose or paniculate with dichotomous or trichotomous branching, closely contracted and aggregate or expanded and large. Flowers pentamerous, tetracyclic. Calyx quincuncial, the sepals suborbicular, ciliate at margin, shortly united at base; petals thick, oblong, hirtellous or glabrous, usually white or greenish white. Stamens 20, 5 opposite sepals the longest, distally tri-

Schistostemon 637

furcate and triantheriferous, 5 opposite petals a little shorter, entire with a single anther, and 10 intermediate the shortest, also with a single anther; filaments flattened, glabrous and united at the base in a tube to the middle; anthers ovate, ovatelanceolate, or obovate with 2 unilocular thecae, these ellipsoid, laterally attached, the connective very thick, acuminate or obtuse, the lateral anthers of the trifurcate stamens or those of the shortest stamens occasionally sterile. Disk cupular. Ovary 5locular, initially 5-carpellate; ovules pendulous; style columnar, long or often short; stigma stellate, flat-capitate, the 5 rays glutinose. Fruit drupaceous, rather large, ellipsoid or globose, the exocarp subcoriaceous, thin to very thick; endocarp woody, seldom 5-celled, rather filled by fibrous-cavernous tissue with round, unequal, resinous cavities, surrounding 1 or 2 oblong seeds; endocarp surface ± bullate or almost smooth marked with 5 thin furrows; exocarp becomes ± carnose in some species and is edible. Colombia, Venezuela, Guyana, French Guiana, Suriname, Peru, Brazil; ca. 9 species, 4 in Venezuela, all in the flora area. Key to the Species of Schistostemon 1.

1. 2(1).

2.

3(2). 3.

Leaf blades broadly elliptic, orbicular, or obovate, very obtuse or retuse at apex, sessile or shortly petiolate; fruit globose, 3.6–5 cm wide .............. .................................................................................................... S. retusum Leaf blades ovate, elliptic, or oblong, ± apiculate, acute at apex, clearly pseudopetiolate; fruit ellipsoid or oblong, narrower than above ......... 2 Leaf blades oblong, ca. 4 times as long as wide, coriaceous smooth, only main veins slightly visible or obsolete on lower surface; fruit oblong and pointed; young branchlets and petioles minutely hirtellous; connective of the anthers lanceolate ...................................... S. oblongifolium Leaf blades ovate or elliptic, 2(–3) times as long as wide, rigid-coriaceous, glabrous; fruit broadly oblong, not pointed; connective of the anther ovate or obovate, obtuse ........................................................................ 3 Leaf blades dark greenish, strongly, minutely reticulate on both surfaces; young branchlets glabrous; petals 4–4.5 mm long ...... S. auyantepuiensis Leaf blades brownish-greenish, weakly veined, laxly reticulate on upper surface, veins hardly noticed on lower surface; young branchlets minutely hirtellous; petals ca. 3 mm long ................................ S. fernandezii

Schistostemon auyantepuiensis Cuatrec., Contr. U.S. Natl. Herb. 35: 15. 1961. Small leafy shrub to 2 m tall. Lower montane cloud forests, scrub on rocky tepui slopes, 900–1100 m; Bolívar (Auyán-tepui near Guayaraca). Endemic. Schistostemon fernandezii Cuatrec., Phytologia 68: 261. 1990. Sprawling shrub or low tree 1–4.5 m tall. Riparian habitats, rocky tepui slopes, 400– 1000 m; Bolívar (Guayaraca, near Urimán along Río Caroní at Laguna Capaura). En-

demic. ŠFig. 544. Schistostemon fernandezii is very similar to S. auyantepuiensis, from which it questionably differs mainly by its smaller petals and a weaker leaf venation. Schistostemon oblongifolium (Benth.) Cuatrec., Contr. U.S. Natl. Herb. 35: 148. 1961, "oblongifolius." —Humirium oblongifolium Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 103. 1853. —Sacoglottis oblongifolia (Benth.) Urb. in Mart., Fl. Bras. 12(2): 447, pl. 93. 1877. —Merecure de ardita, Yurí de caracol.

638

H UMIRIACEAE

Shrub 2–4 m tall, treelet, or tree to 20 m or more tall; leaves elongate, leathery; fruits usually oblong, pointed. Seasonally flooded riparian forests, 50–200 m; Amazonas (Río Atabapo, Río Casiquiare, upper Río Negro, Río Sipapo, Samariapo, Santa Bárbara, San Carlos de Río Negro, Tamatama). Brazil (Amazonas). ŠFig. 545.

Schistostemon retusum (Ducke) Cuatrec., Contr. U.S. Natl. Herb. 35: 156. 1961, "retusus." —Sacoglottis retusa Ducke, Arq. Inst. Biol. Veg. 4: 26, 29, pl. 2 fig. d. 1938. Small or medium-sized tree. Evergreen lowland and low, open forests, 100–200 m; Amazonas (Caño San Miguel, Río Pasimoni, San Carlos de Río Negro). Colombia (Vaupés), Brazil. ŠFig. 543.

Fig. 543. Schistostemon retusum

Schistostemon 639

Fig. 544. Schistostemon fernandezii

Fig. 545. Schistostemon oblongifolius

640

H UMIRIACEAE

6. VANTANEA Aubl., Hist. Pl. Guiane 1: 572. 1775. Lemniscia Schreb., Gen. Pl. 1: 358. 1789. Helleria Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 38. 1824. Shrubs or small to large evergreen trees. Leaves simple, coriaceous or subcoriacous, petiolate or sessile, often with glandular impressions or marks. Inflorescences cymose-paniculate, usually monochasial of dichotomic aspect. Flowers pentamerous. Calyx quincuncial with sepals ± united to a subentire margin and cupular; petals free, usually thick, rigid-oblong or linear, slightly attenuate upward, the aestivation contorted. Stamens 50–120 or more, rarely fewer; filaments glabrous, usually thin, flexuous, united at base into a ring; anthers connate near the base, the connective thick, oblong-ovate, acuminate, acute, or obtuse; thecae elliptic, lateral, bilocular, each cell dehiscent through a longitudinal cleft. Disk cylindriccupular. Ovary 5-locular, the cells biovulate, the anatropous ovules superposed; style erect, as long as the stamens or longer; stigma 5-lobed, ± glutinous. Fruit a drupe, to 8 cm long, spherical, ovoid, ellipsoid, or oblong; exocarp ± thick and carnose, subcoriaceous when dry; endocarp woody, usually compact, without resiniferous cavities, dehiscent at germination by longitudinal, subequal, oblong valves or opercules. Seeds usually only 1 per fruit, rarely 2 or 3. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; ca. 19 species, 3 in Venezuela, all in the flora area. Key to the Species of Vantanea 1.

1.

2(1).

2.

Flowers large, 3–4 cm long; petals and stamens red, purplish, or pinkish; ovary glabrous; disk glabrous; drupe ellipsoid, ca. 6 cm long, the endocarp woody, somewhat rugose, 5-sulcate; leaf blades 6–14 × 2.7–6 cm; petioles 6–12 mm long ............................................................ V. guianensis Flowers small, < 2 cm long; petals white; ovary glabrous or long-hirsutevillous; disk glabrous or hirsute-tomentose; drupe 2.5–5 × 2.2–3 cm, ellipsoid; leaf blades 3–10 × 1.6–5.5 cm; petioles 1–2 mm or 5–15 mm long ................................................................................................................ 2 Intrastaminal disk glabrous; ovary glabrous; petals 12–14 mm long; leaf blades 3–8.5 × 1.6–4.5 cm, green; petioles 1–2 mm long; drupes 3.5–5 × 2.2–3 cm, ellipsoid .......................................................................... V. minor Intrastaminal disk hirsute-tomentose; ovary long-hirsute-villous; petals 7–8 mm long; leaf blades 5–10 × 2–5.5 cm, coriaceous, usually reddishshining on lower surface; petioles 5–15 mm long; drupes 2.5–2.8 × 2.2– 2.5 cm, subglobose .................................................................... V. parviflora

Vantanea guianensis Aubl., Hist. Pl. Guiane 572. 1775. —Lemniscia guianensis (Aubl.) J.F. Gmel., Syst. Nat. 2(1): 817. 1791. Lemniscia floribunda Willd., Sp. Pl. 2: 1172. 1800 [1799]. Medium-sized to large tree to 30 m tall; bark rough, brownish, thick; wood yellowish; inflorescences showy; flowers large, red. Riparian forests, nonflooded premontane for-

ests, 100–200 m; Amazonas (Río Yatúa, western base of Sierra de la Neblina). Guyana, French Guiana, Suriname, Ecuador, Brazil (Acre, Amazonas, Pará). Vantanea minor Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 99. 1853. Low shrub 1–4 m tall or a medium-sized tree 6–10(–15) m tall. Upland gallery forests, upland savannas, rocky shrublands, (400–)

HYDROCHARITACEAE 641

800–1400 m; Bolívar (Gran Sabana). Adjacent Guyana. ŠFig. 546. Vantanea parviflora Lam., J. Hist. Nat. 1: 145. 1792. —Conori. Vantanea cupularis Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 6: 83. 1910. Small to large tree to 28 m tall. Nonflooded evergreen lowland forests, 100–200 m; Amazonas (Río Baría, Río Negro). Guyana, French Guiana, Peru, Brazil (Amazonas, Pará).

Fig. 546. Vantanea minor

HYDROCHARITACEAE by Robert R. Haynes and Lauritz B. Holm-Nielsen Annual or perennial aquatic herbs of fresh, brackish, or marine waters, growing entirely submersed, or with both submersed and floating leaves, or with submersed stolons and emergent leaves, without milky juice, glabrous or less often pubescent. Roots fibrous, few to many, nonseptate, at nodes or bases of stems. Stems either rhizomatous, creeping with an abbreviated, erect axis at the nodes, or erect, leafy, elongate, without teeth or tubers, and rarely with turions (compact stem tips that break off and survive any unfavorable season before beginning growth into new plants). Leaves basal, alternate, opposite, or whorled, entire or serrate, sessile or petiolate, 1–many-veined; stipules sometimes present, forming a tubular sheath around the stem; venation parallel, connected by perpendicular or ascending cross veins, the infravaginal scales membranous. Inflorescence axillary, terminal, or scapose, solitary or cymose, subtended by a spathe consisting of a bifid bract or a pair of opposite bracts, the spathe often persistent, sessile or pedunculate, often ridged or winged. Flowers unisexual, often with rudiments of the opposite sex, or bisexual (plants monoecious, dioecious, or hermaphroditic), actinomorphic or rarely slightly zygomorphic. Perianth mostly of 6 separate parts differentiated into sepals and petals, rarely only 3 sepals; sepals mostly green, valvate, persistent; petals mostly colored, imbricate or convolute, deciduous. Androecium with stamens absent or 2–many in 1 or more whorls, the inner often staminodal; anthers basifixed, 2-locular, dehiscing by parallel vertical slits; filaments slender, rarely absent; pollen spherical, rarely united into slender chains. Gynoecium with carpels absent or 2–5 united; ovary infe-

642

H YDROCHARITACEAE

rior, linear, lanceolate, or ovate, unilocular, the placentation parietal or the parietal zones ill-defined and placentation then laminar; ovules numerous, bitegmic, anatropous; style filiform; stigma equaling the carpels. Fruit berry-like, submersed, linear, lanceolate, or ovate, opening by decay of the pericarp. Seeds many, fusiform, elliptic, ovate, or globose; testa smooth, papillose, or spiny; embryo straight; endosperm absent in mature seed. Native mainly to waters of the tropical and subtropical regions of the world, but also present in temperate areas; 16 genera and ca. 100 species, 2 genera and 2 species in the flora area. Key to Genera of Hydrocharitaceae 1. 1.

Leaves cauline, linear, sessile, serrate, without aerenchyma on lower surface; plants submersed; flowers floating ...................................... 1. Elodea Leaves basal, oval to reniform, petiolate, entire, with aerenchyma on lower surface; plants floating or stranded at edge of water; flowers erect ....................................................................................... 2. Limnobium

1. ELODEA Michx., Fl. Bor.-Amer. 1: 20. 1803. Perennial, glabrous, dioecious or hermaphroditic, submersed in fresh waters, readily propagated by seeds, stem fragments, or turions. Roots smooth, slender, pale, unbranched. Stems erect, rooting at lower nodes, branched or unbranched. Leaves simple, sessile, linear to linear-lanceolate, in whorls of 3–7, or rarely opposite, 1-veined, serrate, the midrib without a dorsal prickle. Inflorescence solitary, axillary; spathes bisexual or unisexual, sessile, usually narrowed toward base, cylindric to elliptic-spatulate, 1-flowered. Flowers bisexual or unisexual, usually projected to surface of water by the elongating hypanthium base. Sepals 3, herbaceous, green; petals 3, membranous, white to pale blue, separate, elliptic, clawed. Stamens 3–9, or reduced to 3 staminodes; anthers oblong to ellipsoid; filaments subulate to lanceolate, separate or the 3 inner united halfway to apex. Carpels 3, the locules 1, the placentation parietal; styles 3; stigmas 3, bifid. Fruit ovoid to lance-ellipsoid, beaked. Seeds 3–8, cylindric to fusiform. Western Hemisphere; 6 species, 1 in Venezuela. The six species of Elodea are separated into subgenus Elodea and subgenus Apalanthe Planch. The subgenera can be separated by subgenus Elodea with five species, having unisexual flowers, the staminate flowers with 6–9 stamens and some filaments united. Subgenus Apalanthe on the other hand, with E. granatensis, has bisexual flowers with 3 stamens having separate filaments. Only subgenus Apalanthe occurs in the flora area. Two species of subgenus Elodea, E. potamogeton (Bertero) Espinoza and E. callitrichoides Casp., occur in western and southern South America, but not in Venezuela. The taxonomy of Elodea is based on differences in floral structure. Without flowers, plants are difficult to determine. Care should be taken in all cases to collect flowering specimens. Elodea granatensis Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 150. 1809 [1813]. —Udora granatensis (Bonpl.) Spreng., Syst. Veg. 4(2): 25. 1827. —Apalanthe granatensis (Bonpl.) Planch., Ann. Mag.

Nat. Hist. ser. 2, 1: 87. 1848. —Philotria granatensis (Bonpl.) Vict., Contr. Lab. Bot. Univ. Montréal 18: 42. 1931. Elodea orinocensis Rich., Mém. Cl. Sci. Math. Inst. Natl. France 12(2): 75. 1814.

Limnobium 643

—Philotria orinocensis (Rich.) Vict., Contr. Lab. Bot. Univ. Montréal 18: 42. 1931. Plant to 26 cm long. Submersed in small lakes and quiet backwaters of rivers, 100– 300 m; Bolívar (La Paragua), Amazonas (Río Negro). Colombia, Guyana, Suriname, Brazil, Bolivia. ŠFig. 547.

Fig. 547. Elodea granatensis

2. LIMNOBIUM Rich., Mém. Cl. Sci. Math. Phys. Inst. Natl. France 12(2): 66. 1814. Perennial, glabrous, monoecious plants growing emersed or with stems floating in fresh waters, readily propagated by seeds or stolons. Roots 1 per node, branched, fibrous, with large root hairs. Stems dimorphic, short ones erect, unbranched, bearing leaves in a rosette and elongate ones stoloniferous, branching. Leaves dimorphic, the lower scale-like, 2 at base of each rosette, membranous, ovate, the upper leaves differentiated, emergent or floating, petiolate, stipulate, laminate; stipules membranous, ovate, sheathing the developing apex; petiole stout, sometimes inflated; blade elliptic to orbicular, the apex obtuse to acuminate, the base reniform or cordate, the abaxial surface smooth or emergent or with aerenchymous tissue on floating leaves. Inflorescence unisexual, cymose; staminate inflorescences sessile or pedunculate, with up to 25 flowers, with 2 ovate bracts, the lower shorter than the upper one; pistillate inflorescences mostly sessile, rarely short-pedunculate, with 1–6 flowers, with 1 or 2 separate ovate bracts. Flowers pedicellate, projected above the surface of water; staminate flowers withering after anthesis, the sepals 3, separate, narrowly to widely elliptic, the apex reflexed at anthesis, greenish white to yellowish, the stamens in 1–6 whorls of 3, inserted on a column of fused filament bases, the anthers 4-loculed, dehiscing lengthwise, the pollen spherical, yellow, separate; pistillate flowers epigynous, the pedicels reflexed after anthesis, the sepals 3, separate, narrowly to widely elliptic, spreading at anthesis, greenish white, the staminodes 2–6, in 1 whorl, the ovary of 3–9 united carpels, unilocular, ellipsoid to oblong, the styles equaling the carpels, each split into 2 filiform stigmas, the stigmas much shorter than the style, with unicellular papillae, the ovules 5 to numerous, the placentation parietal. Fruit ellipsoid to spherical, beaked, developing in mud or under water, the pericarp splitting irregularly. Seeds few to numerous, ellipsoid, with a short micropylar beak, the funiculus persistent; testa covered with cylindric, blunt trichomes. Eastern U.S.A., Mexico, Central America, West Indies, South America except Chile; 2 species, 1 in Venezuela. The second species, Limnobium spongia (Bosc) Steud., is restricted to the eastern U.S.A.

644

H YDROCHARITACEAE

Limnobium laevigatum (Humb. & Bonpl. ex Willd.) Heine, Adansonia n.s. 8: 315. 1968. —Salvinia laevigata Humb. & Bonpl. ex Willd., Sp. Pl. 5: 7. 1810. —Hydromystria laevigata (Humb. & Bonpl. ex Willd.) Díaz-Mir. & Philcox., Bot. J. Linn. Soc. 83: 321. 1981. Plant to 30 cm tall. Widespread along margins of lakes and slow-moving water of small rivers, 0–100 m; widespread in Delta Amacuro. Scattered through northern Venezuela; Central Mexico, Central America, West Indies, South America except Chile. ŠFig. 548.

Fig. 548. Limnobium laevigatum

HYDROPHYLLACEAE by James S. Miller Perennial or annual herbs or less commonly shrubs (Eriodictyon, Hydrolea, Wigandia), sometimes spiny, often hairy, the hairs often from a basal cystolith, sometimes stinging. Leaves alternate or rarely opposite, simple and entire to pinnately or palmately divided, exstipulate. Inflorescences terminal or axillary, cymose, often helicoid, rarely with the flowers solitary. Flowers actinomorphic or rarely slightly zygomorphic, hypogynous. Calyx deeply (4)5-lobed, rarely 10–12-lobed (Codon), the lobes imbricate, often with appendages between the lobes; corolla sympetalous, 5(6–10)-lobed, imbricate or rarely convolute. Stamens the same number as and alternate with the corolla lobes; filaments adnate to the corolla tube; anthers bithecal, dehiscing longitudinally. Ovary superior, 2-carpellate, usually unilocular, the placentation parietal, rarely axile; styles 1 or 2; ovules 2–many, anatropous or amphitropous. Fruits capsular, loculicidally or rarely septicidally or irregularly dehiscent; seeds with abundant to scant, oily, carnose endosperm. Cosmopolitan (greatest diversity in dry areas of the western United States); ca. 20 genera and 250 species, 1 genus and 2 species in the flora area.

Hydrolea 645

Fig. 550. Hydrolea spinosa

Fig. 549. Hydrolea elatior

646

H YDROPHYLLACEAE

1. HYDROLEA L., Sp. Pl. ed. 2, 1: 238. 1762. Perennial or annual herbs or shrubs, the stems sometimes spiny, glabrous or often with glandular hairs. Leaves alternate, simple, entire. Inflorescences axillary or terminal, cymose or corymbose. Flowers bisexual, actinomorphic or slightly zygomorphic. Calyx 5-lobed, divided to nearly the base, persistent and often accrescent, the lobes unequal, imbricate. Corolla blue or rarely white, campanulate to rotate, 5lobed. Stamens 5, usually included in the corolla tube, sometimes shortly exserted, glabrous; filaments adnate to the corolla tube, dilated at the base; anthers bithecal, sagittate, dorsifixed. Ovary superior, 2-locular, the placentation axile; styles 2(3–5); stigmas clavate to capitate; ovules numerous. Fruits capsular, loculicidally, septicidally, or irregularly dehiscent; seeds small, numerous. Southern U.S.A., Central America, West Indies, South America as far south as Argentina, Africa, Asia; ca. 20 species, 2 in Venezuela, both in the flora area. Key to the Species of Hydrolea 1. 1.

Plants without spines, usually glabrous or nearly so ...................... H. elatior Plants with spines in the leaf axils, usually glandular pubescent ............. ..................................................................................................... H. spinosa

Hydrolea elatior Schott in Spreng., Syst. Veg. 4(2): 404. 1827. Hydrolea multiflora Mart. ex Choisy in DC., Prodr. 10: 180. 1846. Hydrolea minima Brand, Notizbl. Bot. Gart. Berlin-Dahlem 10: 120. 1927. Herb to 60 cm tall; flowers blue. Mud flats along rivers, disturbed places, 0–200 m; Delta Amacuro (Caño Mánamo), Bolívar (Movtaro, Río Orinoco between Río Parhueña and Río Horeda,), Amazonas (San Carlos de Río Negro). Apure, Cojedes, Guárico, Portuguesa, Zulia; Mexico, Honduras, Colombia, Ecuador, Guyana, Brazil, Paraguay, Argentina. ŠFig. 549. Hydrolea spinosa L., Sp. Pl. ed. 2, 238.

1762. —Nama spinosa (L.) Kuntze, Revis. Gen. Pl. 2: 434. 1891. Herb or suffrutescent to 1 m tall; flowers blue. Disturbed areas, along streams, rivers, lagoons, wet savannas, 0–500 m; Delta Amacuro (Laguna Ataguía near San José de Macareito), Bolívar (Ciudad Bolívar, Ciudad Piar, El Dorado, El Manteco, El Palmar, lower Río Cuyuni, Represa Guri, middle and upper Río Paragua, Upata), Amazonas (20 km north of Puerto Ayacucho). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Falcón, Guárico, Lara, Mérida, Monagas, Portugesa; widespread in Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay. ŠFig. 550.

ICACINACEAE by Richard A. Howard and Rodrigo Duno de Stefano Trees, shrubs, or lianas, generally all parts pubescent or glabrous, the pubescence persistent or deciduous, rarely with lepidote-stellate trichomes or clustered trichomes. Large tubers rarely present. Leaves alternate, simple, exstipulate; petioles canaliculate; blades coriaceous to subcoriaceous, rarely membranaceous; apex longacuminate to obtuse, rarely mucronate or emarginate; base acute or attenuate, rarely cordate; margins entire, slightly revolute, or the juvenile leaves spinosetipped or -margined (not in flora area); pore-cavities rarely present in the abaxial surface (not in flora area); the secondary veins joined together in a series of mar-

I CACINACEAE 647

ginal loops, free or indistinct, alternate, rarely opposite. Inflorescences axillary, extra-axillary, terminal, or cauliflorous, cymose or racemose; bracts 1–3, rarely ebracteate (not in South American taxa); pedicels generally articulated near the end. Flowers actinomorphic, rarely zygomorphic, bisexual or unisexual, hypogynous. Calyx (4)5-lobed or -toothed, campanulate, fleshy, pubescent without, imbricate. Petals (4)5(6), free, rarely united, pubescent or glabrous, valvate, reflexed; the apex acute, inflexed, sometimes extended into an appendage. Stamens (4)5(6), alternate with the petals; filaments fleshy, filiform, or broad and flattened, occasionally with an adaxial appendage and covered with claviform trichomes; anthers 2(4)-locular, dorsi- or basifixed, longitudinally dehiscent, introrse or lateral; connective expanded, sometimes extended into an appendage. Disk present or absent, basal, free (not in South American taxa), sometimes reniform or a differentiated fleshy sterile ring at the base of the ovary. Pistil with the ovary pubescent or glabrous, 1(2 or 3)locular, rarely pubescent inside, rudimentary in staminate flowers; ovule 2 per locule, pendent from near the apex, collateral or superposed; style 1, terminal or rarely excentric, rarely incurved, additional rudimentary styles sometimes present; stigma capitate or papillose. Fruit a drupe, ovoid, oblong, globose, or rarely flattened and with a large oblong fleshy appendage whitish on concave side; endocarp sometimes with ribs. Seed 1; embryo commonly minute; endosperm copious, nonruminate. Pantropics; ca. 60 genera and ca. 451 species, 7 genera and 16 species in the flora area. Two additional genera, Calatola and Citronella, occur in Venezuela north of Río Orinoco. Poraqueiba sericea is locally cultivated for the edible oily mesocarp and the starchy endosperm of the seed. The tubers of Leretia cordata are rich in starch. Key to the Genera of Icacinaceae 1.

1.

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

4(3).

4.

Plants covered by a diminutive lepidote-stellate indument; foliage turning black when dry; petals joined at the base forming a short tube, the apex of the lobes extended in a long clavate appendage; stamens adnate to the corolla ............................................................. 2. Dendrobangia Plants covered by indument but not lepidote-stellate, sometimes glabrous; foliage rarely turning black when dry; petals free, the apex not extented into an appendage; stamens free from the corolla ................ 2 Lianas, rarely shrubs with scandent branches; petioles with a neat abscission line at the base ............................................................................... 3 Trees or shrubs with scandent branches; petioles without a neat abscission line at the base ............................................................................... 5 Plants covered with malpighiaceous trichomes; inflorescences generally axillary; fruits glabrous outside ................................................... 5. Leretia Plants covered by indument but without malpighiaceous trichomes; inflorescences axillary, supra-axillary or terminal; fruits pubescent outside ................................................................................................................ 4 Plants covered by hispid trichomes; secondary, tertiary, and quaternary venation conspicuous; inflorescences 1–3 spiciform racemes; flowers not articulated; fruits ellipsoid, flattened laterally .......... 6. Pleurisanthes Plants covered by indument but without hispid trichomes, leaves nearly glabrous; tertiary and quaternary venation almost indistinct; inflorescence a single umbelliform panicle; fruits ovoid .................. 1. Casimirella

648

5(2).

5.

6(5).

6.

I CACINACEAE

Secondary veins joined together near the margin; filaments with an appendage covered with claviform trichomes; fruits oblong and flattened with a large fleshy appendage covering the concave side, and strongly five-ribbed on the convex surface .......................................... 3. Discophora Secondary veins free near the margin, sometimes indistinct; filaments exappendiculate; fruits ovoid to globose, never flattened and covered with a fleshy appendage ........................................................................ 6 Secondary veins conspicuous or indistinct, tertiary venation always indistinct; flowers with petals barbate, not longitudinally furrowed within; fruits compressed dorsoventrally, 1.5–2 cm long, mesocarp thin, locules 2 or 3 ................................................................................ 4. Emmotum Secondary and tertiary veins conspicuous; flowers with petals glabrous or slightly pubescent, longitudinally furrowed within; fruits oblong or ellipsoid, 4.5–7 cm long, mesocarp thick, fleshy, 1-locular ...... 7. Poraqueiba

1. CASIMIRELLA Hassl., Repert. Spec. Nov. Regni Veg. 12: 249, fig. 251. 1913. Humirianthera Huber, Bull. Soc. Bot. Genève sér. 2, 6: 184. 1914. Vines or shrubs with scandent branches; generally all parts with fascicles of trichomes, rarely simple trichomes. Large tubers present. Leaves chartaceous to subcoriaceous; blade narrowly oblong, narrowly elliptic, or nearly orbiculate (not in flora area), pubescent or glabrous, the apex obtuse or acute, the base obtuse or cor-

Fig. 551. Casimirella ampla

Dendrobangia 649

date; secondary veins 4–6 pairs, alternate, joined together in a series of marginal loops. Inflorescence axillary, supra-axillary, or terminal, 1(2 or 3) umbelliform panicles, to 20 cm long; pedicels articulated near the end. Flowers actinomorphic, bisexual, 5(6)-parted. Calyx deeply lobed, unequal, pubescent without; petals free, pubescent on both sides. Stamens with filiform filaments, glabrous; anther sacs globose or oblong; connective lineal, extending into an appendage. Disk absent. Pistil with the ovary hirsute, 1-locular, pubescent inside; style 1, incurved, 1 or 2 terminal rudimentary styles; stigma capitate. Fruit ovoid, endocarp woody, pubescent inside, exocarp pubescent. Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; 7 species, 1 in Venezuela. Casimirella ampla (Miers) R.A. Howard, Brittonia 44: 168. 1992. —Leretia ampla Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 364. 1859. —Mappia ampla (Miers) Engler in Mart., Fl. Bras. 12(2): 51. 1872. —Humirianthera ampla (Miers) Baehni, Candollea 7: 182. 1936. Humirianthera duckei Huber, Bull. Soc. Bot. Genève, sér. 2, 6: 184. 1914. Liana, often high-climbing, or shrub with scandent branches; leaf blade 4.5–20 × 1.5–

10 cm, narrowly elliptic or narrowly oblong, almost glabrous. Evergreen lowland forests, 50–100 m; Amazonas (Isla Ratón, near Puerto Ayacucho, near San Carlos de Río Negro). Colombia, French Guiana, Ecuador, Peru, Brazil. ŠFig. 551. The tubers weigh 5–20 kilos and are rich in starch. In Brazil, they are eaten after washing out the bitter-tasting compounds.

2. DENDROBANGIA Rusby, Mem. Torrey Bot. Club 6: 19. 1896. Clavapetalum Pulle, Recueil. Trav. Bot. Néerl. 9: 148. 1912. Asterolepidion Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 207. 1922. Trees 5–25 m tall, all parts lepidote-stellate. Leaves subcoriaceous or coriaceous, turning black when dry; blades elliptic, ovate, narrowly ovate, oblong, or narrowly oblong, the apex acute, base acute, or attenuate; secondary veins 6–8(–20) pairs, alternate, slightly joined together in a series of marginal loops, conspicuous or indistinct. Inflorescence axillary, paniculate; pedicels very short, articulated at the end. Flowers actinomorphic, bisexual, 5-parted. Calyx campanulate, lobed, lepidotestellate without, imbricate. Corolla gamopetalous, lobed, glabrous, the apex extended into a long clavate appendage. Stamens with filiform filament, very short, glabrous, adnate to the corolla; anther dorsifixed, sacs oblong. Disk absent. Pistils lepidote-stellate or glabrous; ovary, 1-locular; style short; stigma minute. Fruit oblong; mesocarp thin and fleshy; exocarp lepidote-stellate, becoming glabrous. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela. Dendrobangia boliviana Rusby, Mem. Torrey Bot. Club 6: 19. 1896. Clavapetalum surinamense Pulle, Recueil Trav. Bot. Néerl. 9: 148. 1912. Asterolepidion elatum Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 207. 1922. —Clavapetalum elatum (Ducke) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 16. 1925. Dendrobangia tenuis Ducke, Bol. Técn. Inst. Agron. N. 4: 15. 1945.

Tree 15–40 m tall; leaf blade 7–20 × 2.5–9 cm, ovate, narrowly ovate, or narrowly oblong. Evergreen lowland forests, 100–200 m; Bolívar (Río Cuyuní), Amazonas (base of Cerro Cuao, base of Cerro Huachamacari, Río Cunucunuma, Río Ventuari). Aragua, Carabobo, Sucre, Yaracuy; Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 552.

650

I CACINACEAE

Fig. 552. Dendrobangia boliviana

3. DISCOPHORA Miers, Ann. Mag. Nat. Hist. ser. 2, 10: 118. 1852. Kummeria Mart., Fl. Bras. 12(2): 52. 1872. —Kummeria Mart., Herb. Fl. Bras. no. 1276. 1837, nom. nud. Shrubs or trees 5–10 m tall, all parts slightly or very pubescent. Leaves subcoriaceous to coriaceous; blades oblong, narrowly oblong, ovate, or lanceolate, the apex acute or acuminate, the base acute or attenuate; secondary veins 7–13 pairs, alternate, joined together in a series of marginal loops, very near the margin. Inflorescence axillary, supra-axillary or cauliflorous, thyrsoid; pedicels articulate at the end. Flowers actinomorphic, bisexual or unisexual, 5-parted. Calyx campanulate, slightly lobed, or mucronate, slightly pubescent without. Petals free, glabrous. Stamens with fleshy filament, with adaxial swelling appendage midway along its length, covered with claviform trichomes; anthers attached basally, divergent at base. Disk a pulviniform tissue around the ovary, reniform. Pistil with the ovary glabrous, cylindrical or angled, slightly compressed, 1-locular abortive and rudimentary in staminate flowers; style not evident; stigma fleshy. Fruit flattened, slightly curved, with large oblong, fleshy, whitish appendage on the concave side, on convex side the endocarp with 2–4 pairs of primary ribs and one median rib. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, 2 in Venezuela, both in the flora area.

Emmotum 651

Key to the Species of Discophora 1. 1.

Leaves densely pubescent with long and erect trichomes; blades narrowly ovate to lanceolate; secondary veins 10–13 ................................. D. froesii Leaves glabrous, if pubescent, trichomes shorter and more adpressed than above; blades oblong, narrowly oblong, ovate, or narrowly ovate; secondary veins 7–10 ............................................................. D. guianensis

Discophora froesii Pires, Bol. Técn. Inst. Agron. N. 38: 28, figs. 12i–o, 14. 1960. Tree 8–9 m tall; leaf blades 20–30 × 8–13 cm, lanceolate, very pubescent; secondary veins 10–13. Evergreen lowland forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Colombia, Ecuador, Peru, Brazil.

green lowland to montane and riparian forests, 100–1700 m; Bolívar (Gran Sabana, Macizo del Chimantá, Uaiparú), Amazonas (Río Casiquiare, Río Negro basin). Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 553.

Discophora guianensis Miers, Ann. Mag. Nat. Hist. ser. 2, 10: 119. 1852. Kummeria brasiliensis Mart., Fl. Bras. 12(2): 52. 1872. Discophora panamensis Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 222. 1929. Tree 5–20 m tall; leaf blade 10–30 × 4–13 cm, oblong, narrowly oblong, or ovate. Ever-

Fig. 553. Discophora guianensis

4. EMMOTUM Desv. ex Ham., Prodr. Pl. Ind. Occid. 29. 1825. Pogopetalum Benth., Trans. Linn. Soc. London 18: 680. 1841. Trees or shrubs, generally all parts pubescent, the pubescence gray to ferruginous or golden. Leaves coriaceous, bicolored; blades narrowly ovate, ovate, narrowly elliptic, elliptic, or nearly orbiculate, the apex acuminate, acute, or mucronate, base acute, obtuse, or attenuate; margins entire, slightly revolute; secondary veins 2–13 pairs, alternate, free near the margin, sometimes indistinct. Inflorescence axillary, paniculate; pedicels short, articulated at the end. Flowers actinomorphic, bisexual, 5-parted. Calyx campanulate, fleshy, lobed, pubescent without; petals free, pubes-

652

I CACINACEAE

cent without, rarely glabrous, densely lanate within, or 2 clusters of trichomes at the base and the apex (not in flora area). Stamens with fleshy filaments, flattened, glabrous; anthers attached basally or dorsally, elongate; connective broadened and cordate, sometimes extended (not in flora area). Disk represented by a differentiated ring of glabrous tissue at the base of the ovary, sometimes absent. Pistils with the ovary hirsute, or glabrous, 2- or 3-locular; style sometimes very short, sometimes eccentric, glabrous; stigma capitate, rarely 3-lobed. Fruits with heavy stony endocarp. Fertile seeds 1–3. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; ca. 10 species, 7 in Venezuela, all in the flora area. Key to the Species of Emmotum 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5. 6(4).

6.

Secondary veins poorly developed, often barely visible, with 2−4(–6) pairs of veins; mature leaves mostly glabrous ............................................... 2 Secondary veins well developed, with 5–13 pairs of veins; mature leaves pubescent or glabrous ............................................................................ 4 Leaves 2–4.5 × 2–3 cm, obovate, the apex acute or rounded, the base attenuate ............................................................................................ E. celiae Leaves 5–13 × 3–4.5 cm, ovate or elliptic, the apex short- or long-acuminate, the base rounded or attenuate ..................................................... 3 Peduncle of the inflorescence < 1 cm long ..................................... E. glabrum Peduncle of the inflorescence with a large peduncle ca. 3 cm long ............ ............................................................................................... E. yapacanum Ovary hirsute; leaves pubescent, ovate, narrowly ovate, oblong, narrowly oblong, or elliptic .................................................................................... 5 Ovary glabrous; leaves glabrous or pubescent, oblong, ovate, elliptic, or nearly orbiculate .................................................................................... 6 Lower surface of leaf blades slightly pubescent with very short trichomes ...............................................................................................E. conjunctum Lower surface of leaf blades very pubescent with long and slightly crisped trichomes ......................................................................... E. fulvum Leaves elliptic or nearly orbiculate; blades drying uniformly green-brown, lower surface pubescent, secondary veins 9–13 pairs, the apex rounded or mucronate ....................................................................... E. floribundum Leaves ovate or oblong; blades drying dark brown, chestnut, or tan below or the lower surface chestnut or tan, pale pubescent or glabrous, secondary veins 6–9 pairs, the apex acuminate ..................... E. acuminatum

Emmotum acuminatum (Benth.) Miers, Ann. Mag. Nat. Hist. ser. 2, 10: 178. 1852. —Pogopetalum acuminatum Benth., Trans. Linn. Soc. London 18: 685. 1841. Emmotum nudum R.A. Howard, J. Arnold Arbor. 23: 485. 1942. Tree 4–17 m tall; leaf blade 8–17 × 3.5–11 cm, ovate or oblong, the apex acuminate. Riparian forests, flooded black-water swamps,

100–200 m; Amazonas (Río Atabapo, Río Casiquiare, Río Negro, Río Sipapo, Santa Bárbara del Orinoco). Colombia, Brazil. Emmotum celiae R.A. Howard, Mem. New York Bot. Gard. 29: 63. 1978. Shrub to 5 m tall. Upland scrub, 500– 1300 m; Amazonas (Cerro Coro Coro, Cerro Parú, Cerro Yutajé). Endemic.

Emmotum 653

Fig. 554. Emmotum conjunctum

Fig. 555. Emmotum fulvum

654

I CACINACEAE

Emmotum conjunctum R.A. Howard, J. Arnold Arbor. 23: 491. 1942. —Uairatombepe. Shrub or tree 2–30 m tall; leaf blade 7–18 × 3–8 cm, ovate, narrowly ovate, oblong, narrowly oblong, elliptic, or narrowly elliptic, the apex acuminate or acute. Savannas, shrublands on white sand, riparian forests, upland scrub, 300–1100 m; Bolívar (widespread). Guyana. ŠFig. 554. Collections from Gran Sabana, including the type of Emmotum conjunctum, have previously been included under Emmotum fagifolium Ham., a species from lowland Guyana, Suriname, French Guiana, and Brazil, but E. conjunctum has more secondary veins. Emmotum floribundum R.A. Howard, J. Arnold Arbor. 23: 487. 1942. Shrub or tree 3–10 m tall; leaf blade 9–15 × 4.5–10 cm, elliptic or nearly orbicular, the apex mucronate. Riparian forests along black-water river and bana-like scrub, 100– 200 m; Amazonas (Caño San Miguel, Pimichín to Yavita, Río Atabapo, Río Baría, Río Guainía, Río Temi, San Carlos de Río Negro, Santa Bárbara del Orinoco, Santa Rosa de Ucata). Colombia, Peru, Brazil. Emmotum fulvum R.A. Howard, J. Arnold Arbor. 23: 493. 1942. —Aporu-ban-yek (Arekuna).

Tree 15–35 m tall; leaf blade 8–16 × 4–7 cm, narrowly ovate, the apex acuminate. Montane forests along sandstone escarpments, 800–1700 m; Bolívar (Caco-tepui west of San Ignacio, Cerro Venamo, Macizo del Chimantá, Ptari-tepui, Roraima-tepui). Endemic. ŠFig. 555. Emmotum glabrum Benth. ex Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 366. 1859. —Epoca-po-yek (Arekuna). Emmotum argenteum Gleason, Bull. Torrey Bot. Club. 58: 385. 1931. Emmotum ptarianum Steyerm., Fieldiana, Bot. 28: 341. 1952. Shrub or tree 2–20 m tall; leaf blades 5– 13 × 2.5–6 cm, ovate or elliptic, the apex short- or long-acuminate, pubescent when young, mostly glabrescent with age. Savannas, shrublands, forests, high-tepui scrub, 100–1800 m; widespread in Bolívar and Amazonas. Guyana, Peru, Brazil (Amazonas: Serra Aracá). Emmotum yapacanum R.A. Howard, Mem. New York Bot. Gard. 29: 66. 1978. Small tree to 3 m tall; leaf blades 6–9 × 4.5–7 cm, ovate or elliptic. Upland scrub, 800–1000 m; Amazonas (Cerro Yapacana). Endemic. Emmotum yapacanum is very similar to E. glabrum.

5. LERETIA Vell., Fl. Flumin. 99. 1825 [1829]. Lianas or shrubs with scandent branches, all parts covered with malpighiaceous trichomes. Leaves subcoriaceous, or coriaceous; blades narrowly ovate, narrowly oblong, or elliptic, the apex acute or acuminate, the base attenuate or cordate; secondary veins 6–8 pairs, alternate, joined together in a series of marginal loops. Inflorescence axillary, much-branched cymes, to 15 cm long; pedicels articulated near the end. Flowers actinomorpic, bisexual, 5-parted. Calyx lobed, pubescent without; petals free, pubescent without, densely lanate within. Stamens with filaments filiform, glabrous; anthers attached basally. Pistils with the ovary hirsute, 1locular; style curved, glabrous, usually with 2 aborted styles; stigma capitate. Fruit a drupe, ovoid-ellipsoid, slightly flattened; endocarp thin, sparsely long pubescent inside, exocarp glabrous. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 1 species. Leretia cordata Vell., Fl. Flumin. 99. 1825 [1829]. —Mappia cordata (Vell.) Engler in Mart., Fl. Bras. 12(2): 51. 1872. Leretia vellozii Miers, Ann. Mag. Nat. Hist. ser. 2, 9: 392. 1852.

Leretia nitida Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 365. 1859. —Mappia nitida (Miers) Engler in Mart., Fl. Bras. 12(2): 51. 1872. Mappia poeppigiana Baill., Adansonia 11:

Pleurisanthes 655

175. 1874. —Leretia poeppigiana (Baill.) Sleumer, Notizbl. Bot. Gart. BerlinDahlem 15: 245. 1940. Liana, often high-climbing; leaf blade 8– 20 × 4–10 cm, narrowly ovate, narrowly oblong, or elliptic. Evergreen lowland forests, 50–100 m; Delta Amacuro (Río Cuyubini). Other countries as in the genus. ŠFig. 556.

Fig. 556. Leretia cordata

6. PLEURISANTHES Baill., Adansonia 11: 201. 1874. Martia Valeton, Crit. Overz. Olacin. 259. 1886, non Spreng. 1818, nor others. Valentonia T. Durand, Index Gen. Phan. 64. 1888. Vines or shrubs with scandent branches, older stems bilaterally flattened, all parts pubescent with hispid trichomes. Leaves subcoriaceous or coriaceous; blades oblong, broadly oblong, elliptic, broadly elliptic, or nearly orbicular, the apex acute, obtuse, or emarginate, the base obtuse, rounded, or cordate; margins entire or dentate; secondary veins 7–9 pairs, alternate, joined together in a series of marginal loops, tertiary and quaternary venation conspicuous. Inflorescence axillary, supraaxillary, or terminal, spiciform racemes, rachis commonly flattened. Pedicels to 10 mm long, not articulated. Flowers actinomorphic, bisexual, 5-parted. Calyx lobed, pubescent without. Petals free, slightly fleshy, pubescent without. Stamens with filiform filament, glabrous; anthers oblong, basifixed. Disk absent. Pistils with the ovary hirsute, 1-locular; style minute or well developed; stigma capitate. Fruit oblong, compressed, persistently pubescent. Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 6 species, 2 in Venezuela, both in the flora area. Key to the Species of Pleurisanthes 1. 1.

Leaf blades 8–16 × 4–14 cm, broadly oblong, the apex rounded or emarginate, the base cordate, the margins dentate .................... P. emarginata Leaf blades 6–17 × 3–7 cm, oblong or elliptic, the apex mucronate or obtuse, the margins entire .................................................................. P. sp. A

Pleurisanthes emarginata Tiegh., Bull. Soc. Bot. France 44: 117. 1897. Vine or tree with decumbent branches.

Lateritic outcrops, ca. 500 m; Amazonas (Caño Colorado near Puerto Ayacucho). Suriname, French Guiana, Brazil.

656

I CACINACEAE

Pleurisanthes sp. A. —Cuyubi, Uña de murciélago. Liana. Riparian forests, 100–400 m; Amazonas (Río Cataniapo, Río Yureba, Sierra de la Neblina). Endemic. ŠFig. 557.

Fig. 557. Pleurisanthes sp. A

7. PORAQUEIBA Aubl., Hist. Pl. Guiane 123. 1775. Trees 5–25 m tall, all parts densely sericeous, becoming glabrous. Leaves coriaceous; blades elliptic, ovate, or obovate, the apex acute or acuminate, base attenuate or obtuse; margins entire, slightly revolute; secondary veins 4–9 pairs, alternate, free near the margin or slightly joined together in a series of marginal loops. Inflorescence axillary, paniculate, branched from the base; pedicels very short, articulated. Flowers actinomorphic, bisexual, 5-parted. Calyx campanulate, lobed, fleshy, pubescent or glabrous without. Petals free, very fleshy, pubescent or glabrous without, pubescent within, edges incurved, prominently longitudinally furrowed inside. Stamens with fleshy filaments, flattened, glabrous; anthers erect; connective broadened, extended to an inflexed apex. Disk absent. Pistil with the ovary glabrous, 1locular; style terminal; stigma capitate. Fruit ovoid to oblong, more or less oblique; mesocarp fleshy; endocarp smooth. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Poraqueiba 1.

1.

Leaf blades 10–18 × 6–10 cm, elliptic or obovate; corolla glabrous without; inflorescence barely exceeding the petiole; fruit with poor flavor ................................................................................................... P. paraensis Leaf blades 15–30 × 8–15, ovate or narrowly ovate; corolla sericeous without; inflorescence twice as long as petiole; fruit of good flavor, quite edible ................................................................................................. P. sericea

Poraqueiba paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 116. 1925. —Turi. Tree 10–15 m tall; leaf blade apex obtuse

or mucronate. Evergreen lowland forests, 100–300 m; Bolívar (Río Paragua), Amazonas (Río Casiquiare, Río Negro). Brazil. ŠFig. 558.

Poraqueiba 657

Fig. 558. Poraqueiba paraensis

Fig. 559. Poraqueiba sericea

658

I CACINACEAE

Poraqueiba sericea Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 172. 1849. —Capibare, Caniba, Madi, Yumari, Yurí. Poraqueiba acuminata Miers, Ann. Mag. Nat. Hist. ser. 3, 4: 365. 1859. Tree 10–30 m tall; leaf blade apex acute or acuminate. Evergreen lowland forests, often

cultivated or escaped near habitations, 100– 300 m; Amazonas (along Río Atabapo, Río Casiquiare, Rio Guainía). Colombia, Peru, Brazil. ŠFig. 559. Poraqueiba sericea is cultivated for its edible fruits, which are variable in size, color, and flavor.

IRIDACEAE by Peter Goldblatt and James E. Henrich Evergreen or seasonal geophytic perennials, sometimes annual (Sisyrinchium species), with rhizomes, bulbs or corms, or evergreen woody shrubs with secondary growth (Cape genera Klattia, Nivenia, and Witsenia). Leaves parallel-veined, mostly distichous, isobilateral and unifacial, usually equitant and ensiform, rarely terete (bifacial or square in some Old World genera), the blade flat or plicate, with open or closed sheathing base. Flowering stems aerial, often branched (or partly to entirely subterranean at anthesis in Crocus, Iris species, etc., from other areas), terete, angled, or winged; inflorescence either an umbel enclosed in a pair of large opposed spathes (rhipidium) and the flowers pedicellate and exserted serially, or a spike or panicle of sessile flowers (in Old World Ixioideae). Flowers typically large and showy, bisexual, actinomorphic or zygomorphic (many Ixioideae). Perianth of 6 petaloid tepals in 2 whorls, all alike, or the 2 whorls differentiated, occasionally the inner whorl lacking, free (most Iridoideae) or united into a tube; nectaries perigonal (many Iridoideae) on the inner tepals (most New World species) or the outer tepals (many Old World genera), nectaries septal in many Ixioideae. Stamens 3 (2 in the Australian Diplarrhena), opposite the outer tepals; filaments inserted on the tepals, either free and symmetrically disposed (or unilateral and arcuate in Ixioideae) or partly to entirely connate; anthers basifixed to sub-basifixed, 2-thecate, extrorse to latrorse, opening by longitudinal slits (by apical pores in Cobana and Sessilanthera). Gynoecium of 3 united carpels; ovary inferior (superior in Isophysis), 3-locular with axile placentation (unilocular with parietal placentation in Hermodactylus); style single below, 3-lobed to 3-branched, the branches sometimes again divided or expanded and variously petaloid with the stigma on the abaxial surface below the apex (many Iridoideae); ovules many to few (occasionally 1 or 2) in each locule. Fruit a loculicidal capsule, rarely indehiscent. Seeds often large, globose to angular (winged in some Old World genera); testa reticulate to smooth, sometimes fleshy or arillate. Cosmopolitan, but most abundant and diversified in South Africa; ca. 65 genera and ca. 1780 species, 4 genera and 9 species in the flora area.

Cipura 659

Key to the Genera of Iridaceae 1. 1. 2(1). 2. 3(1). 3.

Rootstock a rhizome, a short erect crown with fibrous roots; leaves flat or subterete ................................................................................................ 2 Rootstock a bulb; leaves pleated in folds (plicate), appearing flat in some dried specimens ..................................................................................... 3 Tepals subequal and not differentiated, without a zone of nectaries ............................................................................................ 3. Sisyrinchium Tepals unequal and the inner strongly geniculate and with a zone of nectaries on the claw or base of the limb ....................................... 4. Trimezia Flowering stem with a large leaf subtending the inflorescence spathes; spathes nearly sessile, crowded at the stem apex ....................... 1. Cipura Flowering stem without a leaf subtending the inflorescence spathes; spathes stalked on an open branched stem ................................ 2. Cypella

1. CIPURA Aubl., Hist. Pl. Guiane 38, t. 13. 1775. Perennials with bulbs, tunics brown to blackish, papery or glutinous. Leaves plicate, lanceolate to linear, 2–4 basal and 1 subterminal on the flowering stem. Flowering stems scapose, with a conspicuous subterminal cauline leaf extending above the 1–several apical rhipidia; spathes unequal, the inner about twice as long as the outer. Pedicel exserted or included in the spathes. Flowers actinomorphic, yellow, white, or blue-purple with yellow nectar guides. Tepals basally connate for 1–2 mm, unequal, the outer larger, spreading, the inner erect to ascending, sometimes imbricate and partly concealing the stamens and style, with or without nectaries. Staminal filaments free, weak and slender, basally thickened and ± contiguous; anthers adhering to the style, dehiscence extrorse to latrorse. Style thickened above, the branches prominent and terminally stigmatic, or lacking and then the stigmatic lobes sessile on the style. Capsule obovoid to ellipsoid, truncate, included or exserted. Seeds angular. Neotropics; ca. 6 species, 3 in Venezuela, all in the flora area. Key to the Species of Cipura 1. 1.

2(1). 2.

Flowers yellow; capsules and often the ovary exserted; inner tepals with a narrow claw ................................................................................ C. rupicola Flowers blue to purple or white, often with a yellow band of nectaries on the inner tepals; ovary and capsules included; inner tepals without a narrow claw ............................................................................................ 2 Bulbs 1.5–2 cm in diameter; tunics papery .................................. C. paludosa Bulbs ca. 3 cm in diameter; tunics glutinous ...................................... C. sp. A

Cipura paludosa Aubl., Hist. Pl. Guiane 38, t. 13. 1775. —Cebolleta. Plant 16–27 cm tall; flowers pale bluish or white. Savannas in wet or marshy places, often seasonally flooded, 50–400 m; Bolívar (southwest of Caicara, northwest of

El Manteco, Maripa-Aripao, Río Paragua), Amazonas (vicinity of Puerto Ayacucho). Widespread elsewhere in Venezuela; Neotropics. ŠFig. 560. Cipura rupicola Goldblatt & Henrich, Ann.

660

I RIDACEAE

Fig. 560. Cipura paludosa

Fig. 561. Cypella linearis

Sisyrinchium 661

Missouri Bot. Gard. 74: 337. 1987. Plant 12–40 cm tall; flowers yellow. Dry stony savannas, granitic hills, outcrops in shallow soil, ca. 100 m; Amazonas (near Puerto Ayacucho). Adjacent Colombia.

Cipura sp. A Plant 40–50 cm tall; flowers purple. Open savannas, ca. 100 m; Amazonas (near Rincones de Chacorro 30 km north of Puerto Ayacucho). Adjacent Colombia.

2. CYPELLA Herb., Bot. Mag. t. 2637. 1826. Seasonal perennials with bulbs, tunics usually brown to blackish, papery. Leaves few, plicate, lanceolate to linear. Flowering stems branched. Flowers usually shades of yellow or blue to purple, patterned in contrasting colors. Tepals free, unequal, the outer larger, broadly clawed and ± widely cupped below, the inner bearing nectaries in the lower half, partly enclosed by folds. Staminal filaments free, weak and slender, thickened below, or sturdy; anthers attached at least apically to the opposed style branches. Style branches well developed, often compressed, usually divided above into prominent acute crests and with a transverse stigmatic surface on the abaxial surface at the base of the crests, the stigma also occasionally with short paired crests in the center. Capsules obovoid to cylindric, truncate. Seeds angular. Locally in Mexico and Cuba, South America; ca. 20 species, 1 in Venezuela. Cypella linearis (H.B.K.) Baker, Handbk. Irid. 65. 1892. —Moraea linearis H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 321. 1815 [1816]. —Larentia linearis (H.B.K.) Klatt, Abh. Naturf. Ges. Halle 12: 362. 1882. —Mentolito. Plant 12–22 cm tall, the stems branched; flowers blue to purplish. Open savannas, grasslands, lajas, Mauritia palm savannas,

50–400 m; Bolívar (Agua Amena along Caicara to Puerto Ayacucho road, El Tigre on lower Río Cuchivero, base of Los Pijguaos, near Maripa and Aripao, Río Paragua), Amazonas (Puerto Ayacucho). Anzoátegui, Aragua, Barinas, Carabobo, Cojedas, Distrito Federal, Guárico, Miranda, Portuguesa; Brazil, Bolivia, Paraguay. ŠFig. 561.

3. SISYRINCHIUM L., Sp. Pl. 954. 1753. Seasonal perennials or annuals with a persistent rhizome or a short erect crown. Roots fibrous or thickened, sometimes fleshy. Leaves flat or terete, lanceolate to linear. Flowering stems simple or variously branched, compressed and sometimes winged, comprising 1 or several internodes. Flowers actinomorphic, usually yellow (in the flora area) or blue to purple with a yellow center, occasionally whitish, or pink. Tepals subequal, diverging from the base or cupped and then often including the stamens and style. Staminal filaments free or partly to entirely united; anthers spreading to coherent. Style either long and dividing into 3 short branches, or short with 3 long slender branches, these extending between the stamens. Capsule globose to cylindric, glabrous or sparsely to densely papillose. Seeds globose, usually blackish. Widespread in North America, Greenland, Central America, South America; ca. 100 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Sisyrinchium 1. 1.

Flowering stems consisting of a single long leafless internode; capsules ± pear-shaped .......................................................................... S. tinctorium Flowering stems bearing leaves, sometimes branched; capsules globose ................................................................................................ S. vaginatum

662

I RIDACEAE

Sisyrinchium tinctorium H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 324. 1815 [1816]. Plant 15–30 cm tall; flowers yellow, stellate; stems broadly winged, unbranched, and leafless; leaves basal only, soft-textured; capsules pendent. Generally in wet sites, ca. 100 m; Amazonas (La Esmeralda). Uncommon elsewhere in Venezuela; widespread and common in Central America, Colombia, Ecuador, Peru. This species is common outside of the flora area along streams, seeps, and wet savannas, usually above 1000 m. It has been collected only once in the flora area, and its occurrence needs confirmation. Sisyrinchium vaginatum Spreng., Syst. Veg. 1: 166. 1825 [1824]. —Kurariyusu. Souza marchio Vell., Fl. Flumin. 258. 1825 [1829]; Fl. Flumin. Icon. 7: t. 1. 1827 [1831]. —Sisyrinchium marchio (Vell.) Steud., Nomencl. Bot. ed. 2, 2: 596. 1841 [1840]. Sisyrinchium alatum Hook., Icon. Pl. 1: t. 219. 1837. Plant 15–45 cm tall; leaves evenly placed along the stems; flowers yellow, stellate. Open savannas, rocky sites, ca. 50–1400 m; Bolívar (middle Río Paragua), Amazonas (Caño Iguana). Widespread elsewhere in Venezuela; widespread in tropical South America. ŠFig. 562.

Fig. 562. Sisyrinchium vaginatum

4. TRIMEZIA Salisb. ex Herb., Edward’s Bot. Reg. 30: misc. 88. 1844. Evergreen perennials with erect rhizomes sheathed by brownish, fibrous to sometimes fleshy (or glutinous in some South American species) leaf bases. Leaves flat, lanceolate to linear, basal and cauline. Flowering stem ± terete, usually branched. Flowers actinomorphic, yellowish, usually with contrasting dark brownish markings on the claws. Tepals free, broadly clawed, the outer larger, the inner with nectaries in the lower half, partly concealed by a geniculate fold. Staminal filaments free, slender and weak, thickened basally; anthers affixed to the style branches. Style slender below, thickened above, dividing toward the anther apex into 3 branches, each either bearing paired acute crests with a transverse stigma, or bilobed to truncate and apically stigmatic. Capsules globose to cylindric, truncate. Seeds rounded to angular. Mexico, Central America, West Indies, south to Paraguay and Argentina; ca. 15 species, 3 in Venezuela, all in the flora area.

Trimezia 663

Fig. 564. Trimezia chimantensis Fig. 563. Trimezia fosteriana

664

I RIDACEAE

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

Stem bearing a single leaf in the middle part and sometimes branched; leaves soft-textured and with a distinct midrib ................ T. martinicensis Stem bearing 2 or more short, mostly sheathing leaves and never branched; leaves coriaceous and without an apparent midrib ............ 2 Leaves rich green, 2–3 mm wide; stems 80–120 cm tall....... T. chimantensis Leaves glaucous, 4–8 mm wide; stems (15–)25–125 cm tall ....... T. fosteriana

Trimezia chimantensis Steyerm., Ann. Missouri Bot. Gard. 71: 301. 1984. Plant 80–120 cm tall; flowers yellow, without brown spots; stems unbranched; leaves rich green, without midribs, 2–3 mm wide. Open rocky tepui summits, ca. 2000 m; Bolívar (Macizo del Chimantá ). Endemic. ŠFig. 564. Trimezia fosteriana Steyerm., Fieldiana, Bot. 28: 160. 1951. Plant 15–125 cm tall; flowers yellow with brown spots in the center; stems unbranched; leaves glaucous, without midribs, 4–8 mm wide. Wet upland savannas on white

sand, 1300–2100 m; Bolívar (Gran Sabana and associated tepuis). Endemic. ŠFig. 563. Trimezia martinicensis (Jacq.) Herb., Edward’s Bot. Reg. 30: misc. 88. 1844. —Iris martinicencis Jacq., Enum. Syst. Pl. 12. 1760. Plant 30–60 cm tall; flowers yellow with brown markings; stems sometimes branched; leaves soft-textured, with a conspicuous midrib. Savannas, Mauritia palm swamps, along streams, 200–800 m; Bolívar (scattered). Widespread elsewhere in Venezuela; widespread in West Indies and northeastern South America.

IXONANTHACEAE by Nelson Ramírez and Paul E. Berry Trees or shrubs. Leaves alternate, simple; margins entire or toothed; stipules small or lacking. Inflorescences axillary or terminal cymes, thyrses, or racemes. Flowers small, generally bisexual, actinomorphic or nearly so, usually 5-merous. Sepals imbricate, distinct or connate only at the base; petals distinct, imbricate or convolute. Stamens 5–20; filaments folded in bud, expanded at the base but not connate, free or basally adnate to the well-developed, annular to cupular, intrastaminal nectary disk; anthers short, 4-sporangiate and 2-thecal, longitudinally dehiscent. Ovary superior, of (2–)5 carpels united to form a compound, plurilocular ovary with axile-apical placentation and a terminal style, the ovary sometimes divided into locelli or unilocular at the very top with partitions not reaching the summit; ovules 1 or 2 per locule, pendulous, the micropyle directed upward and outward; style terminal, folded in bud; stigma ± capitate. Fruit a septicidal or sometimes loculicidal capsule, with or without a persistent central column. Seeds arillate or winged, with little or no endosperm. Pantropics; 5 genera and ca. 30 species, 2 genera and 8 species in the flora area. Key to the Genera of Ixonanthaceae 1.

Ovary 2-locular; floral pedicel articulated; fruit an elongate, slightly asymmetrical, 2-valved capsule; seeds partly covered at the upper end by a 2-lobed membranaceous aril ...............................................Cyrillopsis

Cyrillopsis

1.

665

Ovary 5-locular; floral pedicel not articulated; fruit a prismatically cylindric, elongate, 5-valved capsule; seeds with a conspicuous, thin, lateral wing ....................................................................................... Ochthocosmus

1. CYRILLOPSIS Kuhlm., Arch. Jard. Bot. Rio de Janeiro 4: 356. 1925. Shrubs or trees to 15 m tall. Stems glabrous, young ones densely lenticellate. Leaves subcoriaceous, 3–20 × 1.5–7 cm, ovate-lanceolate, abruptly narrowed and shortly acuminate at the apex, long attenuate at the base, congested toward the stem apices, shiny on upper surface, opaque on lower surface; margin entire, lateral veins 5–10 per side and prominent on lower surface; petioles 1–15 mm long; stipules minute, lateral and fugaceous. Inflorescence a fascicle of 1–6 lax, delicate racemes in upper leaf axils, each raceme 1.5–5 cm long, with small, triangular bracts; pedicels filiform, articulated near the base, with fugaceous bracteoles 2–4 mm long. Calyx lobes imbricate, broadly ovate, ca. 1 × 0.75 mm; petals imbricate, free, persistent in fruit, ca. 3 times longer than the sepals, broadly obovate, briefly retuse at the apex. Stamens 5, free, alternate with the petals; filaments persistent, united to a shallow annular disk; anthers basifixed and introrsely dehiscent. Ovary 2-locular; ovules 2 per locule; style filiform, elongating in fruit. Fruit an elongate, loculicidal, 2-valved capsule, slightly flattened or asymmetrical. Seeds partly covered at the upper end by a membranaceous aril. Endemic to the Guayana Shield in Venezuela, Guyana, French Guiana, and northern Amazonian Brazil; 2 species, both in the flora area.

Seed Fig. 565. Cyrillopsis micrantha

Fruit

666

I XONANTHACEAE

Fig. 566. Cyrillopsis paraensis

Ochthocosmus 667

Key to the Species of Cyrillopsis 1. 1.

Leaves 3–7 cm long, the apex retuse to rounded ...................... C. micrantha Leaves 10–20 cm long, the apex shortly attenuate ..................... C. paraensis

Cyrillopsis micrantha (Steyerm.) P.E. Berry & N. Ramírez, Novon 5: 227. 1995. —Ochthocosmus micranthus Steyerm., Ann. Missouri Bot. Gard. 75: 319, fig. 5. 1988. —Wontai (Taurepán). Shrub or small tree 1–6 m tall; fruits to 15 mm long and 5 mm thick, brilliant olive or dark green before dehiscing. Locally common in shrublands, 1000–1150 m; Bolívar (Ayavaparú north of confluence of Río Karaurín and Río Yuruaní and 10–15 km southwest of

Wadakapiapué-tepui). Endemic. ŠFig. 565. This species is known only from the type locality, where it is closely associated with populations of Ochthocosmus roraimae. Cyrillopsis paraensis Kuhlm., Arch. Jard. Bot. Rio de Janeiro 4: 357. 1925. Tree to 15 m tall; flowers greenish white. Evergreen lowland forests, 100–200 m; Amazonas (Piedra Arauicaua). Guyana, French Guiana, Brazil (Amazonas, Pará). ŠFig. 566.

2. OCHTHOCOSMUS Benth., London J. Bot. 2: 366. 1843. Small glabrous shrubs or trees to 30 m tall; trunk 10 cm or more in diameter, sparsely to much branched. Leaves subcoriaceous to coriaceous, variously shaped, apex obtuse or broadly rounded and emarginate or retuse, base rounded, obtuse, or acute; margin entire or remotely to conspicuously crenate or crenulate, the sinus of each crenation with a caducous glandular tooth; pinnately veined, lateral veins 4–11 per side. Inflorescence usually in axils of distal leaves or subterminal, of few- to many-flowered, solitary or fasciculate, pedunculate, simple racemes or panicles. Flowers fragrant; pedicels 1–15 mm long, with bracts ca. 0.4 mm long at the base. Sepals 5, slightly connate at the base, reflexed after anthesis, rounded at the apex, entire to glandular-crenulate, green to rose-red; petals 5, contorted, persistent, ovate-oblong, rounded at apex, white to pink. Stamens 5, alternate with the petals; filaments dilated at the base where united with the shallow annular disk within the base of the petals; anthers introrsely dehiscent. Ovary ovoid, 5-ridged and 5-locular; ovules 2 per locule; style simple, filiform, equal to or to 3 times longer than the ovary. Fruit a prismatically cylindric, elongate capsule, pointed at apex, 5-valved. Seeds obliquely oblong and curved at the basal end, with a conspicuous, pale, thin wing attached at the distal end. Southeastern Colombia (Guainía), southern Venezuela, Amazonian Brazil, northeastern Bolivia; 7 species, 6 in Venezuela, all in the flora area. Ochthocosmus parvifolius Hallier f. is treated here as a species of uncertain position (incertae sedis) and is not included in the treatment; see P.E. Berry & N. Ramírez (Novon 5: 227–229. 1995) for further discussion. The seventh species, O. barrae Hallier f., is known from Brazil and Bolivia. Key to the Species of Ochthocosmus 1. 1.

Leaves entire, or with 1–3 barely discernible sinuses along each margin, < 7 cm long ............................................................................................. 2 Leaves slightly to markedly crenate and often with small glandular teeth in the sinuses, or if appearing entire, then with 4 or more slight sinuses along margins or else the blade > 7 cm long ......................................... 3

668

2(1).

2.

3(1). 3.

4(2).

4.

5(4).

5.

I XONANTHACEAE

Inflorescences shorter or longer than the leaves, 2–8 cm long; pedicels 1– 5(–10) mm long at anthesis; leaves firm- to stiff-coriaceous, 1–7 cm long, the margin not revolute ............................................... O. multiflorus Inflorescences much longer than the leaves, 5–12 cm long; pedicels 5– 15 mm long at anthesis; leaves stiff-coriaceous, 1–4 cm long, the margin slightly revolute .............................................. O. longipedicellatus Leaf blades elliptic, more than twice as long as wide, 3–7 × 1.5–3 cm ................................................................................................ O. attenuatus Leaf blades generally broadest above the middle (usually obovate), either less than twice as long as wide or else longer or wider than above, with the apex emarginate or at least more rounded than the base ............. 4 Inflorescences shorter than the nearest leaves; leaves obovate, cuneate to attenuate at the base; 4 sepals ± equal, rounded, with a glandular margin, the fifth sepal longer, ligulate-oblong, and eglandular ......... O. berryi Inflorescences usually longer than the nearest leaves; leaves suborbicular to broadly oblong, elliptic, or obovate, the base obtuse to rounded, acute, or cuneate; sepals either all ± equal, rounded, and with a glandular margin, or else strongly unequal, the outer three rounded and with a glandular margin, the inner two nearly twice as long and eglandular ................................................................................................................ 5 Leaf blades smaller than 6 × 3 cm, the petioles 1–3.5 mm long; sepals strongly unequal, the inner 2 nearly twice as long as the outer 3, outer sepal margins glandular-crenate, the inner ones eglandular ................ .............................................................................................. O. floribundus Leaf blades either > 6 cm long or > 3 cm wide, the petioles 1–10 mm long; sepals equal or subequal in length, margins all glandular-crenate .................................................................................................. O. roraimae

Ochthocosmus attenuatus Steyerm. & Luteyn, Brittonia 32: 142. 1980. Shrub to 3 m tall; leaves erect. Sparse shrublands, 1000–1100 m; Bolívar (northern Gran Sabana near San Rafael de Camoirán around km 175 of main road south of El Dorado and near bridge crossing Río Maremán). Endemic. ŠFig. 569. Ochthocosmus berryi Steyerm., Ann. Missouri Bot. Gard. 71: 313. 1984. Tree to 8 m tall; the fifth sepal eglandular, contrasting to the other much shorter ones. Montane forests, 900–1100 m; Amazonas (Cerro Aratitiyope, Cerro Parú, Sierra Parima 5–8 km north of Simarawochi). Adjacent Brazil (Roraima: Auaris). ŠFig. 571. Florally, this species is similar to Octhocosmus floribundus, but appears to have larger, less coriaceous leaves; further study may show that the two species should be combined.

Ochthocosmus floribundus Gleason, Brittonia 3: 166. 1939. Shrub or tree 2–15 m tall. Montane forests, shrublands, and savannas on sandy and rocky soils, (300–)700–1400 m; Bolívar (lower slopes of Auyán-tepui above Guayaraca, Cerro Guaiquinima, Cerro Venado, base of Cerro Zumbador, Río Aza basin ca. 25 km west southwest of Chiguao). Endemic. ŠFig. 567. Ochthocosmus longipedicellatus Steyerm. & Luteyn, Brittonia 32: 141. 1980. Slender shrub 1–5 m tall, with long, virgate stems; leaves erect, stiffly coriaceous. Scrub on rocks or sand, 1000–1400 m; Bolívar (kms 147 and 175 south of El Dorado, 20–25 km northwest of Ilú-tepui). Guyana (upper Mazaruni River basin). ŠFig. 570. Ochthocosmus multiflorus Ducke, Trop. Woods 50: 33. 1937.

Ochthocosmus 669

Subshrub 20–50 cm tall to shrub or tree 0.7–5(–8) m tall; branches wiry, sparsely leafy. Savannas on white sand, 100–200 m; southwestern Amazonas. Southeastern Colombia (Guainía), northwestern Brazil (Amazonas); 3 varieties, all in the flora area. Key to the Varieties of O. multiflorus 1. Leaf blades nearly equally wide throughout their length, usually linear to linearoblong, 5–7 times longer than broad, 15– 34 × 2–5 mm .......... var. angustifolius 1. Leaf blades not equally wide throughout their length, usually narrower at one end or another, oblong-oblanceolate, linear-oblanceolate, or linear-oblong, 1.5–5 times longer than broad, 12–80 × 5–30 mm ........................................................ 2 2. Leaves mainly 1.5–3 times longer than broad, 10–60 × (15–)20–30 mm; secondary or tertiary venation usually ± evident on upper surface .............................. ................................... var. multiflorus 2. Leaves mainly 2–5 times longer than broad, 10–40 × 5–12 mm; secondary and tertiary venation not visible on upper surface ................... var. canaripoensis O. multiflorus var. angustifolius Steyerm. & Luteyn, Brittonia 32: 141. 1980. 100–200 m; Amazonas (Canaripó, around Cerro Yapacana). Endemic. O.

multiflorus var. canaripoensis Steyerm. & Luteyn, Brittonia 32: 141. 1980. 100–200 m; Amazonas (Canaripó, lower Río Ventuari). Endemic. O. multiflorus var. multiflorus 100–200 m; western Amazonas. Colombia (Guainía), Brazil (Amazonas). ŠFig. 568. Ochthocosmus roraimae Benth., London J. Bot. 2: 366. 1843. Shrub or tree 2–20(–30) m tall. Riparian forests, cloud forests, shrublands on sandy and rocky soil, savannas, 800–1300 m. Venezuela, Guyana; 2 varieties, both in the flora area.

Key to the Varieties of O. roraimae 1. Leaves ovate, 7–12 × 4–7 cm, the margins entire or nearly so; panicles large, 11–14 cm long, 10–12 cm wide .......................... ................................. var. grandifolius 1. Leaves generally more obovate and smaller than above, usually with noticeably crenate or toothed margins; panicles smaller than above ...... var. roraimae O. roraimae var. grandifolius (Steyerm.) Steyerm. & Luteyn, Brittonia 32: 135. 1980. —Ochthocosmus grandifolius Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 421. 1966. Tree ca. 8 m tall; panicles 11–14 × 12– 14 cm. Humid upland forests, 900–1200 m; Bolívar (Cerro Venamo). Endemic. ŠFig. 572. This variety will most likely also be found in adjacent Guyana. O. roraimae var. roraimae Small to large tree 2–20 m tall. Riparian forests, shrublands, scrubby savannas, 800– 1300 m; Bolívar (south and east from Kavanayén through much of the Gran Sabana to El Paují and the lower slopes of Roraimatepui). Guyana. This variety is quite variable in leaf size and texture throughout its range. Most material of this variety was formerly treated as Ochthocosmus roraimae var. parvifolius, but these specimens have all been placed now under var. roraimae (see P. E. Berry & N. Ramírez, Novon 5: 228– 229. 1995). Some specimens from the southern Gran Sabana have flowers similar to O. floribundus but leaves consistent with O. roraimae. These include Huber 11955 (MO, MYF, VEN) and Huber & Fernández 11944 (MO, MYF, US, VEN), both from an unusual shrubland southwest of Wadakapiapué-tepui near the junction of the Río Karaurín and Río Yuruaní that is also the type locality of Cyrillopsis micrantha. Another specimen with floribundus-like flowers is Huber & Alarcón 9636 (MO, MYF), from the vicinity of Perai-tepui, 55 km west of Santa Elena de Uairén.

670

I XONANTHACEAE

Fig. 567. Ochthocosmus floribundus

Fig. 569. Ochthocosmus attenuatus

Fig. 568. Ochthocosmus multiflorus var. multiflorus

Fig. 570. Ochthocosmus longipedicellatus

Ochthocosmus 671

Fig. 571. Ochthocosmus berryi

Fig. 572. Ochthocosmus roraimae var. grandifolius

672

J UNCACEAE

JUNCACEAE by Carlos A. Vargas Perennial, usually rhizomatous herbs or rarely annuals, sometimes forming cushions in high-Andean bogs. Rhizome sympodial, creeping, sometimes branching-ascending or stoloniform, sometimes rich in starch. Culm (stem) generally erect, sometimes procumbent, terete or hollow, rarely flattened, smooth or longitudinally ridged, frequently strongly aerenchymatous, unbranched or sparingly branched, leafy at the base or leafless. Leaves usually spirally arranged or rarely distichous and densely arranged in cushion-forming genera, linear, subulate, filiform, or rarely lanceolate, flat, grooved, angular, terete or laterally compressed in cross section, parallel-veined, sometimes completely reduced (the culms then comprising the main photosynthetic tissue), with an open and sometimes auriculate or closed sheath, air canals common, sometimes partitioned by distinct septa. Inflorescence terminal, sometimes pseudolateral, compound, cymose or racemose, sometimes corymbose or anthelate, usually in heads or spike-like clusters, rarely reduced to a single terminal or lateral flower. Flowers sessile or pedicellate, 3-merous, generally small, actinomorphic, bisexual or unisexual. Tepals 6, in 2 whorls of 3, glumaceous, green, strawcolored, castaneous, or almost black, equal or almost equal, free. Stamens 6, in 2 whorls of 3, opposite the perianth segments, inner whorl sometimes reduced; filaments filiform or somewhat flattened and widened at the base; anthers 4-locular, sometimes much longer than their filaments, oblong to linear, basifixed, obtuse or mucronate, dehiscing by 2 longitudinal lateral slits, at anthesis twisting around their own axis to the right, in some genera the connectives prolonged into mucronate apical appendages; pollen in tetrahedral tetrads, all grains functional. Ovary superior, syncarpous, 1- or 3-locular; style 1, to 10 mm long; stigmas 3, terete or sometimes tapering distally, twining with adaxial papillae; placentation basal or often parietal or axile. Fruit a round or trigonous, 3-lobed loculicidal or circumscissile capsule. Seeds usually many per capsule, sometimes 3; outer seed coat hyaline, whitish or light brown, sometimes with tail-like appendages, often with distinct sculpturing; inner seed coat brown to castaneous or yellow. Cosmopolitan except Antarctica, 6 genera and ca. 300 species, 1 species in the flora area. Two genera (not in the flora area) normally included in Juncaceae are here excluded from the family. Prionium is now considered a monotypic family, Prioniaceae, and Oxychloe is considered a member of the Cyperaceae (Munro and Linder, Syst. Bot. 23(1): 43–55. 1998). 1. JUNCUS L., Sp. Pl. 325. 1753. Cephaloxys Desv., J. Bot. (Desvaux) 1: 321. 1809. Juncastrum Fourr., Ann. Soc. Linn. Lyon, ser. 2, 17: 172. 1869. Annual or perennial, rhizomatous, glabrous herbs. Culms usually erect, rarely procumbent or ascending. Leaves alternate, scale-like on the rhizomes, cataphyllous (with reduced blades), and/or foliar on the base of the culm, and bract-like in the inflorescence; sheaths open, usually conspicuously auriculate at the junction with the blade; blade glabrous, linear, in cross section flat with raised margins and slightly channeled above, grooved, round to elliptic, or completely flat. Inflorescence compound or decompound, cymose or racemose, often anthelate; bracts decreasing in size from the base upwards, the lower bract often conspicuously different from the

Juncus 673

Fig. 573. Juncus densiflorus

674

J UNCACEAE

remaining ones. Flowers bisexual, sometimes clasped by two bracteoles on the pedicel. Tepals equal or subequal, lanceolate, entire, persistent. Stamens 3 or 6; filaments filiform or flat, sometimes widened at the base; anthers obtuse, linear or oblong. Ovary sessile; stigmas 3, filiform, twining, papillose. Capsule 1- or 3-locular. Seeds many, ellipsoid, oblong or ovoid, smooth, rugose, or sometimes reticulate. Cosmopolitan except Antarctica; ca. 220 species, 11 in Venezuela, 1 of these in the flora area. Juncus densiflorus H.B.K., Nov. Gen. Sp. 1: 238 (quarto ed.) 1815 [1816]. Juncus cyperinus Wild. ex. Spreng., Syst. Veg. 2: 106. 1825, nom. in syn. —Juncus densiflorus var. cyperinus (Willd.) Buchenau, Abh. Naturwiss. Vereine Bremen 6: 397. 1879. Herb to 1 m tall. Moist tepuis meadows and white-sand savannas, in humid places, (100–) 1200–2200 m; Bolívar (Gran Sabana, Macizo del Chimantá [Apacará-tepui]), Amazonas

(Guarinuma). Lara, Mérida; Colombia, Brazil, Paraguay, Argentina, Uruguay. ŠFig. 573. Juncus densiflorus is widely distributed in the Andean highlands of Venezuela and Colombia (1500–2700 m). The label of the type collection (Humboldt & Bonpland s.n., P) states the locality as Guarinuma, which is a lowland site (100 m) along the Río Atabapo in Amazonas state. No other collections from this are known, so there is some doubt if the label information is correct.

KRAMERIACEAE by Lois Brako Perennial shrubs, subshrubs, or herbs; probably all hemiparasitic. Woody stems terete with brown to black, striate or fissured bark, young herbaceous stems terete and green, variously covered with trichomes. Leaves alternate, simple and entire, linear to ovate, acute or mucronate apically, sessile with minute petioles or with petioles the same length as the leaves, one species (Krameria cytisoides Cav.) with trifoliolate leaves. Flowers zygomorphic, solitary and axillary, in terminal racemes or open panicles, basically pentamerous. Rachis and flowering stalk variably pubescent, peduncles and pedicles separated from each other near the base by a pair of opposite linear or lanceolate, leaf-like branchlets. Sepals petaloid, unequal, the outer 3 commonly larger than the inner 2, the lowermost sepal geniculate or saccate in some species, pubescent; petals 4 or 5, small, dimorphic, the two lower ones orbicular to cuneate, fleshy, ca. 2–3 mm in diameter, appressed laterally to the ovary or flaring outward from the base, upper petals variously clawed or lanceolate, clustered at the base of the upper side of the ovary forming a “flag,” either free or fused. Stamens 3 or 4, inserted at the juncture of the upper petals and the ovary or on the connate portion of the upper petals, curved with stout filaments; anthers tubular, about the same diameter as the filaments, dehiscing by membranous pores. Ovary superior, ovoid, densely pubescent, 2-carpellate, but appearing unicarpellate due to the ontogenetic suppression of one carpel, the remaining carpel unilocular, with 2 anatropous ovules emerging from the adaxial side, only one maturing into a seed; styles curved upward and terminating in a minute stigma with two barely distinguishable lobes. Fruit globose or a slightly flattened, spiny dry structure, often designated as a nut or a single-sided capsule. Seeds one per fruit (but many fruits empty), globose with a gray-brown seed coat, lacking endosperm; cotyledon orbicular, ventrally flattened.

Krameria 675

Southern U.S.A., Mexico, Guatemala, Honduras, Costa Rica, West Indies, all South American countries except Uruguay, mainly in dry regions; 1 genus and 17 species, 2 species in the flora area. This treatment is largely extracted from B. B. Simpson (Krameriaceae. In Flora Neotropica Monograph 49: 108 pp. 1989). 1. KRAMERIA Loefl., Iter Hispan. 195. 1758. Characters and distribution as in family; 17 species, 2 in Venezuela, both in the flora area. Key to the Species of Krameria 1. 1.

Plants upright shrubs, 0.3–1.5 m tall; leaves lanceolate, petioles > 1.5 mm long ................................................................................................... K. ixine Plants sprawling procumbent subshrubs; leaves linear-lanceolate, petioles < 1.0 mm long ................................................................. K. spartioides

Fig. 574. Krameria ixine

676

K RAMERIACEAE

Krameria ixine Loefl., Iter Hispan. 195. 1758. Krameria ixina L., Sp. Pl. ed. 2, 1: 177. 1762. Krameria linifolia Willd. ex Schult. & Schult. f., Mantissa 3: 303. 1827. Krameria arida O. Berg, Bot. Zeitung (Berlin) 14: 762. 1856. Krameria lanceolata O. Berg, Bot. Zeitung (Berlin) 14: 765. 1856, nom. illeg. Upright shrub 0.3–1.5 m tall; flowers in lateral and terminal racemes, pink. Shrub savannas, xerophytic scrub, 50–400 m; Bolívar (near El Miamo, Morichal Santa Isabel, between San Félix and Ciudad Bolívar, vicinity of Upata). Anzoátegui, Barinas,

Distrito Federal, Falcón, Guárico, Lara, Nueva Esparta, Sucre; Mexico, Central America, West Indies, Colombia, Guyana. ŠFig. 574. Krameria spartioides Klotzsch ex O. Berg, Bot. Zeitung (Berlin) 14: 761. 1856. Krameria evolvuloides Triana & Planch. ex Bennett in Mart., Fl. Bras. 13(3): 72. 1874, as synonym. Sprawling, procumbent subshrub to 0.6 m tall, reaching 1 m in diameter; flowers in terminal racemes, pink-purple. Lowland and upland xeric savannas, 50–900 m; Bolívar (Caicara, vicinity of Roraima-tepui). Anzoátegui; Guyana, Colombia, Brazil (Roraima).

LACISTEMATACEAE by Luther J. Raechal Shrubs or small trees. Leaves alternate, simple, stipulate, entire (Lacistema) or serrate (Lozania). Inflorescences axillary, slender spike-like racemes or catkin-like spikes, the flowers very small, each flower subtended by a basal scale-like or large bract. Flowers bisexual or with bisexual and staminate ones in the same inflorescence (plants hermaphroditic or andromonoecious). Sepals (0)2–6; petals absent. Disk fleshy to membranaceous, annular or semilunate, concave to cup-shaped, sometimes lobed. Stamen 1, attached to the interior of the disk. Ovary superior, 2- or 3-carpellate, unilocular with parietal placentas; style very short to lacking, or elongate; stigmas 3. Fruit capsular. Endosperm present. Neotropics; 2 genera and 14 species, 1 species in the flora area. 1. LACISTEMA Sw., Prodr. 12. 1788. Description as for the family, except leaves mostly entire, bracts conspicuous, and inflorescence spike-like. Mexico, Central America, Jamaica, tropical and subtropical South America; 11 species, 1 in Venezuela. Lacistema aggregatum (Bergius) Rusby, Bull. New York Bot. Gard. 4: 447. 1907. —Piper aggregatum Bergius, Acta Helv. Phys.-Math. 7: 131, t. 10. 1772. Lacistema elongatum Schnizl. in Mart., Fl. Bras. 4(1): 282. 1857. Lacistema coriaceum A. DC., Prodr. 16(2): 593. 1868. Lacistema orinocense Baehni, Candollea

8: 43. 1940. Shrub or small understory tree 2–10 m tall; flowers green or yellowish green, spikes usually at least 1 cm long. Forests, forest edges, edges of savannas, often along streams or rivers, common in distubed areas, 0–1200 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Neotropics. ŠFig. 575.

Lacistema 677

Fig. 575. Lacistema aggregatum

678

L AMIACEAE

LAMIACEAE by Raymond M. Harley Herbs, shrubs, or, rarely, trees, usually glandular and aromatic. Stems often 4angled. Leaves exstipulate, usually simple and opposite. Inflorescence thyrsoid, the cymes borne in the axils of bracts or upper leaves and often much modified by reduction or contraction to form false whorls (verticillasters), or forming terminal spikes, heads, capitula, or, rarely, by reduction, racemes. Flowers bisexual or functionally pistillate in gynodioecious plants, or, rarely, plants dioecious. Bracts similar to or often markedly different from the leaves, in capitulate inflorescences often forming an involucre, bracteoles usually present. Calyx usually 5-lobed, the lobes subequal to 2lipped, with 3 posterior and 2 anterior lobes, but often modified in various ways and sometimes the upper or both lips entire; calyx 5–20-veined, corolla gamopetalous, zygomorphic and bilabiate, the posterior (upper) lip indistinctly 2-lobed to entire, falcate or straight, flat to concave or hooded, anterior (lower) lip usually ± 3-lobed, longer or shorter than the posterior, the median lobe flat to concave or (in Hyptidinae) cymbiform, compressed; sometimes the corolla ± actinomorphic. Stamens 4, adnate to the corolla, didynamous, or 2 fertile with 2 staminodes usually present, included or exserted from the corolla and often oriented toward the upper or lower lip of the corolla; thecae 1- or 2-celled, parallel or divergent, sometimes confluent, the connective sometimes extended or greatly elongated (Salvia). Ovary superior, bicarpellate and 4-ovulate, deeply 4-lobed, each lobe with a single ovule. Style usually gynobasic, shortly bifid above. Fruit of 4 (rarely fewer) usually dry nutlets, mucilaginous or not when wetted. Seeds with scanty or no endosperm. Cosmopolitan, especially frequent in warm-temperate to tropical savanna and submontane regions; ca. 230 genera and over 5000 species, 9 genera and 30 species in the flora area. Recent studies indicate that many genera traditionally included in the Verbenaceae would be better placed within the Lamiaceae. However, as a new overall treatment has not yet been published, the present work maintains the traditional delimitation of these two families. Key to the Genera of Lamiaceae 1. 1. 2(1). 2. 3(2). 3. 4(2). 4.

Calyx 2-lipped, the lips entire, closed in fruit, with a rounded scale-like appendage (the scutellum) on the upper lip ......................... 9. Scutellaria Calyx toothed, without a scale-like appendage on the upper lip, though sometimes the broad upper lobe with decurrent wings ....................... 2 Stamens held under the upper lip of the corolla; calyx teeth 8–10 .......... 3 Stamens directed toward the lower lip of the corolla; calyx teeth 5, if ± obsolete then with a dense tuft of white hairs in the throat ............. 4 Corolla orange (yellow); leaves simple, toothed ............................. 5. Leonotis Corolla pink; leaves deeply lobed ................................................. 6. Leonurus Calyx strongly zygomorphic, the upper tooth broad and decurrent in a wing along the calyx tube .......................................................... 8. Ocimum Calyx not strongly zygomorphic ................................................................ 5

Eriope 679

5(4).

5. 6(5). 6. 7(6). 7. 8(7). 8.

Nutlets concave-cymbiform, with an involute, fimbriate margin on inner face; viscid herb with blue-violet flowers in small heads ....................... .......................................................................................... 7. Marsypianthes Nutlets ovoid or flattened but not as above; plant without the above combination of characters ............................................................................ 6 Upper stem with pruinose and often swollen internodes ..............2. Hypenia Stems never swollen or pruinose ............................................................... 7 Inflorescence a slender raceme of bluish flowers; calyx in fruit turbinate, with a conspicuous tuft of white hairs in throat ........................... 1. Eriope Inflorescence cymose or capitulate ............................................................ 8 Flowers in lax panicles, corolla > 10 mm long ..................... 3. Hyptidendron Flowers more densely clustered, corolla < 10mm .............................4. Hyptis

1. ERIOPE Humb. & Bonpl. ex Benth., Labiat. Gen. Sp. 142. 1833. Trees, shrubs, or subshrubs with a woody, often lobed, underground stock. Stems erect, often with setose trichomes below, sometimes (not in species from the flora area) the flowering shoots with internodes glabrous, covered with waxy bloom, inflated and fistulose above (see Hypenia). Leaves simple. Inflorescence terminal, often branched, raceme-like. Flowers in uniflorous cymes, remote or congested in the axils of slender caducous bracts; bracteoles paired, linear, often very inconspicuous, persistent at the base of calyx. Pedicel-like axes often deflexed in fruit. Calyx shortly campanulate at anthesis, sometimes densely hairy in throat, 5-toothed with broadly deltoid teeth, the 3 upper often ± connate, in fruit tubular, campanulate or turbinate, with horny texture, the throat sometimes closed by dense tuft of trichomes; corolla 2-lipped, violet-blue or sometimes pink or white, the upper lip 2lobed or emarginate, with pattern of darker lines near base; the lower lip 3-lobed, the outer laterally spreading, the inner cymbiform, compressed, with laciniate or dentate margin, hinged at base, at first enclosing the declinate stamens, to provide an explosive pollination mechanism, later deflexed; corolla tube constricted near base, widening above and becoming campanulate. Stamens 4, declinate; anthers with 2 parallel, confluent thecae; posterior pair of stamen-filaments densely clothed with long trichomes. Stylopodium present at base of style, long and tapering, distinctly overtopping the ovary. Nutlets ± flattened, sometimes winged. Colombia, Venezuela, French Guiana, Brazil, Bolivia, Paraguay; ca. 30 species, 2 in Venezuela, 1 of these in the flora area. Eriope crassipes Benth., Labiat. Gen. Sp. 144. 1833. Eriope nudiflora Kunth ex Benth., Labiat. Gen. Sp. 144. 1833. Colombia, Venezuela, French Guiana, Brazil, Bolivia, Paraguay; 3 subspecies, 1 in Venezuela. The other two subspecies have more restricted distributions: subsp. trichopoda (Briq.) Harley is known from Paraguay

and subsp. cristalinae Harley from Goiás, Brazil. E. crassipes subsp. crassipes Musky-scented subshrub 50(–80) cm tall, with several unbranched stems from swollen, woody, underground stock. Savannas, 50–200 m; Amazonas (Samariapo). Apure, Barinas; eastern Colombia, French Guiana, Brazil, Bolivia. ŠFig. 576.

680

L AMIACEAE

Fig. 576. Eriope crassipes subsp. crassipes

Fig. 577. Hypenia salzmannii

Hyptidendron 681

2. HYPENIA (Mart. ex Benth.) Harley, Bot. J. Linn. Soc. 98: 91. 1988. —Hyptis Jacq. sect. Hypenia Mart. ex Benth., Labiat. Gen. Sp. 136. 1833. Shrubs or perennial herbs with virgate stems often densely setose below, the upper internodes glabrous, with a waxy bloom and usually swollen and fistulose. Leaves simple, often coriaceous. Inflorescence a terminal, often raceme-like thyrse. Flowers borne in uniflorous, rarely 2- or 3-flowered cymes on long pedicel-like axes, in the axils of slender, caducous bracts, or cymes congested; bracteoles paired, often inconspicuous, persistent at the base of calyx. Calyx 5-toothed, the teeth subequal; corolla blue, purplish, pink, red, or yellow, sometimes white, tubular, weakly constricted at the base, the upper lip 2-lobed, the lower lip 3-lobed, the outer spreading laterally, the median lobe compressed-cymbiform, hinged at base, at first enclosing the declinate stamens to form an explosive pollination mechanism, later becoming reflexed. Stamens 4, declinate; anthers each with 2 parallel, confluent thecae. Style lacking stylopodium. Nutlets somewhat flattened. Venezuela, Guyana, Brazil, Bolivia, Paraguay; ca. 20 species, 1 in Venezuela. Most species are characterized by wax-covered stems, fistulose below the inflorescence. It seems probable that this is an adaptation to deter ants from reaching the flowers to steal nectar and inadvertently trigger the delicate explosive pollination mechanism on which cross-pollination depends. Hypenia salzmannii (Benth.) Harley, Bot. J. Linn. Soc. 98: 91. 1988. —Hyptis salzmannii Benth., Labiat. Gen. Sp. 138. 1833. Virgate herb or subshrub, stems setose below, the upper internodes glabrous, with

whitish gray waxy bloom, usually swollen, fistulose; flowers on long slender pedicel-like axes; corolla sky-blue. Open, well-drained areas, 50–400 m; Bolívar (Ciudad Bolívar, Ciudad Piar, Puerto Ordaz). Guyana, Brazil (Roraima and northeastern states). ŠFig. 577.

3. HYPTIDENDRON Harley, Bot. J. Linn. Soc. 98: 90. 1988. Hyptis sect. Buddleioides Benth., Labiat. Gen. Sp. 132. 1833. Hyptis sect. Umbellaria Benth., Labiat. Gen. Sp. 133. 1833. Trees or shrubs. Leaves simple, often coriaceous. Inflorescences bracteate, often pedunculate, cymose panicles in the axils of the sometimes reduced upper leaves. Bracteoles present. Calyx ± turbinate to tubular with 5 subequal teeth; corolla lilac, violet-blue, purplish, or pink, 2-lipped, tubular to subcampanulate, not constricted at the base, the upper lip 2-lobed, the lower 3-lobed, the outer spreading laterally, the median lobe compressed-cymbiform, hinged at base, at first enclosing the stamens to form an explosive pollination mechanism, later becoming reflexed. Stamens 4, declinate; anthers 2-thecous, the thecae parallel, confluent. Style jointed above the nutlets, the lower part (stylopodium) persistent. Nutlets ovoid or sometimes flattened and winged. Colombia, Venezuela, Guyana, Peru, Brazil, Bolivia; ca. 16 species, 1 in Venezuela. Hyptidendron arboreum (Benth.) Harley, Bot. J. Linn. Soc. 98: 93. 1988. —Hyptis arborea Benth. in A. DC., Prodr. 12: 132. 1848. Shrub or small tree 1–8 m tall; flowers with corolla violet-blue or purple, in showy

panicles, subtended by whitish or purpletinged bracts. Forest margins, secondary scrub, savannas, 600–1300 m; Bolívar (widespread). Colombia, Guyana, Peru, Brazil, Bolivia. ŠFig. 578.

682

L AMIACEAE

Fig. 578. Hyptidendron arboreum

4. HYPTIS Jacq., Collectanea. 1: 101. 1786 [1787], nom. cons. Mesosphaerum P. Browne, Civ. Nat. Hist. Jamaica: 257. 1756. Often aromatic shrubs, subshrubs, perennial, or, rarely, annual herbs of various habit. Stems erect to prostrate. Leaves simple to lobed or rarely compound. Inflorescence thyrsoid, of bracteate, cymose clusters of flowers, forming congested or rarely diffuse panicles, or verticillasters or capitula, these often surrounded by an involucre of bracts and sometimes aggregated to form pseudoracemes or spikes. Bracteoles present. Calyx usually 10-veined, with 5 usually ± subequal teeth, these sometimes unequal or obsolete; calyx tube straight or curved, usually strongly accrescent in fruit, rarely inflated. Corolla often white, sometimes pink-spotted on upper lip, otherwise pink, purplish, or blue, 2-lipped, the upper lip 2-lobed or emarginate, the lower lip 3-lobed, the outer lobes spreading, the median lobe cymbiform and usually compressed, with thickened hinge at base, at first enclosing the declinate stamens to form an explosive pollination mechanism, later becoming reflexed; corolla tube cylindrical, usually widening slightly above. Stamens 4, declinate; anthers 2-thecous, the thecae parallel, confluent. Stylopodium absent or rarely present. Nutlets usually ovoid, but sometimes elongate or flattened. Southern U.S.A., Mexico, Central America, West Indies, South America, a few species extending to Africa, Asia, Australasia, and the Pacific; ca. 290 species, 32 species in Venezuela, 22 of these in the flora area.

Hyptis 683

Key to the Species of Hyptis 1. 1.

2(1).

2.

3(1).

3.

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

5.

6(4). 6. 7(6). 7. 8(7).

8.

9(7).

Flowers not arranged in heads, but sometimes grouped in congested cymes that form racemes or spikes; involucral bracts absent .............. 2 Flowers arranged in heads subtended by an involucre of filiform to ovate bracts (sometimes reflexed or obscured in fruit); heads usually pedunculate, rarely sessile in the leaf axils or forming a usually interrupted spike ....................................................................................................... 3 Fruiting calyx campanulate, the tube ≥ 5 mm long, the throat not bearded; corolla bluish purple, the tube at least 4 mm long; nutlets > 2 mm long; stems with scattered, spreading, rather coarse trichomes mixed with shorter; ones. leaves often viscous and foetid ..................... ................................................................................................ H. suaveolens Fruiting calyx tubular, the tube ≤ 4 mm, the throat conspicuously bearded with white trichomes; corolla dull pinkish, the tube < 2 mm long; nutlets < 1.5 mm long; stems uniformly gray-green-tomentose, with shorter trichomes; leaves weakly aromatic, not viscous ........ H. pectinata Flowers in congested, few–15-flowered cymes, surrounded by an involucre of erect, narrowly elliptic bracts, tapered at each end; calyx narrowly cylindrical, elongating in fruit to 4.5–10 mm; corolla blue; leaves distinctly petiolate, with truncate to cuneate or weakly attenuate base .................................................................................................. H. mutabilis Flowers in many-flowered heads, the involucral bracts spreading, the heads pedunculate or rarely sessile and then borne in the axils of the leaves or forming terminal spikes; calyx not as above; corolla white or pinkish, if bluish then leaves ± sessile .................................................. 4 Capitula sessile or rarely with peduncles ≤ 4 mm long and then the stems densely hairy with long, coarse trichomes ............................................ 5 Capitula distinctly pedunculate with at least some peduncles > 5 mm long ................................................................................................................ 6 Lower surface of leaves and involucral bracts densely covered with sessile glands, sometimes obscured by trichomes; capitula in the axils of leaflike bracts, at least below; calyx teeth 2–4 mm long ................. H. hirsuta Lower surface of leaves and involucral bracts with few or no sessile glands; capitula subtended by inconspicuous bracts; calyx teeth ≤ 2 mm long ......................................................................................... H. guanchezii Leaves deeply bipinnatifid; slender, usually pink-tinged herb of damp places ......................................................................................... H. laciniata Leaves simple, usually toothed or sometimes shallowly lobed, but not divided ....................................................................................................... 7 Calyx tube strongly curved in fruit, the mouth oblique ........................... 8 Calyx tube straight .................................................................................... 9 Leaves gray-green, elliptic to oblanceolate, ≤ 10 mm wide, with narrowly attenuate base; flower heads with peduncles < 12 mm long; calyx tube minutely tomentose in fruit ................................................. H. microphylla Leaves green, sometimes purple-tinged, ovate with cordate to truncate base, midstem leaves > 10 mm wide; flower heads with peduncles at least 15 mm long, and some usually much longer; calyx tube, on dorsal side, covered distally with short, broad-based trichomes ...... H. recurvata Bracts of involucre filiform at least above, often curved. ....................... 10

684

L AMIACEAE

9. Bracts not filiform above, flattened to the tip ......................................... 11 10(9). Leaves sessile, ovate-oblong to narrowly linear, 2–15 mm wide and often with dilated base, the lower surface usually densely hairy, the margin crenate, sometimes recurved; capitula compact, usually densely hairy; corolla usually bluish ................................................................. H. dilatata 10. Leaves petiolate, elliptic-lanceolate, 1.3–3 cm wide, with cuneate base, sparsely puberulous on both surfaces, the margin rather coarsely serrate, plane; capitula rather lax and distinctly cymose, not densely hairy; corolla white ....................................................................... H. huberi 11(9). Style jointed above the nutlets, the basal part (stylopodium) persistent in fruit; involucral bracts ovate, ≥ 2 mm broad ...................................... 12 11. Style not jointed, falling in entirety as fruit ripens; involucral bracts various, often < 2 mm wide ........................................................................ 13 12(11). Slender, usually decumbent herb, rooting at the lower nodes; lower surface of leaves subglabrous to sparsely hairy; capitula axillary, ca. 1 cm diameter, subtended by bracts similar to the leaves; calyx teeth delicate, to 1.5 mm long ............................................................... H. atrorubens 12. Robust herb or shrub, erect and usually 1 m tall or more; lower surface of leaves gray-green tomentose; capitula in terminal panicles or racemes, 1.5–2 cm diameter, subtended by reduced but leaf-like bracts; calyx teeth rigid, 2–3 mm long ........................................................ H. brachiata 13(11). Calyx tube with a dense ring of trichomes within, above the nutlets .... 14 13. Calyx tube without a ring of trichomes within ....................................... 17 14(13). Leaves < 4 cm long ................................................................................... 15 14. At least some leaves > 4 cm long, often much longer ............................. 16 15(14). Leaves elliptic, somewhat coriaceous, the lower surface often densely appressed-hairy, the trichomes sometimes restricted to the veins; stems uniformly appressed-hairy; petiole indistinct, very short; fruiting capitula rufous, 15–20 mm diameter ...................................... H. lantanifolia 15. Leaves ovate, thin, the lower surface glabrous or thinly hairy on the veins; stem trichomes usually restricted to the angles; petiole distinct, slightly winged toward the cuneate to truncate leaf base; fruiting capitula greenish, or sometimes purple- tinged, to 14 mm diameter ..................................................................................................... H. luticola 16(14). Leaves coriaceous, linear to linear-lanceolate, ≥ 5 times as long as broad, veins strongly impressed on upper surface; petiole indistinct; involucral bracts densely appressed-hairy with pale trichomes, capitula pale in fruit, to 2 cm diameter, on peduncles to 3 cm long; calyx tube in fruit 5–6 mm long ............................................................................... H. conferta 16. Leaves thin, lanceolate to broadly ovate, ≤ 4 times as long as broad, veins not impressed; petiole distinct; involucral bracts sparsely hairy; capitula small at first, becoming dark brown in fruit, to 3 cm diameter, on peduncles to 6 cm long; calyx tube in fruit 6–8 mm long ......... H. capitata 17(13). Calyx teeth deltoid, as broad as long ............................................. H. parkeri 17. Calyx teeth subulate, sometimes broadly so, or acicular, longer than broad ..................................................................................................... 18 18(17). Calyx teeth subulate, sometimes broadly so; fruiting capitula ca. 15 mm long, if less, then leaves coriaceous and rugose .................................. 19 18. Calyx teeth acicular; fruiting capitula to 12 mm diameter; leaves never coriaceous nor rugose .......................................................................... 21

Hyptis 685

19(18). Herbs; leaves thin, elliptic-lanceolate, 6–10 cm long, never rugose; calyx teeth broadly subulate, with rather blunt tip, 1–1.5 mm long ............... ................................................................................................... H. lacustris 19. Shrubs, sometimes of reduced stature; leaves coriaceous, elliptic, to 5 cm long; calyx teeth narrowly subulate, but not filiform, 1–3.5 mm long, if less than 2 mm, then leaves rugose on upper surface ........................ 20 20(19). Leaves narrowing to a weakly cordate to auriculate base; petiole to 2 mm long; capitula 10–12 mm diameter; involucral bracts 4–5.5 × 0.75– 1.5 mm; calyx at anthesis with tube 2.5–3 mm long, the teeth 1–2 mm; corolla tube 4.5 mm; nutlets (?always) hairy at apex ...... H. colubrimontis 20. Leaves variable in shape with a cuneate to attenuate base; petiole 2–3.5 mm long; capitula ca. 15 mm diameter; involucral bracts 6.5–8 × 0.3–1.5 mm; calyx at anthesis with tube 3–4 mm long, the teeth 2–3.5 mm long; corolla tube 6 mm long; nutlets glabrous at apex .................... H. pyriformis 21(18). Cauline leaves rather dark green, appearing glabrous, but usually with minute, sparse, appressed trichomes on upper surface and restricted to the veins on the lower surface; leaf margin usually regularly serrate; calyx teeth 1–1.5 mm long ..................................................... H. lanceolata 21. Cauline leaves pale green, the lower surface very sparsely hairy with long spreading trichomes; leaf margin often irregularly serrate or dentate; calyx teeth (1–)1.5–2 mm long .................................................. H. brevipes Hyptis atrorubens Poit., Ann. Mus. Natl. Hist. Nat. 7: 466, t. 27, fig. 3. 1806. —Mesosphaerum atrorubens (Poit.) Kuntze, Revis. Gen. Pl. 2: 525. 1891. Semiprostrate to erect herb, often growing through surrounding vegetation, 15–50 cm tall; flowers in capitula; corolla white or pale lilac with pink spots on upper lip. Wet savannas, damp forest margins and by water, 50–1300 m; Bolívar (widespread), Amazonas (Puerto Ayacucho area, lower Río Ventuari). Monagas, Portuguesa; Mexico, Central America, West Indies, Colombia, Ecuador, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, West Africa. ŠFig. 582. Hyptis brachiata Briq., Annuaire Conserv. Jard. Bot. Genève 2: 215. 1898. Branched subshrub 0.1–2 m tall; flowers in capitula; corolla white. Savanna margins, shrub patches within savannas, 50–200 m; Bolívar (Caicara, Río Parhueña), Amazonas (around Puerto Ayacucho eastward to upper Río Parucito). Barinas, Guárico, Mérida, Miranda, Portuguesa, Táchira; Costa Rica, Panama, Colombia. ŠFig. 581. Hyptis brachiata together with Hyptis lutescens Pohl ex Benth. from Brazil, forms part of a complex of biotypes requiring further study to determine species limits.

Hyptis brevipes Poit., Ann. Mus. Natl. Hist. Nat. 7: 465. 1806. —Mesosphaerum brevipes (Poit.) Kuntze, Revis. Gen. Pl. 2: 525. 1891. Hyptis brevipes var. serrata Briq., Annuaire Conserv. Jard. Bot. Genève 2: 226. 1898. Erect herb 0.3–1 m tall; flowers borne in capitula; corolla white, the upper lip with pink spots. Cultivated areas, moist savannas, forest margins, 0–200 m; Delta Amacuro (Sacupana, Tucupita), Bolívar (Reserva Forestal Imataca). Northwestern Venezuela; Mexico, Central America, West Indies, tropical South America. ŠFig. 586. Some collections from the Guayana region that were previously incorrectly determined as Hyptis pulegioides Pohl ex Benth. belong here. Hyptis capitata Jacq., Collectanea 101. 1787. —Mesosphaerum capitatum (Jacq.) Kuntze, Revis. Gen. Pl. 2: 525. 1891. Usually robust, erect herb to 2 m tall; flowers in capitula; calyx accrescent and turning dark brown in fruit; corolla white. Open and somewhat disturbed areas, 0–300 m; Bolívar (between Río Caura and Río Parágua). Venezuelan Andes (to 1200 m); U.S.A. (Florida), Mexico, Central America,

686

L AMIACEAE

West Indies, western South America south to Peru, probably introduced into tropical Asia and the Pacific. Hyptis colubrimontis Epling & Játiva, Mem. New York Bot. Gard. 17(1): 229. 1967. Shrub to 50 cm tall or more; leaves coriaceous, dark green; flowers in capitula; corolla white. Wet areas on sandstone outcrops, 1100–1600 m; Amazonas (Cerro Duida, Cerro Parú). Endemic. This species has, until now, been confused with Hyptis pyriformis Epling & Játiva (see R. M. Harley, Kew Bull. 53: 973–976. 1998). Hyptis conferta Pohl ex Benth., Labiat. Gen. Sp. 112. 1833. —Mesosphaerum confertum (Pohl ex Benth.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Robust herb or shrub, with erect stems 0.5–1.5 m tall; leaves somewhat coriaceous, linear to linear-lanceolate; flowers in capitula; corolla white. Mexico, Central America, West Indies, Colombia, Venezuela, Brazil, Bolivia, Paraguay; 2 varieties, 1 in Venezuela. Some collections have been misdetermined as Hyptis savannarum Briq., a species of Central America and western South America, which does not occur in the flora area. Hyptis conferta var. conferta is apparently restricted to Central Brazil. It differs from var. angustata by its shorter and broader leaves. H. conferta var. angustata (Briq.) Pool & Harley, Novon 4: 45. 1944. —Hyptis jurgensenii var. angustata Briq., Annuaire Conserv. Jard. Bot. Genève. 2: 220. 1898. Hyptis ferruginea Benth., Labiat. Gen. Sp. 113. 1833. Hyptis excelsa M. Martens & Galeotti, Bull. Acad. Roy. Bruxelles 11, 2: 188. 1844. —Mesosphaerum excelsum (M. Martens & Galeotti) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Hyptis lundii Benth. in A. DC., Prodr. 12: 111. 1848. Hyptis constricta Briq., Annuaire Conserv. Jard. Bot. Genève. 2: 220. 1898. Hyptis jurgensenii Briq., Annuaire Conserv. Jard. Bot. Genève. 2: 219. 1898.

Hyptis conferta var. angustifolia Benth. ex Epling, Spec. Nov. Regni Veg. Beih. 85: 124. 1936, nom. illeg. Hyptis belizensis Lundell, Contr. Univ. Michigan Herb. 8: 83. 1942. Savannas, 50–500 m; Bolívar (northern part of state and lower slopes of Auyántepui), northern Amazonas. Central and northeastern Venezuela; other distribution as in species. ŠFig. 588. Hyptis dilatata Benth. in A. DC., Prodr. 12: 103. 1848. —Mesosphaerum dilatatum (Benth.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Hyptis crenata var. angustifolia Benth. ex Seem., Bot. Voy. Herald 187. 1854. Hyptis lanicephala Epling & Játiva, Mem. New York Bot. Gard. 17(1): 228. 1967. Aromatic herb to 1 m tall, often smaller; leaves oblong, linear-oblong or rarely narrowly linear; flowers in densely hairy capitula; corolla lavender-blue or white. Moist savannas, Mauritia palm swamps, 50–500 m; northern and western Bolívar, Amazonas (around Puerto Ayacucho, Río Ventuari). Apure, Guárico, Portuguesa; Panama, Colombia, northern Brazil. ŠFig. 595. This species is closely related to the more southern Hyptis crenata Benth. from Brazil, with which it may prove conspecific. It differs mainly in its oblong leaves, which are relatively narrower than those of H. crenata. Hyptis lanicephala is a less hirsute form with extremely narrow leaves, often to 4 mm wide, apparently endemic to the Río Manapiare region of Amazonas. Although it may merit formal recognition at infraspecific rank, intermediates exist between this and H. dilatata. Typical material of the latter has also been determined as H. lanicephala, further adding to the confusion. Hyptis guanchezii Harley, Ann. Missouri Bot. Gard. 76: 1169. 1989. Erect herb to 80 cm tall, branching from a central rootstock; leaves ovate-lanceolate to narrowly ovate; flowers in sessile capitula; corolla white. Forests on granitic outcrops, 100–200 m; Amazonas (Cerro Uchonhua ca. 60 km southeast of Puerto Ayacucho). Endemic. Hyptis hirsuta H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 318, t. 161. 1817 [1818].

Hyptis 687

—Mesosphaerum hirsutum (H.B.K.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Often robust, spreading herb, to 1 m tall, much branched, the stems hirsute with spreading trichomes; flowers in sessile or subsessile capitula in the axils of the leaves; corolla white, the upper lobe pink-spotted. Savannas, evergreen lowland forests, 200– 900 m; Bolívar (Gran Sabana, east of Río Caroní). Distrito Federal, Sucre; Colombia (Meta), Guyana, Suriname, Brazil, Bolivia, Paraguay. ŠFig. 596.

stream sides, forests, sometimes as a weed of cultivation, 100–1400 m; Delta Amacuro (Río Amacuro), eastern Bolívar, Amazonas (Sierra Parima). Belize to Panama, West Indies, Guyana, Suriname, French Guiana, Brazil, apparently introduced into tropical Africa. ŠFig. 592.

Hyptis huberi Harley, Kew Bull. 41: 147. 1986. Erect, unbranched herb to 70 cm tall; leaves elliptic-lanceolate, sparsely puberulous, flowers in capitula; corolla white. Savannas, 200–400 m; Amazonas (Río Manapiare basin). Endemic. ŠFig. 594.

Hyptis lantanifolia Poit., Ann. Mus. Natl. Hist. Nat. 7: 468, t. 29. 1806. —Mesosphaerum lantanifolium (Poit.) Kuntze, Revis. Gen. 2: 525. 1891. Decumbent to erect, usually weakstemmed, perennial herb to 1 m tall; flowers in capitula; corolla white, often with pale lilac lobes, the upper lip pink-spotted. Savannas, 100–1200 m; Bolívar (Cerro Jaua, Gran Sabana), Amazonas (Puerto Ayacucho). Northern Venezuela; widespread in the Neotropics. ŠFig. 584.

Hyptis laciniata Benth., J. Bot. (Hooker) 2: 49. 1840. —Mesosphaerum laciniatum (Benth.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Slender perennial herb to 30 cm tall with procumbent to erect stems; leaves 2-pinnatifid; flowers in capitula; corolla white or pinkish, with spotted upper lip. Sandy river banks, moist savannas, 50–200 m; Bolívar (Río Orinoco), Amazonas (Puerto Ayacucho, Río Orinoco, Río Parguaza). Guárico; Colombia, Guyana, Brazil. ŠFig. 580.

Hyptis luticola Epling, Brittonia 7: 142. 1951. Slender herb with weak, decumbent stems to 1 m tall, usually smaller; flowers in capitula; corolla white. Seasonally flooded savannas, 50–200 m; Amazonas (San Juan de Manapiare). Apure, Guárico; Colombia (Meta). ŠFig. 587. This species is possibly only a slender form of the southern Hyptis lorentziana O. Hoffm., which occurs from southern Brazil and Bolivia to Argentina.

Hyptis lacustris St.-Hil. ex Benth., Labiat. Gen. Sp. 107. 1833. —Mesosphaerum lacustre (St.-Hil. ex Benth.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Sprawling herb to ca. 1 m tall; leaves narrowly elliptic-lanceolate; flowers in capitula in the axils of the leaves; corolla white. Evergreen lowland forests, 50–200 m; Bolívar (Río Caura), Amazonas (Río Casiquiare, Río Padamo). Amazonian Colombia and Peru, Brazil (Amazonas and Mato Grosso), with a second center of distribution in southern Brazil.

Hyptis microphylla Pohl ex Benth., Labiat. Gen. Sp. 82. 1833. —Mesosphaerum microphyllum (Pohl ex Benth.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Much-branched subshrub or herb to 1.5 m tall, with grayish stems; leaves small, gray-green, elliptic to oblanceolate; flowers in capitula; corolla white. Depressions in savannas, often near water, 50–300 m; Bolívar (Ciudad Piar, Tumeremo). Barinas, Guárico, Monagas; Cuba, tropical South America south to Paraguay and Bolivia. ŠFig. 589.

Hyptis lanceolata Poir. in Lam., Encycl. suppl. 3: 114. 1813. —Mesosphaerum lanceolatum (Poir.) Kuntze, Revis. Gen. Pl. 2: 526. 1891. Herb 1–2 m tall, the stems glabrous to thinly hairy; leaves elliptic-lanceolate; flowers in capitula; corolla white. Wet places,

Hyptis mutabilis (Rich.) Briq., Bull. Herb. Boissier 4: 788. 1896. —Nepeta mutabilis Rich., Actes Soc. Hist. Nat. Paris 1: 110. 1792. —Mesosphaerum mutabile (Rich.) Kuntze, Revis. Gen. Pl. 2: 525. 1891.

688

L AMIACEAE

Hyptis spicata Poit., Ann. Mus. Natl. Hist. Nat. 7: 474, t. 28, fig. 2. 1806. Hyptis canescens H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 321. 1817 [1818]. —Hyptis mutabilis var. canescens (H.B.K.) Briq., Bull. Herb. Boissier 4: 788. 1896. Hyptis polystachya H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 321. 1817 [1818]. Hyptis micrantha Pohl ex Benth., Labiat. Gen. Sp. 120. 1833. Robust, erect herb 1–2 m tall; leaves ovate to rhomboid, the lower surface often tomentose; flowers in small compact cymes surrounded by an involucre of bracts, and forming a long, terminal spike-like thyrse; corolla bluish purple or white. In disturbed and waste places, 50–400 m; northeastern Bolívar, Amazonas (Puerto Ayacucho). Widespread in northern Venezuela; frequent throughout the Neotropics, often as a weed, from U.S.A. (Florida), Mexico and the West Indies, south to Argentina, introduced in South Africa. ŠFig. 593. Material from the flora area appears to fall within var. canescens (H.B.K.) Briq., which has more hirsute leaves. However, it is still unclear if varieties merit recognition, as intermediates occur between varieties.

Hyptis pectinata (L.) Poit., Ann. Mus. Natl. Hist. Nat. 7: 474, t. 30. 1806. —Nepeta pectinata L., Syst. Nat. ed. 10, 2: 1097. 1759. —Mesosphaerum pectinatum (L.) Kuntze, Revis. Gen. Pl. 2: 525. 1891. Robust, erect, perennial or annual herb to 2 m tall or more; leaves ovate, the lower surface gray-green; flowers in short cincinnate cymes, condensed to form a terminal spikelike thyrse; calyx with white trichomes in throat and short filiform teeth; corolla very small, dull pink. Roadsides and disturbed areas through forests and savannas, 0–800 m; Delta Amacuro (Pedernales, Sacupana), scattered in northern and eastern Bolívar. Frequent and weedy elsewhere in Venezuela; widespread in the Neotropics from U.S.A. (Florida) and Mexico southwards, introduced in Africa and Asia. ŠFig. 585.

Hyptis parkeri Benth., Labiat. Gen. Sp. 108. 1833. —Mesosphaerum parkeri (Benth.) Kuntze, Revis. Gen. Pl. 2: 525. 1891. Hyptis verbenifolia J.A. Schmidt ex Mart., Fl. Bras. 8(1): 110. 1858, “verbenaefolia.” —Hyptis parkeri var. verbenifolia (J.A. Schmidt ex Mart.) Epling, Bull. Torrey Bot. Club 58: 465. 1931, “verbenaefolia.” Decumbent to ascending herb, rooting at the nodes; leaves elliptic-lanceolate to elliptic-ovate with serrate margin; flowers in capitula; calyx pale green; corolla white. River margins, wet, sandy soil, 100–900 m; Bolívar (Cerro Guaiquinima), Amazonas (San Carlos del Río Negro area). Eastern Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil, Bolivia. ŠFig. 579. Var. verbenifolia (J.A. Schmidt ex Mart.) Epling has shorter leaves than the typical variety. However both varieties, with intermediates, have been observed growing together in Brazil, and there is no good reason for recognizing them as two taxa.

1. Lower surface of leaves densely hairy, obscuring the sessile glands, the upper surface rugose ............ var. pyriformis 1. Lower surface of leaves glabrous, except for appressed trichomes on the veins, sessile glands clearly visible, the upper surface not rugose .... var. yutajeensis

Hyptis pyriformis Epling & Játiva, Mem. New York Bot. Gard. 17: 229. 1967. Shrub 1–5 m tall; leaves coriaceous; flowers in capitula; corolla white. Wet areas among sandstone outcrops, 1100–2100 m; 2 varieties, both restricted to the flora area. Endemic. Key to the Varieties of H. pyriformis

H. pyriformis var. pyriformis 1700–1800 m; Bolívar (Cerro Guanay), Amazonas (Caño Guaviarito, near the Río Manapiare). Endemic. ŠFig. 590. H. pyriformis var. yutajeensis Harley, 53: 975. 1998. Hyptis colubrimontis Epling & Játiva, Mem. New York Bot. Gard. 17(1): 229. 1967, pro parte. 1100–2100 m; Amazonas (Cerro Yutajé). Endemic. ŠFig. 591. Hyptis recurvata Poit., Ann. Mus. Natl. Hist. Nat. 7: 467, t. 28. 1806. —Mesosphaerum recurvatum (Poit.) Kuntze, Revis. Gen. Pl. 2: 527. 1891.

Hyptis 689

Hyptis recurvata var. megacephala Benth. in A. DC., Prodr. 12: 90. 1848. Weak-stemmed, weedy herb, often sprawling on surrounding vegetation; leaves ovate to ovate-elliptic; flowers in capitula; corolla white, the upper lip pink-spotted. Moist savannas, often by water and along disturbed forest margins, 50–800 m; northern Bolívar (Ciudad Piar, Nueva Fortuna, Playa Blanca to Río Uaiparú), Amazonas (near Puerto Ayacucho, Río Casiquiare, San Carlos de Río Negro). Widespread elsewhere in Venezuela; Mexico, Central America, West Indies, South America southward to Brazil and Bolivia. ŠFig. 583. Hyptis suaveolens (L.) Poit., Ann. Mus. Natl. Hist. Nat. 7: 472, t. 29. 1806. —Ballota suaveolens L., Syst. Nat. ed.

Fig. 579. Hyptis parkeri

10, 2: 1100. 1759. —Mesosphaerum suaveolens (L.) Kuntze, Revis. Gen. Pl. 2: 525. 1891. Usually robust, erect annual to perennial herb, of variable habit, to 2 m tall; leaves viscid and with a strong odor; flowers in axillary cymes, sometimes condensed to form a terminal spike-like thyrse; calyx accrescent in fruit, widened above and with a ring of white trichomes at throat, the calyx teeth subulate; corolla violet-blue; nutlets usually 2 per calyx, rather large, flattened. Often weedy in disturbed or cultivated areas, forest margins or clearings, 50–1000 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, West Indies, south to Brazil and Bolivia, widely introduced as a weed in the Old World Tropics.

Fig. 580. Hyptis laciniata

690

L AMIACEAE

Fig. 581. Hyptis brachiata

Fig. 582. Hyptis atrorubens

Fig. 583. Hyptis recurvata

Hyptis 691

Fig. 584. Hyptis lantanifolia

Fig. 585. Hyptis pectinata

Fig. 586. Hyptis brevipes

692

L AMIACEAE

Fig. 587. Hyptis luticola

Fig. 588. Hyptis conferta var. angustata

Fig. 589. Hyptis microphylla

Hyptis 693

Fig. 590. Hyptis pyriformis var. pyriformis

Fig. 591. Hyptis pyriformis var. yutajeensis

694

L AMIACEAE

Fig. 593. Hyptis mutabilis

Fig. 592. Hyptis lanceolata

Hyptis 695

Fig. 595. Hyptis dilatata Fig. 594. Hyptis huberi

Fig. 596. Hyptis hirsuta

696

L AMIACEAE

5. LEONOTIS (Pers.) R. Br., Prodr. 504. 1810. —Phlomis subgenus Leonotis Pers., Syn. 2: 127. 1807. Robust annual to perennial herbs or shrubs. Leaves simple. Inflorescence composed of 3–11 dense, ± spherical, many-flowered verticillasters. Bracts usually leaflike, bracteoles spinescent. Calyx tubular, 10-veined, 8–10-toothed, teeth usually spinescent, the posterior tooth usually much longer than the others; corolla tubular, deeply 2-lipped, usually appearing orange due to a dense indumentum of orange-colored trichomes; upper lip entire, hooded, almost as long as the tube, the lower lip 3lobed, soon withering; stamens 4, inserted near the throat of the corolla and directed under the upper lip, the anterior pair of stamens longer; thecae 2, divaricate. Style with only the ventral branch developed. Nutlets glabrous, 3-angled. Tropical Africa and Asia (one species a pantropical weed); ca. 10 species, 1 in Venezuela. Leonotis nepetifolia (L.) R. Br. in W.T. Aiton, Hortus Kew ed. 2, 3: 409. 1811. —Phlomis nepetaefolia L., Sp. Pl. 586. 1753. Annual or short-lived perennial herb 1–3 m tall; leaves deeply crenate, petiolate; inflo-

rescences terminal, of 2–5 many-flowered verticillasters of large, orange flowers. Weed of waste places, 0–200 m; Delta Amacuro (Tucupita). Common in northern Venezuela; pantropical weed. ŠFig. 597.

6. LEONURUS L., Sp. Pl. 584. 1753. Annual to perennial herbs. Leaves digitately lobed or undivided, the margin coarsely toothed or pinnatifid. Inflorescence of many-flowered verticillasters in the axils of leafy bracts, verticillasters remote or often crowded near apex of flowering shoot to form an interrupted spike-like inflorescence. Bracteoles linear, subulate. Calyx 5–10-veined, with spinose teeth. Corolla pink, strongly 2-lipped, the upper lip hooded, hairy on back; lower lip 3-lobed. Stamens 4, directed under the upper corolla lip; anthers with 2 thecae, parallel or divergent. Nutlets trigonous with truncate apex. Temperate to subtropical Eurasia, 2 species introduced into the New World; ca. 6 species, 1 in Venezuela. Leonurus cardiaca has been introduced into temperate areas in North America. Leonurus japonicus is a frequent tropical weed, except in Africa, and is a rare temperate introduction. Leonurus japonicus Houtt., Nat. Hist 9: 336, t. 57, fig. 1. 1778. Leonurus heterophyllus Sweet, Brit. Fl. Gard. 2: t. 197. 1827. Leonurus sibiricus auctt. non L. 1753. Annual or short-lived perennial herb; leaves divided. Waste places, 0–200 m; Delta Amacuro (Sacupana), probably in other areas within the flora area. Frequent elsewhere in Venezuela; also introduced in temperate North America, Mexico, West Indies, southward to Brazil, tropical Asia (native), the Mascarenes, absent from Africa. ŠFig. 598.

In his revision of Leonurus sect. Cardiochilium V. Krecz. & Kuprian., Krestovskaya (Novosti Sist.-Vyssh. Rast. 24: 156– 158. 1987, and 25: 131–134. 1988) has proposed that the correct name for the taxon introduced into South America is L. japonicus Houtt. (= L. heterophyllus Sweet). According to this author, Leonurus sibiricus L. is a species of restricted distribution in eastern Asia. A full evaluation of relevant material in the Kew Herbarium has confirmed this.

Leonurus 697

Fig. 597. Leonotis nepetifolia

Fig. 598. Leonurus japonicus

698

L AMIACEAE

7. MARSYPIANTHES Mart. ex Benth., Labiat. Gen. Sp. 53. 1832, spelled Marsypianthus in generic conspectus, but corrected in text and index. Usually viscid and aromatic perennial herbs or subshrubs. Stems erect or procumbent to ascending. Leaves soft. Inflorescences axillary, in subglobose, cymose, many- or rarely few-flowered heads, the flowers shortly pedicellate and subtended by long, curved, linear to subulate bracteoles. Calyx campanulate to broadly funnelshaped; calyx teeth subequal, broadly lanceolate to deltate, spreading in fruit, not densely hairy in throat; corolla violet-blue, 2-lipped, with slender tube and ± spreading lobes, the upper lip 2-lobed or emarginate; the lower lip 3-lobed, the outer spreading, the inner saccate, hinged at base and at first enclosing the declinate stamens to form an explosive pollination mechanism, later deflexed. Stamens 4, declinate; anthers 2-thecous with parallel thecae. Nutlets ovate, smooth on outer surface, the inner concave-cymbiform with involute, fimbriate margin. Mainly Central Brazil to Paraguay and Argentina, but one species extending throughout the Neotropics; ca. 6 species, 1 in Venezuela. Marsypianthes chamaedrys (Vahl) Kuntze, Revis. Gen. Pl. 524. 1891. —Clinopodium chamaedrys Vahl, Symb. Bot. 3: 77. 1794. Marsypianthes hyptoides Mart. ex Benth., Labiat. Gen. Sp. 64. 1833. Prostrate to ascending, usually viscid herb; leaves petiolate; heads of violet-blue flowers. Damp savannas, forest clearings, 0– 200 m; Delta Amacuro (Santa Catalina), northern Bolívar. Widespread elsewhere in Venezuela; throughout the Neotropics, often as a weed, from Mexico and the West Indies, southward to northern Argentina. ŠFig. 599.

Fig. 599. Marsypianthes chamaedrys

8. OCIMUM L., Sp. Pl. 597. 1753. Annual to perennial aromatic herbs or small shrubs. Leaves simple. Inflorescence interrupted, spike-like, composed of verticillasters in the axils of bracts in the upper part of the flowering stems. Bracteoles absent. Calyx strongly 2-lipped, usually deflexed in fruit on a curved pedicel; upper lip of calyx larger than the lower, entire, ovate-orbicular with margins decurrent along the calyx tube; lower lip 4toothed with acuminate teeth; corolla strongly 2-lipped, the upper lip 4-lobed, the lower entire or toothed. Stamens 4, declinate along the lower lip of the corolla; filaments of the upper pair appendaged in some species; anthers 2-thecous. Nutlets ovoid, mucilaginous when wetted. Pantropics; ca. 65 species, 4 in Venezuela (3 of them introduced from the Old World), 1 of these in the flora area.

Scutellaria 699

Ocimum campechianum Mill., Gard. Dict. ed. 8, Ocimum no. 5. 1768. Ocimum micranthum Willd., Enum. Pl. 630. 1809. Strongly aromatic, branched, and usually bushy herb to ca. 50 cm; leaves ovate, petiolate; corolla very small, white or purplish. Savannas, roadsides, forest clearings, 0–300 m; scattered in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; throughout tropical America. ŠFig. 600.

Fig. 600. Ocimum campechianum

9. SCUTELLARIA L., Sp. Pl. 598. 1753. Perennial herbs or subshrubs, lacking aromatic odor, of varied habit. Leaves, at least the lower ones, usually petiolate. Inflorescence a raceme or spike, the flowers opposite or sometimes borne irregularly, or rarely flowers solitary, the flowers arising singly in the axils of leaf-like, reduced or highly differentiated bracts. Calyx 2lipped, accrescent in fruit, the tube ventricose-campanulate, the upper part usually with a rounded, concave, scale-like appendage (the scutellum); the lips entire, rounded, closed in fruit, the upper part of the calyx eventually deciduous, the lower part persistent, borne on a rigid and usually curved pedicel; corolla with a long, usually sigmoid or arcuate tube dilated above, 2-lipped, the upper lip galeate. Stamens 4, ascending under the upper lip and included within it, the anterior pair longer, 1thecous, posterior pair with 2 convergent thecae; style branches unequal, the dorsal reduced. Nutlets depressed-globose to ovoid, borne on an elongate, peg-like gynophore. Cosmopolitan; ca. 300 species, ca. 5 in Venezuela, 1 of these in the flora area.

700

L AMIACEAE

Scutellaria purpurascens Sw., Prodr. 89. 1788. Small herb to ca. 50 cm tall, with several erect stems arising from a short rhizome bearing a fascicle of fleshy roots; flowers borne in racemes; corolla tubular, purplish. Open, somewhat weedy areas, 50–200 m; Bolívar (south of Ciudad Bolívar). Scattered in northern Venezuela; Central America, West Indies, northern South America to Brazil. ŠFig. 601. Fig. 601. Scutellaria purpurascens

LAURACEAE by Henk van der Werff and Jens G. Rohwer Shrubs to tall trees, mostly evergreen, only Cassytha an herbaceous parasite. Indument of simple trichomes. Leaves alternate, sometimes clustered at tips of branches, rarely opposite, reduced to scales in Cassytha; exstipulate; blades entire or rarely lobed, pinnately or sometimes tripli- or triveined, often coriaceous. Inflorescence axillary, paniculate or rarely spicate, sometimes protected by large bracts. Flowers small, usually white, green, or yellow, often fragrant, bisexual or unisexual (plants hermaphroditic or dioecious). Tepals in 2 whorls of 3 (rarely 2 in Old World), equal to strongly unequal; if unequal, the outer whorl usually smaller than the inner, rarely larger. Stamens in 4 whorls of 3, rarely all fertile, the innermost whorl nearly always staminodial or lacking, usually the outer 3 whorls functional, but sometimes only 6 or 3 functional stamens; stamens of the third whorl with 2 glands near their base, rarely all stamens with glands. Anthers with 2- or 4-celled, the outer 6 stamens usually with introrse cells, the inner 3 with extrorse cells. Ovary unilocular, anatropous, ovule 1. Fruit a 1-seeded berry, which may be unprotected, with persistent tepals at the base, or partly or entirely enclosed by the accrescent floral tube. Mainly tropics, best represented in Neotropics and Asia; ca. 53 genera and ca. 3000 species, 15 genera and 142 species in the flora area. Many species yield good quality timber. Persea americana is widely cultivated for its edible fruit. Many species contain volatile oils, and a few species produce commercial products such as cinnamon (Cinnamomum verum), camphor (Cinnamomum camphora), bay leaves (Laurus, Umbellularia), and rosewood oil (Aniba rosaeodora).

L AURACEAE 701

The following generic key and list of fruit and vegetative characters apply only to species from the Venezuelan Guayana. Because the key is based on stamen characters, pistillate specimens will be difficult to identify to genus. This problem with Endlicheria, Ocotea, and Rhodostemonodaphne cannot easily be solved; one can only identify these specimens by comparison with reliably identified material. This applies to fruiting or sterile specimens as well. Some of the genera can be readily identified by fruit or vegetative characters. Following the Key to the Genera is a short list of fruit characters and one of vegetative characters. Key to the Genera of Lauraceae by Henk van der Werff 1.

Twining herbaceous parasite, the leaves reduced to small scales ................................................................................................... 4. Cassytha 1. Shrubs or trees with normal leaves ........................................................... 2 2(1). Outer 6 stamens 4-celled (if only 6 stamens present, all 4-celled) ........... 3 2. Outer 6 stamens 2-celled (if only 6 or 3 stamens present, all 2-celled) .............................................................................................................. 11 3(2). Tepals unequal, the outer ones 2/3 or less the size of the inner ones ........ 4 3. Tepals equal or subequal ............................................................................ 5 4(3). Leaves clustered at tips of branches; stamens sessile or nearly so .................................................................................................. 15. Sextonia 4. Leaves evenly distributed along the branches; stamens with well-developed filaments, these longer than the anthers ........................... 12. Persea 5(3). Flowers unisexual ...................................................................................... 6 5. Flowers bisexual ......................................................................................... 7 6(5). Anther cells arranged in an arc; filaments poorly differentiated from anthers; flowers densely pubescent ....................... 14. Rhodostemonodaphne 6. Anther cells arranged in 2 horizontal rows; filaments well differentiated, narrower than anthers; flowers variously pubescent or glabrous ...................................................................................................... 11. Ocotea 7(5). Outer 6 stamens with the lower pair of cells lateral, the other pair introrse and the filament about as long as the anther .... 13. Pleurothyrium 7. Outer 6 stamens with all cells introrse or if one pair seemingly lateral, then filament several times longer than anther ................................... 8 8(7). Anther cells arranged in an arc; anthers and often inner face of tepals papillate ................................................................................ 10. Nectandra 8. Anther cells arranged in two horizontal rows; anthers not papillate ...... 9 9(8). Staminodia lacking or linear ........................................................... 11. Ocotea 9. Staminodia present, the tips cordate or sagittate .................................. 10 10(9). Leaves tripliveined; stamens 9; cupule small, with persistent tepals ........................................................................................... 5. Cinnamomum 10. Leaves pinnately veined; stamens 6 or 9; cupule lacking or as in Cinnamomum .............................................................................. 12. Persea 11(2). Flowers unisexual ..................................................................... 6. Endlicheria 11. Flowers bisexual ....................................................................................... 12

702

L AURACEAE

12(11). 12. 13(12). 13. 14(13). 14. 15(14).

Fertile stamens 6 .............................................................................. 1. Aiouea Fertile stamens 3 or 9 .............................................................................. 13 Fertile stamens 3 ..................................................................................... 14 Fertile stamens 9 ..................................................................................... 17 Anther cells introrse ........................................................................ 8. Licaria Anther cells extrorse ................................................................................ 15 Leaves clustered near tip of branches; cupule small, plate-like; staminodia absent ............................................................................... 9. Mezilaurus 15. Leaves evenly distributed along twigs; cupule cup-shaped; staminodia present .................................................................................................. 16 16(15). Stamens of whorl I fertile; anther cells extrorse, opening upwards; leaves and flowers glabrous ..................................................................... 1. Aiouea 16. Stamens of whorl III fertile; anther cells apical or, if extrorse, opening downwards; most species variously pubescent ........................... 8. Licaria 17(13). Filaments of the inner 3 stamens fused into a column .............. 7. Kubitzkia 17. Filaments of the inner 3 stamens free .................................................... 18 18(17). Tepals much longer than the short floral tube; leaves subopposite; fruit without cupule .................................................................. 3. Beilschmiedia 18. Tepals much shorter than the long floral tube; leaves alternate or clustered at tips of branches; fruits subtended by cupule .................. 2. Aniba Fruiting Characters • Fruit without cupule or persistent tepals: Persea, Beilschmiedia. • Fruit without cupule, but with persistent tepals: Persea; check also Cinnamomum, which has a small cupule with persistent tepals. • Cupule with double margin: Licaria, Kubitzkia, a few species of Ocotea. • Cupule small, plate-like: Mezilaurus, Ocotea. • Cupule shallow, becoming flat, with a lobed margin: Endlicheria rubriflora. • Cupule very large, 3–5 cm diameter: Licaria cannella, Ocotea. Vegetative characters • Leaves opposite: Licaria, check also Beilschmiedia. • Leaves clustered at tips of branches: Mezilaurus, Aniba, Endlicheria, Sextonia rubra, less clearly in Licaria cannella. 1. AIOUEA Aubl., Hist. Pl. Guiane 310. 1775. by Henk van der Werff Trees or shrubs. Leaves alternate, simple, firmly chartaceous, sometimes with a cartilaginous margin, usually glabrous, pinnately veined or tripliveined, axillary tufts of hair at the base of the lateral veins rarely present. Inflorescences axillary or subterminal, usually slender and glabrous, paniculate. Flowers small; tepals 6, usually erect at anthesis, equal; stamens 9, 6, or rarely 3, the inner 3 extrorse and with 2 glands at their base; staminodia usually present, 3, with a cordate or sagittate apex; ovary usually immersed in the floral tube; ovule 1. Fruit an ellipsoid or sub-

Aiouea 703

globose 1-seeded berry, seated on a shallow cupule; tepals deciduous in fruiting stage. Southern Mexico, Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay; ca. 20 species, ca. 8 in Venezuela, 5 of these in the flora area. The genus, as here circumscribed, is not homogeneous. Some of the Andean and Central American species have little in common with the species found in Guayana and Brazil, but a better classification is currently not available. Key to the Species of Aiouea 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Fertile stamens 3 ....................................................................................... 2 Fertile stamens 6 ....................................................................................... 3 Connective of fertile stamens greatly prolonged and tepaloid beyond anther cells, the cells lateral-basal ........................................ A. myristicoides Connective of fertile stamens not prolonged, not tepaloid; the cells introrse-lateral ...................................................................... A. benthamiana Anther cells on outside of anthers opening toward the tepals .............. ................................................................................................ A. guianensis Anther cells on inner side of anthers opening toward center of flower ................................................................................................................ 4 Flowers glabrous; tepals ca. as long as the floral tube; floral tube not constricted ................................................................................. A. maguireana Flowers puberulous or pubescent; tepals ca. half as long as the floral tube; floral tube constricted at apex ....................................................... A. laevis

Aiouea benthamiana Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 48. 1889. Tree to 15 m tall. Black-water areas, 100– 600; Bolívar (Río Suapure), Amazonas (Río Casiquiare, Río Negro). Colombia (Vaupés), Brazil (Amazonas: Río Negro). The collections from Bolívar, which are all recent, have more pronounced venation on the upper surface of leaf, but are best placed here. Aiouea guianensis Aubl., Hist. Pl. Guiane 310. 1775. Tree or shrub. Montane forests, ca. 1000 m; Bolívar (Sierra de Pakaraima). TrinidadTobago, Guyana, Suriname, French Guiana, Brazil. Aiouea laevis (Mart.) Kosterm., Recueil Trav. Bot. Néerl. 35: 84. 1938. —Cryptocarya laevis Mart., Flora 21(2) Beibl.: 64, 1838. Tree to 15 m tall. Evergreen lowland forests, 100 m; Bolívar (between El Imperial and Montaña de Araiba). Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil.

Collected only once in the flora area, this species is close to Aiouea guianensis Aubl., which differs only in having extrorse anther cells. Aiouea maguireana (C.K. Allen) S.S. Renner in Kutitzki & S.S. Renner, Fl. Neotrop. Monogr. 31: 103. 1982. —Endlicheria maguireana C.K. Allen, Mem. New York Bot. Gard. 10(5): 68. 1964. Tree. Evergreen lowland and montane forests, 100–1100 m; Amazonas (base and slopes of Sierra de la Neblina, Río Yatúa, near San Carlos de Río Negro). Brazil (Amazonas). ŠFig. 602. Aiouea myristicoides Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 48. 1889. Aiouea pernitida C.K. Allen, Mem. New York Bot. Gard. 12(3): 102. 1965. Tree. Lower montane forests, 400–500 m; Bolívar (Río Blanco, tributary of Río Icabarú). Brazil (Amapá, Pará, Maranhão). The stamens with almost basal cells are characteristic.

704

L AURACEAE

Fig. 602. Aiouea maguireana

2. ANIBA Aubl., Hist. Pl. Guiane 327. 1775. by Henk van der Werff Trees, rarely shrubs. Wood yellow, sometimes fragrant. Leaves alternate or clustered at the tips of the branches, sometimes with a yellow-brown, papillose lower surface. Inflorescences axillary, often congested near tips of branches, usually many-flowered. Flowers bisexual, small; tepals 6, equal, rarely unequal, erect at anthesis. Stamens usually 9, all 2-celled, the filaments often wooly pubescent, the anthers glabrous; rarely only the 6 outer stamens fertile; staminodes present or absent. Berry 1-seeded, about 1/3 included in the often warty cupule. Neotropics; ca. 40 species, 17 in Venezuela, 15 of these in the flora area. Key to the Species of Aniba 1. 1. 2(1). 2.

Leaves evenly distributed along branches ................................................ 2 Leaves clustered at tips of branches ......................................................... 9 Lower surface of leaf minutely papillose, with an opaque, yellow-brown cast ......................................................................................................... 3 Lower surface of leaf smooth, without an opaque, yellow-brown cast ..... 6

Aniba 705

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

8.

9(1). 9. 10(9). 10. 11(10). 11. 12(11).

12.

13(9). 13. 14(13).

14.

Filaments of stamens narrower than the anthers; connective projecting beyond anther cells, the cells lateral-introrse ........................ A. burchellii Filaments of stamens about as wide as the anthers; connective not extending beyond anther cells, the cells introrse ..................................... 4 Flowers 3–3.5 mm long; leaves coriaceous, margins inrolled ..................... ....................................................................................... A. cinnamomiflora Flowers 1.5–2 mm long; leaves coriaceous or chartaceous, margins inrolled or not ......................................................................................... 5 Leaves coriaceous, the margins inrolled; upper surface of leaf usually shiny; in forests on white sand .............................................. A. rosaeodora Leaves chartaceous, the margins plane; upper surface of leaf usually opaque; nonflooded forests on sand or clay ........................... A. panurensis Outer tepals smaller than inner ones, usually about half the size of the inner ones ................................................................................... A. canelilla Tepals equal or at most slightly subequal ................................................. 7 Fertile stamens 6, representing the 2 outermost whorls ............. A. kappleri Fertile stamens 9, representing the 3 outer whorls ................................. 8 Flowers sparsely puberulous, the floral tube smooth outside, not longitudinally wrinkled; flowers abruptly widened above the floral tube, tepals ovate-orbicular, longer than the floral tube ................... A. citrifolia Flowers densely tomentellous, the floral tube outside with 6 longitudinal wrinkles at anthesis (disappearing in young fruit); flowers not abruptly widened above the floral tube; tepals ovate, shorter than the floral tube ....................................................................................................... A. affinis Leaves glabrous below or with few appressed hairs along midvein ...... 10 Leaves pubescent or minutely puberulent below, the pubescence not restricted to veins, but present on lamina as well ................................. 13 Outer tepals smaller than inner ones; leaves gradually narrowed toward base, at base abruptly rounded ............................................ A. megaphylla Tepals equal; leaf base not abruptly narrowed, but acute, angustate, or cuneate ................................................................................................. 11 Leaf base cuneate, gradually narrowed into the petiole; filaments of stamens as wide as the anthers ................................................. A. guianensis Leaf base acute, not cuneate; filaments of stamens narrower than the anthers ..................................................................................................... 12 Flowers sparsely puberulous, the floral tube smooth outside, not longitudinally wrinkled; flowers abruptly widened above the floral tube, tepals ovate-orbicular, longer than the floral tube ................... A. citrifolia Flowers densely tomentellous, the floral tube outside with 6 longitudinal wrinkles at anthesis (disappearing in young fruit); flowers not abruptly widened above the floral tube; tepals ovate, shorter than floral tube ....................................................................................................... A. affinis Pubescence on lower surface of leaf consisting of mostly erect hairs; these sometimes sparse ................................................................................. 14 Pubescence on lower surface of leaf fine and appressed, mostly covering the lamina surface ............................................................................... 15 Petioles swollen toward base, becoming gray-corky with age; twigs 3–4 mm thick, gray-corky; pubescence on lower surface of leaf rather sparse; tertiary venation on lower surface of leaf raised ................ A. taubertiana Petioles not swollen toward base, rufous-tomentellous; twigs 6–7 mm

706

L AURACEAE

thick, ferruginous-tomentellous, not gray-corky; pubescence on lower surface of leaf denser; tertiary venation raised ..................... A. ferruginea 15(13). Young twigs and petioles densely rufous-tomentose; cupules verrucose ................................................................................................. A. williamsii 15. Young twigs and petioles not densely rufous tomentose; cupules various .............................................................................................................. 16 16(15). Leaf base acute; leaves 3.5–5.5 times longer than wide; lateral veins 11– 17 per side .................................................................................... A. excelsa 16. Leaf base abruptly rounded; leaves 2.5–3.5 times longer than wide; lateral veins 15–25 per side ................................................. A. hostmanniana Aniba affinis (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 63. 1889. —Aydendron affine Meisn. in A. DC., Prodr. 15(1): 92. 1864. Small tree. Evergreen lowland forests, 100–200 m; Amazonas (between Tamatama and Caño Manaviche along Río Orinoco). Brazil (Amazonas: Rio Negro basin). Aniba burchellii Kosterm., Recueil Trav. Bot. Néerl. 35: 895. 1938. Shrub or tree to 25 m tall. Forests on sandy soil, 100–200 m; Amazonas (Río Mawarinuma at base of Sierra de la Neblina). Amazonian Brazil (Acre, Amazonas, Minas Gerais, Pará). The protruding connectives of the anthers and the narrow filaments are the best characters for this species. Aniba canelilla (H.B.K.) Mez., Jahrb. Königl. Bot. Gart. Berlin 5: 53. 1889. —Cryptocarya canelilla H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 192, t. 645. 1825. Tree to 30 m tall. Evergreen lowland forests, 100–400 m; Bolívar (Urimán), Amazonas (La Esmeralda). Guyana, Peru, Brazil, Bolivia. Aniba cinnamomiflora C.K. Allen, Mem. New York Bot. Gard. 10(5): 49. 1964. Shrub or small tree. Montane forests, 1500–1800 m; Amazonas (upper slopes of Cerro Aracamuni and Cerro Sipapo). Lowland forests in Apure, Aragua, and Mérida (some of these records might be erroneous). Aniba citrifolia (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 74. 1889. —Aydendron citrifolium Nees, Syst. Laur. 257. 1836. Aydendron trinitatis Meisn. in A. DC., Prodr. 15(1): 91. 1864. —Aniba trinitatis

(Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 68. 1889. Tree to 30 m tall. Evergreen lowland and lower montane forests, 100–900 m; Delta Amacuro (El Palmar, near border with Bolívar), Bolívar (upper Río Caroní, Santa Elena de Uairén). Trinidad, Guyana, French Guiana, Brazil (Amapá, Amazonas, Maranhão, Pará). ŠFig. 603. Aniba excelsa Kosterm., Recueil Trav. Bot. Néerl. 35: 887. 1938. Tree to 30 m tall. Evergreen lowland forests, 100–300 m; Delta Amacuro (El Palmar), Bolívar (El Paraíso on San Félix to Upata road, near Upata). Guyana. ŠFig. 604. A Venezuelan collection was cited by Allen as a paratype of Aniba murçana C.K. Allen. Kubitzki included it (as well as a paratype from Guyana) in Aniba excelsa and placed the Brazilian holotype and a paratype in Aniba williamsii O.C. Schmidt, a species with a denser, longer pubescence on the lower surface of leaf. Kubitzki’s treatment is followed here. Aniba ferruginea Kubitzki in Kubitzki & S.S. Renner, Fl. Neotrop. Monogr. 31: 32. 1982. Tree to 16 m tall. Nonflooded forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Endemic. Aniba guianensis Aubl., Hist. Pl. Guiane 327. 1775. Tree to 25 m tall. Riparian forests, 50–100 m; Amazonas (9 km from Puerto Ayacucho). Zulia; Colombia, Suriname, French Guiana, Peru, Brazil. ŠFig. 606. The only collection known from the flora area is in fruit and the identification is not certain.

Aniba 707

Fig. 603. Aniba citrifolia

Fig. 604. Aniba excelsa

708

L AURACEAE

Fig. 605. Aniba hostmanniana

Fig. 606. Aniba guianensis

Beilschmiedia 709

Aniba hostmanniana (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 67. 1889. —Aydendron hostmannianum Nees, Linnaea 21: 499. 1848. Aniba pittieri O.C. Schmidt, Repert. Spec. Nov. Regni Veg. 31: 169. 1937. Tree to 20 m tall. Seasonally flooded forests, 100–200 m; Amazonas (Río Baría area). Aragua, Distrito Federal, Yaracuy; Guyana, Suriname, French Guiana, Peru, Brazil. ŠFig. 605. Aniba kappleri Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 52. 1889. Aniba riparia auctt. non (Nees) Mez.: sensu C.K. Allen, Mem. New York Bot. Gard. 15: 58. 1966. Tree to 25 tall. Evergreen lowland forests, 100–200 m; Delta Amacuro (El Palmar, Río Cuyubini). Guyana, Suriname, French Guiana, Brazil. Aniba kappleri hybridizes with A. riparia (Nees) Mez. Hybrids have been reported by Kubitzki from Delta Amacuro (El Palmar); these have a variable number of anther cells in the stamens of the third whorl, even within the same flower. Aniba megaphylla Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 67. 1889. Small tree. Lower montane forests, 100– 500 m; Amazonas (base of Cerro Duida), Bolívar (Sierra de Maigualida). Guyana, Suriname, French Guiana, Peru, Brazil. Aniba panurensis (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 58. 1889. —Aydendron panurense Meisn. in A. DC., Prodr. 15(1): 89. 1864. —Laurel amarillo.

Tree to 10 m tall. Evergreen lowland forests, 100–400 m; Bolívar (east of El Crucero, Río Caroní, Río Gavilán east of Puerto Ayacucho, Río Parguaza, Río Suapure), Amazonas (Isla Ratón, Río Pasimoni, San Carlos de Río Negro, base of Sierra de la Neblina). Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Aniba rosaeodora Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 109. 1930. Fragrant tree to 25 m tall. Evergreen lowland forests, 100–400 m; Amazonas (Cañon Grande along Río Mawarinuma, San Carlos de Río Negro, Sierra de la Neblina). Guyana, Suriname, French Guiana, Peru, Brazil. This species is not always easy to separate from Aniba panurensis. Aniba taubertiana Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 65. 1889. —Laurel amarillo, Tau-wan-yek (Arekuna). Aniba simulans C.K. Allen, Mem. New York Bot. Gard. 10(5): 51. 1964. Tree to 20 m tall. Evergreen lowland to montane forests, 100–1200 m; Bolívar (northeast of El Paují, near Kavanayén along Río Acanán, La Escalera, base of Ptarítepui), Amazonas (base of Sierra de la Neblina, Yavita to Maroa road). Colombia, Suriname, French Guiana, Brazil. Aniba williamsii O.C. Schmidt, Repert. Spec. Nov. Regni Veg. 31: 169. 1933. Large tree. Evergreen lowland forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Suriname, French Guiana, Ecuador, Peru, Brazil.

3. BEILSCHMIEDIA Nees, Pl. Asiat. Rar. 2: 69. 1831. by Henk van der Werff Trees. Leaves alternate or subopposite near tips of branches. Inflorescences axillary, paniculate. Flowers bisexual. Tepals 6, equal, erect at anthesis, united at the base and falling as a unit in old flowers. Stamens usually 9, all 2-celled, the outer 6 with introrse cells, the inner 3 with extrorse cells; staminodia 3. Fruit a 1-seeded berry, seated on a woody, slightly swollen pedicel; tepals not persistent in fruiting stage. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, French Guiana, Brazil, most diverse in tropical Africa and Asia; ca. 100 species, 2 or 3 in Venezuela, 1 of these in the flora area.

710

L AURACEAE

Fig. 607. Beilschmiedia curviramea

Beilschmiedia curviramea (Meisn.) Kosterm., Recueil Trav. Bot. Néerl. 35: 853. 1938. —Aydendron curvirameum Meisn. in A. DC., Prodr. 15(1): 90. 1864. —Aguacatillo moisés.

Tree to 45 m tall; flowers yellow-green; fruit to ca. 5 cm long. Evergreen lowland forests, 100–600 m; Delta Amacuro (near El Palmar), Bolívar (El Tigre, La Soledad). Guyana. ŠFig. 607.

4. CASSYTHA L., Sp. Pl. 35. 1753. by Henk van der Werff Parasitic, herbaceous vines, with small haustoria; stems filiform, containing chlorophyll. Leaves reduced to minute scales. Flowers sessile or pedicellate, in heads or in spicate or racemose inflorescences; floral tube shallow; tepals 6, the outer 3 smaller than the inner 3. Fertile stamens 9, those of the third whorl with 2 basal glands, fourth whorl reduced to staminodia; anthers 2-celled. Fruit enclosed in the floral tube, tepals persisting. Pantropics (mostly Australia), extending into subtropics; ca. 20 species, 1 in Venezuela. Cassytha filiformis L., Sp. Pl. 35. 1753. —Bejuco de oro. Vining parasite, similar to Cuscuta. Moist savannas, often near Mauritia palm swamps, disturbed areas, 0–1300 m; Delta Amacuro (Pedernales), Bolívar (widespread), Amazonas (Río Atabapo, Río Negro). Widespread in northern Venezuela; pantropics, extending into subtropics. ŠFig. 608.

Fig. 608. Cassytha filiformis

Cinnamomum 711

5. CINNAMOMUM Schaeff., Bot. Exped. 268. 1760. by Henk van der Werff Trees or shrubs. Leaves simple, entire, alternate, chartaceous or coriaceous, usually tripliveined, pubescence variable. Inflorescences axillary or clustered on leafless spurs, shorter or longer than the leaves, paniculate, usually many-flowered. Flowers usually greenish to yellowish, small, sometimes with unpleasant odor. Tepals 6, equal, erect at anthesis, rarely spreading. Stamens 9, the outer 6 with introrse cells, 4-locellate; the inner 3 with extrorse cells, 4-locellate or infrequently 2celled; staminodia 3, with a cordate or sagittate apex. Floral tube shallow. Ovary superior, glabrous, 1-ovulate. Fruit a round, 1-seeded berry, subtended by a small cupule, the tepals deciduous or (partly) persistent. Central America, West Indies, Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 150 species, 2 in Venezuela, both in the flora area. Cinnamomum is a poorly understood genus, and the description given above applies only to the neotropical species. Possibly the genus is polyphyletic. Confusion (in the New World) with Persea and Ocotea makes it difficult to give the exact distribution and number of species, but the highest diversity is in Central America and southern Brazil. Cinnamomum is poorly represented in northern South America. The neotropical species have frequently been placed in Phoebe, but seem closer to Cinnamomum. Key to the Species of Cinnamomum 1. 1.

Leaves obtuse, pinnately veined; axillary tufts of hairs lacking ................ ....................................................................................... C. semecarpifolium Leaves acute, tripliveined; lowermost lateral veins with an axillary tuft of hairs ........................................................................................ C. triplinerve

Fig. 609. Cinnamomum semecarpifolium

712

L AURACEAE

Cinnamomum semecarpifolium (Meisn.) Kosterm., Reinwardtia 6: 23. 1961. —Oreodaphne semecarpifolia Meisn. in A. DC., Prodr. 15(1): 120. 1864. —Phoebe semecarpifolia (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 217. 1889. —Persea sprucei Kosterm., Reinwardtia 7: 143. 1965. Large tree. Evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, Río Pasimoni, Sierra de la Neblina). Endemic. ŠFig. 609. The generic placement of this distinct species is uncertain. It is quite unlike any other Cinnamomum species. The common names cited by Mez (Paraturi, Itauba) have been applied to Mezilaurus species, but the flowers are different from that genus. Additional flowering and fruiting material is needed. The wood of this species is prized for making canoes.

Cinnamomum triplinerve (Ruiz & Pav.) Kosterm., Reinwardtia 6: 24. 1961. —Laurus triplinervis Ruiz & Pav., Fl. Peruv. 4: t. 369. 1804. Persea cinnamonifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 160. 1817. —Phoebe cinnamonifolia (H.B.K.) Nees, Linnaea 21: 488. 1848. Phoebe filamentosa C.K. Allen, Mem. New York Bot. Gard. 15: 69. 1966. —Cinnamomum filamentosum (C.K. Allen) Kosterm., Reinwardtia 7: 461. 1969. Tree to 15 m tall. Semideciduous forests, 100–200 m; Delta Amacuro (Serranía de Imataca). Anzoátegui, Apure, Barinas, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Central America, West Indies, South America.

6. ENDLICHERIA Nees, Linnaea 8: 37. 1833. by Henk van der Werff Trees or shrubs. Leaves alternate or whorled, entire. Inflorescence axillary, paniculate, usually many-flowered. Flowers unisexual (plants dioecious); tepals 6, in 2 rows of 3, equal, often reflexed or fully spreading at anthesis. Staminate flowers with 9 2-celled stamens (except E. anomala), the 6 exterior with introrse cells, the 3 interior with extrorse cells and often longer than the 6 exterior; 3 interior stamens with 2 glands near the base of the filaments; staminodia lacking; pistillode present. Pistillate flowers with 9 staminodia, the glands of the inner 3 stamens rather conspicuous, the stigma large. Fruit a 1-seeded berry; cupule usually cup-shaped, rarely plate-like or almost lacking. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 40 species, ca. 18 in Venezuela, 15 of these in the flora area. Key to the Species of Endlicheria 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

Leaves whorled; leaf scars clustered along stems .................................... 2 Leaves evenly distributed along twigs; leaf scars not clustered .............. 7 Lower surface of leaves glabrous , often glaucous; inflorescences and flowers glabrous ........................................................................... E. arunciflora Lower surface of leaves pubescent; inflorescences and/or flowers not glabrous ....................................................................................................... 3 Cupules ca. 2 × 2 cm, densely golden-brown-pubescent; young twigs golden-brown-tomentose ........................................................... E. chalisea Cupules ca. 1 × 1 cm, glabrous or nearly so; young twigs pale or darkbrown-tomentose, never golden-brown-tomentose ............................... 4 Petioles 2–5 cm long; secondary veins departing from midrib at ca. 90° or leaves pruinose-glaucous on lower surface ........................................... 5 Petioles 0.5–1.5(–2) cm long; secondary veins departing from midrib at

Endlicheria 713

5(4).

5.

6(4). 6. 7(1). 7. 8(7). 8. 9(8). 9. 10(9). 10. 11(10). 11. 12(10). 12.

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

acute angles; lower surface of leaves not glaucous-pruinose ............... 6 Secondary veins departing from midrib at 90° angle; lower surface of leaves not glaucous-pruinose; inflorescence bracts persisting at anthesis; floral tube not constricted at apex; flowers densely pilose outside ................................................................................ E. directonervia Secondary veins departing from midrib at acute angle; lower surface of leaves glaucous-pruinose; inflorescence bracts not persisting at anthesis; floral tube constricted at apex; flowers loosely strigose outside ............................................................................................. E. macrophylla Petioles 1–1.5(–2) cm long; base of leaf acute ............................ E. multiflora Petioles 0.5(–1) cm long; base of leaf rounded to cordate ...... E. dictifarinosa Inner 3 stamens with 4-celled anthers; fruits subtended by a thickened pedicel with the bases of the tepals persisting ........................ E. anomala All stamens with 2-celled anthers; fruits subtended by a variously shaped cupule ..................................................................................................... 8 Leaves with shiny, sericeous pubescence on lower surface (check also E. anomala); leaves subtripliveined ......................................... E. bracteolata Leaves without shiny, sericeous pubescence; leaves pinnately veined ................................................................................................................ 9 Twigs and leaves glabrous or nearly so; cupule a slightly conical plate, to 4 cm diameter ......................................................................... E. rubriflora Twigs and leaves variously pubescent; cupule cup-shaped, or almost plate-like but then less than 1 cm diameter ....................................... 10 Pubescence on lower surface of leaf consisting of straight, appressed hairs (rather sparse in E. gracilis) ............................................................... 11 Pubescence on lower surface of leaf consisting of straight, erect hairs, or curled, ± erect hairs ............................................................................. 12 Leaves elliptic; reticulation raised on lower surface of leaf; tall tree to 30 m .................................................................................................... E. nilssonii Leaves ovate; reticulation not raised; shrub or small tree ............ E. gracilis Tepals of staminate flowers at anthesis erect, the stamens not readily visible; inner surface of tepals glabrous ......................... E. aff. szyszylowiczii Tepals of staminate flowers at anthesis spreading or reflexed, the stamens readily visible; inner surface of tepals glabrous or pubescent .............................................................................................................. 13 Inner surface of tepals glabrous; shrub on granitic outcrops along the Río Orinoco ..................................................................................... E. reflectens Inner surface of tepals completely covered by pubescence; not known from granitic outcrops along the Río Orinoco .............................................. 14 Outer stamens with short filaments, these much narrower than the anthers; trees to 25 m tall ........................................................... E. canescens Outer stamens with filaments as wide as or wider than anthers; (clambering) shrubs .............................................................................. E. vinotincta

Endlicheria anomala (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 133. 1889. —Goeppertia anomala Nees, Syst. Laur. 370. 1836. —Laurel de rebalse, Laurel orillero. Ocotea simulans C.K. Allen, Mem. New York Bot. Gard. 10(5): 99. 1964. Tree to 20 m tall; flowers reddish yellow

or reddish green (an unusual color for Lauraceae). Riparian forests, 100–400 m; Bolívar (Río Parguaza), Amazonas (Puerto Ayacucho, Río Casiquiare, Río Cataniapo, Río Siapa, Río Sipapo, San Carlos de Río Negro). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

714

L AURACEAE

Endlicheria arunciflora (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 131. 1889. —Ampelodaphne arunciflora Meisn. in A. DC., Prodr. 15(1): 81. 1864. Small tree to 6 m tall. Flooded lowland forests, along black-water streams, 100–200 m; Amazonas (upper Río Orinoco and Río Negro and their tributaries). Peru, Brazil. Two collections from along the Río Negro (fruiting and sterile) probably represent an undescribed, related species. It has narrower leaves and more lateral veins than Endlicheria arunciflora, but the material is not sufficient to allow a description. Endlicheria bracteolata (Meisn.) C.K. Allen, Mem. New York Bot. Gard. 10(5): 64. 1964. —Goeppertia sericea var. bracteolata Meisn. in A. DC., Prodr. 15(1): 174. 1864. —Laurel rebalsero. Tree to 10 m tall. Riparian forests, 100– 800 m; Bolívar (Macizo del Chimantá [Chimantá-tepui], upper Río Caura), common in Amazonas. Guyana, Suriname, Brazil. Endlicheria canescens Chanderbali, Novon 6: 328. 1996. Tree to 35 m tall. Montane forests, ca. 1200 m; Bolívar (between San Ignacio and San Francisco). Guyana, Suriname, Ecuador, Peru. Endlicheria chalisea Chanderbali, Novon 6: 329. 1996. Tree ca. 8 m tall. Evergreen lowland forests, 100–200 m; Amazonas (Río Atacavi, upper Río Casiquiare, San Carlos de Río Negro). Guyana, French Guiana, Peru, Brazil, Bolivia. Endlicheria directonervia C.K. Allen, Mem. New York Bot. Gard. 10(5): 60. 1964. Small tree to 10 m tall. Nonflooded lowland forests, 100–200 m; Amazonas (base of Cerro Sipapo, Yavita). Endemic. Endlicheria directonervia belongs to a widely distributed complex in need of study. The flora area specimens had been annotated as Endlicheria tessmannii O.C. Schmidt, a Peruvian species, which has a quite different indument and, pending further study, E. directonervia is maintained here as a distinct species.

Endlicheria dictifarinosa C.K. Allen, Mem. New York Bot. Gard. 12(3): 108. 1965. —Laurel rebalsero. Tree to 20 m tall, but mostly smaller. Riparian forests, ca. 100–300 m; Bolívar (Cedeño, Río Caura), Amazonas (Río Cunucunuma). Endemic. Most collections of this species had been identified as Endlicheria multiflora, which is here considered a distinct species. Allen (Mem. New York Bot. Gard. 10(5): 59. 1964) named specimens now included here E. cocuirey Kosterm., a Peruvian species close to E. dictifarinosa, but with a different indument. In 1965, when describing E. dictifarinosa, she used the name E. cocuirey for material here included in E. multiflora (Miq.) Mez. Endlicheria gracilis Kosterm., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 42: 528. 1937. Small tree or shrub. Savannas, semideciduous forests, 100–1100 m; Bolívar (Kavanayén area), Amazonas (Culebra). Colombia, Suriname. Endlicheria macrophylla (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 128. 1889. —Ampelodaphne macrophylla Meisn. in A. DC., Prodr. 15(1): 81. 1864. Shrub or small tree to 5 m tall. Shrub islands in wet savannas, 100–200 m; Amazonas (near Cerro Cucurito and Cerro Yapacana). Brazil. Endlicheria macrophylla is close to E. multiflora, but differs in having broader leaves with a truncate or rounded base and in having the leaves pruinose or glaucous below. Endlicheria multiflora (Miq.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 130. 1889. —Goeppertia multiflora Miq., Natuurk. Verh. Holl. Maatsch. Wetensch. Haarlem 7: 203. 1851. —Laurel de babilla. Endlicheria levelii C.K. Allen, Mem. New York Bot. Gard. 10(5): 62. 1964. Tree to 10 m tall. Seasonally flooded forests, 100–200 m; scattered in Delta Amacuro, Bolívar, and Amazonas. Guyana, Suriname, French Guiana, Brazil. ŠFig. 610. Endlicheria nilssonii C.K. Allen, Mem. New York Bot. Gard. 10(5): 66. 1964.

Endlicheria 715

Fig. 610. Endlicheria multiflora

716

L AURACEAE

Tree to 20 m tall. Montane forests, ca. 1200 m; Bolívar (between Luepa and Cerro Venamo). Endemic. More material is needed to ascertain the relationships and generic placement of this species.

Small tree. Evergreen lowland forests, 100–200 m; Amazonas (Río Siapa). Apure; Colombia, Peru, Brazil.

Endlicheria reflectens (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 126. 1889. —Goeppertia reflectens Nees, Linnaea 21: 514. 1848. Shrub or small tree. Granitic outcrops, 100–300 m; Bolívar (near Agua Amena 265 km southwest of Caicara). Colombia (across the Río Orinoco from Puerto Ayacucho), Guyana, French Guiana, Brazil.

Endlicheria aff. szyszylowiczii Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 121. 1889. Tree. Evergreen lowland forests, 100–200 m; Amazonas (along river near base camp at Sierra de la Neblina). Peru, Brazil. The flora area collection is included in Endlicheria szyszylowiczii with hesitation; the type is from the mountains in northern Peru, and the few lowland collections have a slightly different indument. More collections are needed.

Endlicheria rubriflora Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 494. 1889. Endlicheria wurdackiana C.K. Allen, Mem. New York Bot. Gard. 10(5): 67. 1964.

Endlicheria vinotincta C.K. Allen, Mem. New York Bot. Garden 10(5): 63. 1964. Shrub to 2 m tall. Open forests, tepui slopes, 500–1600 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Endemic.

7. KUBITZKIA van der Werff, Taxon 35: 165. 1986. Systemonodaphne Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 78. 1889, not as to type. by Henk van der Werff Small tree. Leaves alternate, simple. Inflorescences axillary, few-flowered; flowers reddish. Tepals 6, equal, pubescent on the inner surface. Stamens 9, all 2-

Fig. 611. Kubitzkia macrantha

Licaria 717

celled, the outer 6 with slender, densely pubescent filaments, the cells nearly apical and lateral-introrse; inner 3 stamens with the filaments fused into a column. Berry subtended by a cup-shaped cupule; tepals persistent and enlarged, becoming reflexed, the cupule thus with a double margin; stamens persisting on the cupule. Venezuela, Guyana, Suriname, French Guiana, Brazil (to south of Rio Amazon); 1 or 2 species, 1 in Venezuela. Kubitzkia macrantha (Kosterm.) van der Werff, Taxon 35: 165. 1986. —Systemonodaphne macrantha Kosterm., Bol. Tecn. Inst. Agron. N. 28: 73. 1954. Tree to 12 m tall. Upland forests, 800–900 m; Bolívar (near El Paují). Amazonian Brazil. ŠFig. 611. The record by C.K. Allen of Systemono-

daphne mezii Kosterm. [= Kubitzkia mezii (Kosterm.) van der Werff], based on a sterile Steyermark collection from Río Cuyubini, Delta Amacuro, is based on a misidentification. The differences between K. macrantha and K. mezii are weak and the two may turn out to be synonymous once more material becomes available.

8. LICARIA Aubl., Hist. Pl. Guiane 313. 1775. by Henk van der Werff Shrubs or medium-sized, rarely tall, trees. Leaves alternate or opposite, rarely somewhat clustered at tips of branches, entire. Inflorescences paniculate (in flora area), sometimes by reduction racemose or capitate. Flowers bisexual; tepals 6, equal or unequal, at anthesis mostly erect or incurved. Stamens 3, 2-celled, with 2 glands at the base; locelli mostly apical and opening toward the style or away from it; staminodia present or absent. Fruit a 1-seeded berry; cupule usually distinctly cup-shaped and with a double margin. U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 40 species, ca. 13 in Venezuela, 12 of these in the flora area. Key to the Species of Licaria 1. 1. 2(1).

2.

3(1).

3.

4(3). 4.

Tepals reflexed at anthesis ........................................................................ 2 Tepals erect or incurved at anthesis .......................................................... 3 Inflorescences short, to 2 cm long; leaves obovate; stamens with extrorseapical locelli; pedicel of cupule to 1 cm long; in nonflooded forests ................................................................................................... L. vernicosa Inflorescences 2–8 cm long; leaves lanceolate or elliptic; stamens with locelli on the inner surface; pedicel of cupule 2–5 cm long; in flooded forests ..................................................................................... L. armeniaca Tepals unequal, the outer 3 smaller than the inner 3; locelli on outer surface of stamens, the valves attached at the base of the locelli; cupule shallow, the enlarged, persistent tepals reflexed, the stamens and outer staminodia persistent ............................................... L. macrophylla Tepals equal or slightly subequal; locelli apical, or nearly so, or on the inner surface, the valves usually attached near the tip of the locelli; cupule various, but never with persistent, reflexed tepals and persistent stamens and staminodia ........................................................................ 4 Leaves alternate, rarely somewhat clustered near tips of twigs ............. 5 Leaves opposite and evenly distributed along twigs; lower leaves sometimes alternate ....................................................................................... 8

718

L AURACEAE

5(4).

Twigs glabrous; anther locelli slit-like, at inner surface of anthers; leaves somewhat clustered at tips of branches, the tertiary venation raised; cupule large, to 4 cm long and 5 cm wide ................................. L. cannella 5. Twigs variously pubescent; anther locelli pore-like, apical or apical-extrorse; leaves not clustered; tertiary venation not or only slightly raised; cupule, when known, not more than 2 cm long ........................ 6 6(5). Young leaves fuscous-tomentose on lower surface; flowers wider than long, tomentose .......................................................................... L. trinervis 6. Young leaves with appressed pubescence; flowers as long as wide or longer than wide, the pubescence of fine, straight hairs ..................... 7 7(6). Flowers several times longer than wide; floral tube conspicuously constricted near the apex; leaves lanceolate ............................ L. dolichantha 7. Flowers nearly as long as wide; floral tube very slightly constricted; leaves elliptic ......................................................................... L. brasiliensis 8(4). Young twigs and lower surface of leaf ferruginous-tomentose .... L. tomentosa 8. Twigs and leaves with appressed pubescence to subglabrous .................. 9 9(8). Anthers at anthesis not exserted; tepals incurved or erect .................... 10 9. Anthers at anthesis exserted; tepals erect .............................................. 11 10(9). Tepals incurved at anthesis; flowers, pedicels, and lower surface of leaf densely puberulous, the hairs minute; flowers 2.5–3.5 mm long .......... ............................................................................................. L. chrysophylla 10. Tepals erect at anthesis; flowers, pedicels, and lower surface of leaf glabrous or with some coarse, reddish, appressed hairs; flowers 1–2 mm long ................................................................................................ L. debilis 11(9). Leaves acuminate, the acumen 1–2 cm long; lower surface of leaf with some appressed, rather coarse hairs; flowers strigose, 2–4 mm long ................................................................................................. L. polyphylla 11. Leaves acute; lower surface of leaf with minute appressed pubescence; flowers densely puberulous, 1.5–2 mm long ....................... L. oppositifolia Licaria armeniaca (Nees) Kosterm., Recueil Trav. Bot. Néerl. 34: 584. 1937. —Evonymodaphne armeniaca Nees, Syst. Laur. 264. 1836. Small tree to 5 m tall. Seasonally flooded forests, 100–200 m; Amazonas (Isla Hormiga, Río Casiquiare, San Carlos de Río Negro). Eastern Colombia, Guyana, Peru, Brazil. Licaria armeniaca is widespread in flooded forests in the Amazon basin.

Río Nichare basin), Amazonas (near Yutajé). Peru, Brazil. The Amazonas collection is placed here with some hesitation. The collection is in fruit, and fruiting specimens of Licaria brasiliensis had not been collected earlier. The rather large leaves (to 14 cm long), and the free stamens point to this species; the smooth cupule excludes L. guianensis. However, the papillae on the lower surface of the leaf are not well developed.

Licaria brasiliensis (Nees) Kosterm., Recueil Trav. Bot. Néerl. 34: 601. 1937. —Acrodiclidium brasiliense Nees, Pl. Laur. Expos. 13. 1833. Tree. Evergreen lowland and secondary forests, 100–200 m; Bolívar (Río Tabaro in

Licaria cannella (Meisn.) Kosterm., Recueil Trav. Bot. Néerl. 34: 538. 1937. —Aydendron cannella Meisn. in A. DC., Prodr. 15(1): 90. 1864. Tree to 20 m tall or more. Swamp forests along black-water rivers, 100–500 m; Delta

Licaria 719

Amacuro (Serranía de Imataca), Amazonas (Río Baría, Río Guainía, Río Negro). Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ŠFig. 612. Herbarium specimens always have a slightly revolute leaf margin. Licaria chrysophylla (Meisn.) Kosterm., Recueil Trav. Bot. Néerl. 34: 601. 1937. —Acrodiclidium chrysophyllum Meisn. in A. DC., Prodr. 15(1): 87. 1864. —Laurel marrón. Tree to 30 m tall. Evergreen lowland forests, 100–400 m; Bolívar (Río Caroní, Río Nichare basin, near Upata). Guyana, Suriname, French Guiana, adjacent Brazil. Licaria debilis (Mez) Kosterm., Recueil Trav. Bot. Néerl. 34: 596. 1937. —Acrodiclidium debile Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 99. 1899. Tree to 15 m tall. Evergreen lowland forests, 0–200 m; Delta Amacuro (Río Amacuro, upstream from San Victor, Serranía de Imataca). Sucre; Guyana, Suriname, French Guiana, Brazil (Pará). Licaria dolichantha Kurz, Ph.D. thesis Univ. Hamburg 158, 1983. Tree to 20 m tall. Evergreen lowland forests, 50–100 m; Amazonas (Isla Ratón). Brazil (Amazonas: vicinity of Manaus). Licaria macrophylla (A.C. Sm.) Kosterm., Recueil Trav. Bot. Néerl. 34: 583. 1937. —Acrodiclidium macrophyllum A.C. Sm., Bull. Torrey Bot. Club 58: 101. 1931. Licaria schultesii C.K. Allen, Mem. New York Bot. Gard. 12(3): 104. 1965. Tree to 20 m tall; cupules shallow with reflexed tepals; petioles long (to 3.5 cm). Forested tepui slopes, ca. 600 m; Amazonas (Cerro Huachamacari). Peru (vicinity of Iquitos), Brazil (Amazonas: Río Vaupés). Licaria oppositifolia (Nees) Kosterm., Recueil Trav. Bot. Néerl. 34: 597. 1937. —Acrodiclidium oppositifolium Nees, Linnaea 21: 500. 1848. Licaria martiniana auct. non (Mez) Kosterm.: sensu C.K. Allen, Mem. New York Bot. Gard. 15: 64. 1966.

Tree to 25 m tall. Evergreen lowland forests, 100–300 m; Bolívar (south of El Dorado, north of El Palmar, Río Toro). Reported from Barinas; Colombia, Guyana, Brazil (Amazonas). ŠFig. 614. Licaria polyphylla (Nees) Kosterm., Recueil Trav. Bot. Néerl. 34: 584. 1937. —Nectandra polyphylla Nees, Syst. Laur. 322. 1836. —Aro de atarraya rebalsero, Laurel. Acrodiclidium guianense var. caudatum Meisn. in A. DC., Prodr. 15(1): 173. 1864. Acrodiclidium meissneri Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 98. 1889. —Licaria meissneri (Mez) Kosterm., Recueil Trav. Bot. Néerl. 34: 597. 1937. Small tree or shrub. Seasonally flooded forests, ca. 100 m; Amazonas (Río Negro). Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, Pará). ŠFig. 613. Two recent fruiting collections are placed here with some hesitation. The twigs are used for making net hoops, etc. Licaria tomentosa van der Werff, Ann. Missouri Bot. Gard. 76: 462. 1989. Ocotea roraimae auct. non Mez 1920: sensu C.K. Allen pro parte. Tree to 10 m tall; leaves opposite, ferruginous-tomentose. Forested tepui slopes, 1100– 1700 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Endemic. Licaria trinervis van der Werff, Ann. Missouri Bot. Gard. 76: 960. 1989. Shrub or small tree to 5 m tall; leaves alternate, rufous-tomentose. Savannas with patches of forests, 1400–1600 m; Amazonas (Cerro Aracamuni). Endemic. Licaria vernicosa (Mez) Kosterm., Recueil Trav. Bot. Néerl. 34: 604. 1937. —Ocotea vernicosa Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 331. 1889. —Laurel paraguito. Licaria simulans C.K. Allen, Mem. New York Bot. Gard. 10(5): 55. 1964. Small tree. Evergreen lowland forests, 100–200 m; Bolívar (near Kilómetro 88). Guyana, Suriname, French Guiana, Brazil (Pará, also reported from Bahia).

720

L AURACEAE

Fig. 612. Licaria cannella

Fig. 613. Licaria polyphylla

Fig. 614. Licaria oppositifolia

Mezilaurus 721

Fig. 615. Mezilaurus sprucei

Fig. 616. Mezilaurus lindaviana

722

L AURACEAE

9. MEZILAURUS Taub., Biol. Centralbl. 50: 21. 1892. by Henk van der Werff Shrubs to tall trees. Leaves alternate, congested at apex of branches; petioles often swollen at base. Inflorescence axillary, few- to many-flowered, forming a compound raceme or flowers clustered at tips of inflorescence branchlets. Tepals 6, equal or nearly so, small, scale-like, usually erect at anthesis. Fertile stamens 3, representing the third whorl, 2-celled; filaments free or connate; staminodia usually lacking, staminal glands present in one species. Fruit an ellipsoid berry, seated on a small, plate-like cupule. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia (mostly Amazon basin); 16 species, 3 in Venezuela, all in the flora area. Several species have very hard wood that is used for construction. Key to the Species of Mezilaurus 1. 1. 2(1). 2.

Leaves pubescent ...................................................................... M. lindaviana Leaves glabrous .......................................................................................... 2 Leaf apex acute; pedicels 10–15 mm long ...................................... M. sprucei Leaf apex obtuse or rounded; pedicels 2–4 mm long ...................... M. itauba

Mezilaurus itauba (Meisn.) Taubl. ex Mez, Arbeiten Königl. Bot. Gart. Breslau 1: 112. 1892. —Acrodiclidium itauba Meisn. in A. DC., Prodr. 15(1): 86. 1864. —Parature negro. Acrodiclidium anacardioides Meisn. in A. D.C., Prodr. 15(1): 86. 1864. —Mezilaurus anacardioides (Meisn.) Taub. ex Mez, Arbeiten Königl. Bot. Gart. Breslau 1: 112. 1892. Large tree to 35 m tall. Evergreen lowland forests, 100-200 m; Amazonas (Río Casiquiare, Río Negro, San Carlos de Río Negro, San Fernando de Atabapo). Suriname, French Guiana, Peru, Brazil, Bolivia. The wood is hard, heavy, but easy to work and is used for houseposts, boats, and doors. It is said to be very decay-resistant. Mezilaurus lindaviana Schwacke & Mez,

Arbeiten Königl. Bot. Gart. Breslau 1: 112. 1892. Mezilaurus wurdackiana C.K. Allen, Mem. New York Bot. Gard. 10(5): 56. 1964. Tree to 25 m tall; leaves pubescent. Semidecoduous forests, 100–200 m; Bolívar (La Vergareña). Guyana, Brazil. ŠFig. 616. Mezilaurus sprucei (Meisn.) Taub. ex Mez, Arbeiten Königl. Bot. Gart. Breslau 1: 112. 1892. —Acrodiclidium sprucei Meisn. in A. DC., Prodr. 15(1): 86. 1864. —Parature. Mezilaurus maguireana C.K. Allen, Mem. New York Bot. Gard. 10(5): 58. 1964. Tree to 20 m tall. Seasonally flooded riparian forests, 100–200 m; Amazonas (Río Casiquiare, Río Guianía, Río Mawarinuma, Río Negro, Río Orinoco). Peru, Brazil. ŠFig. 615. The trunks are used for making canoes.

10. NECTANDRA Rol. ex Rottb., Acta Lit. Univ. Hafn. 1: 279 1778, nom. cons. by Jens G. Rohwer Trees or rarely shrubs. Leaves alternate (rarely opposite, but not in the flora area). Inflorescence axillary to foliage leaves, rarely to cataphylls, thryrsoid, usually many-flowered. Flowers bisexual; tepals 6, in 2 rows of 3, equal or the inner ones slightly smaller, papillose on inside, ± spreading at anthesis. Stamens 9, ± papillose, with 4 separate pollen sacs (rarely the lower pollen sacs ± lateral), the 3 interior with all pollen sacs extrorse or with the upper pair lateral, with 2 glands at the base; sta-

Nectandra 723

minodia subulate, clavate or slightly capitate, often glandular on inside, free or ± united with the filaments of the inner stamens. Receptacular tube shallow to deeply urceolate, pistil well developed, style distinct to absent, stigma conspicuous. Fruit a 1-seeded berry, globose to rather elongate; cupule shallowly bowl-shaped to deeply cup-shaped. U.S.A. (Florida), central Mexico, Central America, West Indies, Trinidad-Tobago, South America except Chile; 114 species, 20 in Venezuela, 12 of these in the flora area. Key to the Species of Nectandra 1. 1.

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

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

5.

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

7. 8(5).

8.

Indument on lower leaf surface predominantly ± erect ........................... 2 Indument on lower leaf surface either extremely sparse or predominantly appressed; occasionally with some ± erect trichomes in addition to a dense cover of appressed trichomes ...................................................... 3 Leaf base with large reflexed lobes; flowers large, (6.5–)8–14(–16) mm diameter .................................................................................. N. reticulata Leaf base without reflexed lobes; flowers small, 2.5–4(–5.5) mm diameter ...................................................................................................... N. pearcei Anthers not prolonged beyond the pollen sacs, in all three whorls apically rounded to slightly emarginate, with distinct filaments ...................... 4 Anthers distinctly prolonged beyond the pollen sacs, especially in the second whorl with a ± triangular tip (if anther apically ± rounded, then filament extremely short) ...................................................................... 5 Young twigs covered with extremely short, erect, brown trichomes; style distinct, about as long as the ovary ........................................ N. cuspidata Young twigs covered with appressed white trichomes; style very short ........................................................................................... N. martinicensis Style short, usually reaching < 40% the length of the ovary (if almost half as long as the ovary, then young twigs with an inconspicuous grayish indument, if any) ................................................................................... 6 Style long, usually reaching > 60% the length of the ovary (if ca. half as long as the ovary, then young twigs with a conspicuous reddish indument ................................................................................................ 8 Indument on young twigs persistent as a ± continuous grayish cover ...................................................................................................... N. lineata Indument on young twigs becoming sparse within a few centimeters from the terminal bud .................................................................................... 7 Leaves with 6–9 pairs of secondary veins, their axils on lower leaf surface (within the area) usually ± barbellate; stamens 1–1.4 mm long ........................................................................................................ N. hihua Leaves with 9–14 pairs of secondary veins, their axils on lower leaf surface glabrous; stamens 0.5–0.8 mm long .............................. N. sanguinea Young twigs with almost exclusively short trichomes, most of them < 0.1 mm long, very few (if any) longer than 0.3 mm, either forming a brownish to grayish cover, or not perceptible to the naked eye ..................... 9 Young twigs with both short and longer trichomes, many of them longer than 0.3 mm, at least partly curled, indumentum perceptible to the naked eye, yellowish to reddish ............................................................... 10

724

L AURACEAE

9(8).

Inflorescences and twigs with a dense indument (epidermis obscured); at least some mature leaves > 5.5 cm wide, rarely > 3.5 times longer than wide ............................................................................................. N. globosa 9. Inflorescences and twigs with a sparse to moderately dense indument (epidermis visible); mature leaves to 4.5 cm wide, rarely < 4 times longer than wide ...................................................................... N. pichurim 10(8). Lower surface of young leaves with a dense, ± golden indument .... N. aurea 10. Lower surface of young leaves ± sparsely hairy, indument inconspicuous .............................................................................................................. 11 11(10). Young leaves densely hairy on upper surface, indument on young twigs becoming sparse within a few centimeters from the terminal bud ......................................................................................................... N. fulva 11. Young leaves subglabrous on upper surface, indument on young twigs subpersistent ........................................................................ N. ruforamula Nectandra aurea Rohwer, Ann. Missouri Bot. Gard. 75: 1525. 1988. —Laurel negro. Small to medium-sized tree to 15 m tall; leaves narrow, smooth, with golden indument on lower surface. Gallery forests, 50– 300 m; Bolívar (Las Trincheras, Río Orinoco northeast of Puerto Paez, Temblador, 20 km east of Túriba). Apure. Nectandra cuspidata Nees, Syst. Laur. 330. 1836. —Guada-yek (Arekuna), Laurel amarillo, Laurel eucalipto, Laurelito, Laurel rastrojero. Nectandra olivacea Lasser, Bol. Técn. Minist. Agric. 3: 17. 1942. Shrub or tree to 30 m tall, capable of flowering when as little as 2 m tall; leaves lanceolate, with a fine acumen. Various habitats, frequently in secondary forests, 0–2000 m; Delta Amacuro (Río Amacuro, Río Toro), Bolívar (Cerro Marutaní, Perai-tepui, Río Nichare, Sierra de Lema), Amazonas (Isla Ratón, Mavaca, San Carlos de Río Negro, San Fernando de Atabapo). Apure, Barinas, Carabobo, Trujillo; southern Mexico, Central America, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay. Nectandra fulva Rohwer, Ann. Missouri Bot. Gard. 75: 1527. 1988. Tree to 20 m tall; leaves with gland dots. Evergreen lowland to lower montane forests, 100–200 m; Amazonas (base of Sierra de la Neblina near Río Mawarinuma). Endemic.

Nectandra globosa (Aubl.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 415. 1889. —Laurus globosa Aubl., Hist. Pl. Guiane 364. 1775. —Canau, Laurel blanco, Tabau-yek (Arekuna). Nectandra pisi Miq., Stirp. Surinam. Select. 199. 1850 [1851]. Nectandra vaga Meisn. in A. DC., Prodr. 15(1): 153. 1864, including varieties. Tree to 30 m tall. Riparian forests, lower montane forests, 0–800 m; Delta Amacuro (Caño Araguao, Río Amacuro), Bolívar (Cerro Marutaní, Río Venamo), Amazonas (Río Coro Coro, San Carlos de Río Negro). Guyana, Suriname, French Guiana, Brazil. ŠFig. 617. Nectandra hihua (Ruiz & Pav.) Rohwer, Fl. Neotrop. Monogr. 60: 196. 1993. —Laurus hihua Ruiz & Pav., Fl. Peruv. 4: t. 366. 1804. Nectandra willdenoviana Nees, Syst. Laur. 290. 1836. Nectandra bredemeyeriana Nees, Linnaea 21: 505. 1848. Nectandra leucantha var. attenuata Meisn. in A. DC., Prodr. 15(1): 151. 1864. Nectandra leucantha var. guianensis Meisn. in A. DC., Prodr. 15(1): 151. 1864. Tree to 30 m tall. Evergreen lowland and riparian forests, 50–200 m; Delta Amacuro (Río Toro), Amazonas (Río Orinoco below San Fernando de Atabapo). Barinas, Carabobo, Distrito Federal, Monagas, Portuguesa, Sucre, Yaracuy; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia.

Nectandra 725

Nectandra lineata (H.B.K.) Rohwer, Fl. Neotrop. Monogr. 60: 209. 1993. —Ocotea lineata H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 165. 1817. Tree to 30 m tall. Evergreen lowland or riparian forests, 50–100 m; Amazonas (San Fernando de Atabapo). Zulia; Central America, Colombia, Ecuador, northern Peru, possibly Bolivia. Although the type of this species is labeled as coming from San Fernando de Atabapo, it is odd that this is more than 500 km from the nearest confirmed locality. Nectandra martinicensis Mez, Mitt. Bot. Vereins Kreis Freiburg 47/48: 421. 1888. —Laurel blanco. Nectandra glandulifolia Lasser, Bol. Técn. Minist. Agric. 3: 16. 1942. Tree to 20 m tall. Semideciduous forests, 200–300 m; Bolívar (17 km east of El Callao, La Camilera). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Yaracuy, Zulia; Mexico, Central America, Colombia, Trinidad-Tobago, Ecuador. Nectandra pearcei Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 444. 1889. —Laurel verde. Nectandra kaburiensis Kosterm., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 25: 22. 1936. Tree to 30 m tall; leaves obovate, slightly bullate, somewhat glaucous on lower surface. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (Río Caura, Río Tabaro in Río Nichare basin, Santa Elena de Uairén, northeast of Upata). Anzoátegui, Apure; Colombia, Trinidad, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ŠFig. 618. Nectandra pearcei is very close to N. cissiflora Nees, which differs mainly by having an appressed indument. Nectandra pichurim (H.B.K.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 449. 1889. —Ocotea pichurim H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 166. 1817. —Laurel. Shrub or tree to 25 m tall; leaves narrow and elongate; cupules deeply campanulate; fruits elongate, 16–24 mm long, 6–8 mm di-

ameter. Gallery forests, swamp forests, 0– 300 m; Delta Amacuro (Boca Macareo, Caño Manamito, Río Grande), Bolívar (Río Nichare, Río Tonoro in Río Paragua basin), Amazonas (Río Casiquiare). Apure, Barinas, Cojedes, Guárico; Amazonian Colombia. Mez confused this species with Nectandra cuspidata, a species with similar leaves but very different flowers and cupules. Nectandra reticulata (Ruiz & Pav.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 404. 1889. —Laurus reticulata Ruiz & Pav., Fl. Peruv. 4: t. 348. 1804. Nectandra pittieri Lasser, Bol. Técn. Minist. Agric. 3: 13. 1942. Tree to 40 m tall. Evergreen lowland to montane forests, 100–1800 m; Bolívar (Río Nichare basin), Amazonas (known from Brazilian side of Sierra de la Neblina). Southern Mexico, central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Nectandra ruforamula Rohwer, Ann. Missouri Bot. Gard. 75: 1529. 1988. —Laurel de aparo. Small tree to 10 m tall; young twigs with reddish indument; upper surface of leaves grayish green with impressed reticulation, lower surface reddish brown with elevated reticulation. Evergreen lowland forests, 100–200 m; Amazonas (base of Cerro Sipapo, Cucurital de Caname, San Carlos de Río Negro, below San Fernado de Atabapo, Yavita to Pimichín road). Endemic. Nectandra sanguinea Rol. ex Rottb., Acta Lit. Univ. Hafn. 1: 279. 1778. Nectandra guianensis Meisn. in A. DC., Prodr. 15(1): 160. 1864. Tree to 25 m tall; leaves smooth, oblong, with numerous inconspicuous secondary veins. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (upper Río Caroní, upper Río Suapure, base of Roraimatepui, Serra do Sol along Brazilian border, near Túriba). Guyana, Suriname, Brazil. Nectandra sanguinea has long been misinterpreted in the sense of N. salicifolia (H.B.K.) Nees, which does not occur in South America.

726

L AURACEAE

Fig. 617. Nectandra globosa

Fig. 618. Nectandra pearcei

Ocotea 727

11. OCOTEA Aubl., Hist. Pl. Guiane 780. 1775. by Henk van der Werff Shrubs or trees, evergreen or rarely deciduous. Leaves alternate (rarely clustered at tips of branches), entire, mostly elliptic, but sometimes ovate or obovate, often with small gland dots, pinnately veined or rarely tripliveined, chartaceous to coriaceous. Inflorescences borne in the axils of leaves or deciduous bracts, sometimes the inflorescences grouped together at the leafless tips of branches and appearing terminal, but always with a terminal bud indicating that the inflorescences are lateral; inflorescence paniculate (rarely spicate), mostly many-flowered, 2–3 times branched, the flowers cymosely arranged. Flowers small, mostly white, green, or yellow, very rarely reddish, bisexual or unisexual (plants hermaphroditic or dioecious). Tepals 6, equal, at anthesis erect, spreading, or reflexed, ovate. Stamens in 4 whorls of 3, the outer 3 whorls functional, the inner whorl staminodial or lacking; filaments usually present, about as long as anthers, rarely the anthers subsessile; all anthers with 4 cells, these arranged in 2 horizontal rows and usually filling the entire anther; rarely anthers with a sterile tip; 2 outer whorls of stamens with cells introrse, inner whorl with extrorse cells or the upper pair lateral; inner 3 stamens with 2 glands at the base of the filament. Pistillate flowers with all stamens small and staminodial; in staminate flowers a well-developed pistillode may be present or lacking. Ovary (partly) enclosed in the floral tube. Fruit a 1-seeded berry, always subtended by a cupule. Cupule variable, from deeply cup-shaped to almost lacking, then with a thickened, lenticellate pedicel, or plate-like; margin usually single (rarely double-rimmed) and entire (sometimes lobed or toothed due to persistent tepals). Neotropics, tropical Africa, Madagascar, 1 species in Canary Islands; ca. 400 species, 70–80 in Venezuela, 56 of these in the flora area. Several species are highly valued for their wood. Ocotea rodiei (R.H. Schomb.) Mez., which was reported by Pittier et al. (Catalogo de la Flora Venezolana 1: 311. 1945) is excluded. Pittier lists this species as collected by Schomburgk on Roraima-tepui. However, the Schomburgk collection was made farther east, in Guyana, and specimens from the flora area were not seen for this study. The following key to the species is primarily based on floral characters and is of little help with the identification of fruiting specimens. Because some species have striking fruit characters, an aid to identification of fruiting collections precedes the key to the species. Aid to Identification of Fruiting Collections of Ocotea Species • Cupule very large, > 3 cm diameter: O. julianii, O. megacarpa. • Cupule with a double margin and cup-shaped (this character is typical for Licaria; one needs to check dried stamens on the cupule; if 4celled, it is an Ocotea): O. cujumary (strongly developed double margin); O. cymbarum and O. javitensis (double margin not very pronounced). • Cupule with double margin, but plate-like: O. floribunda.

728

L AURACEAE

• Cupule with persistent tepals, but not double-margined: O. adenotrachelium, O. basirecurva , O. depauperata, O. gracilis, O. huberi, O.nilssonii, O. perrobusta O. tomentosa; sometimes tepals persist in O. cowaniana, O. erectifolia, and O. glaucophylla. • Cupule shallow, almost plate-like or very small; pedicel swollen and lenticellate: O. fendleri, O. leucoxylon, and O. oblonga. Key to the Species of Ocotea 1. 1. 2(1).

Flowers bisexual ......................................................................................... 2 Flowers unisexual .................................................................................... 14 Stamens densely papillate, tepaloid, the connective clearly prolonged beyond the anther cells ............................................................. O. cymbarum 2. Stamens not or slightly papillate, the connective not prolonged beyond the anther cells ...................................................................................... 3 3(2). Flowers glabrous or nearly so .................................................................... 4 3. Flowers variously pubescent ..................................................................... 6 4(3). Inflorescences axillary, not grouped together; leaf apex rounded or subacute; shrubs or trees; Amazonas state ................................................. 5 4. Inflorescences grouped at leafless lateral spurs or at the tips of twigs; leaf apex acute or acuminate; trees; Bolívar state ...................... O. fasciculata 5(4). Leaves to 16 × 10 cm; filaments of stamens pubescent; twigs of herbarium specimens not strikingly light colored ....................................... O. liesneri 5. Leaves to 10 × 5 cm; filaments of stamens glabrous; twigs of herbarium specimens strikingly light colored ..................................... O. esmeraldana 6(3). Leaves gradually narrowed toward the base; petioles flat (rarely slightly canaliculate); cupule a small disk on a swollen pedicel; flowers without staminodia; lateral veins equidistant throughout the leaf .................. 7 6. Leaves abruptly narrowed at the base or rounded to subcordate; petioles distinctly canaliculate; cupule a well-developed cup (if disk-shaped, then 3–4 cm diameter); flowers with staminodia; basal 2 pairs of lateral veins often closer together than veins in the rest of the leaf ............... 9 7(6). Leaf tips rounded; leaves glabrous; pit domatia present in vein axils on lower surface of leaf ............................................................... O. tillettsiana 7. Leaf tips acute; leaves pubescent on lower surface; pit domatia lacking .... 8 8(7). Hairs on lower surface of leaf appressed; leaves usually to 3 cm wide ..................................................................................................... O. oblonga 8. Hairs on lower surface of leaf erect; leaves to 6 cm wide .............. O. fendleri 9(6). Dried fruit to 6 × 4 cm; cupule at maturity a shallow cup, ca. 4 cm diameter ........................................................................................ O. megacarpa 9. Dried fruit to 3.5 × 3 cm; cupule cup-shaped, rarely exceeding 2 cm diameter ....................................................................................................... 10 10(9). Young leaves ferruginous-tomentose on lower surface, glabrescent with age; petioles thick, to 5 mm long ......................................................... 11 10. Young leaves glabrous or with appressed hairs on lower surface; petioles not swollen, usually > 10 mm long ...................................................... 12 11(10). Cupule cup-shaped, ca. 3 cm diameter; fruit to 3.5 × 3 cm; leaf base rounded to subcordate ................................................................ O. julianii

Ocotea 729

11. 12(10). 12. 13(12). 13. 14(1). 14. 15(14). 15. 16(15). 16. 17(15). 17. 18(17). 18. 19(18). 19. 20(17). 20. 21(20). 21. 22(21). 22. 23(21). 23. 24(23). 24. 25(24). 25. 26(14).

Cupule funnel-shaped, ca. 1–1.3 cm diameter; fruits to 1.5 × 1 cm; leaf base truncate to rounded ........................................................... O. revoluta Twigs solid; leaves with varying amounts of long, appressed hairs on lower surface ............................................................................ O. aciphylla Twigs hollow; leaves glabrous or with minute, appressed hairs on lower surface .................................................................................................. 13 Leaf margins revolute; cupule funnel-shaped, deep, the margin simple ................................................................................................ O. laticostata Leaf margins flat; cupule shallow, double-margined .................. O. javitensis Twigs, leaves, and terminal buds glabrous ............................................. 15 Twigs, leaves, and/or terminal buds pubescent ...................................... 26 Inflorescence and/or outside of flowers pubescent .................................. 16 Inflorescence and outside of flowers glabrous ........................................ 17 Leaf tips blunt or rounded; tepals persisting as teeth on margin of cupule; petioles thick, to 0.5 cm long ................................................. O. perrobusta Leaf tips acute; tepals not persisting on cupule; petioles 1–2 cm long ................................................................................................ O. myriantha Inflorescences repeatedly branched, many-flowered; leaves usually > 15 cm long (see also O. liesneri) ..................................................................... 18 Inflorescences spicate, few-flowered or rarely lower branches with a few flowers present; leaves usually < 12 cm long ...................................... 20 Petiole ca. 1 cm long; leaf base not recurved nor inrolled or decurent on petiole ..................................................................................... O. crassifolia Petiole shorter; leaf base inrolled, decurrent, or revolute ...................... 19 Leaf base recurved, tepals of staminate flowers conspicuously reflexed at anthesis; leaf apex shortly acute ......................................... O. basirecurva Leaf base inrolled; tepals of staminate flowers spreading, but not reflexed; leaf apex rounded ........................................................... O. neblinae Leaves with well-defined petiole, 1–2 cm long .......................... O. cowaniana Leaves sessile or subsessile, the short petiole, if present, swollen and dark colored .................................................................................................. 21 Leaf base acute ......................................................................................... 22 Leaf base obtuse, rounded, or cordate ..................................................... 23 Leaves elliptic to narrowly elliptic, widest at the middle; lateral veins ascending under an angle of ± 45° ................................. O. roseopedunculata Leaves obovate to narrowly obovate, widest above the middle; lateral veins spreading under an angle of ± 80° ...................................... O. glabra Venation immersed on upper surface of leaf; leaves sessile with cordate base .......................................................................................... O. erectifolia Venation raised on upper surface of leaf; leaf base variable .................. 24 Leaves to 5 cm long ................................................................ O. glaucophylla Leaves 10 cm or more long ...................................................................... 25 Leaf base obtuse; inflorescences shorter than leaves, ca. 5 cm long .... ...................................................................................................... O. venosa Leaf base cordate; inflorescences exceeding leaves, ca. 15 cm long ..... ....................................................................................................... O. huberi Flowers sessile or at least pedicels shorter than flowers; inflorescences usually longer than leaves; flowers with same kind and degree of pu-

730

L AURACEAE

26.

27(26). 27. 28(27). 28. 29(28). 29. 30(29). 30. 31(30). 31.

32(31). 32. 33(32).

33. 34(26). 34. 35(34).

35.

36(34). 36. 37(36).

bescence as pedicels; twigs angled; cupule deeply cup-shaped .......... 27 Flowers pedicellate, pedicels as long as or longer than flowers; length of inflorescences variable; flowers with same kind and density of pubescence as pedicels or less pubescent than pedicels; twigs terete or angled; cupule various ......................................................................... 34 Leaf bases inrolled and decurrent on the petiole ...................... O. tomentella Leaf bases not inrolled and decurrent on the petiole ............................. 28 Lower surface of leaf with a dense, mostly golden, sericeous indument, this completely covering the surface ..................................... O. guianensis Lower surface of leaf without a dense sericeous pubescence, the surface at least partly visible ............................................................................... 29 Hairs on lower surface of leaf erect ............................................ O. glomerata Hairs on lower surface of leaf appressed or absent ................................ 30 Pubescence on young twigs and inflorescence of pale hairs; terminal bud pale-tan sericeous, slender, several times longer than wide ..... O. longifolia Pubescence on young twigs and inflorescences (reddish) brown; terminal bud mat-brown pubescent, about twice as long as wide .................... 31 Inflorescences to 20 cm long, much longer than leaves; cupule cupshaped, with double margin .................................................... O. cujumary Inflorescences scarcely longer than leaves, usually not exceeding 10 cm; cupule (when known) without double margin (in O. flavantha many inflorescences clustered at the tip of stems, this inflorescence complex to 20 cm long; the position of the terminal bud shows that what appears to be the main axis of the inflorescence is in reality a leafless twig with many lateral, short inflorescences) ..................................................... 32 Leaves waxy on lower surface; mature leaves to 6 cm wide ...... O. bracteosa Leaves not waxy on lower surface; mature leaves 3–4 cm wide ............ 33 Leaf bases not recurved, gradually narrowed into the petiole, the petiole not distinct; inflorescences restricted to the upper 2 cm of the twigs ............................................................................................. O. canaliculata Leaf bases recurved, the petioles distinct, ca. 1 cm long; inflorescences mostly on leafless parts of twigs, these 5–15 cm long ............ O. flavantha Lower surface of leaves with tufts of hairs or pits in the axils of the basal lateral veins ......................................................................................... 35 Lower surface of leaves without tufts of hairs or pits in the axils of basal lateral veins ......................................................................................... 36 Tepals sparingly pubescent, pubescence usually denser on pedicels and inflorescence; cupule a rather deep cup without persistent petals; nonflooded forests ........................................................ O. schomburgkiana Tepals densely pubescent, pubescence as dense as or denser than on pedicels and inflorescence; cupule a shallow cup with persistent tepals; nearly always in vicinity of streams ................................................ O. bofo Lower surface of leaf white- to gray-tomentose, the tomentum covering the leaf surface completely ..................................................... O. tomentosa Lower surface of leaf not tomentose; if dense indument present, then consisting of appressed hairs .................................................................... 37 Tepals glabrous outside, densely pubescent inside; lateral veins leaving midvein at very blunt angles (70–90°); leaf apex rounded; lower surface of leaf covered with minute papillae ................................. O. wurdackiana

Ocotea 731

37. Without above combination of characters ............................................... 38 38(37). Style densely pubescent, also in staminate flowers well developed; cupule a flat, double-rimmed disk ..................................................... O. floribunda 38. Style glabrous or at most with a few hairs; cupule not a flat, doublerimmed disk ......................................................................................... 39 39(38). Leaves with a thin wax covering on lower surface, glaucous, this often turning violet upon drying .................................................................. 40 39. Leaves without a thin wax layer, not glaucous, not turning violet upon drying ................................................................................................... 43 40(39). Twigs, leaves, and flowers glabrous; pedicels slender, 5–7 mm long ... ....................................................................................................... O. debilis 40. Twigs, leaves, or flowers pubescent; pedicels shorter, thicker ............... 41 41(40). Tepals pubescent outside, the pubescence covering the entire surface; leaves often shiny on upper surface; pubescence on twigs and leaves consisting of gray hairs; style in staminate flowers with a few hairs ................................................................................................. O. splendens 41. Tepals glabrous or very sparingly pubescent outside, the surface clearly visible; leaves not shiny on upper surface; pubescence on twigs and leaves of brown or reddish hairs; style always glabrous .................... 42 42(41). Twigs angular; leaves oblong or oblong-elliptic ........................ O. leucoxylon 42. Twigs terete; leaves elliptic ................................................. O. aff. amazonica 43(39). Twigs angular, with a reddish or reddish brown indument; fine reticulation not raised on upper surface of leaf; cupule plate-like, the pedicel swollen and often lenticellate in fruit ................................................. 44 43. Twigs terete, lacking reddish indument; upper surface of leaf with reticulation raised or not; cupule, when known, cup-shaped, never plate-like with a swollen, lenticellate pedicel ..................................................... 46 44(43). Leaves chartaceous; plants mostly of lower elevations, but also in the Gran Sabana .......................................................................... O. leucoxylon 44. Leaves coriaceous; plants restricted to tepuis ........................................ 45 45(44). Lower surface of leaf with venation prominently raised ................. O. atrata 45. Lower surface of leaf with venation immersed or slightly raised ......... ................................................................................................... O. nilssonii 46(43). Reticulation raised both on upper and lower surfaces of leaf ................ 47 46. Reticulation not raised or raised only on either the upper or lower surface of leaf, never on both ........................................................................... 49 47(46). Style and stigma not present in staminate flowers; leaves elliptic, the tip rounded or bluntly acute ......................................................... O. duidensis 47. Style and stigma present in staminate flowers; leaves elliptic and blunt to slightly acute, or ovate-elliptic and acuminate ................................... 48 48(47). Leaves ovate-elliptic, acuminate; petioles 1–2 cm long ............. O. yutajensis 48. Leaves elliptic, apex blunt or slightly acute; petioles less than 1 cm long ............................................................................................ O. ceanothifolia 49(46). Stamens 1.5–2 mm long, filaments very short, anthers narrowly triangular to lanceolate, with a sterile tip; leaf base rounded to cordate; inflorescences to 3 cm long; leaves with short (< 1 cm) petioles, 15–25 cm long ............................................................................... O. adenotrachelium 49. Stamens shorter than ca. 1.5 mm, filaments better developed, anthers without a sterile tip; leaf base various, but not cordate; inflorescences

732

L AURACEAE

50(49). 50. 51(50). 51. 52(51). 52. 53(50). 53.

54(53). 54. 55(54). 55.

usually > 3 cm long or petioles longer and leaves shorter than above .............................................................................................................. 50 Young twigs and inflorescences glabrous or with very few appressed hairs .............................................................................................................. 51 Young twigs and/or inflorescences strigose or pubescent ....................... 53 Leaves ovate or narrowly ovate, the base rounded or obtuse; branches zig zag ....................................................................................... O. depauperata Leaves elliptic, the base acute; branches straight .................................. 52 Inflorescences to 3 cm long, with < 10 flowers; tepals reflexed at anthesis ................................................................................................. O. pauciflora Inflorescences 5–10 cm long, usually with > 15 flowers; tepals erect at anthesis ............................................................................................ O. cernua Inflorescences to 2 cm long, much shorter than the leaves; leaves ovate, widest below the middle; tepals persisting on cupule ............... O. gracilis Inflorescences longer, at least half as long as leaves; leaves broadly to narrowly elliptic, widest at the middle; tepals not persisting on cupule .............................................................................................................. 54 Leaves elliptic to broadly elliptic, to 20 × 8 cm; reticulation weakly raised on both surfaces ........................................................................ O. puberula Leaves (narrowly) elliptic, to 12 × 5 cm, reticulation weakly raised on upper surface ............................................................................................ 55 Upper surface of leaf usually drying olive-green, the lower surface lighter green; cupule clearly cup-shaped ..................................... O. sanariapensis Upper and lower surfaces of leaf usually concolorous; cupule a very shallow cup or plate-like, the pedicel thickened............................. O. neesiana

Ocotea aciphylla (Nees & Mart. ex Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 243. 1889. —Oreodaphne aciphylla Nees & Mart. ex Nees, Linnaea 8: 43. 1833. —Cavi-si-yek (Arekuna), Kanau, Sasafrás. Oreodaphne costulata Nees, Linnaea 21: 520. 1848. —Ocotea costulata (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 244. 1889. Ocotea roraimae Mez, Repert. Spec. Nov. Regni Veg. 16: 307. 1920. Ocotea aciphylla var. chimantaensis C.K. Allen, Mem. New York Bot. Gard. 10(5): 76. 1964. Ocotea maguireana C.K. Allen, Mem. New York Bot. Gard. 10(5): 79. 1964. Ocotea sericiflora C.K. Allen, Mem. New York Bot. Gard. 10(5): 77. 1964. Ocotea finium C.K. Allen, Mem. New York Bot. Gard. 12(3): 111. 1965. Ocotea fulvifolia C.K. Allen, Acta Bot. Venez. 2: 212, fig. 10. 1967. Tree. Evergreen lowland forests, tepui for-

ests, 100–1700 m; Bolívar (widespread), Amazonas (Cerro Marahuaka, Cerro Yapacana, San Carlos de Río Negro, Sierra de la Neblina, Sierra Parima). Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia. ŠFig. 619. Several species have been recognized within the concept of Ocotea aciphylla adopted here, but their differences are too weak to maintain them. Included here is Cardona 2496 (VEN), which is annotated as the type of “O. cardonae Lasser,” a species apparently never published. Ocotea adenotrachelium (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 304. 1889. —Oreodaphne adenotrachelium Nees, Syst. Laur., p. 431. 1836. —Bejuco de raya. Leaves large, acuminate, with rounded base; inflorescences and petioles short; tepals remain on the cupule in young fruiting stage. Evergreen lowland forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Amazonian Brazil.

Ocotea 733

Ocotea aff. amazonica (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 359. 1889. —Oreodaphne amazonica Meisn. in A. DC., Prodr. 15(1): 123. 1864. —Laurel blanco. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (Río Caura, Río Icabarú), Amazonas (Cerro Arauricaua, Río Mawarinuma, vicinity of San Carlos de Río Negro). Peru, Brazil, possibly Guyana. The application of this name is uncertain; specimens placed here have previously been identified as Ocotea rubrinervis, which is likely to be a synonym of O. bofo. Ocotea atrata C.K. Allen, Mem. New York Bot. Gard. 10(5): 91. 1964. Tree. Montane forests, 1700–2100 m; Amazonas (Cerro Asisa, Cerro Sipapo). Colombia (Caquetá: Cerro Chiribiquete). Ocotea basirecurva C.K. Allen, Mem. New York Bot. Gard. 10(5): 88. 1964. Small tree to 10 m tall. Montane forests, scrub, 1000–1300 m; Amazonas (Cerro Yutajé, along Río Coro Coro). Endemic. Ocotea bofo H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 161. 1817. Small tree to 15 m tall; upper surface of leaf with fine, raised reticulation and dark gland dots. Frequently collected along streams at base of granitic outcrops, 50–200 m; Bolívar (Río Orinoco, Río Tabaro in Río Nichare basin), Amazonas (Puerto Ayacucho). Anzoátegui, Apure, Guárico, Portuguesa, Táchira; Panama, Colombia, Ecuador, Peru, Brazil. The name Ocotea rubrinervis Mez has been applied to this species and may prove to be a synonym of O. bofo. Ocotea bracteosa (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 356. 1889. —Oreodaphne bracteosa Meisn. in A. DC., Prodr. 15(1): 114. 1864. Evergreen lowland forests, 100–500 m; Bolívar (Río Carrao), Amazonas (La Esmeralda). Brazil. Ocotea canaliculata (Rich.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 361. 1889. —Laurus canaliculata Rich., Actes Soc.

Hist. Nat. Paris 1: 168. 1792. —Laurelillo. Tree to 30 m tall. Evergreen lowland forests, 100–500 m; Delta Amacuro, Bolívar, Amazonas (Río Pasimoni). Guyana, Suriname, French Guiana, Brazil. Ocotea ceanothifolia (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 365. 1889. —Mespilodaphne ceanothifolia Nees, Syst. Laur. 236. 1836. Large tree. Evergreen lowland forests, 100–300 m; Bolívar (vicinity of Upata). Guyana, Brazil. This species is poorly known. The collection from Upata is included because of its lateral veins leaving the midrib under a blunt angle, the leaves somewhat clustered near the branch tips, and the slightly raised reticulation. Ocotea cernua (Nees) Mez, Mitt. Bot. Vereins Kreis Freiburg 47/48: 422. 1888. —Oreodaphne cernua Nees, Syst. Laur. 422. 1836. Oreodaphne caudata Nees, Linnaea 21: 515. 1848. —Ocotea caudata (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 378. 1889. Ocotea punctulata auct. non Mez. 1889: sensu C.K. Allen, Mem. New York Bot. Gard. 10(5): 103. 1964. Shrub or small tree. Mostly collected near savannas, 100–900 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Barinas, Portuguesa; Central America, West Indies, tropical South America, Brazil. In this species the basal lateral veins are often more strongly developed than the others. Collections from southern Amazonas state don’t show this character very well and are difficult to separate from Ocotea myriantha when sterile or fruiting. Ocotea cowaniana C.K. Allen, Mem. New York Bot. Gard. 10(5): 94. 1964. Shrub to 2 m tall. Montane scrub, ca. 2000 m; Amazonas (summit area of Cerro Parú). Endemic. Ocotea crassifolia (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 336. 1889. —Oreodaphne crassifolia Nees, Linnaea 21: 520. 1848.

734

L AURACEAE

Tree to 10 m tall. Montane forests, ca. 1400 m; Bolívar (Macizo del Chimantá [Sarvén-tepui]). Guyana (Mount Roraima).

Ocotea gracilis, but differs in its coriaceous leaves with scarcely visible venation and scarcely loop-connected lateral veins.

Ocotea cujumary Mart., Reise Bras. 3: 1128. 1831. Lower montane forests, 400–900 m; Bolívar (Río Icabarú region). Amazonian Peru, Brazil. This is the only Ocotea species in Venezuela with a pronounced double rim of the cupule; fruiting collections are therefore frequently misidentified as Licaria.

Ocotea duidensis Moldenke, Bull. Torrey Bot. Club 58: 363. 1931. Shrub. Shrubby savannas, 500–2100 m; Bolívar (Gran Sabana), Amazonas (Cerro Duida, Cerro Sipapo). Endemic. Most collections known are from Amazonas state, and it is not certain if they all belong in the same taxon. Ocotea duidensis possibly extends south to the Serra Aracá in Amazonas, Brazil.

Ocotea cymbarum H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 166. 1817. —Licaria cymbarum (H.B.K.) Pittier, Bol. Soc. Venez. Ci. Nat. 7: 135. 1941. —Nectandra cymbarum (H.B.K.) Nees, Syst. Laur. 305. 1836. —Sasafrás. Nectandra barcellensis Meisn. in A. DC., Prodr. 15(1): 155. 1864. —Ocotea barcellensis (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 237. 1889. Tree to 25 m tall. Evergreen lowland and riparian forests, 50–300 m; widely scattered in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Mérida; Colombia, Brazil. ŠFig. 622. The wood is aromatic and is used for construction of boats and houses; the oil which accumulates in the wood of older trunks is used medicinally. Ocotea debilis Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 370, 1889. —Coeppertia caudata Meisn. in A. DC., Prodr. 15(1): 175. 1864. Ocotea directoramea C.K. Allen, Mem. New York Bot. Garden 10: 103. 1964. Shrub or small tree to 5 m tall. Shrubby white-sand savannas, 100–200 m; Amazonas (Caño San Miguel, Río Guainía). Adjacent Colombia, Brazil. The slender pedicels and caudate leaves with glaucous lower surface allow easy identification. Ocotea depauperata C.K. Allen, Mem. New York Bot. Gard. 12(3): 115. 1965. A somewhat clambering shrub with zigzag branches. Savannas, 400–600 m; Bolívar (Río Ichún, Río Kanarakuni). Endemic. Ocotea depauperata is probably close to

Ocotea erectifolia (C.K. Allen) van der Werff, Ann. Missouri Bot. Gard. 75: 419. 1988. —Phoebe erectifolia C.K. Allen, Mem. New York Bot. Gard. 23: 860. 1972. Ocotea budowskiana Bernardi, Candollea 30: 256. 1975. Shrub or small tree to 3 m tall. Montane scrub, 1400–2200 m; Bolívar (upper slopes and summit area of Cerro Jaua). Adjacent Brazil. ŠFig. 625. Ocotea esmeraldana Moldenke, Bull. Torrey Bot. Club 58: 362. 1931. —Cureo barero, Laurel hediondo. Nectandra spumea Kubitzki, Acta. Amaz. 11: 307. 1981. Shrub or small tree. White-sand savannas (banas), 100–600 m; Amazonas (common in white-sand areas). Adjacent Colombia and Brazil. ŠFig. 621. The light-colored bark is characteristic on herbarium specimens. Ocotea fasciculata (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 248. 1889. —Oreodaphne fasciculata Nees, Linnaea 21: 521. 1848. —Caracamate. Tree to 15 m tall. Riparian forests, lower montane forests, 50–600 m; Bolívar (Altiplanicie de Nuria, lower Río Caura). Guyana, Suriname, French Guiana, Brazil (Pará). Ocotea fendleri (Meisn.) Rohwer, Mitt. Staatsinst. Allg. Bot. Hamburg 20: 152. 1986. —Gymnobalanus fendleri Meisn. in A. DC., Prodr. 15(1): 142. 1864. Ocotea glandulosa Lasser, Bol. Técn. Minist. Agric. 3: 9. 1942.

Ocotea 735

Tree. Evergreen montane forests, 1000– 1700 m; Bolívar (Macizo del Chimantá [southwestern slope of Toronó-tepui]). Venezuelan Coastal Cordillera. Ocotea flavantha van der Werff, Ann. Missouri Bot. Gard. 76: 464. 1989. Tree to 30 m tall. Seasonally dry forests, ca. 1100 m; Amazonas (basin of Río Coro Coro, west of Cerro Yutajé). Endemic. Ocotea floribunda (Sw.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 325. 1889. —Laurus floribunda Sw., Prodr. 65. 1788. —Heburu (Warao), Nasinaba (Warao). Ocotea wachenheimii Benoist, Bull. Mus. Hist. Nat. (Paris) 30: 150. 1924. Tree to 20 m tall. Riparian and lowland swamp forests, evergreen lowland to lower montane forests, 0–1200 m; Delta Amacuro (Caño Guiniquina), northern Bolívar, Amazonas (Cerro Yutajé). Distrito Federal, Mérida, Monagas, Táchira, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. Ocotea glabra van der Werff, Ann. Missouri Bot. Gard. 76: 466. 1989. Shrub 2–3 m tall. Montane scrub, 2000– 2500 m; Bolívar (summits of Carrao-tepui and Karaurín-tepui). Endemic. Ocotea glaucophylla Moldenke, Bull. Torrey Bot. Club 58: 364. 1931. Shrub 2–3 m tall. Montane scrub, 1100– 1200(–2000) m; Bolívar (Macizo del Chimantá [Toronó-tepui]), Amazonas (slopes of Cerro Duida). Endemic. This species is poorly known, and the specimens from Bolívar state are only tentatively included here. Ocotea glomerata (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 294. 1889. —Oreodaphne glomerata Nees, Linnaea 21: 515. 1848. Tree to 25 m tall. Common in lowland to lower montane forests, 0–700 m; Delta Amacuro (Tucupita), northern Bolívar, Amazonas (San Carlos de Río Negro). Cojedes, Falcón, Lara, Miranda, Portuguesa, Táchira, Zulia; Guyana, Suriname, French Guiana, Peru, Brazil.

Ocotea gracilis (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 327. 1889. —Oreodaphne gracilis Meisn. in A. DC., Prodr. 15(1): 128. 1864. Ocotea gracilis is a rather common shrub characterized by clambering habit, short inflorescences, venation with well-marked, loop-connected lateral veins, and cupule with persistent tepals. White-sand savannas, 100–200 m; Amazonas (San Carlos de Río Negro). Colombia, Peru, Brazil. ŠFig. 628. Ocotea guianensis Aubl., Hist. Pl. Guiane 781, t. 310. 1775. Tree to 20 m tall; lower surface of leaf densely sericeous. Lowland forests to tepui summits, 100–2200 m; Bolívar (Ptari-tepui), Amazonas (Atabapo, San Carlos de Río Negro). Mérida, Táchira, Zulia; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ŠFig. 627. Ocotea huberi van der Werff, Ann. Missouri Bot. Gard. 76: 467. 1989. Shrub 2–3 m tall. Montane scrub, 1700– 2200 m; border area between Bolívar (Cerro Guanay) and Amazonas (Cerro Coro Coro). Endemic. Ocotea javitensis (H.B.K.) Pittier, Cat. Fl. Venez. 1: 311. 1945. —Laurus javitensis H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 169. 1817. Tree to 35 m tall; twigs hollow. Evergreen lowland forests, 100–200 m; Amazonas (Tamatama, Yavita). Mérida, Táchira; eastern Ecuador and Peru. Ocotea julianii van der Werff, Ann. Missouri Bot. Gard. 76: 468. 1989. —Phoebe steyermarkiana C.K. Allen, Mem. New York Bot. Gard. 10(5): 74. 1964. Shrub or small tree. Montane scrub, 1900–2400 m; Bolívar (Macizo del Chimantá [upper parts of Chimantá-tepui]). Endemic. ŠFig. 623. A sterile collection from Cerro Jaua is tentatively placed here. Ocotea laticostata C.K. Allen, Mem. New York Bot. Gard. 10(5): 111. 1964. Small tree. Montane scrub, 1100–1300 m; Amazonas (Cerro Duida, Cerro Marahuaka). Endemic.

736

L AURACEAE

Two fruiting collections from Cerro Yutajé are tentatively placed here. They differ in their inrolled leaf margins and pubescent leaves and may represent an undescribed species. Ocotea leucoxylon (Sw.) Laness., Pl. Util. Col. Franç. 158. 1886. —Laurus leucoxylon Sw., Prodr. 65. 1788. —Kanauyek, Muneu-yek (Arekuna). Ocotea lasseriana C.K. Allen, Mem. New York Bot. Gard. 10(5): 90. 1964. Ocotea duotincta C.K. Allen, Mem. New York Bot. Gard. 12(3): 119. 1965. Ocotea persulcata C.K. Allen, Mem. New York Bot. Gard. 12(3): 20. 1965. Tree to 20 m tall. Montane and lowland forests, 100–1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Lara, Falcón, Nueva Esparta, Táchira; Central America, West Indies, Colombia, Guyana, Peru, Brazil. ŠFig. 620. In the sense used here, this is a variable species. The extremes seem quite distinct within the Venezuelan Guayana, but often cannot be separated from species in adjoining areas, such as Ocotea acutangula (Miq.) Mez or O. froesii A.C. Sm. Seemingly distinct entities are connected by intermediates with other forms (e.g., O. persulcata, with large, densely appressed pubescent leaves in eastern Delta Amacuro and Bolívar states; a similar pubescence type occurs on specimens from southern Amazonas state). Two collections from Sierra de la Neblina (1800 and 2000 m) are placed here with hesitation; their venation suggests O. atrata, but the leaves are chartaceous. The two collections differ greatly in leaf size from each other, and I do not wish to treat them as a distinct taxon. Ocotea leucoxylon in the wide sense occurs in a variety of habitats. Ocotea liesneri van der Werff, Ann. Missouri Bot. Gard. 76: 468. 1989. Small tree. Montane forests, ca. 800 m; Amazonas (Sierra de la Neblina). Endemic. Ocotea longifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 164. 1817. Ocotea opifera Mart., Reise Bras. 3: 1128. 1831. Tree to 15 m tall. Evergreen lowland forests, 100–200 m; Amazonas (Puerto Ayacucho, base of Sierra de la Neblina). Mérida,

Táchira; Colombia, Ecuador, Peru, Brazil, Bolivia. Several collections from the base of Sierra de la Neblina are provisionally placed here; they have thinner leaves and less pubescent inflorescences than typical material. Ocotea megacarpa van der Werff, Ann. Missouri Bot. Gard. 76: 469. 1989. Tree 10–15 m tall. Montane forests, 1100– 1300 m; Amazonas (Cerro Marahuaka). Endemic. Ocotea myriantha (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 328. 1889. —Mespilodaphne myriantha Meisn. in A. DC., Prodr. 15(1): 120. 1864. —Laurel espina, Laurel hediondo, Palo amarillo. Tree to 15 m tall. In a variety of habitats, forests along stream, nonflooded forests, forests on white sand, thickets of Leopoldinia, savannas, shrubby caatinga, 100–500 m; Amazonas (Río Casiquiare, Río Negro, San Carlos de Río Negro, base of Sierra de la Neblina). Adjacent Colombia and Brazil. The type collection shows waxy lower surface of leaves, but this wax deposit is very poorly visible on recent specimens collected in alcohol. Rather easily confused when fruiting with Ocotea cernua sensu lato; differences are glabrous versus pubescent terminal buds; immersed versus raised midrib on lower surface of leaf; and nearly flat versus distinctly canaliculate petioles. Ocotea neblinae C.K. Allen, Mem. New York Bot. Gard. 10(5): 87. 1964. Tree to 10 m tall. Lower montane forests, 200–700 m; Amazonas (Cerro Yapacana, Sierra de la Neblina). Endemic. Ocotea neesiana (Miq.) Kosterm., Meded. Bot. Mus. Herb. Utrecht 25: 16. 1936. —Nectandra neesiana Miq., Linnaea 18: 745. 1845. Oreodaphne florulenta Meisn. in A. DC., Prodr. 15(1): 125. 1864. —Ocotea florulenta (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 309. 1889. Shrub or small tree to 10 m tall; cupule a very shallow cup, almost plate-like (see discussion under Ocotea sanariapensis). Occasional along lowland black-water streams, 100–400 m; Amazonas (Río Negro, Río Pasimoní, Río Yatúa, San Carlos de Río Ne-

Ocotea 737

gro). Colombia, Guyana, Suriname, French Guiana, Brazil. Ocotea nilssonii C.K. Allen, Mem. New York Bot. Gard. 10(5): 92. 1964. Shrub or tree to 20 m tall. Upper slopes and summits of tepuis, 900–1700 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Jaua, Chivatón east of Kavanayén, Kukenán-tepui, Macizo del Chimantá [Chimantátepui], Uei-tepui). Endemic. Ocotea oblonga (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 367. 1889. —Mespilodaphne oblonga Meisn. in A. DC., Prodr. 15(1): 107. 1864. Ocotea cuprea (Meisn.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 299. 1889. Tree to 32 m tall. Lowland and montane forests, 100–1100 m; Bolívar (Río Icabarú, Río Paramichi), Amazonas (Cerro Yutajé, Río Coro Coro, vicinity of San Carlos de Río Negro). Costa Rica, Panama, Guyana, Ecuador, Peru, Brazil. Ocotea pauciflora (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 371. 1889. —Oreodaphne pauciflora Nees, Syst. Laur. 409. 1836. Oreodaphne diospyrifolia var. incompta Meisn. in A. DC., Prodr. 15(1): 126. 1864. Shrub or small tree to 6 m tall. Lowland forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Guyana, Peru, Brazil, Bolivia. This species is very similar to Ocotea cernua, and sterile fruiting or pistillate material cannot be identified with certainty. Ocotea perrobusta (C.K. Allen) Rohwer, Mitt. Staatsinst. Allg. Bot. Hamburg 20: 170. 1986. —Aiouea perrobusta C.K. Allen, Acta Bot. Venez. 2: 209. 1967. Shrub or small tree to 5 m tall; flowers not yet collected. Tepui-summit thickets, 1600–1900 m; Bolívar (summit of Auyán-tepui). Endemic. Ocotea puberula (Rich.) Nees, Syst. Laur. 472. 1836. —Laurus puberula Rich., Actes Soc. Hist. Nat. Paris 1: 108. 1792. Oreodaphne martiniana Nees, Syst. Laur. 415. 1836. —Ocotea martiniana (Nees) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 344. 1889. Tree to 20 m tall. Widely scattered in lowland forests, 100–500 m; Bolívar (El Palmar,

El Paují), Amazonas (La Esmeralda, San Carlos de Río Negro, base of Sierra de la Neblina). Mexico to Argentina. This species is variable and poorly understood. Ocotea revoluta Moldenke, Bull. Torrey Bot. Club. 58: 365. 1931. Small tree. Montane scrub, 800–1600 m; Amazonas (Cerro Duida). Endemic. Ocotea roseopedunculata van der Werff, Ann. Missouri Bot. Gard. 76: 470. 1989. Shrub to 2 m tall. Tepui scrub, ca. 2000 m; Bolívar (Macizo del Chimantá [near summit of Acopán-tepui]). Endemic. ŠFig. 626. Ocotea sanariapensis Lasser, Bol. Técn. Minist. Agric. 3: 12. 1942. Tree to 4 m tall. Occasional along streams and rivers, ca. 50–300 m; Bolívar (Río Parguaza), Amazonas (Caño Guaca, Río Atabapo, San Fernando, San Juan). Apure; adjacent Colombia, probably Brazil. This species can closely resemble Ocotea neesiana, but O. sanariapensis has deeply cup-shaped cupules and leaves often with a green lower surface lighter than the olivegreen upper surface. Also, the venation is more raised on the upper surface of the leaf in O. neesiana. Ocotea sanariapensis is closely related to O. schomburgkiana, but lacks domatia. Ocotea schomburgkiana (Nees) Mez, Jahrb. Königl. Bot. Gard. Berlin 5: 337. 1889. —Oreodaphne schomburgkiana Nees, Linnaea 21: 269. 1848. —Laurel paraguito, Laurel tigrito. Ocotea castanea C.K. Allen, Mem. New York Bot. Gard. 12(3): 114. 1965. Ocotea subalveolata C.K. Allen, Mem. New York Bot. Gard. 15: 83. 1966. Small tree to 15 m tall. Nonflooded forests, 100–1000 m; Delta Amacuro (Río Toro), Bolívar (Gran Sabana, Río Acañán, Río Carún, Río Caura). Guyana. The wood is hard and used for timber. Ocotea splendens (Meisn.) Baill., Hist. Pl. 2: 446. 1872. —Oreodaphne splendens Meisn. in A. DC., Prodr. 15(1): 129. 1864. —Laurel blanco. Ocotea globifera Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 328. 1889.

738

L AURACEAE

Fig. 619. Ocotea aciphylla

Fig. 620. Ocotea leucoxylon

Ocotea 739

Fig. 621. Ocotea esmeraldana

Fig. 622. Ocotea cymbarum

740

L AURACEAE

Fig. 623. Ocotea julianii

Fig. 625. Ocotea erectifolia

Fig. 624. Ocotea wurdackiana

Ocotea 741

Fig. 626. Ocotea roseopedunculata

Fig. 627. Ocotea guianensis

Fig. 628. Ocotea gracilis

742

L AURACEAE

Ocotea dissimilis C.K. Allen, Mem. New York Bot. Gard. 15: 86. 1966. Tree to 15 m tall. Evergreen lowland forests, 100–200 m; Amazonas (near San Carlos de Río Negro). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. Ocotea tillettsiana C.K. Allen, Mem. New York Bot. Gard. 12(3): 109. 1965. Tree to 20 m tall. Montane forests, 1600– 1700 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Guyana. Ocotea tomentella Sandwith, Bull. Misc. Inform. Kew 49: 130. 1935. —Laurel blanco. Tree to 30 m tall. Evergreen lowland forests, 100–300 m; Delta Amacuro (near El Palmar). Guyana. Ocotea tomentosa van der Werff, Ann. Missouri Bot. Gard. 76: 470. 1989. Shrub or small tree. Savannas, 100–200 m; Amazonas (Río Guainía, Río Negro). Endemic. Ocotea venosa Gleason, Bull. Torrey Bot. Club 58: 365. 1931. Small tree. Montane scrub, ca. 2200 m; Amazonas (summit of Cerro Duida). Endemic. A fruiting collection from Cerro Mara-

huaka (at 1100 m) probably belongs here, but the species is known with certainty only from the type. Ocotea wurdackiana C.K. Allen, Mem. New York Bot. Gard. 10(5): 81. 1964. Shrub to 6 m tall. Montane scrub, (1200–) 1700–2100 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Acopán-tepui, Chimantátepui]). Brazil (Amazonas: Serra Aracá). ŠFig. 624. Ocotea yutajensis C.K. Allen, Mem. New York Bot. Gard. 10(5): 105. 1964. Small tree; petioles long. Tepui slope forests and scrubs, 500–1200 m; Amazonas (Cerro Yutajé). Endemic. Ocotea sp. A Montane scrub, 2100–2200 m; Bolívar (Tereké-Yurén-tepui), (Murisipán-tepui) Endemic. Known from a fruiting collection and a sterile collection, this species, with lenticellate cupules and angular twigs, belongs to the Ocotea leucoxylon group. It differs from the other species in this group (O. atrata, O. leucoxylon, O.nilssonii, O. tomentosa) in the densely ferruginous-tomentose lower surface of the leaves. Flowers are needed for a description.

12. PERSEA Mill., Gard. Dict. Abr. ed. 4. 1754. by Henk van der Werff Trees or shrubs. Leaves alternate, evenly distributed along branches, rarely clustered near tip of branches, entire, often coriaceous, glabrous or with appressed or erect pubescence. Inflorescences axillary, usually as long as or longer than the subtending leaves, many-flowered, paniculate. Tepals 6, in 2 rows of 3, the rows equal, subequal or strongly unequal, usually erect at anthesis. Stamens in 4 whorls of 3; the outermost 2 whorls (I & II) always fertile and 4-celled (very rarely 2-celled); whorl III usually fertile and 4-celled, but occasionally staminodial or 2-celled; whorl IV always staminodial, the staminodia usually conspicuous; filaments of stamens in whorl III each with 2 globose glands at their base. Fruit pear-shaped or globose, usually subtended by persistent tepals (tepals rarely deciduous); fruiting pedicel very rarely swollen. Neotropics; ca. 80 species, 15–20 in Venezuela, 13 of these in the flora area. The generic boundaries of Persea are not well defined. Whether the paleotropical segregates, Machilus, Nothaphoebe, and Alseodaphne should be recognized as genera or merged into Persea is not yet clear. Persea americana is widely cultivated for its edible fruit, the avocado.

Persea 743

Key to the Species of Persea Tepals equal or subequal, the outer ones > 3/4 the size of the inner ones ... 2 Tepals unequal, the outer ones < 3/4 the size of the inner ones, often much less .......................................................................................................... 5 2(1). Twigs hollow; tepals 3.5–4 mm long, densely pubescent on the outside; fruits globose, ca. 1.3 cm diameter; pedicels swollen in fruit, tepals persistent in fruit; only known from flooded forests .................... P. fluviatilis 2. Twigs solid; tepals larger or not densely pubescent; fruits pear-shaped (poorly known in P. steyermarkii); pedicels not swollen in fruit; tepals deciduous in fruit ................................................................................... 3 3(2). Tepals 5–6 mm long; lower surface of leaf variously pubescent; widely cultivated for its edible fruit and often escaping from cultivation ............. ................................................................................................. P. americana 3. Tepals ca. 3 mm long; lower surface of leaf glabrous or nearly so; native ................................................................................................................ 4 4(3). Tepals sparsely pubescent, the surface of the tepals clearly visible; leaves to 10 cm long ........................................................................ P. steyermarkii 4. Sepals densely pubescent, the surface of the tepals completely covered by the pubescence; leaves to 18 cm long ............................... P. areolatocostae 5(1). Flowers with 6 fertile stamens, the inner 6 reduced to staminodia ........ 6 5. Flowers with 9 fertile stamens, only the inner 3 reduced to staminodia .... 8 6(5). Petioles to 1.5 cm long; highland species (above 800 m) ............. P. fastigiata 6. Petioles > 2.5 cm long; lowland species ..................................................... 7 7(6). Indument on inflorescences mostly erect; lateral veins 8–10 pairs; lower leaf surface glabrous or with scattered, ± erect hairs ............ P. fluviatilis 7. Indument on inflorescences appressed; lateral veins 5 or 6 pairs; lower leaf surface sparsely appressed, pubescent to almost glabrous ............. ..................................................................................... P. pseudofasciculata 8(5). Lower surface of leaf densely appressed-pubescent, sericeous, the epidermis of leaf not visible ............................................................................. 9 8. Lower surface of leaf tomentose, sparsely appressed-pubescent, or glabrous, but never densely sericeous ..................................................... 11 9(8). Leaves broadly ovate to ovate-elliptic, 6–12 cm wide; inflorescences longer than subtending leaves, 15–30 cm long; lateral veins prominently raised on lower surface of leaf .................................. P. grandiflora 9. Leaves elliptic or narrowly elliptic, to 4 cm wide; inflorescences subequal to leaves, to 10 cm long; lateral veins immersed or slightly raised on lower surface of leaf ............................................................................. 10 10(9). Pubescence on lower surface of leaf copper-colored, sericeous; leaves 10– 15 cm long; inflorescence to 15 cm long .............................. P. perseiphylla 10. Pubescence on lower surface of leaf light brown, sericeous; leaves 4–6 cm long; inflorescence few-flowered, slender, ca. 5 cm long .......... P. maguirei 11(8). Lower surface of leaf completely covered by tomentose pubescence ..... 12 11. Lower surface of leaf sparsely appressed-pubescent to nearly glabrous ... 13 12(11). Ovary and style pubescent; leaf pubescence sericeous-tomentose; inner tepals ca. 8 mm long ............................................................. P. grandiflora 12. Ovary and style glabrous; leaf pubescence tomentose; inner tepals to 1. 1.

744

L AURACEAE

5 mm long ................................................................................... P. jenmanii 13(11). Outer tepals 1/3–1/4 as long as inner tepals; tips of inner tepals breaking off in fruiting stage, the subtending tepals all appearing the same length; leaf base (acute-)obtuse or sometimes rounded; leaves often > 10 cm wide ............................................................................................. P. caerulea 13. Outer tepals 1/2 as long as inner tepals, the tips not breaking off in fruiting stage; leaf base acute; leaves ≤ 6 cm wide .......................................... 14 14(13). Inner tepals ca. 4 mm long; lateral veins raised on lower surface of leaf; petioles 2–2.5 cm long; pubescence on lower surface of leaf very sparse, best visible along the midvein .................................................... P. croizatii 14. Inner tepals ca. 6 mm long; lateral veins immersed on lower surface of leaf; petioles 1.5–2 cm long; pubescence on lower surface of leaf easily visible under a microscope, covering about 50% of the surface ... P. croatii Persea americana Mill., Gard. Dict. ed. 8. 1768. —Aguacate. Tree to 30 m tall. Cultivated and naturalized near human habitations, 0–1000 m; probably widely scattered in Delta Amacuro and Bolívar, rarer in Amazonas. Native of Central America, widely cultivated for its edible fruit. Persea areolatocostae (C.K. Allen) van der Werff, Ann. Missouri Bot. Gard. 76: 471. 1989. —Phoebe areolatocostae C.K. Allen, Mem. New York Bot. Gard. 10(5): 75. 1964. —Cinnamomum areolatocostae (C.K. Allen) Kosterm., Reinwardtia 10: 441. 1988. Tree to 15 m tall. Lower montane forests, 600–700 m; Amazonas (Sierra de la Neblina). Endemic. ŠFig. 629. This species is closely related to the poorly known P. steyermarkii and differs mainly in having densely pubescent flowers and larger leaves. Persea caerulea (Ruiz & Pav.) Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 171. 1889. —Laurus caerulea Ruiz & Pav., Fl. Peruv. 4: t. 350. 1804. Persea lignitepala Lasser, Bol. Soc. Venez. Ci. Nat. 9: 177. 1944. Tree. Montane forests, 1300–1500 m; Bolívar (Cerro Uananapán near Uraday). Aragua, Barinas, Distrito Federal, Mérida; Monagas; widely distributed from Honduras to Bolivia. The only flora area collection (Cerro Uananapán cited in Mem. New York Bot. Gard. 14: 36. 1966) was not seen for this treatment, but it is considered likely that Persea caerulea does occurs in the flora area.

Persea croatii van der Werff, Ann. Missouri Bot. Gard. 76: 471. 1989. Small tree. Montane forests, 1800 m; Amazonas (Sierra de la Neblina). Endemic. Persea croizatii van der Werff, Ann. Missouri Bot. Gard. 76: 472. 1989. Known only from the type collection made by Leon Croizat along the upper Río Orinoco, Amazonas state. Exact locality, habit, and habitat are unknown. Endemic. Persea fastigiata Kopp, Mem. New York Bot. Gard. 10(5): 71. 1964. Persea fastigiata var. pilosa Kopp, Mem. New York Bot. Gard. 10(5): 72. 1964. Shrub or tree to 20 m tall. Occasional on tepui slopes, 800–2400 m; Bolívar (Auyántepui, Cerro Jaua, Macizo del Chimantá [Chimantá-tepui], Ptarí-tepui), Amazonas (Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Endemic. ŠFig. 630. Kopp also described Persea fastigiata var. sericea Kopp, in which stamens in whorl III are fertile, not sterile. The material of this taxon is poor (type from Sierra de la Neblina, 2000 m elevation), and it may very well be an aberrant specimen of P. fastigiata. A second collection of this taxon (from 1800 m elevation, Sierra de la Neblina) has diseased flowers. Persea fluviatilis van der Werff, Ann. Missouri Bot. Gard. 76: 474. 1989. Tree to 10 m tall. Seasonally flooded forests along black-water rivers, ca. 100 m; Amazonas (Río Baría, Río Temi). Endemic. ŠFig. 632. A fruiting collection cited by Kopp as Persea nivea is placed here in P. fluviatilis;

Persea 745

no material of P. nivea from the Venezuelan Guayana was found. Persea grandiflora Kopp, Mem. New York Bot. Gard. 10(5): 73. 1964. Tree to 10 m tall. Montane forests, 700– 1300 m; Amazonas (Cerro Marahuaka, Sierra de la Neblina). Endemic. ŠFig. 633. This species is close to Persea jenmanii and best recognized by its pubescent style and ovary. Persea jenmanii Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 162. 1889. Shrub or tree to 12 m tall. Montane forests, along streams, 900–1800 m; Bolívar (Auyán-tepui, Cerro Jaua, Cerro Venamo, near Santa Elena de Uairén), Amazonas (Cerro Duida). Guyana. ŠFig. 631. The two collections from Cerro Duida, identified as Persea jenmanii by Kopp, probably don’t belong here, but the material is insufficient for description of a new taxon. Persea maguirei Kopp, Mem. New York Bot. Gard. 10(5): 73. 1964. Shrub. Montane shrublands, ca. 1400 m; Amazonas (Cerro Duida). Endemic. ŠFig. 634. Persea perseiphylla (C.K. Allen) van der Werff, Ann. Missouri Bot. Gard. 76: 961.

1989. —Ocotea perseiphylla C.K. Allen, Mem. New York Bot. Gard. 10(5): 109. 1964. Shrub. Open scrub with bromeliads and lichens, 600–1500 m; Amazonas (Cerro Aracamuni, Sierra Unturán). Colombia (Guainía, Vaupés). Persea pseudofasciculata Kopp, Mem. New York Bot. Gard. 14(1): 85. 1966. Tree to 12 m tall. Nonflooded forests on sandy soil, 100–1100 m; Amazonas (Cerro Yutajé, base of Sierra de la Neblina). Colombia, Brazil (Amazonas), Bolivia. The lack of flowers makes it difficult to assign the specimens to one of the varieties recognized by Kopp; the variety parviflora Kopp is the most likely one. Persea steyermarkii C.K. Allen, J. Arnold Arbor. 26: 286. 1945. Medium-sized tree. Edge of a small granitic outcrop, ca. 100 m; Amazonas (Río Temi). Guatemala, El Salvador. Because the species is otherwise only known from a few collections in the mountains of Guatemala and El Salvador, determination of the single collection from the flora area must be considered doubtful. More collections are needed to determine the correct name of this species.

Fig. 629. Persea areolatocostae

746

L AURACEAE

Fig. 630. Persea fastigiata

Fig. 631. Persea jenmanii

Fig. 632. Persea fluviatilis

Persea 747

Fig. 633. Persea grandiflora

Fig. 634. Persea maguirei

748

L AURACEAE

13. PLEUROTHYRIUM Nees, Syst. Laur. 349. 1836. by Henk van der Werff Shrubs or trees. Leaves alternate or congested near tips of branches, usually elliptic, glabrous or densely tomentellous, the lateral veins often curving upwards and connected with the upper lateral vein. Inflorescences axillary, paniculate. Flowers bisexual; tepals 6, equal or the inner 3 with a narrower base. Stamens 9, all 4celled, the outer 6 with lateral or lateral-introrse cells, the inner 3 with extrorse cells; inner 3 stamens each with 2 glands near the base of the filaments, these sometimes growing outwards between the outer stamens, and becoming fused, forming a conspicuous glandular mass in the flowers; staminodia lacking or inconspicuous. Style glabrous or pubescent; cupule large, cup-shaped, often warty. Central America, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 20 species, 4 or 5 in Venezuela, 1 of these in the flora area. Pleurothyrium amapáense C.K. Allen, Mem. New York Bot. Garden 15: 92. 1966. Liana(?). Forests on hummocks, ca. 100

m; Amazonas (upper Río Baría). Brazil (Amapá: Río Araguari). The collector noted “liana” as habit, although this is highly unusual for Lauraceae.

14. RHODOSTEMONODAPHNE Rohwer & Kubitzki, Bot. Jahrb. 107: 135. 1985. by Henk van der Werff Trees or shrubs. Leaves alternate, entire. Inflorescences axillary, paniculate. Flowers unisexual (plants dioecious), tepals 6, equal, erect or spreading at anthesis. Staminate flowers with 9 4-celled stamens, the locelli in a weak arc or in a straight line; outer 6 stamens with the locelli introrse or the outermost locelli lateral; inner 3 stamens with extrorse locelli and 2 glands attached near base of filaments; filaments of all stamens very short; pistillode very small or lacking. Pistillate flowers with well-developed ovary and style, the staminodes sometimes with opened locelli, but not producing pollen. Fruit a 1-seeded berry, cupule (when known) rather large and deep. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 10 species, 4 in Venezuela, all in the flora area. Key to the Species of Rhodostemonodaphne 1. 1. 2(1). 2.

3(1). 3.

Shrubs or small trees to 4 m tall; lower surface of leaf with dense, ferruginous tomentum ...................................................................................... 2 Trees to 30 m tall; lower surface of leaf with appressed or sparse, erect pubescence, never with ferruginous tomentum ................................... 3 Inflorescences longer than subtending leaves; leaves to 10 cm long, ovate, the main veins on lower surface of leaf conspicuously raised ....... R. celiana Inflorescences shorter than subtending leaves; leaves to 7 cm long, elliptic to narrowly elliptic; venation on lower surface of leaf scarcely raised ........................................................................................ R. steyermarkiana Pubescence on twigs, inflorescences, and flowers finely appressed, yellowish brown ..................................................................................... R. grandis Pubescence on twigs, inflorescences, and flowers finely erect, ferruginous brown ...................................................................................... R. kunthiana

Rhodostemonodaphne 749

Rhodostemonodaphne celiana (C.K. Allen) Rohwer, Mitt. Staatsinst. Allg. Bot. Hamburg 20: 85. 1986. —Ocotea celiana C.K. Allen, Mem. New York Bot. Gard. 10(5): 106. 1964. Shrub to 2 m tall, with weak twigs. Rocky slopes and cliffs, 1200–2000 m; Amazonas (slopes and summit of Cerro Yutajé). Endemic. Rhodostemonodaphne grandis (Mez) Rohwer, Mitt. Staatsinst. Allg. Bot. Hamb. 20: 84. 1986. —Endlicheria grandis Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 124. 1889. —Nectandra grandis (Mez) Kosterm., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 25: 17. 1936. Tree to 30 m tall. Evergreen lowland to montane forests, 50–1200 m; Delta Amacuro (Río Amacuro), Bolívar (Reserva Forestal de Imataca, between San Ignacio and San Francisco de Yuruaní), Amazonas (Río Manapiare). Colombia, Guyana, Ecuador, Peru, Brazil. This species is very similar to some Endlicheria species, and pistillate material especially may be hard to determine. In an upcoming revision of Rhodostemonodaphne by S. Madriñán, this taxon will probably have a different name. Rhodostemonodaphne kunthiana (Nees) Rohwer, Mitt. Staatsinst. Allg. Bot.

Hamburg 20: 84. 1986. —Acrodiclidium kunthiana Nees, Syst. Laur. 269. 1836. —Nectandra kunthiana (Nees) Kosterm., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 25: 19. 1936. Nectandra meyeriana Lasser, Bol. Soc. Venez. Ci. Nat. 11: 184. 1948. Pleurothyrium cowanianum C.K. Allen, Mem. New York Bot. Gard. 10(5): 121. 1964. Tree to 30 m tall. Riparian forests, evergreen lowland to montane forests, 100–1300 m; Bolívar (Cerro Venamo, Río Nichare basin), Amazonas (Caño Yureba, Cerro Huachamacari, Río Mawarinuma). Venezuelan Coastal Cordillera; Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 635. Rhodostemonodaphne steyermarkiana (C.K. Allen) van der Werff, Ann. Missouri Bot. Gard. 76: 474. 1989. —Ocotea steyermarkiana C.K. Allen, Acta Bot. Venez. 2: 216. 1967. Ocotea ferruginea auct. non (Meisn.) Mez 1889: sensu Bernardi, Candollea 30: 261. 1975. Shrub or small tree to 4 m tall. Montane streamside forests, tepui shrublands, 1400– 2300 m; Bolívar (Auyán-tepui, Cerro Sarisariñama). Endemic.

Fig. 635. Rhodostemonodaphne kunthiana

750

L AURACEAE

15. SEXTONIA van der Werff, Novon 7: 436. 1997. by Henk van der Werff Trees. Twigs thick, corky. Leaves alternate, clustered near the tips of the branches, pinnately veined, without domatia. Inflorescences axillary, paniculate-cymose or racemose. Flowers bisexual. Tepals 6, erect, unequal, the outer 3 smaller than the inner 3. Stamens 9, all 4-celled, the outer 6 with the cells in a shallow arc, the inner 3 with the cells placed in 2 pairs; filaments short, as wide as the anthers. Staminodia 3, present. Receptacle deep, enclosing the ovary. Fruit initially covered by the cupule, at maturity either largely exserted or remaining enclosed in the cupule. South America; 2 species, 1 in Venezuela. Sextonia, previously included in Ocotea, is recognized because of its unequal tepals, clustered leaves, profound differences in wood and bark anatomy, and different embryology from Ocotea species. Sextonia rubra (Mez) van der Werff, Novon 7: 438, fig. 1I-K. 1997. —Ocotea rubra Mez, Jahrb. Königl. Bot. Gart. Berlin 5: 258. 1889. —Nectandra rubra (Mez) C.K. Allen, Mem. New York Bot. Gard. 10: 120. 1964. —Caobillo, Mana-choroyek (Arekuna), Paci-ra-ru-yek (Arekuna). Tree to 40 m tall. Montane forests, 900–

1600 m; Bolívar (Cerro Venamo, near Kavanayén, Ptarí-tepui, upper Río Cuyuní). Guyana, Suriname, French Guiana, Brazil. The wood is of fine quality and is used for making furniture. Vegetatively, Sextonia rubra resembles Mezilaurus due to its clustered leaves with a rounded apex, but the flowers and fruits are quite different.

LECYTHIDACEAE by Scott A. Mori and Ghillean T. Prance Small to large trees. Leaves alternate, simple, exstipulate or with minute, caducous stipules, without pellucid punctations. Flowers actinomorphic or zygomorphic. Petals usually free (absent or fused together in subfamilies Foetidioideae and Napoleonaeoideae). Stamens numerous, connate at base into staminal ring, the ring actinomorphic or prolonged on one side into a strap-like structure that arches over summit of ovary, some apetalous species with outermost stamens modified into peta-

L ECYTHIDACEAE 751

loid corona (Asteranthos brasiliensis); intrastaminal disc present or absent. Ovary inferior to half inferior. Fruits fibrous berries, dry drupes, or woody circumscissile capsules. Seeds 1–many, with or without cotyledons. Pantropics; 20 genera and ca. 300 species, 8 genera and 35 species in the flora area. The Lecythidaceae, sensu lato, is a pantropical family of small to very large trees. The family includes four subfamilies, only two of which, the Napoleonaeoideae and Lecythidoideae, are found in the Neotropics. Both subfamilies are represented in the Venezuelan Guayana. The Napoleonaeoideae comprises three genera, with two genera and 11 species in West Africa and a single genus and species, Asteranthos brasiliensis, in the upper Río Negro region of Colombia, Venezuela, and Brazil. The systematic positions of the Napoleonaeoideae and of Asteranthos brasiliensis within the Napoleonaeoideae are currently under study, and considerable evidence is accumulating to align Asteranthos with the West African Scytopetalaceae and treat this group as a subfamily of the Lecythidaceae (C. M. Morton, S. A. Mori, G. T. Prance, K. G. Carol, and M. W. Chase. Amer. J. Bot. 84: 530–540. 1997). The Lecythidoideae, with 10 genera and ca. 200 species, are limited to the Neotropics. Key to Genera of Lecythidaceae 1. 1. 2(1).

2.

3(2).

3.

4(1).

4.

5(4).

Flowers actinomorphic ............................................................................... 2 Flowers zygomorphic ................................................................................. 4 Flowers solitary in leaf axils, petals fused into a radiate corona; fruits with persistent, annular calyx; embryo J-shaped, embedded in ruminate endosperm ..................................................................... 2. Asteranthos Flowers in multi-flowered inflorescences, infrequently with a single flower, with 6–8 separate petals; fruits without persistent, annular calyx; embryo not J-shaped, without endosperm ..................................... 3 Flower buds oblong, the flowers < 2.5 cm diameter at anthesis; stamens < 50, the anthers dehiscing via lateral slits; ovules on lower 1/2 of septum; fruits dehiscent, the seeds freely falling from fruit; embyro with undifferentiated cotyledons .................................................... 1. Allantoma Flower buds globose, the flowers 5–20 cm diameter at anthesis; stamens 500–1200, the anthers dehiscing via apical pores; ovules on upper 1/2 of septum; fruits indehiscent, the seeds released by deliquescence of pericarp or by fragmentation of opercular region; embryo with plano-convex, fleshy cotyledons ............................................................... 7. Gustavia Inflorescences from trunk; flowers with all of hood appendages bearing anthers; fruits round, indehiscent; seeds embedded in pulp which oxidizes bluish green when exposed to air ................................ 5. Couroupita Inflorescences not from trunk; flowers usually with none of hood appendages bearing anthers, less frequently the proximal with and the distal appendages without anthers; fruits usually not round (except for B. excelsa), dehiscent; seeds not embedded in pulp .................................. 5 Buds enclosed by calyx except for horizontal slit at apex; calyx with 2 lobes at anthesis; style > 10 mm long; fruits functionally indehiscent, with

752

5.

6(5).

6.

7(6).

7.

L ECYTHIDACEAE

small, inwardly falling operculum; seeds with thick, bony testa, remaining inside fruit at maturity .......................................... 3. Bertholletia Buds not enclosed by calyx; calyx with 6 lobes at anthesis; style usually < 10 mm long; fruits dehiscent, with freely falling opercula; seeds without thick, bony testa, usually falling or removed from fruit at maturity ................................................................................................................ 6 Androecial hood coiled inward, then with outwardly extended flap at apex of coil; fruits cylindric or campanulate; seeds with wing around circumference; embryo with 2 foliaceous cotyledons ......................... 4. Couratari Androecial hood flat, or if coiled inward without outwardly extended flap at apex of coil; fruits not cylindric or campanulate; seeds without wings; embryo undifferentiated ............................................................ 7 Androecial hood usually forming complete coil inward, with blunt-tipped appendages at apex of coil, these differentiated from more abundant, echinate hood appendages; ovary usually 2-, less frequently 4-locular; seeds usually with lateral arils, less frequently aril basal or completely surrounding testa ................................................................ 6. Eschweilera Androecial hood flat or expanded at apex but not forming complete coil inward, the hood appendages not differentiated into blunt-tipped appendages; ovary 4-locular; seeds usually with basal arils ......... 8. Lecythis

1. ALLANTOMA Miers, Trans. Linn. Soc. London 30(2): 170, 291. 1874. Small to medium-sized trees. Leaves entire, glabrous. Inflorescences racemes or once-branched arrangements of racemes, terminal or subterminal. Flowers nearly actinomorphic, oblong, 15–22 mm long; receptacle campanulate; sepals (5)6, united at base, with triangular, small lobes at apex; petals 6, oblong; androecium urceolate, only slightly asymmetrical, the apex divided into 8–10 inwardly reflexed lacinae with anthers at apex of most of them, with additional stamens inserted sparsely over interior of androecium on short, thick, reflexed filaments, stamens ca. 30, all anthers fertile; ovary (3)4- or 5-locular, with numerous ovules in each locule; style short, with rounded, undivided stigmatic surface. Fruits woody capsules, tubular-cylindric to cylindric-campanulate, straight or slightly curved, always longer than broad, the operculum dehiscing when fruits still on tree. Seeds narrowly linearelongate, with poorly developed basil aril, not winged, the testa verrucose. Río Orinoco, Río Negro, and Rio Amazonas watersheds of Venezuela and Brazil; 1 species. Allantoma lineata (Mart. ex O. Berg) Miers, Trans. Linn. Soc. London 30(2): 297, t. 65, figs. 4, 5. 1874. —Couratari lineata Mart. ex O. Berg in Mart., Fl. Bras. 14(1): 508, t. 7, 77. 1858. —Tabarí. Usually a canopy tree; leaves with rounded to truncate bases, venation eucamptodromous, the tertiary veins finely percurrent, forming 135° angle with midrib; flowers oblong; androecium nearly symmetric, the anthers inserted on upper margin and all over

interior; fruits cylindric, much longer than broad; seeds with poorly developed, basal aril. Lowland forests along rivers and streams, 100–200 m; Amazonas (Caño Morocoto below San Fernando del Atabapo, base of Cerro Yapacana, Río Autana, upper Río Baría, Río Cuao, between San Carlos de Río Negro and Solano). Brazil (Amazonas: Río Negro and Rio Amazonas basins). ŠFig. 636. The seeds are edible, and the bark is used for making cigarette paper.

Allantoma 753

Fig. 636. Allantoma lineata

754

L ECYTHIDACEAE

2. ASTERANTHOS Desf., Mém. Mus. Hist. Nat. 6: 9, t. 3. 1820. Medium-sized trees. Leaves entire, with minute stipules, visible only on youngest leaves. Inflorescences solitary in leaf axils. Flowers actinomorphic, 3.5–4.5 cm diameter; receptacle conical-campanulate; calyx accrescent to receptacle, radiate, circular; corolla corona-like, with 24–28 conspicuous veins from center to margin, pleated and unfolding like a parasol, the margins irregularly dentate and ciliate; fertile stamens numerous, free, inserted in several rows attached in circle around base; anthers 2-locular, longitudinally dehiscent, basally attached; style erect and thin; stigma 5–8-lobed; ovary semi-inferior, 5–8-locular, with 4 ovules in each locule. Fruit an oblong-pyramidal, tardily dehiscent capsule, semi-inferior, ± 2 cm long, with enlarged persistent calyx persisting around middle, the fruiting calyx coriaceous, part of fruit below calyx smooth, unribbed, 0.5 cm long, part above calyx 1.5 cm long, deeply 6-ribbed and tapered to pointed apex, unilocular, with single cone-shaped seed, with abundant ruminate endosperm; embryo curved on lower portion to form J-shape, with 2 small, membranous cotyledons at apex. Endemic to the Guayana Shield in Colombia, Venezuela, and Brazil; 1 species. Asteranthos brasiliensis Desf., Mém. Mus. Hist. Nat. 6: 9, t. 3. 1820. Tree to 25 m tall; flowers with radiate, corona-like, yellow corolla; fruits with circular, persistent calyx; seeds with abundant, ruminate endosperm. Periodically flooded, lowland forests on white sand, 100–200 m; Amazonas (Río Guaínia, San Carlos de Río Negro, road between Yavita and Maroa). Colombia

(upper Río Negro basin), Brazil (Amazonas: upper Rio Negro basin). ŠFig. 637. According to Bruce Nelson (personal communication 1986), this species descends the Rio Negro to at least Ecunaí (ca. 70 km south of Tapuruquara) in Brazil, which roughly corresponds to the southern limit of the Guayana Shield outcrops along the Rio Negro.

Fig. 637. Asteranthos brasiliensis

Bertholletia 755

3. BERTHOLLETIA Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 122, t. 36. 1808 [1807], spelling variant: Barthollesia. Canopy or emergent trees. Lower surface of leaves minutely papillate. Inflorescences terminal and in axils of uppermost leaves. Flowers zygomorphic. Calyx lobes 2, the apex of the lobes often obscurely 3-dentate; petals 6; androecium with appendages of hood swept inward but not forming full coil. Ovary 4-locular; style long, geniculate. Fruit round, falling from tree at maturity with seeds inside, the pericarp woody and hard, the diameter of operculum < 1.5 cm, the operculum falling into fruit at dehiscence. Seeds triangular in cross section, without well-developed funicle Fig. 638. Bertholletia excelsa

756

L ECYTHIDACEAE

and aril, the testa very woody and hard, the embryo undifferentiated; cotyledons present only as two inconspicuous scales. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 1 species. Bertholletia excelsa Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 122, t. 36. 1808 [1807]. —Almendra, Jibia, Juvia, Matamatá de altura. Bertholletia nobilis Miers, Trans. Linn. Soc. London 30(2): 197, t. 37, figs. 4–7. 1874. Canopy or emergent tree; leaves large, 17–36 × 6–16 cm, with finely papillate abaxial surface; flowers with 2 calyx lobes and long, geniculate styles; fruits round, woody, falling to ground with seeds inside at maturity. Lowland, nonflooded forests, 100– 200 m; Amazonas (upper Río Negro). Wide-

spread in Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, cultivated in tropical areas outside its native range, e.g., Trinidad, Ghana, and Malaysia. ŠFig. 638. This plant provides the well-known Brazil nut of commerce, which is harvested mainly from wild trees throughout its range. The wood provides excellent timber, but is little used because of the value of the tree as a commercial source of seeds. The seeds are used locally for food and cooking oil.

4. COURATARI Aubl., Hist. Pl. Guiane 723, t. 290. 1775, fruit only, spelling variants: Curatari, Curataria. Lecythopsis Schrank, Denkschr. Königl. Akad. Wiss. München 7: 241. 1821. Emergent or, less frequently, canopy trees. Inflorescences in terminal or axillary racemes or paniculate arrangements of racemes. Flowers zygomorphic, appearing with or without leaves. Calyx lobes 6; petals 6. Androecium markedly asymmetrical, the hood spirally coiled inward, then folded outward to form second hood over first, the second hood either thickened and fleshy, smooth to transversely plaited or not thickened but covered with numerous slender, sterile appendages, with a few clavate, sterile appendages in hoods of both types; staminal ring with 10– 65 stamens. Ovary 3-locular; style very short. Fruits dehiscent, cylindrical to campanulate, always longer than broad, woody or thin and coriaceous. Seeds surrounded by a wing, the embryo with foliaceous cotyledons. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 20 species, 5 in Venezuela, all in the flora area. Key to the Species of Couratari 1.

1.

2(1).

2.

Lower surface of leaves with stellate pubescent; flowers with 15–25 stamens; fruits long (to 12 cm) and broad (to 6 cm), the exocarp usually with conspicuous white lenticels ........................................... C. guianensis Lower surface of leaves glabrous; flowers with 30–45 stamens; fruits short (usually < 6 cm) and narrow (usually < 5 cm), the exocarp without conspicuous lenticels .............................................................................. 2 Leaves present at flowering, the blades with conspicuous striations; flowers with pedicels < 10 mm long; petals white; androecial hood echinate externally; fruits with pericarp ca. 2 mm thick .......................... C. stellata Leaves absent at flowering, the blades without conspicuous striations; flowers with pedicels > 10 mm long; petals pink to purple; androecial hood not echinate externally; fruits with pericarp ca. 1 mm thick ...... 3

Couratari 757

3(2). 3.

4(3). 4.

Leaf blades coriaceous, elliptic, the apices acute or bluntly acuminate; petioles thick, 10–14 mm long ............................................... C. sandwithii Leaf blades usually chartaceous to thinly coriaceous, elliptic to narrowly oblong, the apices acute to finely acuminate; petioles slender, 5–25 mm long ......................................................................................................... 4 Petioles 15–25 mm long, the blades oblong to ovate; fruit narrowly cylindrical-campanulate ................................................................. C. multiflora Petioles 5–11 mm long, the blades oblong to oblong-lanceolate; fruit broadly campanulate ............................................................... C. oligantha

Couratari guianensis Aubl., Hist. Pl. Guiane 724, t. 290. 1775, fruit only. —Cachimbo, Capa de tabaco, Sawana (Yekwana), Tapa tabaco. Couratari pulchra Sandwith, Bull. Misc. Inform. Kew 1932: 217. 1932. Canopy or emergent tree, often with welldeveloped buttresses; leaves with dense covering of stellate hairs on petioles and lower surface; flowers present when tree is leafless, pink to purple; androecial hood not echinate externally; fruits long-cylindric, the exocarp with conspicuous, white lenticels. Common in lowland, nonflooded forests, near sea level to 500 m; Delta Amacuro (eastnortheast of El Palmar), Bolívar (Altiplanicie de Nuria, Amaruay-tepui, near El Palmar, Kilómetro 88, La Paragua, Los Patos south of El Manteco, Río Tabaro near Río Nichare), Amazonas (Río Mawarinuma). Monagas; Costa Rica, Panama, Colombia (west of the Andes), Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 639. The inner bark is used for making cigarette paper. Couratari multiflora (Sm.) Eyma, Polygon. Guttif. Surinam 60. 1932. —Lecythis multiflora Sm. in Rees, Cycl. 20: 8. 1812. —Karipon, Tampipio. Lecythis fagifolia Miq. ex O. Berg, Linnea 27: 451. 1856. —Allantoma fagifolia (Miq. ex O. Berg) Miers, Trans. Linn. Soc. London 30(2): 298. 1874. —Couratari fagifolia (Miq. ex O. Berg) Eyma, Polygon. Guttif. Surinam 62. 1932. Canopy or emergent tree; leaves glabrous, relatively small, usually with long, slender petioles, flowering when tree is without leaves; flowers pink to purple; androecial hood not echinate externally; fruits relatively thin and slender. Common in lowland,

nonflooded forests, near sea level to 400 m; Delta Amacuro (Serranía de Imataca), Bolívar (upper Río Caroní, Serranía de Imataca, northeast of Upata). Guyana, Suriname, French Guiana, Amazonian Brazil. ŠFig. 641. Couratari oligantha A.C. Sm., Amer. J. Bot. 26: 411. 1939. Tree to 30 m tall, flowering when tree is leafless; flowers pink; androecial hood not echinate externally. Grows along river banks, usually above flood level, 100–200 m; Amazonas (Maroa, Río Yatúa between Río Yaciba and Piedra Arauicaua). Guyana, Colombia, Peru, Brazil. Couratari oligantha is similar to Couratari multiflora, but the leaves have shorter petioles and the fruits are broader. Couratari sandwithii Prance, Brittonia 33: 19, fig. 4. 1981. Tree to 15 m tall, probably flowering when tree is leafless; flowers unknown. Lowland forests along rivers, ca. 100 m; Bolívar (Río Parguaza between El Carmen and Raudal Maraca). Suriname. Couratari sandwithii is similar to C. multiflora and C. oligantha. Additional collections are needed to establish the differences among these species. Couratari stellata A.C. Sm., Amer. J. Bot. 25: 410. 1939. Emergent tree, usually with buttresses; leaves glabrous, with longitudinal striations on adaxial surface; flowers when leaves are present, the petals white; androecial hood yellow, echinate externally. Common in nonflooded, lowland forests, 50–100 m; Amazonas (Río Cataniapo). Widespread in Guyana, Suriname, French Guiana, and Amazonian Brazil (central and western). ŠFig. 640.

758

L ECYTHIDACEAE

Fig. 639. Couratari guianensis

Fig. 640. Couratari stellata

Couratari 759

Fig. 641. Couratari multiflora

760

L ECYTHIDACEAE

Fig. 642. Couroupita guianensis

Eschweilera 761

5. COUROUPITA Aubl., Hist. Pl. Guiane 708. 1775. Pontopidana Scop., Intr. Hist. Nat. 195. 1777. Elshotzia Neck., Elem. Bot. 2: 256. 1790, non Elsholtzia Willd. 1790. Pekea Juss., Gen. Pl. 249. 1798, pro parte. Pontopidana Steud., Nomencl. Bot. ed. 2, 2: 381. 1841. Medium- to large-sized trees. Leaves born in apical whorls of 7–40, usually with hirsute pubescence in axils of secondary veins. Inflorescences cauliflorous or rarely ramiflorous on oldest branches, usually racemes or infrequently oncebranched paniculate arrangements of racemes, the growth of rachises indeterminate. Flowers zygomorphic. Sepals 6; petals 6. Androecial hood flat, the appendages anther bearing. Ovary 6-locular, the ovules numerous, attached to bilamellar placenta along entire length of locule. Fruit indehiscent, dropping from tree at maturity, round or nearly round, the pericarp fragile, often cracking when fruit hits ground. Seeds numerous, ovate to lenticular, 10–15 mm long, embedded in pulp that oxidizes bluish green when exposed to air, the dry pulp breaking into 6 segments, these crescent-shaped when viewed laterally, wedge-shaped in cross section, the testa hirsute to slightly hairy on exterior; cotyledons foliaceous. El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 3 species, 1 in Venezuela. Couroupita guianensis Aubl., Hist. Pl. Guiane 708, t. 282. 1775. —Mamey hediondo, Muco, Taparo de monte. Couroupita venezuelensis R. Knuth, Repert. Spec. Nov. Regni Veg. 35: 341. 1934. Medium-sized tree; leaves in terminal clusters; inflorescences ramiflorous or mostly cauliflorous; flowers ca. 6 cm diameter; petals yellow abaxially, red adaxially; androecial hood flat, the appendages all anther-bearing; fruits round, falling from tree at maturity, the pericarp thin, often breaking open upon impact. Lowland forests, 50–300 m; Delta

Amacuro (Caño Güiniquina, east-northeast of El Palmar, between Tucupita and La Horqueta), Bolívar (between Puerto Ordaz and San Félix, Represa Guri). Anzoátegui, Barinas, Carabobo, Cojedes, Mérida, Miranda, Monagas, Zulia; Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 642. Couroupita guianensis was collected in Colombia along the Río Orinoco just below the mouth of Río Atabapo and might eventually be found in the flora area in Amazonas state. It is cultivated widely in botanical gardens and as an ornamental throughout the tropics.

6. ESCHWEILERA Mart. ex DC., Prodr. 3: 393. 1828. Jugastrum Miers, Trans. Linn. Soc. London 30(3): 275. 1874. Neohuberia Ledoux, Lecointea 1: 3. 1963. Understory, canopy, or emergent trees. Inflorescences terminal, axillary, ramiflorous, or cauliflorous, racemes, spikes, or paniculate arrangements of racemes or spikes. Flowers zygomorphic. Calyx lobes 6; petals 6. Androecium zygomorphic, the appendages forming complete coil, antherless. Ovary usually 2-, infrequently 4locular, the ovules usually basal, i.e., attached to floor of locule. Fruits dehiscent, woody. Seeds often with lateral aril; embryo undifferentiated. Mexico, Honduras, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Brazil; 90 species, 22 in Venezuela, 17 of these in the flora area. This genus has the widest range and greatest number of species of all neotropical Lecythidaceae. New species are still being discovered and species concepts refined in the genus; therefore this treamtent must be considered provisional.

762

L ECYTHIDACEAE

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

Flowers sessile ............................................................................................ 2 Flowers pedicellate .................................................................................... 3 Largest leaf blades 34–38 × 11–13 cm, with 20–23 pairs of lateral veins; petiole 20–28 mm long; inflorescences branched; flowers with calyx lobes 12–15 × 14–18 mm; fruits with calycine ring inserted at middle, the calyx lobes persistent but scarcely thickened; seeds unknown but probably not wedge-shaped, or completely surrounded by aril, or with apical germination ................................................................ E. rionegrense 2. Largest leaf blades 7.5–17 × 4–8.5 cm, with 10–15 pairs of lateral veins; petiole 5–12 mm long; inflorescences unbranched; flowers with calyx lobes 3 × 4 mm; fruits with calycine ring inserted well above middle, the calyx lobes persistent as woody, thickened knobs; seeds wedgeshaped, completely surrounded by aril, with lateral germination .................................................................................................. E. tenuifolia 3(1). Twigs, especially the younger ones, distinctly winged ...................... E. alata 3. Twigs not winged ........................................................................................ 4 4(3). Bracts and bracteoles persistent at anthesis ............................................ 5 4. Bracts and bracteoles not persistent at anthesis ...................................... 6 5(4). Leaf blades chartaceous, the upper surface not shiny, often undulate, the margins not revolute; petioles 9–18 mm long; inflorescences unbranched; flowers with pedicels 25–40 mm long; petals white .................................................................................................. E. bracteosa 5. Leaf blades coriaceous, the upper surface often shiny, not undulate, the margins revolute; petioles absent to < 5 mm long; inflorescences branched; flowers with pedicels < 5 mm long, petals yellowish cream .................................................................................................... E. revoluta 6(4). Calyx lobes usually ≥ 7 mm wide, imbricate for more than 1/2 their length, not or only slightly gibbous at base abaxially ....................................... 7 6. Calyx lobes usually < 7 mm wide, imbricate for < 1/2 their length, usually gibbous at base abaxially ....................................................................... 9 7(6). Flowers and fruits not turning bluish green when cut or bruised; inflorescence axis and pedicels with dense, yellowish pubescence ... E. roraimensis 7. Flowers and fruits turning bluish green when cut or bruised; inflorescence axis and pedicels without dense, yellowish pubescence ............. 8 8(7). Bark not scalloped; calyx lobes without undulate margins, as long as wide; fruits truncate to pedicel directly below calycine ring, the external surface rough, with vermiculate pattern ........................ E. decolorans 8. Bark scalloped; calyx lobes with undulate margins, longer than wide; fruits rounded to pedicel, the external surface smooth or rough, but without vermiculate pattern .................................................. E. laevicarpa 9(6). Inflorescences unbranched ...................................................................... 10 9. Inflorescences branched ........................................................................... 13 10(9). Plants of periodically flooded, riparian habitats; leaf blades coriaceous; flowers with style well differentiated from summit of ovary; fruits narrowly turbinate, the operculum with narrow umbo ............... E. parvifolia 10. Plants usually of nonflooded habitats; leaf blades chartaceous; flowers usually with style poorly differentiated from summit of ovary; fruits

Eschweilera 763

11(10). 11. 12(11).

12.

13(9). 13. 14(13). 14. 15(14). 15. 16(14).

16.

broadly turbinate or globose, the operculum not umbonate or with broad umbo........................................................................................... 11 Flowers ≥ 3.5 cm diameter; calyx lobes > 6 mm long; androecial hood with 3 inward coils ......................................................................... E. pedicellata Flowers < 3.5 cm diameter; calyx lobes < 6 mm long; androecial hood with 2 inward coils ....................................................................................... 12 Leaf blades with tertiary venation salient on upper and lower surfaces, with 7–10 pairs of lateral veins; flowers with white petals, the pedicels > 10 mm long; fruits broadly turbinate, 3.5–6 cm diameter, with distinct, very woody calyx lobes, the pericarp ca. 5 mm thick ......... E. collina Leaf blades with tertiary venation salient on lower surface only, with 10– 13 pairs of lateral veins; flowers with yellow petals, the pedicels < 10 mm long; fruits globose, 2–3 cm diameter, without woody calyx lobes, the pericarp 2–3 mm thick ............................................................ E. parviflora Upper surface of leaf shiny; fruits with pericarp rough, brown with lighter brown spots ...................................................................... E. subglandulosa Upper surface of leaf not shiny; fruits with relativey smooth pericarp, brown or blackish but without lighter colored spots .......................... 14 Leaf blades dry reddish brown; axes of inflorescence and pedicels densely ferruginous-pubescent ......................................................................... 15 Leaf blades not drying reddish brown; axes of inflorescence and pedicels glabrous or, if pubescent, the pubescence not ferruginous................. 16 Pedicels 10–13 mm long; petals light yellow ............................ E. neblinensis Pedicels 4–7 mm long; petals white .......................................... E. paniculata Trunks usually buttressed; bark without shallow, scallop-like depressions; flowers 3–3.5 cm diameter; fruits globose, the calycine ring inserted near middle ..................................................................... E. coriacea Trunks usually not buttressed; bark with shallow, scallop-like depressions; flowers 3–3.5 cm; fruits cup-shaped, the calycine ring inserted above middle .......................................................................... E. micrantha

Eschweilera alata A.C. Sm., Amer. J. Bot. 26: 407. 1939. Canopy tree; leaf blades 15–27 × 5–10 cm, elliptic, oblong, or narrowly ovate, with 9–12 pairs of lateral veins; petiole 7–12 mm long; inflorescences unbranched, the pedicels 3–20 mm long; flowers 4–5 cm diameter; calyx lobes 4–6 × 4–7 mm, very widely ovate; petals white, sometimes flushed with rose, red, or purple; androecial hood with double coil, yellowish orange; fruits ca. 2–3 × 3–5 cm, the supracalycine zone erect, the calycine ring inserted below middle, the infracalycine zone truncate, the pedicel persisting as woody knob. Nonflooded forests dominated by Mora gonggrijpii, ca. 300 m; Bolívar (El Pauji). Guyana. Eschweilera bracteosa (Poepp. ex O. Berg) Miers, Trans. Linn. Soc. London 30(2):

274. 1874. —Lecythis bracteosa Poepp. ex O. Berg, Linnaea 27: 455. 1856. —Coco de mono. Small tree; leaf blades 11–28 × 4–12.5 cm, oblong to oblong-elliptic, with 10–14 pairs of lateral veins; petiole 9–18 mm long; inflorescences unbranched, the pedicels 25–40 mm long; flowers 4–5 cm diameter; calyx lobes 6– 10 × 6–8 mm, ovate; petals white; androecial hood with double coil, yellow; fruits 2.5 × 5 cm, very broadly turbinate or cup-shaped, the calycine ring inserted near opercular opening, the infracalycine zone rounded to base, with petiole persistent as woody stalk. Lowland, nonflooded forests, ca. 100 m; Amazonas (San Carlos de Río Negro, road between Yavita and Maroa). Peru, Brazil. ŠFig. 644. Eschweilera collina Eyma, Meded. Bot.

764

L ECYTHIDACEAE

Mus. Herb. Rijks Univ. Utrecht 4: 63, fig. 9-III, pl. 1. 1932. —Cacao grande, Camaji, Majaguillo, Majaguillo negro. Canopy tree to 30 m tall, usually unbuttressed, the bark brown, ± smooth, with occasional vertical cracks and well-developed hoop marks; leaf blades 8–14 × 4–8 cm, widely elliptic to elliptic, with 7–10 pairs of lateral veins; petiole 7–12 mm long; inflorescences unbranched, the pedicels 15–25 mm long; flowers ca. 3 cm diameter; calyx lobes 2–4 × 3–5.5 mm, very widely ovate; petals white; androecial hood with double coil, yellow; fruits 3.5–6 × 3.5–6 cm, broadly turbinate, the calyx persistent as prominent woody knobs. Lowland, nonflooded forests, 100–300 m; Delta Amacuro (east-northeast of El Palmar), Bolívar (Altiplanicie de Nuria, El Paraíso on San Félix to Upata road, near Soledad), Amazonas (Caño Yagua, Culebra, lower Río Casiquiare, Río Mawarinuma, Río Yureba, San Carlos de Río Negro). Guyana, Suriname, French Guiana, central Amazonian Brazil. Eschweilera coriacea (DC.) S.A. Mori, Fl. Neotrop. Monogr. 21(2): 203. 1990. —Lecythis coriacea DC., Prodr. 3: 291. 1828. —Cacao, Pocay yek (Arekuna). Lecythis odora Poepp. ex O. Berg in Mart., Fl. Bras. 14(1): 492. 1858. —Eschweilera odora (Poepp. ex O. Berg) Miers, Trans. Linn. Soc. London 30(2): 273. 1874. Canopy tree to 37 m tall, usually with well-developed buttresses, the bark brown to very dark brown or nearly black, not noticeably fissured; leaf blades 9–26 × 4.5–12.5 cm, elliptic to narrowly elliptic or narrowly obovate to oblanceolate, with 9–16 pairs of lateral veins; inflorescences branched, the axes often somewhat zigzagged, the pedicels 10– 20 mm long, often leaving persistent knob after falling; flowers 3.5–5 cm diameter; calyx lobes 3–9 × 3–7 mm, ovate to very widely ovate; petals white or yellow; androecial hood with double coil, yellow; fruits (excluding operculum) 3–4 × 3.5–8 cm, the supracalycine zone erect, the calycine ring inserted near middle, often contracted directly below calycine ring; seeds with welldevloped lateral funicle-aril. Lowland, nonflooded and periodically flooded forests, 100– 500 m; Delta Amacuro (east-northeast of El Palmar), Bolívar (Amaruay-tepui, near Kilómetro 88, Río Acanán, Río Erebato), Ama-

zonas (Río Coro Coro, Río Matacuni, Río Putaco, Yavita to Maroa road). Eastern Panama, northwestern Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Eschweilera decolorans Sandwith, Bull. Misc. Inform. Kew 1932: 214. 1932. —Cacao. Canopy tree to 30 m tall; leaf blades 8.5– 23 × 3.5–8.5 cm, widely elliptic to elliptic, with 9–14 pairs of lateral veins; petiole 6–12 mm long; inflorescences usually unbranched, the pedicels glabrous, 1.5–13 mm long; flowers 5–6 cm diameter; calyx lobes 4.5–13 × 6– 12 mm, widely to very widely ovate; petals white; androecial hood with double coil, yellow; fruits 4–5 × 7–8 cm, depressed globose, truncate directly below calycine ring, the calycine ring inserted below middle, the pericarp very rough, with vermiculate pattern. Lowland, nonflooded forests, 100–500 m; Delta Amacuro (east-northeast of El Palmar, Río Cuyubini), Bolívar (El Paraíso on San Félix to Upata road, Río Tírica), Amazonas (Río Cataniapo). Colombia, Guyana, Suriname, French Guiana, Brazil. ŠFig. 645. Eschweilera laevicarpa S.A. Mori, Mem. New York Bot. Gard. 44: 32. 1987. —Corrona (Curripaco), Oroma (Yekwana), Warapa (Yanomami). Canopy tree to 30 m tall; leaf blades 9–22 × 5–8 cm, elliptic, with 9–13 pairs of lateral veins; petiole 7–15 mm long; inflorescences usually unbranched, the pedicels glabrous, ca. 12 mm long; flowers ca. 3–5 cm diameter; calyx lobes 12–17 × 11–14 mm, widely oblong; petals white; androecial hood with double coil, white outside, yellow inside; fruits 2.5–5 × 4.4–7 cm, cup-shaped, rounded to base, the calycine ring inserted above middle, the pericarp smooth. Evergreen lowland to lower montane forests, 50–600 m; Bolívar (Río Nichare), Amazonas (Cerro El Danto east of San Juan, Río Parú, Serranía Batata ca. 55 km southeast of Puerto Ayacucho). French Guiana, Colombia, Ecuador, Peru, Brazil. The bark of the tree is used as cigarette or cigar paper. Eschweilera micrantha (O. Berg) Miers, Trans. Linn. Soc. London 30(2): 160. 1874. —Lecythis micrantha O. Berg,

Eschweilera 765

Linnaea 27: 454. 1854. —Coco de mono, Copa de tabaca. Lecythis gracilipes Sagot, Ann. Sci. Nat. Bot. sér. 6, 20: 203. 1885. —Eschweilera gracilipes (Sagot) R. Knuth in Engl., Pflanzenr. IV. 219a(Heft 105): 101. 1939. Eschweilera floribunda Eyma, Polygon. Guttif. Surinam 74, pl. 2. 1932. Canopy tree to 30 m tall, the bark gray to dark brown, with irregular depressions left by sloughing plates; leaf blades 10–21 × 4–7 cm, elliptic, infrequently narrowly ovate, with 9–12 pairs of lateral veins; petiole 5–9 mm long; inflorescences usually branched, the pedicels 6–10 mm long; flowers 1.5–2.5 cm diameter; calyx lobes widely ovate, 1.5–3 × 1.4–2.5 mm; petals usually white, infrequently yellow; androecial hood with double coil, yellow; fruits (excluding operculum and pedicel) 2–3 × 4–5 cm, cup-shaped, the calycine ring inserted above middle. Lowland, usually nonflooded forests, 100–500 m; Bolívar (Río Tonoro), Amazonas (Río Casiquiare, Río Coro Coro, Río Padamo, between San Carlos de Río Negro and Solano, Yavita). Guyana, Suriname, French Guiana, Peru, Brazil. Eschweilera neblinensis S.A. Mori, Fl. Neotrop. Monogr. 21(2): 256, fig. 89. 1990. Understory tree to 12 m tall; leaf blades 17–25 × 7–10.5 cm, usually elliptic, less frequently widely elliptic, drying characteristicly reddish brown; petiole 8–15 mm long; inflorescences once-branched, the pedicels 10–13 mm long; flowers 3–3.5 cm diameter; calyx lobes 4.5–5 × 4–5 mm, widely to very widely ovate; petals light yellow; androecial hood with double coil, yellow; fruits (excluding operculum) 1.7 × 5 cm, cup-shaped, the calycine ring inserted well above middle, the pedicel persistent as woody stalk. Lowland, nonflooded forests, 100–200 m; Amazonas (base of Sierra de la Neblina). Endemic. Eschweilera paniculata (O. Berg) Miers, Trans. Linn. Soc. London 30(2): 265. 1874. —Lecythis paniculata O. Berg in Mart., Fl. Bras. 14(1): 501. 1858. Tree; leaf blades 20–25 × 7–9 cm, oblong, with 12–13 pairs of lateral veins, drying reddish brown; petiole 12–16 mm long; inflorescences branched, the pedicels 4–7 mm long; flowers ca. 2 cm diameter; calyx lobes 5 × 2.5

mm, narrowly ovate; petals white; style geniculate, well-defined from summit of ovary; fruits unknown. Lowland forests, 100–200 m; Amazonas (Río Siapa). Brazil. Eschweilera parviflora (Aubl.) Miers, Trans. Linn. Soc. London 30(2): 260, pl. 60, fig. 19. 1874. —Lecythis parviflora Aubl., Hist. Pl. Guiane 717, t. 285, 287. 1775. —Cacaíto. Eschweilera grata Sandwith, Bull. Misc. Inform. Kew 1932: 216. 1932. Understory tree to 25 m tall; the bark often with vertically disposed lenticels and very shallow cracks; leaf blades 7.5–15.5 × 2.5–7.5 cm, elliptic, with 10–13 pairs of lateral veins; petiole 5–10 mm long; inflorescences unbranched, the pedicels 8–10 mm long; flowers 2–2.5 cm diameter; calyx lobes 2.5–6 × 2–5.5 mm, widely to very widely ovate; petals yellow; androecial hood with double coil, yellow; fruits (excluding operculum) 1.5–2 × 2–3 cm, cup-shaped, the pericarp 2–3 mm thick. Lowland, nonflooded forests, 100–600 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (south of El Dorado, El Paraíso on San Félix to Upata road), Amazonas (Capibara, La Esmeralda, 10 km east of Tamatama). Guyana, Suriname, French Guiana, Brazil. ŠFig. 643. Eschweilera parvifolia Mart. ex DC., Prodr. 3: 293. 1828. —Coco de mono. Lecythis chaffanjonii Benoist, Notul. Syst. (Paris) 3: 179. 1915. —Eschweilera chaffanjonii (Benoist) R. Knuth in Engl., Pflanzenr. IV. 219a(Heft 105): 97. 1939. Tree to 20 m tall; leaf blades ovate to narrowly ovate, with 7–9 pairs of lateral veins; inflorescences unbranched, the pedicels 7–12 mm long; flowers ca. 3 cm diameter; calyx lobes 2.5–4.5 × 1.6–3.7 mm, very widely ovate to ovate; petals usually white, less frequently yellow; androecial hood with double coil, less frequently with incipient triple coil, yellow; fruits turbinate, the calycine ring inserted near opercular opening, often prominent. Lowland, periodically flooded forests along rivers, 50–200 m; Bolívar (Río Parguaza), Amazonas (widespread). Apure, Mérida, Zulia; Guyana, Ecuador, Peru, Brazil, Bolivia. ŠFig. 647. Eschweilera pedicellata (Rich.) S.A. Mori, Mem. New York Bot. Gard. 44: 34. 1987.

766

L ECYTHIDACEAE

—Lecythis pedicellata Rich., Actes Soc. Hist. Nat. Paris 1: 111. 1792. —Cacaíto, Cacao, Jocotouji (Yanomami). Lecythis longipes Poit., Mém. Mus. Hist. Nat. 13: 144, pl. 1. 1825. —Eschweilera longipes (Poit.) Miers, Trans. Linn. Soc. London 30(2): 253, pl. 60, figs. 12, 13. 1874. Understory tree to 20 m tall; leaf blades 9–26 × 4.5–9.5 cm, elliptic to oblanceolate, with 10–13 pairs of lateral veins; petiole 4– 11 mm long; inflorescence unbranched, the pedicels 10–35 mm long; flowers 3.5–5 cm diameter; calyx lobes 6–11 × 4–7 mm, widely to narrowly ovate; petals white, often tinged with pink or yellow; androecial hood with triple coil; fruits 2–4 × 2.5–5 cm, broadly turbinate. Lowland, nonflooded and riparian forests, often found on rocky river banks, 100–400 m; Bolívar (Kilómetro 88, upper Río Caura, Río Samay in Río Icabarú basin), Amazonas (Cerro Huachamacari, Culebra, upper Río Orinoco at Salto Siro Vásquez, Río Padamo, Río Siapa, Río Yureba, Yavita to Maroa road). Guyana, Suriname, French Guiana, Brazil. Eschweilera revoluta S.A. Mori, Fl. Neotrop. Monogr. 21(2): 174. 1990. Tree to 35 m tall, the bark brown, flaky, the inner bark flesh-colored; leaf blades 12– 19 × 5–9 cm, elliptic, with 9–11 pairs of lateral veins; petiole absent or to 5 mm long; inflorescences once-branched, the pedicels ca. 5 mm long, with persistent bracts and bracteoles; flowers with foetid odor, ca. 2.5– 3.5 cm diameter; calyx lobes 4–5 × 5–6 mm, widely to very widely ovate; petals yellowish cream; androecial hood with double coil, lemon-yellow; fruits cup-shaped to very broadly cup-shaped, the calycine ring inserted above middle, the infracalycine zone rounded to base, the pedicel persistent as woody stalk. Lowland, nonflooded forests, 100–200 m; Amazonas (Río Mawarinuma). Endemic. Eschweilera rionegrense S.A. Mori, Fl. Neotrop. Monogr. 21(2): 223, fig. 81. 1990. Canopy tree; leaf blades 34–38 × 11–13 cm, narrowly oblong, with 20–23 pairs of lateral veins; petiole 20–28 mm long. Inflorescences once-branched, spicate, the axes an-

gular, the flowers sessile; flowers 4.5–6 cm diameter; calyx lobes 12–15 × 14–18 mm, very widely ovate; petals white or yellowish tinged with rose; androecial hood forming double coil; fruits globose, the calyx lobes persistent, the supracalycine zone erect, the infracalycine zone round, 5 × 7 cm (excluding operculum); seeds not seen. Amazonas (Río Mawarinuma). Brazil (Amazonas: Rio Negro). Eschweilera roraimensis S.A. Mori, Fl. Neotrop. Monogr. 21(2): 237, fig. 84. 1990. Canopy tree to 30 m tall; the bark brown, with longitudinal fissures, the inner bark red to reddish brown; leaf blades 12–21.5 × 6–11 cm, elliptic to oblong, with 12–16 pairs of lateral veins; petiole 8–20 mm long; inflorescences usually unbranched, the pedicels 5–7 mm long; flowers 4–5 cm diameter; calyx lobes 11–13 × 9–10 mm, widely to very widely oblong or widely to very widely ovate; petals white; androecial hood with double coil, pale yellow; fruits 4–5 × 6–7.5 cm, (excluding pedicel and operculum) broadly turbinate, the calycine ring inserted at or above middle. Nonflooded forests, 700–1100 m; Amazonas (Simarawochi). Brazil (Amazonas: Sierra Parima). Eschweilera subglandulosa (Steud. ex O. Berg) Miers, Trans. Linn. Soc. London 30(2): 266. 1874. —Lecythis subglandulosa Steud. ex O. Berg, Linnaea 27: 459. 1854. —Cacao, Camaji, Cascarare, Coco de mono, Majagüillo, Ollita, Oripopo, Tabarí, Tampipio, Tapa de tabaca. Canopy tree to 35 m tall, frequently buttressed, the bark light gray to dark brown, peeling in large, thin, irregular plates; leaf blades 12–30 × 6–11 cm, elliptic to narrowly ovate, the upper surface smooth and shiny; petiole 10–15 mm; inflorescences oncebranched, the pedicels 5–15 mm long; flowers 2.5–3.5 cm diameter; calyx lobes 4–8 × 3– 6 mm, widely to very widely ovate; petals white or yellow; androecium with double coil, yellow; fruits 2–3.5 × 3–5.5 cm, globose to turbinate, often with prolonged woody knob at base, the calycine ring inserted near middle, the pericarp rough, brown with lighter brown spots. Common in nonflooded and periodically flooded lowland forests,

Eschweilera 767

Fig. 643. Eschweilera parviflora

Fig. 644. Eschweilera bracteosa

768

L ECYTHIDACEAE

Fig. 645. Eschweilera decolorans

Fig. 646. Eschweilera tenuifolia

Eschweilera 769

Fig. 647. Eschweilera parvifolia

Fig. 648. Eschweilera subglandulosa

770

L ECYTHIDACEAE

often found in forest patches and gallery forests in savanna areas, 100–500 m; Delta Amacuro (Serranía de Imataca), northern Bolívar, Amazonas (Boca Mavaca, Caño Caname, Culebra, near Puerto Ayacucho). Northern coastal Venezuela; Trinidad, Guyana, Suriname, northern Brazil. ŠFig. 648. A water extract from the bark is said to be a remedy for colitis and dysentery. Fiber from the bark is used as a cord, and the timber makes good fence posts because it is resistent to rot. Eschweilera tenuifolia (O. Berg) Miers, Trans. Linn. Soc. London 30: 266. 1874. —Lecythis tenuifolia O. Berg in Mart., Fl. Bras. 14(1): 502. 1858. —Coco de mono, Coco de mono rebalsero, Totuma de mono. Jugastrum obtectum Miers, Trans. Linn. Soc. London 30: 276, t. 61, fig. 7, 8. 1874. —Eschweilera obtecta (Miers) Nied. in Engl. & Prantl, Nat. Pflanzenfam. 3(7): 41. 1892. Jugastrum christii Pittier, Arb. Arbust.

Venez. 4/5 [reprinted from Bol. Minist. RR. EE. no. 12]: 66. 1926. Jugastrum sifontesii Pittier, Bol. Soc. Venez. Ci. Nat. 3: 431. 1937. Tree to 15 m tall, the bark gray, fissured; leaf blades 7.5–17 × 4–8.5 cm, elliptic to oblong-lanceolate, with 10–15 pairs of lateral veins; petiole 5–12 mm; inflorescences unbranched, the flowers subsessile; flowers 3.5–5 cm diameter; calyx lobes triangular, ca. 3 × 4 mm; petals 6, usually white or cream, less frequently light yellow; androecial hood with double coil, yellow to golden; fruits depressed globose, the calycine ring inserted 3–12 mm below opercular opening; seeds wedge-shaped, surrounded by aril, germinating laterally. Rivers and streams, 50– 100 m; Bolívar (near Ciudad Bolívar, Río Caura, Río Cuchivero, Río Orinoco at Río Suapure), Amazonas (Río Casiquiare, Río Guainía, Río Yatúa, Samariapo, San Carlos de Río Negro, Yavita). Apure, Guárico; Amazonian Brazil. ŠFig. 646. The bark is reported by some collectors to be medicinal.

7. GUSTAVIA L., Pl. Surin. 17, t. 12. 1775, nom. cons. Japarandiba Adans., Fam. Nat. Pl. 2: 448. 1763. Pirigara Aubl., Hist. Pl. Guiane 487. 1775. Teichmeyeria Scop., Intr. Hist. Nat. 267. 1777. Spallanzania Neck., Elem. Bot. 2: 79. 1790. Pirigaria Span., Linnaea 15: 204. 1841. Understory trees, infrequently to 30 m tall; growth form either single-stemmed or few-branched with clusters of large leaves at ends of thickened branches (pachycaul) or with single dominant trunk and much-branched crown with smaller leaves at ends of slender branches (leptocaul), the leaves often ± congested at branch ends. Inflorescences terminal, axillary, or cauline, solitary or racemose. Flowers actinomorphic, the largest of neotropical members of family, to 20 cm diameter. Calyx entire or 4–6-lobed; petals 6 or 8. Androecium with 500–1210 stamens, all fertile, fused at bases into symmetrical ring, this adnate to bases of petals and summit of ovary; anthers dehiscing by 2 apical pores. Ovary with 4–6(–10) locules, each locule with 7–93 ovules attached to expanded placenta occupying upper 1/2 of septum, the style < 5 mm long. Fruits indehsicent, berry-like, sometimes becoming woody, normally releasing seeds by deliquescence of pericarp, in some species appearing dehiscent because of rotting away of weaker fruit summit. Seeds of 2 types, without welldeveloped funicles or with yellow, expanded, contorted funicles; embryo with large, fleshy, plano-convex cotyledons and minute hypocotyl and plumule. Widely distributed in southwestern Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, northern Ecuador, Peru, Brazil; 41 species, 11 in Venezuela, 6 of these in the flora area.

Gustavia 771

Key to the Species of Gustavia 1.

1.

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

4.

5(2). 5. 6(5). 6.

Inflorescences always terminal; flowers with 6 petals; calyx lobes 6, with inverted Y-shaped thickenings on adaxial surfaces; hypanthium with strongly developed costae, these persistent on fruits ........... G. hexapetala Inflorescences usually cauline, if terminal then with terminal and axillary inflorecences also present on same tree; flowers with 8 petals; calyx lobes 4, 6 or absent; hypanthium without costae or if costae present weakly developed and not persisting on fruits ..................................... 2 Largest leaf blades usually > 30 cm long, usually with > 25 pairs of lateral veins ....................................................................................................... 3 Largest leaf blades usually < 30 cm long, usually with < 25 pairs of lateral veins ....................................................................................................... 5 Plants usually found in periodically flooded forests along rivers; calyx with 6 lobes; hypanthium usually inconspicuously costate ...... G. pulchra Plants usually of nonflooded forests; calyx with 4 lobes; hypanthium not costate .................................................................................................... 4 Leaf blades < 15 cm wide, with conspicuous marginal vein and strongly percurrent tertiary venation; petiole 0–15 mm long; flowers ca. 8 cm diameter ................................................................................. G. acuminata Leaf blades > 15 cm wide, without conspicuous marginal vein and without strongly percurrent tertiary venation; flowers > 10 cm diameter ......... .................................................................................................... G. coriacea Inflorescences cauline; calyx with 6 lobes; hypanthium inconspicuously costate ......................................................................................... G. pulchra Inflorescences cauline, axillary, or terminal; calyx entire or with 4 lobes; hypanthium not costate ......................................................................... 6 Leaf blades cuneate to base; inflorescences axillary, terminal, or cauline, with well-defined rachis, usually with > 1 flower..................... G. augusta Leaf blades slightly auriculate at base; inflorescences axillary, without well-defined rachis, usually with 1 flower ......................... G. poeppigiana

Gustavia acuminata S.A. Mori, Fl. Neotrop. Monogr. 21(1): 185, figs. 54, 56. 1979. —Cachimbo, Palo cachimbo. Usually unbranched, understory tree, < 10 m tall, often flowers when < 5 m; leaf blades 30–70 × 8.5–14 cm, oblanceolate, with 25–40 pairs of relatively closely spaced lateral veins, the tertiary veins more markedly percurrent than in other species, with marginal vein well developed; petioles usually appearing absent because of narrowly decurrent blade, sometimes to 15 mm long; inflorescences cauline; flowers ca. 8 cm diameter; calyx with 4 broadly triangular lobes; petals 8, white; hypanthium not costate; staminal ring yellow, the filaments yellow or purple; fruits without persistent calyx lobes and cos-

tae; seeds with yellow, contorted funicle. Nonflooded forests, 100–200 m; Amazonas (Río Casiquiare, road between San Carlos de Río Negro and Solano). Brazil (Amazonas). Gustavia augusta L., Fl. Surin. 17, t. 12. 1775. —Coco de mono, Falso coco de mono, Guatero, Guatoso, Muerto, Rabo pelado, Rosa de muerto, Vela. Understory tree, often much branched, sometimes shrub-like, to 20 m tall; leaf blades 16–48 × 4–13 cm, narrowly obovate to oblanceolate, with 14–22 pairs of lateral veins; petiole 0–40 mm long; inflorescences terminal, axillary, or cauline; flowers 9–20 cm diameter; calyx entire or with 4 rounded

772

L ECYTHIDACEAE

lobes; petals 8, white, often tinged with pink on outside; hypanthium not costate; staminal ring yellow or white, the upper half of filaments often pink; fruits without persistent calyx and costae; seeds with yellow, contorted funicle. Lowland forests along rivers and on well-drained sites, near sea level to 400(–1000) m; Delta Amacuro (Caño Arature, Caño Corisal, Caño Güiniquina, Cerro Lucupanar, east of Los Castillos), northern Bolívar, Amazonas (Río Casiquiare, Río Cuao). Anzoátegui, Apure, Barinas, Mérida, Monagas, Sucre; Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian and northeastern coastal Brazil, Bolivia, occasionally cultivated in botanical gardens elsewhere. ŠFig. 649. Some of the common names allude to the nauseous odor produced by the rotting wood. Gustavia coriacea S.A. Mori, Fl. Neotrop. Monogr. 21(1): 184, fig. 54. 1979. Small understory tree to ca. 5 m tall; leaf blades 70–85 × 20–22 cm, oblanceolate, with 25–28 pairs of lateral veins; petiole 15–21 mm long; inflorescences from branches or trunk; flowers 17–20 cm diameter; calyx lobes 4, broadly triangular; petals 8(9), white, flushed with pink abaxially; hypanthium without costae; androecium yellow inside, white outside; fruits known only from immature material; seeds with funicle. Lowland forests, 100–200 m; Amazonas (Río Casiquiare, Río Pasimoni, Río Yatúa). Endemic. Gustavia hexapetala (Aubl.) Sm. in Rees, Cycl. 17: 2. 1819. —Pirigara hexapetala Aubl., Hist. Pl. Guiane 490, t. 193. 1775. Understory tree to 20 m tall; leaf blades 10–24 × 3–13 cm, elliptic, oblanceolate, or obovate, with 9–13 pairs of lateral veins; petiole 2–17 mm long; inflorescences terminal; flowers 6–9 cm diameter; calyx lobes 6, with inverted, adaxial, Y-shaped thickenings; petals 6, white; hypanthium costate; staminal ring white on outside yellow on inside, the filaments without pink; fruits with persistent calyx lobes and costae, yellowish gold at maturity; seeds with straight, poorly developed funicle. Lowland, nonflooded forests, periodically flooded forests along rivers, 50–600 m; Bolívar (Río Parguaza), Amazonas (Caño Yagual above mouth of Río

Atabapo, Isla Ratón, Río Cuao, between Tamatama and La Esmeralda). Aragua, Carabobo, Distrito Federal, Falcón, Mérida, Miranda, Táchira, Yaracuy, Zulia; widespread throughout Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia. ŠFig. 651. Gustavia poeppigiana O. Berg in Mart., Fl. Bras. 14(1): 471. 1858. —Carutillo, Cola de pava, Guatero, Guatoso, Koyamo. Gustavia yaracuyensis Pittier, Arb. Arbust. Venez. 4/5 [reprinted from Bol. Minist. RR. EE. no. 12]: 63. 1926. Understory tree to 15 m tall; leaf blades 15.5–41 × 4.5–11.5 cm, narrowly obovate or oblanceolate, with 10–18 pairs of lateral veins; petiole 1–12 mm long; inflorescences axillary, usually with a single flower; flowers 12–18 cm diameter; calyx lobes 4, broadly triangular or rounded; petals white, often tinged with pink on outside; hypanthium without costae; staminal ring white on outside, yellow on inside, the filaments usually white, infrequently with pink toward apex; fruits without persistent calyx and costae; seeds with contorted, fleshy, yellow funicle. Lowland forests along rivers, forests on welldrained sites, near sea level to 300(–1000) m; Delta Amacuro (Río Cuyubini, Curiapo, south of Sacupana, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Río Caura, Río Paragua), Amazonas (Boca Mavaca). Apure, Barinas, Carabobo, Falcón, Lara, Mérida, Sucre, Táchira, Yaracuy, Zulia; Colombia, Guyana, Peru, Brazil, Bolivia. ŠFig. 650. Gustavia pulchra Miers, Trans. Linn. Soc. London 30(2): 182. 1874. —Cachimbo de mono, Coco de mono, Palo de cachimbo de rebalse, Palo de rabipelado, Vara de muerto, Vela de muerto. Gustavia acuta S.A. Mori, Fl. Neotrop. Mongr. 21(1): 180, fig. 53. 1979. Understory tree to 20 m tall; leaf blades 25–61 × 6–13 cm, with 21–29 pairs of lateral veins; petiole 2–40 mm long; Inflorescences cauline; flowers 12–14 cm diameter; calyx with 6, triangular, poorly developed lobes; petals 8, white with flushes of pink on outside; hypathium often obscurely costate; staminal ring often yellow on inside, white

Gustavia 773

on outside, the filaments often pink on upper half; fruits without persistent calyx lobes and costae; seeds black, with fleshy, yellow funicle. Lowland forests mostly along rivers but occasionally found on well-drained sites,

50–300 m; Amazonas (Caño Cupueini near San Fernando de Atabapo, La Esmeralda, Puerto Ayacucho, Río Casiquiare, upper Río Negro, Río Ventuari, Tamatama). Apure; Brazil.

Fig. 649. Gustavia augusta

774

L ECYTHIDACEAE

Fig. 650. Gustavia poeppigiana

Fig. 651. Gustavia hexapetala

8. LECYTHIS Loefl., Iter Hispan. 189. 1758. Bergena Adans., Fam. Pl. 2: 345. 1763. Chytroma Miers, Trans. Linn. Soc. London 30: 229, t. 58. 1874. Cercophora Miers, Trans. Linn. Soc. London 30: 301, pl. 36B. 1874. Holopyxidium Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 152. 1925. Sapucaya R. Knuth in Engl., Pflanzenr. IV. 219a(Heft 105): 73, fig. 4D. 1939. Trees, small to very large, to 50 m tall, with leptocaulous growth form. Bark nearly smooth to deeply fissured, the outer bark frequently laminated. Leaves deciduous or evergreen, medium-sized, never densely congested at branch ends. Inflo-

Lecythis 775

rescences terminal, axillary, or ramiflorous, spicate or racemose. Flowers zygomorphic. Calyx with 6 lobes, these with or without mucilage-bearing ducts; petals 6. Androecial hood flat, dorsiventrally expanded, or forming partial coil, the appendages with or without anthers; staminal ring with 70–1000 stamens; anthers dehiscing via lateral slits; pollen all same color or some anthers with white and others with yellow. Ovary (3)4(–6)-locular, the summit usually truncate; style usually well differentiated, 1.5–11 mm long, erect, slanted toward anterior end of flower, or geniculate, the ovules inserted on lower part of septum. Fruits woody, circumcissile capsules, the base usually remaining in tree after dehiscence. Seeds usually with basal aril, less frequently with lateral aril on lower half, the embryo without well-developed cotyledons. Neotropics; 26 species, 6 in Venezuela, 4 of these in the flora area. Key to the Species of Lecythis 1. 1.

2(1).

2.

3(2).

3.

Flowers with pink to red petals throughout; androecial hood dorsiventrally expanded, the appendages fused; hypanthium rugose .......... L. corrugata Flowers with white or yellow petals, sometimes tinged with pink or purple on the margins, the hood not dorsiventrally expanded, the appendages free; hypanthium not rugose ................................................. 2 Flowers 4–5 cm diameter; androecial hood flat, the proximal appendages with anthers; style < 2 mm long, erect, with expanded annular ring to ward apex; fruits > 7.5 cm diam ................................................ L. zabucajo Flowers 2–3 cm diameter; androecial hood not flat, the appendages antherless; style 2–6.5 mm long, oblique, without expanded annular ring toward apex; fruits < 7.5 cm diameter .......................................... 3 Leaf blades with 16–18 pairs of lateral veins; flowers with calyx lobes 9– 14 × 8–12 mm; style 5–6.5 mm long; ovary with mucilage-filled ducts as seen in cross section ................................................................... L. alutacea Leaf blades with 9–14 pairs of lateral veins; flowers with calyx lobes 3–6 × 2.5–4 mm; style 2–3.5 mm long; ovary without mucilage-filled ducts as seen in cross section ................................................................. L. chartacea

Lecythis alutacea (A.C. Sm.) S.A. Mori, Brittonia 33: 362. 1981. —Eschweilera alutacea A.C. Sm., Amer. J. Bot. 26: 409. 1939. —Comé-yek. Lecythis karuaiensis Steyerm., Fieldiana, Bot. 28: 422. 1952. Tree to 25 m tall; bark reddish brown, with longitudinal fissures; leaf blades 18–26 × 8–11.5 cm, elliptic or narrowly ovate, with 16–18 pairs of lateral veins; petiole 5–13 mm long; inflorescences branched or unbranched, the pedicels lacking; flowers ca. 2–3 cm diameter; calyx lobes 9–14 × 8–12 mm, very widely ovate, with longitudinally oriented mucilage-bearing ducts; petals white- or yellow-tinged; androecial hood with appendages swept inward, free from one another, the

staminal ring with 237–300 stamens, the anthers all same color; style oblique, 5–6.5 mm long; fruits turbinate, incompletely known. Upland forests bordering savannas, 200–700 m; Bolívar (Río Caruay), Amazonas (Cerro Huachamacari, 45 km southeast or Puerto Ayacucho). Guyana, Suriname, Brazil (northern Pará). The bark of Lecythis alutacea serves for cordage and for head bands. Lecythis chartacea O. Berg, Linnaea 27: 450. 1856. —Chytroma chartacea (O. Berg) Miers, Trans. Linn. Soc. London 30: 231. 1874. —Eschweilera chartacea (O. Berg) Eyma, Polygon. Guttif. Surinam 70, fig. 9-II. 1932. —Cachimbo,

776

L ECYTHIDACEAE

Coco de mono, Guacharaco amarillo, Guacharaco blanco, Tutuma de mono. Cercophora anomala Miers, Trans. Linn. Soc. London 30: 301, fig. 36B. 1874. Canopy tree to 35 m tall, with straight unbuttressed boles, often swollen at very base; bark brown, with vertical fissures; leaf blades 6.5–11.5 × 2–6 cm, narrowly to widely elliptic or narrowly to widely oblong, with 9– 14 pairs of lateral veins; petiole 6–12 mm long; inflorescences unbranched or oncebranched, the pedicels 3.5–4.5 mm long; flowers ca. 2.5 cm diameter, calyx lobes 3–6 × 2.5–4 mm, ovate to widely ovate or widely to very widely oblong; petals usually white, infrequently yellow; androecial hood with appendages swept inward, free from one another, the staminal ring with 63–125 stamens, the anthers all same color; style oblique, 2–3.5 cm long; fruits 4–5 × 5 cm, turbinate. Lowland forests along rivers, welldrained forests at margins of savannas, 100– 300 m; Delta Amacuro (east-northeast of El Palmar), Bolívar (Altiplanicie de Nuria), Amazonas (Río Sipapo, San Carlos de Río Negro, Santa Bárbara, Solano). Widespread throughout Guyana, Suriname, French Guiana, Colombia, Peru, Brazil. ŠFig. 653. Lecythis corrugata Poit., Mém. Mus. Hist. Nat. 13: 146, t. 2. 1825. —Eschweilera corrugata (Poit.) Miers, Trans. Linn. Soc. London 30: 253. 1874. —Chytroma corrugata (Poit.) R. Knuth in Engl., Pflanzenr. IV. 219a(Heft 105): 78. 1939. Canopy tree to 35 m tall, usually unbuttressed; bark brown or grayish brown, with shallow, vertical fissures; leaf blades 8– 25 × 4–10 cm, oblong, elliptic, or infrequently oblanceolate, with 11–23 pairs of lateral veins; petiole 10–25 mm long; inflorescences unbranched or once-branched, the pedicels 2–8 mm long; flowers 2.5–3 cm diameter; calyx lobes 2–8 × 2–5 mm, ovate to widely ovate; petals pink, red, violet, or purple; androecial hood flat but dorsiventrally thickened, the appendages fused together, the staminal ring with 150–190 stamens, the anthers on ligular side of ring white, the rest yellow; style oblique, 3–5 mm long; hypanthium rugose; fruits 2.5–5 × 2.5–6 cm, broadly conical, turbinate, or globose, rugose. Venezuela, Guyana, Suriname, French Guiana, eastern Amazonian Brazil; 2 subspecies; both in the flora area.

Key to the Subspecies of L. corrugata 1. Leaf blades with upper surface dull, without prominent, longitudinally oriented striations ................. subsp. corrugata 1. Leaf blades with upper surface shiny, with prominent, longitudinally oriented striations ........................ subsp. rosea L. corrugata subsp. corrugata. —Guacharaco, Guacharaco rojo, Guacharaco rosado. Lowland forests of different types, ranging from swamp to semideciduous forests, 50–500 m; Delta Amacuro (Serranía de Imataca). Guyana, Suriname, French Guiana, eastern Amazonian Brazil. ŠFig. 652. L. corrugata subsp. rosea (Spruce ex O. Berg) S.A. Mori, Brittonia 33: 363, fig. 3. 1981. —Lecythis rosea Spruce ex O. Berg in Mart., Fl. Bras. 14(1): 488. 1858. —Chytroma rosea (Spruce ex O. Berg) Miers, Trans. Linn. Soc. London 30: 242. 1874. —Cabullo, Camaji, Coco de mono, Coquito, Tabarí, Tahuari rosado. Eschweilera conduplicata A.C. Sm., Amer. J. Bot. 26: 408. 1939. Lowland forests, ranging from riparian to semideciduous forests on well-drained sites, 50–400 m; Bolívar (Aparurén, Río Asa, Río Nichare, Río Parguaza), Amazonas (Culebra, northeast of Puerto Ayacucho). Apure, Mérida, Táchira, Zulia; Brazil (Roraima). ŠFig. 655. The fibrous bark of Lecythis corrugata subsp. rosea is used for making cigarette paper and for cordage and head bands. Lecythis zabucajo Aubl., Hist. Pl. Guiane 718. 1775. —Coco de mono, Tinajito. Lecythis davisii Sandwith, Bull. Misc. Inform. Kew 1932: 213. 1932. Lecythis davisii var. gracilipes Eyma, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 4: 77. 1932. Canopy or emergent tree to 55 m tall; bark brown, with deep vertical fissures, the outer bark laminated; leaf blades 6–11.5 × 2– 5.5 cm, narrowly to widely elliptic, with 10– 16 pairs of lateral veins; petiole 3–10 mm long; inflorescences unbranched, the pedicels 3–5 mm long; flowers 4–5 cm diameter; calyx lobes 5–10 × 5–9 mm, widely to very widely ovate; petals white or light green, often

Lecythis 777

Fig. 652. Lecythis corrugata subsp. corrugata

Fig. 653. Lecythis chartacea

778

L ECYTHIDACEAE

Fig. 654. Lecythis zabucajo

Fig. 655. Lecythis corrugata subsp. rosea

Lemna 779

tinged with various degrees of purple on the apices and margins; androecial hood flat, the appendages not swept inward, free from one another, the proximal-most appendages with anthers, the staminal ring with 370–510 stamens, the anthers all same color; style erect, with expanded annular ring toward apex, 1.5–2 mm long; fruits 6–16.5 × 7.5–17.5 cm, globose or turbinate, the largest fruits of the family in the flora area; seeds longitudinally

sulcate, with well-developed cord-like funicle surrounded by large, fleshy aril at base. Lowland, nonflooded forests, 100–700 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Los Patos south of El Manteco, Río Ayaiche near base of Sierra de Lema, Río Cuyuní, Amazonas (San Carlos de Río Negro). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ŠFig. 654.

LEMNACEAE by Lawrence J. Davenport Reduced, thalloid annuals, free-floating on water’s surface or slightly below, the young plants often initially remaining attached to the parent by short stipes. Fronds 0.5–8 × 0.5–4 mm, fleshy to membranous, subglobular to flat, orbicular to linear, symmetrical to asymmetrical, rounded to pointed at apices, with 0–many veins, with 0–3 papules on the dorsal surface, with or without prominent dots; roots 0–many, unbranched, sheathed, the sheaths winged or smooth; prophyllum (ventral scale) present or absent, if present then penetrated by 1–5 roots. Reproduction mostly vegetative, by budding from 1 or 2 lateral pouches or from a basal pouch. Inflorescence, if produced, borne in 1 of the 2 lateral pouches or in 1 or 2 dorsal pouches; lateral inflorescence of 2 or 3 staminate and 1 carpellate flowers, subtended by a membranous spathe; dorsal inflorescence of 1 staminate and 1 carpellate flower, the spathe absent; staminate flowers of 1 stamen; carpellate flowers hypogynous, 1-locular, with a short terminal style. Fruit a utricle. Seeds 1–4. Cosmopolitan (most abundant and diverse in the tropics); 4 genera and ca. 35 species, 4 genera and 4 species in the flora area. Key to the Genera of Lemnaceae 1. 1. 2(1). 2. 3(2). 3.

Fronds with 7–20 roots ................................................................ 2. Spirodela Fronds with 1 root or rootless .................................................................... 2 Fronds with 1 root ............................................................................. 1. Lemna Fronds rootless ........................................................................................... 3 Fronds fleshy, subglobular ................................................................ 3. Wolffia Fronds membranous, flat .............................................................. 4. Wolffiella

780

L EMNACEAE

1. LEMNA L., Sp. Pl. 970. 1753. Free-floating on water’s surface, up to 4 fronds cohering. Fronds 1–5 × 0.5–3 mm, membranous, flat, elliptic to linear, symmetrical to asymmetrical, rounded at apices, with 1–3 veins, with 1–3 papules, without prominent dots; root 1, the sheath smooth or winged; prophyllum absent. Reproductive pouches 2, lateral. Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay, widespread elsewhere except Antarctica; 13 species, 3 in Venezuela, 1 of these in the flora area. Lemna aequinoctialis Welw., Apont. 578. 1858 [1859]. Frond 2–5 × 1–3 mm, elliptic, symmetrical, with 3 veins; root sheath prominently winged. Slow-moving water of swamps, near sea level to 50 m; Delta Amacuro (Tucupita). Widespread in lowland areas of northern Venezuela; U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, widespread elsewhere in subtropics and tropics. ŠFig. 656.

Fig. 656. Lemna aequinoctialis

2. SPIRODELA Schleid., Linnaea 13: 391. 1839. Free-floating on water’s surface, up to 5 fronds cohering. Fronds 3–6 × 3–5 mm, membranous, flat, orbicular to elliptic, symmetrical and rounded at apices or strongly asymmetrical and pointed at one apex, with 5–12 veins, without papules, with or without prominent dots; roots 5–20, the sheaths smooth, covered by the prophyllum, the prophyllum with 1–5 roots penetrating. Reproductive pouches 2, lateral. Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay, widespread elsewhere except Antarctica; 3 species, 1 in Venezuela. Spirodela intermedia W. Koch, Ber. Schweiz. Bot. Ges. 41: 114. 1932. Frond 4–6 × 4–5 mm, the dorsal surface lacking a prominent dot; roots 7–20, with 2 or 3 roots penetrating the prophyllum. Uncommon in ponds, backwaters, and roadside ditches, 0–50 m; Delta Amacuro (La Florida south of Clavellina, Laguna Las Clavellinas, near Tucupita). Scattered in north-coastal Venezuela; Central America, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay. ŠFig. 657. Some Venezuelan specimens have been treated as Spirodela biperforata W. Koch, which is currently considered to be a form of Spirodela intermedia having fewer roots and more asymmetrical fronds.

Fig. 657. Spirodela intermedia

Wolffiella 781

3. WOLFFIA Horkel ex Schleid., Linnaea 13: 389. 1839. Free-floating on water’s surface or slightly below, up to 2 fronds cohering. Fronds 0.5–1.5 × 0.5–1 mm, fleshy, subglobular, symmetrical, rounded at apices, the dorsal surface rounded or flattened, with or without a single papule, without prominent dots; veins and roots absent. Reproductive pouches 2, basal and dorsal. Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Suriname, Ecuador, Brazil, Bolivia, Paraguay, Argentina, Uruguay, widespread elsewhere except Antarctica; ca. 10 species, 2 in Venezuela, 1 of these in the flora area. Wolffia brasiliensis Wedd., Ann. Sci. Nat. Bot. sér. 3, 12: 170. 1849. Frond 1–1.2 mm long, the dorsal surface flattened, with a single papule. Roadside ditches, stream margins, 50–300 m; Bolívar (Tumeremo). Scattered in lowland areas of northern Venezuela; Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Suriname, Ecuador, Brazil, Paraguay, Argentina, Uruguay. ŠFig. 658.

Fig. 658. Wolffia brasiliensis

4. WOLFFIELLA Hegelm., Bot. Jahrb. Syst. 21: 303. 1895. Wolffiopsis Hartog & Plas, Blumea 18: 366. 1970. Free-floating on water’s surface or slightly below, up to 5 fronds cohering. Fronds 3–8 × 1–4 mm, membranous, flat, orbicular to linear, symmetrical, rounded or pointed at apices, without papules or prominent dots; veins and roots absent. Reproductive pouches 2 or 3, 1 basal and 1 or 2 dorsal, the basal pouch marked by a band of longer cells (stipe). Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay, Africa, India; 9 species, 2 in Venezuela, 1 of these in the flora area. Wolffiella welwitschii (Hegelm.) Monod, Mem. Soc. Hist. Nat. Afrique N., Hors. Sér. 2: 229. 1949. —Wolffia welwitschii Hegelm., J. Bot. 3: 114. 1865. Frond 5–6 × 3–4 mm, suborbicular, rounded at apices; basal reproductive pouch wider than deep, the stipe medial; dorsal reproductive pouches 2, on either side of frond midline. Uncommon in swamps and cloud forest lakes, 0–600 m; Delta Amacuro (Tucupita), Bolívar (Altiplanicie de Nuria). Widespread elsewhere in Venezuela; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Africa. ŠFig. 659.

Fig. 659. Wolffiella welwitschii

782

L ENTIBULARIACEAE

LENTIBULARIACEAE by Peter G. Taylor Herbs of wet or damp places, plants modified in various ways for the capture and digestion of small organisms. Leaves rosulate, alternate, or verticillate on stolons, simple or divided into capillary segments. Flowers zygomorphic, bisexual, solitaryscapose, and ebracteate, or in racemes and bracteate (often bibracteolate). Calyx 2-, 4-, or 5-lobed, the lobes spreading; corolla bilabiate, lower lip usually conspicuously spurred, 2–5-lobed. Stamens 2, inserted at base of upper corolla lip. Ovary superior, 1-locular with a free basal placenta; style usually short, ± persistent; stigma bilabiate, upper lip usually smaller than lower one. Ovules 2–many. Fruit capsular, variously valvate, circumscissile, or rarely indehiscent. Seeds usually many, sometimes 1–few, usually very small, diverse. Cosmopolitan; 3 genera and ca. 280 species, 2 genera and 53 species in the flora area. A third genus found in Venezuela, Pinguicula, occurs only in the Andes at high altitudes. Key to the Genera of Lentibulariaceae 1. 1.

Calyx lobes 5 ................................................................................... 1. Genlisea Calyx lobes 2 ............................................................................... 2. Utricularia

1. GENLISEA A. St.-Hil., Voy. Distr. Diam. 2: 428. 1833. Small terrestrial herbs. Leaves of 2 kinds: (1) white, subterranean, root-like, forming inverted Y-shaped traps, and (2) green, rosulate, obovate, or linear-lanceolate and entire. Inflorescence usually glandular, racemose. Flowers yellow or violet, pedicellate, bracteate and bracteolate, with sterile bracts (scales) on the peduncle. Calyx 5-lobed, lobes ovate or deltoid; corolla lower lip always conspicuously spurred, 3-lobed. Ovules many; style short, persistent; stigma 2-lipped, upper lip smaller. Fruit in Guayanan species a globose, multiple-circumscissile capsule. Seeds numerous, very small. Neotropics, tropical Africa, Madagascar; 20 species, 7 in Venezuela, all in the flora area. The Y-shaped traps in Genlisea function on the lobster-pot principle; any organism entering through apertures in the 2 divergent arms is prevented from escaping by numerous inward-pointing hairs. Genlisea is unique in the plant kingdom because of its specialization in trapping only protozoan prey, which are attracted chemically (W. Barthlott, S. Porembski, E. Fischer, and B. Gemmel. Nature 392: 447. April 2, 1998). Key to the Species of Genlisea 1. 1. 2(1).

Corolla violet or purple .............................................................................. 2 Corolla yellow ............................................................................................. 4 Spur of corolla about as long as lower lip; whole plant glabrous; leaves narrowly obovate-spatulate .......................................................... G. glabra

Genlisea 783

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

Spur of corolla 11/2–2 times as long as lower lip; bracts and calyx lobes ciliate; leaves linear-lanceolate .................................................................. 3 Peduncle glabrous, pedicels glabrous or with a few scattered hairs ................................................................................................ G. guianensis Peduncle and pedicels densely glandular-hispid ................. G. sanariapoana Lower lip of corolla as long as or longer than spur; peduncle relatively stout, < 10 cm long ............................................................... G. roraimensis Lower lip of corolla shorter than spur; peduncle slender, often > 10 cm long ......................................................................................................... 5 Spur inflated, cylindrical, with apex very obtuse or rounded .... G. filiformis Spur narrowly conical, with apex acute .................................................... 6 Inflorescence densely covered with short hairs intermixed with long gland-tipped hairs ..................................................................... G. pygmaea Inflorescence glabrous, or with a few scattered hairs or gland-tipped hairs at the base of peduncle and apex of pedicels ............................... G. repens

Genlisea filiformis A. St.-Hil., Voy. Distr. Diam. 2: 430. 1833. Genlisea anfractuosa Tutin, J. Bot. 72: 310. 1934. Herb 4–30 cm tall. Wet savannas, 100– 900 m; Bolívar (near Santa Elena de Uairén), Amazonas (near Puerto Ayacucho, upper Río Ventuari). Belize, Cuba, Colombia, Trinidad-Tobago, Guyana, Brazil (south to São Paulo). ŠFig. 663. Genlisea glabra P. Taylor, Mem. New York Bot. Gard. 17(1): 203. 1967. Herb 10–16 cm tall. Swamps and wet sandy ground among rocks, 2100–2500 m; Bolívar (Aprada-tepui, Macizo del Chimantá). Endemic. ŠFig. 666.

Bolívar (Ilú-tepui, Kukenán-tepui, Macizo del Chimantá, Murisipán-tepui, Ptari-tepui), Amazonas (Cerro Aracamuni, Cerro Duida, San Juan de Manapiare). Trinidad-Tobago, Guyana, Suriname, Brazil. ŠFig. 664. Genlisea repens Benj. in Mart., Fl. Bras. 10: 254. 1847. Herb 5–22 cm tall. Wet savannas, along stream banks and damp soil among rocks, (100–)1900–2500 m; widespread on tepui summits in Bolívar, Amazonas (Cerro Aracamuni, Cerro Autana, Cerro Parú, Sierra de la Neblina). Colombia, Guyana, Suriname, Brazil, Paraguay. ŠFig. 665.

Genlisea guianensis N.E. Br. in Hook., Icon. Pl. 27, pl. 2629. 1900. Herb 10–45 cm tall. In shallow running or still water in streams and pools in savannas and forests, (100–)400–1100 m; Bolívar (Caicara, Gran Sabana), Amazonas (Cerro Aracamuni). Guyana, Brazil (Goiàs, Mato Grosso). ŠFig. 662.

Genlisea roraimensis N.E. Br., Trans. Linn. Soc. London, Bot. 6: 56. 1901. Herb 3–12 cm tall. Wet savannas, moist soil among rocks and on stream banks, 1300–2800 m; Bolívar (Cerro Jaua, Ilú-tepui, Kukenán-tepui, Roraima-tepui), Amazonas (Cerro Autana, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Guyana (Mount Roraima). ŠFig. 661.

Genlisea pygmaea A. St.-Hil., Voy. Distr. Diam. 2: 431. 1833. Genlisea nigrocaulis Steyerm., Bull. Torrey Bot. Club 75: 657. 1948. Genlisea esmeraldae Steyerm., Fieldiana, Bot. 28: 534. 1953. Herb 5–20 cm tall. Wet savannas, banks of streams and pools, (200–)1300–2700 m;

Genlisea sanariapoana Steyerm., Fieldiana, Bot. 28: 534. 1953. Herb 7–30 cm tall. Wet savannas and among rock outcrops, 100–600 m; northwestern Amazonas (near Gavilán, Río Orinoco near Puerto Ayacucho, Río Samariapo, San Juan de Manapiare). Endemic. ŠFig. 660.

784

L ENTIBULARIACEAE

Fig. 660. Genlisea sanariapoana

Fig. 662. Genlisea guianensis

Fig. 661. Genlisea roraimensis

Utricularia 785

Fig. 663. Genlisea filiformis

Fig. 664. Genlisea pygmaea

Fig. 665. Genlisea repens

Fig. 666. Genlisea glabra

2. UTRICULARIA L., Sp. Pl. 18. 1753. Terrestrial, epiphytic, or aquatic herbs, with small bladder-like traps. Leaves in terrestrial and epiphytic species mostly entire, linear or obovate, in aquatic species divided into capillary segments. Inflorescences racemose. Flowers violet, lavender, shades of red, white, or yellow, pedicellate, bracteate and mostly bracteolate, with sterile bracts (scales) usually present on the peduncle; bracts and bracteoles basifixed or basisolute (produced below the point of attachment). Calyx usually 2lobed (4-lobed in some Australian species), the lobes mostly ovate; corolla with lower lip entire or variously 2–5-lobed, usually conspicuously spurred (but spur sometimes much reduced and saccate). Ovules 2–many; style usually short; stigma 2-lipped, the upper lip usually smaller. Fruit a globose or ovoid, valvate, circumscissile, or rarely indehiscent capsule. Seeds 1–many, usually very small. Cosmopolitan (mainly tropical); ca. 220 species, 49 in Venezuela, 46 of these in the flora area. The complex traps consist of a globose or ovoid body (0.2–1.2 mm long), with a hinged door. When set, the trap is laterally compressed and the liquid content under negative pressure. An approaching organism triggers the door to open and is sucked in with inrushing water. The door then closes, and digestion and resetting takes place.

786

L ENTIBULARIACEAE

Key to the Species of Utricularia 1. 1.

2(1). 2. 3(1). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(3). 7. 8(7). 8. 9(8). 9. 10(9). 10. 11(10). 11. 12(11). 12. 13(11). 13. 14(13). 14. 15(14).

Leaves divided into verticillate capillary segments with terminal traps; corolla pink ............................................................................................. 2 Leaves not divided into verticillate segments but entire or dichotomously divided, with lateral traps; corolla various shades of yellow, white, violet or lavender, rarely red or pink ......................................................... 3 Traps with ciliate mouth and a single ventral ciliate appendage .............. .................................................................................................. U. cucullata Traps with mouth not ciliate nor with ventral appendage ....... U. myriocista Calyx lobes, bracts, and bracteoles with margins ± deeply fimbriate; corolla yellow ............................................................................................. 4 Calyx lobes with margins not deeply fimbriate; corolla various shades of yellow, white, violet, or lavender, rarely red or pink ............................ 7 Bracteoles similar to the bract, basisolute and inserted with the bract at the base of the pedicel ........................................................... U. sandwithii Bracteoles much longer than bract, basifixed, similar to the calyx lobes and inserted with them at the apex of the pedicel ............................... 5 Pedicel about as long as calyx; bracteoles auriculate ................. U. fimbriata Pedicel much shorter than calyx; bracteoles not auriculate .................... 6 Uppermost scales much smaller than those below ................. U. longeciliata Uppermost scales as large as or larger than those below .......... U. simulans Bract tubular, encircling pedicel; corolla pale lavender or white ............... ................................................................................................. U. spruceana Bract not tubular; corolla various ............................................................. 8 Bract basisolute (prolonged downward beyond their true origin) ........... 9 Bract basifixed ......................................................................................... 21 Bracteoles present; leaves peltate; corolla pale violet or white .... U. pubescens Bracteoles absent; leaves not peltate; corolla pale violet, white, cream, or yellow ................................................................................................... 10 Lower lip of corolla entire, inflorescence viscid-glandular; corolla pale violet or white ................................................................................... U. viscosa Lower lip of corolla 3-lobed; inflorescence not viscid-glandular; corolla white to yellow, or white with lavender streaks ................................. 11 Spur of corolla very short, scarcely longer than the lower calyx lobe; corolla cream; on rocks in flowing water ................................................ 12 Spur of corolla considerably longer than lower calyx lobe; corolla yellow, white, or white with lavender streaks; habitat various ..................... 13 Leaves divided into capillary segments .................................... U. neottioides Leaves entire, obovate ................................................................ U. oliveriana Lower calyx lobe about 1/2 as wide as the upper lobe; corolla yellow, white, or white with lavender streaks ........................................................... 14 Lower calyx lobe as wide as or wider than upper lobe; corolla yellow ... 17 Upper lip of corolla narrower than upper calyx lobe; corolla yellow ......... ............................................................................................. U. steyermarkii Upper lip of corolla wider than upper calyx lobe; corolla yellow, white, or white with lavender streaks ................................................................ 15 Upper lip of corolla with a porrect (directed outward and forward), hornlike projection from the palate; corolla white ......................... U. mirabilis

Utricularia 787

15. 16(15). 16. 17(13). 17. 18(17). 18. 19(18).

19. 20(19). 20. 21(8). 21. 22(21). 22. 23(22). 23. 24(22). 24. 25(24). 25. 26(25). 26. 27(25). 27. 28(21). 28. 29(28).

Upper lip of corolla without a horn-like projection; corolla yellow, or white with lavender streaks .......................................................................... 16 Leaf blade obovate; corolla white with lavender streaks .... U. heterochroma Leaf blade reniform; corolla yellow .................................... U. aureomaculata Veins of calyx lobes obscure or apparently absent ................. U. trichophylla Veins of calyx lobes distinct or prominent ............................................... 18 Calyx lobes with apex acute, with ca. 5 very prominent veins which extend to and meet at the apex ....................................................... U. triloba Calyx lobes with apex rounded, the veins not extending to or meeting at the apex ................................................................................................ 19 Raceme axis bearing numerous sterile bracts in addition to those which subtend flowers; spur of corolla about 2 times as long as lower lip ...................................................................................................... U. pusilla Raceme axis with all bracts subtending flowers; spur of corolla considerably < 2 times as long as lower lip ...................................................... 20 Corolla 1.5–2 cm long ...................................................................... U. nervosa Corolla < 1 cm long ........................................................................ U. subulata Bracteoles absent; aquatic plants with leaves divided into capillary segments .................................................................................................... 22 Bracteoles present, but may be partly connate with the bract epiphytic or terrestrial plants with leaves not divided into capillary segments ... 28 Peduncle with a whorl of inflated leaf-like organs; corolla yellow or violet .............................................................................................................. 23 Peduncle without a whorl of inflated leaf-like organs; corolla white, cream, yellow, or red to pink ............................................................... 24 Stolons glabrous; corolla yellow, the lower lip 3-lobed, longer than the spur ......................................................................................... U. breviscapa Stolons villous; corolla violet, the lower lip entire, shorter than spur .......................................................................................... U. benjaminiana Corolla red to pink; lowermost flower cleistogamous and deflexed beneath the water ............................................................................... U. hydrocarpa Corolla yellow to white; lowermost flower not cleistogamous and deflexed beneath the water ................................................................................ 25 Spur of corolla very short and saccate; corolla < 3 mm long, white or cream .................................................................................................... 26 Spur of corolla distinct, cylindrical or subulate; corolla > 5 mm long, yellow ........................................................................................................ 27 Peduncle and raceme axis extremely short, the pedicels appearing to arise from the stolons; corolla cream to white .......................... U. olivacea Peduncle and raceme axis normally developed; corolla white .... U. naviculata Fruiting pedicels erect; upper lip of corolla 3-lobed, larger than lower lip ........................................................................................................ U. gibba Fruiting pedicels decurved; upper lip of corolla smaller than lower lip ....................................................................................................... U. foliosa Bracteoles connate with bract for 1/2 or more of its length; corolla violet .............................................................................................................. 29 Bracteoles not or only very shortly connate with bract; corolla violet, lavender, white to yellow, or red .............................................................. 30 Corolla 0.3–1 cm wide, lower lip ± distinctly 3-lobed ............ U. amethystina

788

L ENTIBULARIACEAE

29. Corolla 1.5–2.5 cm wide, lower lip not or scarcely 3-lobed ............ U. tricolor 30(28). Lateral margins of calyx lobes crenulate or denticulate; corolla lavender, violet, white, or yellow .......................................................................... 31 30. Lateral margins of calyx lobes entire; corolla pale violet, violet, white, yellow, or red .............................................................................................. 33 31(30). Lateral margins of calyx lobes minutely, regularly, and acutely denticulate, lobes somewhat wider than long; corolla lavender, violet, or white ................................................................................................. U. calycifida 31. Lateral margins of calyx lobes irregularly crenulate, the lobes longer than wide; corolla pale violet, white, or yellow ............................................ 32 32(31). Lateral margins of calyx lobes crenulate throughout their length; leaves mostly narrowly linear, multiveined; bracteoles much narrower than bracts; corolla usually violet, sometimes yellow ........................ U. hispida 32. Lateral margins of calyx lobes crenulate in the distal 1/2 only; leaves all obovate; bracteoles > 1/2 as wide as the bracts; corolla pale violet or white ......................................................................................... U. schultesii 33(30). Epiphytic plant with tubers; corolla red, white, or pale violet ................ 34 33. Terrestrial or epiphytic plant without tubers; corolla violet, white, or yellow ......................................................................................................... 37 34(33). Corolla bright red ...................................................................................... 35 34. Corolla white or pale violet ....................................................................... 36 35(34). Peduncle with 4 or 5 conspicuous scales; leaf blade narrowly obovate, longer than the petiole ...................................................... U. campbelliana 35. Peduncle with 1 or 2 inconspicuous scales; leaf blade obovate, shorter than the petiole ................................................................................... U. quelchii 36(34). Lower lip of corolla distinctly 3-lobed, much shorter than spur ................. ............................................................................................ U. jamesoniana 36. Lower lip of corolla entire, longer than spur ................................... U. alpina 37(33). Corolla 5–10 cm wide, violet; leaf blade cuneate to reniform, coriaceous ............................................................................................... U. humboldtii 37. Corolla < 1.5 cm wide, violet, white, or yellow; leaf blade linear, membranous ....................................................................................................... 38 38(37). Corolla violet or white ............................................................................... 39 38. Corolla yellow ............................................................................................ 40 39(38). Both calyx lobes with very prominent veins, lower lobe larger than upper and about as long as capsule ....................................................... U. costata 39. Both calyx lobes with obscure veins, upper lobe larger than lower and about 1/2 as long as capsule ................................................... U. tenuissima 40(38). Spur short, broadly conical, wider than long ............................. U. guyanenis 40. Spur subulate, much longer than wide .................................................... 41 41(40). Upper calyx lobe with prominent veins ................................................... 42 41. Upper calyx lobe without prominent veins .............................................. 43 42(41). Upper calyx lobe larger than the lower; fruiting pedicels erect ...... U. juncea 42. Upper calyx lobe smaller than lower; fruiting pedicels decurved ..... U. nana 43(41). Pedicel < 1/2 as long as fruiting calyx; upper corolla lip shorter and narrower than upper calyx lobe ................................................... U. erectiflora 43. Pedicel > 1/2 as long as fruiting calyx; upper corolla lip somewhat to considerably longer or wider than upper calyx lobe ...................................... 44

Utricularia 789

44(43). Upper corolla lip about 2 times as long as upper calyx lobe ...................... ........................................................................................ U. chiribiquitensis 44. Upper corolla lip about as long as upper calyx lobe ............................... 45 45(44). Upper corolla lip 3 times or more as wide as upper calyx lobe; seeds with small testa cells ......................................................................... U. adpressa 45. Upper corolla lip only slightly wider than upper calyx lobe; seeds with testa cells relatively very large .................................................... U. lloydii Utricularia adpressa Salzm. ex A. St.-Hil. & Girard, Compt. Rend. Hebd. Séances Acad. Sci. 7: 870. 1838. Utricularia aureolimba Steyerm., Fieldiana, Bot. 28: 535. 1953. Terrestrial 4–18 cm tall. Wet sandy savannas, 1100–1300 m; Bolívar (Gran Sabana). Central America, Colombia, TrinidadTobago, Guyana, Suriname, French Guiana, Brazil. ŠFig. 708. Utricularia alpina Jacq., Enum. Syst. Pl. 11. 1760. —Orchyllium alpinum (Jacq.) Barnhart, Mem. New York Bot. Gard. 6: 53. 1916. Utricularia montana Jacq., Select. Stirp. Amer. Hist. 7, t. 6. 1780. Epiphyte 15–40 cm tall. On rocks, trees, and shady banks in humid montane forests, 600–1600 m; Bolívar (widespread), Amazonas (Cerro Coro Coro, Cerro Marahuaka, Sierra Parima). Venezuelan Andes and Coastal Cordillera; West Indies, Colombia, TrinidadTobago, Guyana, Suriname, Brazil (Amazonas). ŠFig. 697. Utricularia amethystina Salzm. ex A. St.Hil. & Girard, Compt. Rend. Hebd. Séances Acad. Sci. 7: 870. 1838. —Calpidisca amethystina (Salzm. ex A. St.Hil. & Girard) Barnhart in Gleason, Bull. Torrey Bot. Club 58: 468. 1931. Utricularia modesta DC. in A. DC., Prodr. 8: 17. 1844. —Calpidisca modesta (A. DC.) Barnhart, Bull. Torrey Bot. Club 58: 469. 1931. Utricularia punctifolia Benj., Linnaea 20: 492. 1847. Utricularia versicolor Benj., Linnaea 20: 488. 1847. Utricularia roraimensis N.E. Br., Trans. Linn. Soc. London, Bot. 6: 54. 1901. —Calpidisca roraimensis (N.E. Br.) Gleason, Bull. Torrey Bot. Club 58: 405. 1931.

Utricularia spatulifolia Pilger, Notizbl. Königl. Bot. Gart. Berlin 6: 190. 1914. Utricularia alutacea Tutin, J. Bot. 72: 313. 1934. —Utricularia amethystina f. alutacea (Tutin) Steyerm., Bull. Torrey Bot. Club 75: 658. 1948. Utricularia kaieturensis Steyerm., Bull. Torrey Bot. Club 75: 658. 1948. Utricularia bolivarana Steyerm., Fieldiana, Bot. 28: 538. 1953. Utricularia tepuiana Steyerm., Fieldiana, Bot. 28: 550. 1953. Utricularia turumiquirensis Steyerm., Fieldiana, Bot. 28: 552. 1953. Terrestrial 3–50 cm tall. Wet sandy savannas, swamps, damp soil among rocks, sometimes in light shade, 100–2800 m; widespread in Bolívar (except northern part) and Amazonas. Mérida, Monagas, Sucre; U.S.A. (southern Florida), Central America, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ŠFig. 692. Utricularia amethystina is extremely variable in stature, calyx indument, corolla size, and lobing of the lower lip, but in view of the many intermediate forms it seems best at present to treat this as a single polymorphic species. Utricularia aureomaculata Steyerm., Fieldiana, Bot. 28: 536. 1953. Terrestrial 10–25 cm tall. Dripping sandstone bluffs, 600–2400 m; Bolívar (Amaruaytepui, Ilú-tepui, Macizo del Chimantá [Amurí-tepui], Ptari-tepui). Endemic. ŠFig. 684. Utricularia benjaminiana Oliv., J. Proc. Linn. Soc., Bot. 4: 76. 1860. Aquatic, with emergent inflorescence 3– 25 cm tall. Pools in savannas, 100–1000 m; Bolívar (base of Auyán-tepui), Amazonas (Puerto Ayacucho). Honduras, Nicaragua, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Roraima), tropical Africa, Madagascar.

790

L ENTIBULARIACEAE

Utricularia breviscapa Wright ex Griseb., Cat. Pl. Cub. 161. 1866. Aquatic, with emergent inflorescence 1–10 cm tall. Swamps, flooded savannas, ca. 100 m; Bolívar (Río Parguaza), Amazonas (Puerto Ayacucho). Apure; Cuba, Colombia, Ecuador, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 670.

Aquatic, with emergent inflorescence 2– 10 cm tall. Pools in wet savannas, 100–1000 m; Bolívar (Gran Sabana), Amazonas (near Puerto Ayacucho, upper Río Ventuari, Santa Barbara). Guárico; Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (south to Paraná). ŠFig. 668.

Utricularia calycifida Benj., Linnaea 20: 493. 1847. —Calpidisca calycifida (Benj.) Gleason, Bull. Torrey Bot. Club 56: 404. 1919. Utricularia maguirei Steyerm., Bull. Torrey Bot. Club 75: 659. 1948. Utricularia cuspidata Steyerm., Fieldiana, Bot. 28: 542. 1953. Terrestrial 2–50 cm tall. Wet rocks, by waterfalls and streams, wet sandy tracks in forests, often in shade, 300–1200 m; northeastern and central-eastern Bolívar, Amazonas (Cerro Sipapo). Guyana, Suriname, Brazil (Pará). ŠFig. 694.

Utricularia erectiflora A. St -Hil. & Girard, Compt. Rend. Hebd. Séances Acad. Sci. 7: 870. 1838. Utricularia spicata Sylvén, Ark. Bot. 8(6): 14, pl. 1, figs. 15, 16. 1909. —Stomoisia spicata (Sylvén) Gleason, Bull. Torrey Bot. Club 56: 404. 1929. Terrestrial 5–35 cm tall. Wet savannas, 100–800 m; Bolívar (scattered). Carabobo, Guárico; Belize, Nicaragua, Colombia, Guyana, Suriname, Ecuador, Brazil, Bolivia. ŠFig. 705.

Utricularia campbelliana Oliv., Trans. Linn. Soc. London, Bot. 2: 280. 1887. Utricularia campbelliana var. minor Steyerm., Fieldiana, Bot. 28: 539. 1953. Epiphyte 2–13 cm tall. On tree trunks, in bromeliad tanks, and sometimes on mosscovered rocks in forest or low scrub, 1500– 2600 m; widespread on tepuis in Bolívar and Amazonas. Guyana. ŠFig. 696. Utricularia chiribiquitensis A. Fern., Caldasia 9: 42. 1964. Terrestrial 6–30 cm tall. Wet sandy savannas, often among rock outcrops, ca. 100– 200(–1800) m; Bolívar (Cerro Guanay), northern Amazonas. Eastern Colombia. ŠFig. 706. Utricularia costata P. Taylor, Kew Bull. 41: 7. 1986. Terrestrial 2–7 cm tall. Wet soil and among rocks in savannas, ca. 400 m; Bolívar (base of Macizo del Chimantá). Brazil (Mato Grosso, Pará, Roraima). ŠFig. 700. Utricularia cucullata A. St.-Hil. & Girard, Compt. Rend. Hebd. Séances Acad. Sci. 7: 869. 1838. —Vesiculina cucullata (A. St.-Hil. & Girard) Barnhart, Mem. New York Bot. Gard. 6: 63. 1916. Utricularia ayacuchae Steyerm., Fieldiana, Bot. 28: 538. 1953.

Utricularia fimbriata H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 225. 1817 [1818]. Terrestrial 15–30 cm tall. Wet sandy savannas and among granitic outcrops, 50–600 m; Bolívar (Cerro San Borja along lower Río Suapure), northern Amazonas. Adjacent Colombia. ŠFig. 675. Utricularia foliosa L., Sp. Pl. 18. 1753. Utricularia guianensis Splitg. ex de Vries, Ned. Kruidk. Arch. 1: 323. 1848. Aquatic, with emergent inflorescence 7– 45 cm tall. Pools and swamps in wet savannas, lakes, slow-flowing rivers, 50–400 m; scattered in Delta Amacuro, northern Bolívar, northern Amazonas. Southern U.S.A., Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, tropical and southern Africa, Madagascar. ŠFig. 673. Utricularia gibba L., Sp. Pl. 18. 1753. Utricularia obtusa Sw., Prodr. 14. 1788. Utricularia tenuifolia Benj., Linnaea 20: 304. 1847. Aquatic, with emergent inflorescence 1– 20 cm tall. Lakes, pools, swamps, 50–2100 m; Delta Amacuro (Caño Güiniquina, Caño Simoina west of Isla Cocuina), Bolívar (widespread). Widespread elsewhere in Venezuela; Canada, U.S.A., Mexico, Central America, south in South America to Argentina, Portugal, Africa, Madagascar, Mauritius, tropical

Utricularia 791

Asia, northern Australia, New Zealand, occasionally introduced elsewhere. This is the most widespread species of Utricularia in the world. Utricularia guyanensis DC. in A. DC., Prodr. 8: 11. 1844. Utricularia rubricaulis Tutin, J. Bot. 72: 312. 1934. Terrestrial 5–13 cm tall. Wet savannas, 100–1100 m; Bolívar (Gran Sabana), Amazonas (Caño Cotúa near base of Cerro Yapacana). Belize, Honduras, Puerto Rico, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (southern Bahia). ŠFig. 701. Utricularia heterochroma Steyerm., Fieldiana, Bot. 28: 544. 1953. Terrestrial 5–6 cm tall. Wet rocks by cascades and waterfalls, 1200–2500 m; southeastern Bolívar (Cerro Guanay, Macizo del Chimantá, Ptari-tepui). Endemic. ŠFig. 683. Utricularia hispida Lam., Tabl. Encycl. 1: 50. 1791. —Calpidisca hispida (Lam.) Barnhart, Bull. Torrey Bot. Club 58: 469. 1931. Terrestrial 4–80 cm tall. Wet savannas, 100–1800 m; widespread in Bolívar and Amazonas. Carabobo; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ŠFig. 693. Utricularia humboldtii R.H. Schomb., Flora 23: 416. 1840. —Calpidisca humboldtii (R.H. Schomb.) Gleason, Bull. Torrey Bot. Club 56: 404. 1929. —Orchyllium humboldtii (R.H. Schomb.) Barnhart, Bull. Torrey Bot. Club 58: 469. 1931. Utricularia humboldtii f. cuneata Steyerm., Fieldiana, Bot. 28: 544. 1953. Terrestrial or epiphytic to 1.3 m tall. Usually occurring in water of bromeliad tanks (Brocchinia species), but sometimes on bare wet ground, (100–)1000–2500 m; widespread in Bolívar and Amazonas. Guyana, northern Brazil. ŠFig. 699. Utricularia hydrocarpa Vahl, Enum. Pl. 1: 200. 1804. Utricularia coccinea Benj., Linnaea 20: 490. 1847.

Utricularia fockeana Miq., Stirp. Surinam. Select. 113. 1850 [1851]. Utricularia amazonasana Steyerm., Fieldiana, Bot. 28: 535. 1953. Aquatic, with emergent inflorescence 4– 10 cm tall. Swamps, lakes, wet savannas, ca. 100–300 m; Bolívar (near El Manteco), Amazonas (near Samariapo and Puerto Ayacucho). Anzoátegui, Apure, Aragua, Barinas, Cojedes, Distrito Federal, Monagas, Sucre; Neotropics. Utricularia jamesoniana Oliv., J. Proc. Linn. Soc., Bot. 4: 169. 1860. Utricularia schimperi Schenck in Pringsh., Jahrb. Wiss. Bot. 18: 230. 1887. —Orchyllium schimperi (Schenck) Barnhart, Bull. Torrey Bot. Club 58: 469. 1931. Utricularia concinna N.E. Br., Trans. Linn. Soc. London, Bot. ser. 2, 6: 55. 1901. Epiphyte 1–10 cm tall. On moss-covered tree trunks and branches in forests, (100–) 500–1600 m; widespread in Bolívar (except northern part) and Amazonas. Táchira; Central America, Antilles, Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil. ŠFig. 695. Utricularia juncea Vahl, Enum. Pl. 1: 202. 1804. —Stomoisia juncea (Vahl) Barnhart in Small, Fl. Miami: 171. 1913. Utricularia angulosa Poir. in Lam., Encycl. 8: 273. 1808. Utricularia stricta G. Mey., Prim. Fl. Esseq. 14. 1818. Terrestrial 2–45 cm tall. Wet savannas, swamps, stream banks, 300–1900 m; Bolívar (frequent in Gran Sabana), Amazonas (Cerro Duida). Southeastern U.S.A., Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil (south to Bahia), tropical Africa. ŠFig. 704. Utricularia lloydii Merl, Bull. Torrey Bot. Club 61: 370. 1934. Terrestrial 2–26 cm tall. Periodically flooded sand by river, ca. 100 m; Amazonas (Río Yatúa). Panama, Suriname, Brazil, Bolivia. ŠFig. 707. Utricularia longeciliata DC. in A. DC., Prodr. 8: 23. 1844.

792

L ENTIBULARIACEAE

Polypompholyx schomburgkii Klotzsch in R.M. Schomb., Reis. Br.-Guiana 2: 973, 1160. 1848. Utricularia pectinata Splitg. ex de Vries, Ned. Kruidk. Arch. 1: 325. 1848. Terrestrial 15–30 cm tall. Wet sandy savannas and swamps, 100–2000 m; widespread in Bolívar (except northern part) and Amazonas. Colombia, Guyana, Suriname, northern Brazil. ŠFig. 676. Utricularia mirabilis P. Taylor, Kew Bull. 41: 14. 1986. Terrestrial 7–20 cm tall. Creek beds in tepui meadows, ca. 1500 m; Amazonas (Cerro Sipapo). Endemic. ŠFig. 682. Utricularia myriocista A. St.-Hil. & Girard, Compt. Rend. Hebd. Séances Acad. Sci. 7: 869. 1838. Utricularia magnifica Pilger, Notizbl. Königl. Bot. Gart. Berlin 6: 290. 1914. Aquatic, with emergent inflorescence 3– 10 cm tall. Flooded savannas, 200–1100 m; Bolívar (Gran Sabana), Amazonas (Río Parucito). Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, northern Argentina. ŠFig. 669. Utricularia nana A. St.-Hil. & Girard, Compt. Rend. Hebd. Séances Acad. Sci. 7: 869. 1838. Utricularia dubia Benj., Linnaea 20: 494. 1847. Utricularia arenicola Tutin, J. Bot. 72: 312. 1934. Utricularia arenicola var. kavanayena Steyerm., Fieldiana, Bot. 28: 535. 1953. Terrestrial 1.5–12 cm tall. Wet savannas and stream banks, 100–1400 m; Bolívar (Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá [Chimantá-tepui]) Amazonas (scattered). Carabobo; Guyana, Suriname, French Guiana, Brazil (south to Paraná), Paraguay. ŠFig. 703. Utricularia naviculata P. Taylor, Mem. New York Bot. Gard. 17(1): 226. 1967. Aquatic, with emergent inflorescence 2–5 cm tall. Wet savannas, ca. 100 m; Amazonas (Caño Cumare near San Fernando de Atabapo). Brazil (Pará). ŠFig. 672. Utricularia neottioides A. St.-Hil. & Girard, Compt. Rend. Hebd. Séances

Acad. Sci. 7: 869. 1838. —Avesicaria neottioides (A. St.-Hil. & Girard) Barnhart, Mem. New York Bot. Gard. 6: 56. 1916. Affixed aquatic 3–30 cm tall. On rocks in flowing water, 50–1800 m; Bolívar (extreme northwestern region, Cerro Jaua), central and northern Amazonas. Colombia, Brazil, Bolivia. ŠFig. 679. Utricularia nervosa Weber ex Benj. in Mart., Fl. Bras. 10: 247. 1847. Utricularia flaccida auct. non A. DC.: sensu P. Taylor, Mem. New York Bot. Gard. 17(1): 222. 1967. Terrestrial 15–50 cm tall. Wet savannas, 100–1300 m; Bolívar (between El Dorado and Santa Elena de Uairén), central and northeastern Amazonas (Río Orinoco, Río Ventuari). Colombia, Brazil, Paraguay, Argentina. ŠFig. 688. Utricularia olivacea Wright ex Griseb., Cat. Pl. Cub. 161. 1866. —Biovularia olivacea (Wright ex Griseb.) Kamienski, Zap. Novorossijsk. Obšc. Estestvoisp. 15(Mem. 1): 208. 1890. Aquatic, with emergent inflorescence 0.5– 5 cm tall. Swamps, flooded savannas, 100– 1900 m; Bolívar (vicinity of Canaima, El Paují, near Gavilán), Amazonas (Cerro Yapacana, Sierra de la Neblina). Eastern U.S.A. (New Jersey) south to Cuba, Nicaragua, Trinidad-Tobago, Guyana, Suriname, Brazil, Bolivia. ŠFig. 671. Utricularia oliveriana Steyerm., Fieldiana, Bot. 28: 546. 1953, “oliverana.” Affixed aquatic 2–15 cm tall. On rocks in shallow flowing water, 100–1200 m; Bolívar (northwestern portions, northeastern Gran Sabana), Amazonas (northwestern portions, northeastern Río Siapa). Colombia (Vaupés), Brazil (Goias, Pará). ŠFig. 678. Utricularia pubescens Sm. in Rees, Cycl. 37, Utricularia no. 74. 1819 [1818]. Utricularia connellii N.E. Br., Trans. Linn. Soc. Bot., ser. 2, 6: 53. 1901. Utricularia sciaphila Tutin, J. Bot. 72: 333. 1934. Utricularia subpeltata Steyerm., Fieldiana, Bot. 28: 548.1953 Utricularia venezuelana Steyerm., Fieldiana, Bot. 28: 552. 1953.

Utricularia 793

Terrestrial 2–35 cm tall. By streams and waterfalls, often in shade, 100–2600 m; widespread in Bolívar (except northern part) and Amazonas. Panama, Colombia, Guyana, southern Brazil, tropical Africa, India. ŠFig. 667. Utricularia pusilla Vahl, Enum. Pl. 1: 202. 1804. —Setiscapella pusilla (Vahl) Barnhart ex Britton & P. Wilson, Bot. Porto Rico 6: 191. 1925. Utricularia leptantha Benj., Linnaea 20: 313. 1847. Terrestrial 3–20 cm tall. Moist savannas, swamps, stream banks, rocks by waterfalls, 300–1200 m; Bolívar (near Ciudad Piar, Gran Sabana, Altiplanicie de Nuria). Aragua, Carabobo, Monagas, Táchira; Mexico to Argentina. ŠFig. 686. Utricularia quelchii N.E. Br., Trans. Linn. Soc. London, Bot. 6: 53. 1901. —Orchyllium quelchii (N.E. Br.) Gleason, Bull. Torrey Bot. Club 56: 405. 1929. Epiphytic or terrestrial 7–20 cm tall. In humus or on low trunks and branches in forests or low shrubby vegetation, or on stream banks, 1400–2800 m; Bolívar (widespread), Amazonas (Cerro Huachamacari, Cerro Sipapo, Sierra de la Neblina). Guyana, northern Brazil. ŠFig. 698. Utricularia sandwithii P. Taylor, Mem. New York Bot. Gard. 17(1): 218. 1967. Terrestrial 6–18 cm tall. Wet sandy savannas, 100–1000 m; Bolívar (near Auyántepui), Amazonas (Canaripó at lower Río Ventuari, near Cerro Yapacana). Guyana, Suriname, Brazil (Roraima). ŠFig. 674. Utricularia schultesii A. Fern., Caldasia 9: 39. 1964. Terrestrial 10–27 cm tall. Wet sandy savannas, 50–300(–1000) m; Amazonas (Cerro Aratitiyope, Raudal de Atures, upper Río Ventuari). Colombia. ŠFig. 691. Utricularia simulans Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 194. 1914. Utricularia congesta Steyerm., Fieldiana, Bot. 28: 540. 1953. Utricularia congesta f. deminutiva Steyerm., Fieldiana, Bot. 28: 540. 1953. Utricularia orinocensis Steyerm., Fieldiana, Bot. 28: 548. 1953.

Terrestrial 2–35 cm tall. Wet sandy savannas, 100–1200 m; Bolívar (widespread), Amazonas (Raudal de Atures, Raudal de Maipures, Río Ventuari basin). Anzoátegui, Apure, Guárico; southern U.S.A. (Florida), Belize, Cuba, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, tropical Africa. ŠFig. 677. Utricularia spruceana Benth. ex Oliv., J. Proc. Linn. Soc., Bot. 4: 173. 1860. —Lecticula spruceana (Benth. ex Oliv.) Barnhart, Mem. New York Bot. Gard. 6: 58. 1916. Utricularia resupinata P. Taylor, Mem. New York Bot. Gard. 17(1): 220. 1967, non Greene ex Bigelow 1840. Subaquatic 3–6 cm tall. In sand and mud by riversides, rare, ca. 100–200 m; Amazonas (San Fernando de Atabapo). Northern Brazil. Utricularia steyermarkii P. Taylor, Acta Bot. Venez. 2: 326. 1967. Terrestrial 6–12 cm tall. Moist rock ledges and at base of wet vertical rocks, 1600–1900 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Amurí-tepui]). Endemic. ŠFig. 681. Utricularia subulata L., Sp. Pl. 18. 1753. —Setiscapella subulata (L.) Barnhart in Small, Fl. Miami 170. 1913. Terrestrial 2–50 cm tall. Damp savannas, wet rocky slopes, by streams, in forests, wet habitats generally, 50–2700 m, but mostly at the lower elevations; common and widespread in Bolívar and Amazonas. Widespread elsewhere in Venezuela, but apparently absent from the northeast; pantropics, but scattered and scarce in Asia. ŠFig. 690. Utricularia tenuissima Tutin, J. Bot. 72: 334. 1934. Terrestrial 4–8 cm tall. Wet sandy savannas, 100–2000 m; Bolívar (Macizo del Chimantá [Apacará-tepui], Río Caruay), Amazonas (near Ocamo, Río Atabapo, near San Juan de Manapiare). Colombia, Trinidad-Tobago, Suriname, French Guiana, Brazil. ŠFig. 702. Utricularia trichophylla Spruce ex Oliv., J. Proc. Linn. Soc., Bot. 4: 173. 1860. Subaquatic 10–30 cm tall. Margins of streams and rivers, wet sandy savannas, 50– 1400 m; Bolívar (near Caicara del Orinoco,

794

L ENTIBULARIACEAE

Gran Sabana). Guárico; Belize, Nicaragua, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. ŠFig. 685. Utricularia tricolor A. St.-Hil., Voy. Distr. Diam. 2: 418. 1833. Terrestrial 10–60 cm tall. Wet savannas, 1800 m; Amazonas (Cerro Huachamacari). Táchira; Colombia, southern Brazil, Bolivia, Paraguay, Argentina, Uruguay. ŠFig. 689. The two collections from Amazonas (Maguire et al. 30051, K, NY, VEN; Maguire et al. 30139, K, NY, VEN) are somewhat atypical for the species. Utricularia triloba Benj. in Mart., Fl. Bras. 10: 248. 1847. Terrestrial 5–30 cm tall. Banks of rivers and streams, wet savannas, 100–1300 m;

Bolívar (near El Manteco, Gran Sabana, Macizo del Chimantá, upper Río Cuyuní, middle Río Paragua), Amazonas (Cerro Aracamuni, Cerro Huachamacari, Puerto Ayacucho, Sierra de la Neblina). Táchira; Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina. ŠFig. 687. Utricularia viscosa Spruce ex Oliv., J. Proc. Linn. Soc., Bot. 4: 172. 1860. Terrestrial or sub-aquatic 6–50 cm tall. Damp depressions and shallow pools in savannas, 50–800 m; Bolívar (Caicara del Orinoco, Río Parguaza, Santa Elena de Uairén), Amazonas (near Cerro Cucurito and Caño Yagua). Apure; Belize, Nicaragua, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. ŠFig. 680.

Fig. 667. Utricularia pubescens

Fig. 668. Utricularia cucullata

Fig. 669. Utricularia myriocista

Utricularia 795

Fig. 670. Utricularia breviscapa

Fig. 671. Utricularia olivacea

Fig. 672. Utricularia naviculata

Fig. 673. Utricularia foliosa

796

L ENTIBULARIACEAE

Fig. 674. Utricularia sandwithii

Fig. 676. Utricularia longeciliata

Fig. 677. Utricularia simulans

Fig. 675. Utricularia fimbriata

Utricularia 797

Fig. 678. Utricularia oliveriana

Fig. 680. Utricularia viscosa

Fig. 679. Utricularia neottioides

Fig. 681. Utricularia steyermarkii

Fig. 682. Utricularia mirabilis

798

L ENTIBULARIACEAE

Fig. 683. Utricularia heterochroma

Fig. 686. Utricularia pusilla

Fig. 684. Utricularia aureomaculata

Fig. 685. Utricularia trichophylla

Fig. 687. Utricularia triloba

Utricularia 799

Fig. 688. Utricularia nervosa

Fig. 689. Utricularia tricolor

Fig. 690. Utricularia subulata

800

L ENTIBULARIACEAE

Fig. 691. Utricularia schultesii

Fig. 692. Utricularia amethystina

Fig. 694. Utricularia calycifida

Fig. 693. Utricularia hispida

Utricularia 801

Fig. 695. Utricularia jamesoniana

Fig. 696. Utricularia campbelliana

Fig. 697. Utricularia alpina

Fig. 698. Utricularia quelchii

802

L ENTIBULARIACEAE

Fig. 699. Utricularia humboldtii

Fig. 700. Utricularia costata

Fig. 701. Utricularia guyanensis

Fig. 702. Utricularia tenuissima

Utricularia 803

Fig. 703. Utricularia nana

Fig. 704. Utricularia juncea

Fig. 706. Utricularia chiribiquitensis

Fig. 707. Utricularia lloydii

Fig. 705. Utricularia erectiflora

Fig. 708. Utricularia adpressa

Appendix List of new names published in this volume

Phyllanthus myrsinites subsp. francavillanus (Müll. Arg.) G.L. Webster, comb. & stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllanthus subapicalis subsp. sequoiifolius (Jabl.) G.L. Webster, comb. & stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aldina macrophylla var. yapacanensis (R.S. Cowan) Stergios, comb. & stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ormosia bolivarensis (Rudd) Stirton, comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . Platypodium elegans subsp. maxonianum (Pittier) H.C. Lima, comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taralea cordata var. rigida (Schery) H.C. Lima, comb. & stat. nov. . . . . . . . . . . Chorisepalum sipapoanum (Maguire) L. Struwe & V.A. Albert, comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

804

198 200 248 366 377 416 491

Index Entries in Roman type = accepted names of taxa and vernacular names Entries in italics = synonyms Page numbers in bold = illustrations

—A— Aarana, 576 Abrus, 237 minor, 239 precatorius subsp. africanus, 238, 239 precatorius, 239 pulchellus, 239 subsp. tenuiflorus, 238, 239 tenuiflorus, 239 Acalypha, 81 alopecuroides, 82 arvensis, 82 cuneata, 82 var. obovata, 82 diversifolia, 82, 85 var. carpinifolia, 83 var. leptostachya, 83 hirsutissima, 83 karsteniana, 84 leptostachya, 83 var. carpinifolia, 83 macrophylla, 83 macrostachya, 83, 84 var. hirsutissima, 83 var. macrophylla, 83 var. tristis, 83 obovata, 82 samydifolia, 83 scandens, 83 schiedeana, 83, 85 f. angustifolia, 83

var. angustifolia, 83 tristis, 83 villosa, 83, 85 var. intermedia, 83 var. paniculata, 84 var. tomentosa, 84 Achimenes erinoides, 560 Acidoton, 86 nicaraguensis, 86, 87 venezolanus, 87 Acoi-yerí-yek, 344 Acosmium, 239 nitens, 240, 240 sp. A, 240 Acrodiclidium anacardioides, 722 brasiliense, 718 chrysophyllum, 719 debile, 719 guianense var. caudatum, 719 itauba, 722 kunthiana, 749 macrophyllum, 719 meissneri, 719 oppositifolium, 719 sprucei, 722 Actinostemon, 87 amazonicus, 87, 88 depauperatus, 88 parvifolius, 88 schomburgkii, 88, 88 Adelia, 89 triloba, 89, 89

805

Adenolisianthus, 479 arboreus, 480, 481 virgatus, 485 Adenophaedra, 90 grandifolia, 90, 90 Aeschynomene, 241 americana, 242 var. americana, 242 brasiliana, 242 var. brasiliana, 242 var. carichanica, 242, 244 emerus, 388 evenia, 242, 243 var. serrulata, 242 fluminensis, 242 var. fluminensis, 242 guarica, 242 histrix, 243 var. histrix, 243, 243 paniculata, 243 pratensis, 243 var. caribaea, 243, 244 rudis, 243 scabra, 244, 245 sensitiva, 245 var. sensitiva, 245 Aguacate, 744 Aguacatillo blanco, 592 Aguacatillo moisés, 710 Aiouea, 702 benthamiana, 703 guianensis, 703 laevis, 703 maguireana, 703, 704

806

I NDEX

[Aiouea] myristicoides, 703 pernitida, 703 perrobusta, 737 Ají de morrocoy, 496, 500 Ají de paloma, 605 Ajonjolí sabanero, 291 Ajumma, 92 Aka-yek, 403 Akayu-wajunai, 401 Alchornea, 91 castaneifolia, 91, 94 discolor, 91, 93 glandulosa, 92, 92 var. floribunda, 95 grandiflora, 92 orinocensis, 100 schomburgkii, 91 triplinervia, 92, 93 Alchorneopsis, 94 floribunda, 95, 95 Alcornoque, 264, 330 Alcornoque montañero, 330 Alcornoque sabanero, 264 Alcornoquillo, 327 Aldina, 245 aurea, 247 berryi, 247 discolor, 247 elliptica, 247 heterophylla, 248 kunhardtiana, 248, 251 latifolia, 248 var. auyantepuiensis, 248, 250 var. latifolia, 248 var. pubescens, 248 macrophylla, 248 var. macrophylla, 248, 252 var. yapacanensis, 248 paulberryi, 249, 253 petiolulata, 249 reticulata, 249, 252 sp. A, 249 sp. B, 249 sp. C, 249 sp. D, 250 sp. E, 250 sp. F, 250 sp. G, 250 yapacanensis, 248 Alectoroctonum

caracasanum, 147 Alexa, 254 bauhiniiflora, 258 canaracunensis, 258 confusa, 258 cowanii, 255, 258 herminiana, 256, 258 imperatricis, 257, 258 superba, 258 Alexandra, 254 imperatricis, 258 Algodón, 92 Algodoncillo, 92 Aliso blanco, 91 Allania, 245 Allantoma, 752 fagifolia, 757 lineata, 752, 753 Alloplectus, 544 circinatus, 557 coccineus, 557 congestus, 556 deltoideus, 556 metamorphophyllus, 568 microsepalus, 553 pallidus, 563 savannarum, 544, 545 Almendra, 756 Almendro, 342 Alysicarpus, 258 vaginalis, 259, 259 Amanoa, 95 almerindae, 96, 98 cupatensis, 96, 98 glaucophylla, 96 grandiflora, 99 guianensis, 96, 98 var. grandiflora, 99 oblongifolia, 97, 99 pubescens, 96 steyermarkii, 97, 99 Americana, 218 Amerimnon, 295 brownei, 297 pinnatum, 377 Amiña, 316 Amjadi, 309 Ampelodaphne arunciflora, 714 macrophylla, 714 Amuña, 316 Andira, 259 inermis, 260

subsp. inermis, 260, 260 retusa, 262 sp. A., 262 surinamensis, 261, 262 tervequinata, 262 trifoliolata, 261, 262 Aniba, 704 affinis, 706 burchellii, 706 canelilla, 706 cinnamomiflora, 706 citrifolia, 706, 707 excelsa, 706, 707 ferruginea, 706 guianensis, 706, 708 hostmanniana, 708, 709 kappleri, 709 megaphylla, 709 panurensis, 709 pittieri, 709 riparia, 709 rosaeodora, 709 simulans, 709 taubertiana, 709 trinitatis, 706 williamsii, 709 Anicillo, 296 Anila, 330 bongardiana, 331 Anilillo, 291 Añilito, 331 Anón, 401 Anontaya-yek, 120 Anonti-wa-yek, 111 Anthodon, 596 decussatum, 597, 597 laxiflorus, 617 Anthodus ellipticus, 613 Antidesma triplinervia, 92 Anzuelito, 330 Apalanthe granatensis, 642 Aparisthmium, 99 cordatum, 99, 100 Apodandra, 207 loretensis, 208 Apophragma, 494 tenuifolium, 496 Aporu-ban-yek, 654 Apoykapundek, 445 Araburi, 119

I NDEX

Aramasa, 177 Arapurihusi, 119 Arapurima, 117 Arcyphyllum, 385 Arepillo, 239, 297, 343, 419 Arepito, 379 Arestín, 309 Argythamnia polygama, 143 Aribái-panáru-kusí, 48 Aribái-panáru-kusí-yek, 49 Arisoru, 262 Aristín, 331 Aro de atarraya rebalsero, 719 Aru-ca-yu-dek, 316 Ascra, 436 Asterandra, 191 Asteranthos, 754 brasiliensis, 754, 754 Asterolepidion, 649 elatum, 649 Astrococcus, 100 coriaceus, 151 cornutus, 100, 101 Ateramnus hypoleucus, 150 Athenaea guianensis, 445 Atón-yek, 67 Aubria, 633 Automoyek, 248 Avesicaria neottioides, 792 Aydendron affine, 706 cannella, 718 citrifolium, 706 curvirameum, 710 hostmannianum, 709 panurense, 709 trinitatis, 706 Azara umbellata, 460 —B— Badunuwa, 463 Bainepá, 342 Balaústre, 271 Ballota suaveolens, 689 Banara, 436

glandulosa, 437 guianensis, 437, 438 var. mollis, 437 mollis, 437 nitida, 437, 438 orinocensis, 437 sacupanensis, 438 Barbasco, 196, 280, 338, 355, 420 Barbasco blanco, 338 Barbasco blanco caicareño, 338 Barbasco bravo, 338 Barbasco caicareño, 420 Barbasquillo, 428 Barbieria, 262 pinnata, 263, 263 polyphylla, 263 Barthollesia, 755 Barui de venado, 500 Beilschmiedia, 709 curviramea, 710, 710 Bejuco arepillo, 239 Bejuco colorado, 297 Bejuco de estribo, 297 Bejuco de jabón, 313 Bejuco de morrocoy, 208 Bejuco de murciélago, 342 Bejuco de pereza, 615 Bejuco de rana, 472 Bejuco de raya, 732 Bejuco de sapo, 602 Bejuco de zamuro, 313, 315, 358 Bejuco des amni, 313 Bejuco roblito, 297 Bergena, 774 Bernardia, 101 amazonica, 101, 101 sect. Adenophaedra, 90 Bertholletia, 755 excelsa, 755, 756 nobilis, 756 Besleria, 545 coccinea, 557 flavo-virens, 546 gibbosa, 546 incarnata, 571 laxiflora, 546, 547 parviflora, 548 penduliflora, 547, 548 pulchella, 549 sanguinea, 555

807

serrulata, 558 sp. A, 548 sp. B, 548 sp. C, 548 spectabilis, 558 Bihai marginata, 586 revoluta, 586 Biovularia olivacea, 792 Boborutepe, 171 Boca, 436 Bola de fuego, 43 Borrachero, 470 Bowdichia, 263 brasiliensis, 316 martiusii, 316 racemosa, 316 var. kaieteurensis, 316 virgilioides, 264, 264 Brachycodon, 498 pumilus, 501 ramosissimus, 501 Brachyloma rhodomallos, 561 Brachyloma, 561 Bradburya, 271 angustifolium, 273 brasilianum, 273 macrocarpa, 273 pascuora, 276 plumieri, 276 triquetra, 276 venosum, 276 virginianum, 276 Brasil, 325 Bucare, 323 —C— Cabarí, 280 Cabeza de negro, 64 Cabo de vela, 473 Cabullo, 776 Cacaíto, 765, 766 Cacao, 764, 766 Cacao grande, 764 Cachete de vieja, 605, 613, 617 Cachete de viejo, 613 Cachicamo montañero, 316 Cachimbo, 167, 757, 771, 775

808

I NDEX

Cachimbo de mono, 772 Cachimpapudek, 171 Cachito, 64 Cacho, 105 Cacú, 382 Cadena, 492 Cadillo, 394 Caicareno, 444 Caicareño, 196, 216, 258, 278, 444 Caicareño montañero, 258 Caimito cimarrón, 461 Cajan, 264 Cajanus, 264 cajan, 265, 265 Cajario, 403 Cajoro, 592 Calabazín, 358 Calolisianthus imthurnianus, 483 tatei, 501 tepuiensis, 516 Calopogonium, 266 coeruleum, 266, 267 mucunoides, 266, 267 Calpidisca amethystina, 789 calycifida, 790 hispida, 791 humboldtii, 791 modesta, 789 roraimensis, 789 Camaji, 764, 766, 776 Canalete amarillo, 377 Canau, 724 Canavali, 267 Canavalia, 267 brasiliensis, 268, 268 dictyota, 269 fendleri, 268 grandiflora, 269 maritima, 269 rosea, 269 sericophylla, 269 Candilón, 403 Canelito, 120 Canelito negro, 262, 423 Canelo, 119, 120, 121 Canelón, 119, 120 Caniba, 109, 658 Canjilón de agua, 403 Cañoflote, 92 Caobillo, 750

Capa de tabaco, 757 Caperonia, 102 castaneifolia, 102, 102 Capibare, 658 Caracamate, 734 Caramacate, 206 Caramacate blanco, 459 Caramacatillo, 438 Caraota, 357 Caraota montañera, 330 Caraota rebalsera, 296 Caraotillo, 267 Carcanapire, 117, 120, 123 Carcanapire rebalsero, 116 Carpotroche, 438 amazonica, 440, 467 crispidentata, 440 grandiflora, 439, 440 laxiflora, 465 paludosa, 465 zuliana, 467 Carptotepala, 41 insolita, 45 jenmanii, 45 Carrasposo, 402 Cartán, 271 Caruana, 100, 109 Caruguano, 147 Carurú, 117 Carutillo, 772 Cascarare, 766 Casearia, 440 aculeata, 444, 449 arborea, 444 arguta, 445, 446 celastroides, 448 celtidifolia, 448 commersoniana, 444 decandra, 444, 450 densiflora, 444 grandiflora, 444, 450 guianensis, 445, 450 guidonia, 473 hirsuta, 445, 451 inaequilatera, 447 javitensis, 445, 452 laevigata, 448 lasiophylla, 446 laurifolia, 444 lingua, 448 macrophylla, 446 mariquitensis, 445, 452 mollis, 446, 453

neblinae, 446 negrensis, 461 pitumba, 446, 453 prunifolia, 446 rusbyana, 446 singularis, 447, 452 spinescens, 447, 454 spruceana, 447, 454 sylvestris, 447 var. carpinifolia, 447 var. lingua, 447, 451 var. sylvestris, 448 tremula, 448, 449 ulmifolia, 448, 451 zizyphoides, 448, 453 Casimirella, 648 ampla, 648, 649 Casinga procera, 461 suaveolens, 461 Caspadillo, 218 Cassytha, 710 filiformis, 710, 710 Caucho, 153, 221 Caucho banero, 153 Caucho blanco, 221 Caucho fino, 153 Caucho legítimo, 153 Caucho yapi, 153 Cavi-si-yek, 732 Cayenito, 437 Cazabe, 117 Cazabe de tara, 461 Cebolleta, 659 Cedrito, 459 Celaenodendron, 206 Celianella, 103 montana, 103, 103 Celiantha, 481 bella, 482, 483 chimantensis, 482, 483 imthurniana, 482, 483 Centrolobium, 269 orinocense, 271 paraense, 269 var. orinocense, 270, 271 var. paraense, 271 patinense, 271 Centrosema, 271 angustifolium, 273 brasilianum, 273, 275 var. angustifolium, 273 macrocarpum, 273, 274

I NDEX

molle, 273 pascuorum, 274, 276 plumieri, 274, 276 pubescens, 273, 276 tetragonolobum, 276 triquetrum, 275, 276 venosum, 275, 276 virginianum, 276 Centrosolenia hirsuta, 563 Cephaloxys, 672 Cercophora, 774 anomala, 776 Cereipo, 361 Chaetocalyx, 276 fissa, 277 magniflora, 277 nigrescens, 277 paucifolia, 277 perglandulosa, 277 pubescens, 277 scandens, 277 var. pubescens, 277, 277 vestita, 277 Chaetocarpus, 104 schomburgkianus, 104, 105 stipularis, 105 williamsii, 105 Chaetocrater javitensis, 445 Chamaesyce, 105 dioeca, 106 hirta, 106, 106 hypericifolia, 107 hyssopifolia, 107 prostrata, 106, 107 thymifolia, 107 Chamanare, 400, 401, 402, 403, 404, 405 Chamanare de tierra firme, 399 Chamanare negro, 402 Chamanare orillero, 400 Chaparillo, 344, 367 Chapere, 308 Chaperno, 342 Charo macho, 461 Cheiloclinium, 597 anomalum, 599 belizense, 599, 600 cognatum, 600, 600 diffusiflorum, 599

glaziovii, 599 gleasonianum, 599 habropodum, 599 hippocrateoides, 599 jenmanii, 599 klugii, 599 krukovii, 599 lineolatum, 599 lucidum, 599 meianthemum, 599 obtusum, 599 parviflorum, 599 podostemmum, 599 Chelonanthus, 483 acutangulus, 485 alatus, 485 albus, 485, 487 angustifolius, 485, 487 arboreus, 516 bifidus, 486 campanuloides, 501 cardonae, 500 chelonoides, 485 fruticosus, 481 longistylus, 486 purpurascens, 486 pyriformis, 486 schomburgkii, 486 uliginosus, 486 Chimaco, 240 Chimako, 280 Chimanare, 404 Chinak guaiquin-chinaten, 327 Chorisepalum, 488 acuminatum, 489 breweri, 489 carnosum, 488, 490 ovatum, 489 var. sipapoanum, 491 psychotrioides, 489, 490 var. acuminatum, 489 rotundifolium, 489, 490 sipapoanum, 489, 491 Chrysothemis, 548 dichroa, 549, 549 pulchella, 549, 549 Chytroma, 774 chartacea, 775 corrugata, 776 rosea, 776 Cimbra pote, 599 Cinnamomum, 711

809

areolatocostae, 744 filamentosum, 712 semecarpifolium, 711, 712 triplinerve, 712 Cipura, 659 paludosa, 659, 660 rupicola, 659 sp. A, 661 Clathrotropis, 278 brachypetala, 278, 279 brunnea, 280 colombiana, 280 glaucophylla, 280 macrocarpa, 280 nitida, 280 Clavapetalum, 649 elatum, 649 surinamense, 649 Claytonia nemorosa, 500 Cleidion nicaraguense, 87 Clinopodium chamaedrys, 698 Clitoria, 280 amoena, 282 angustifolia, 273 arborescens, 282 brasiliensis, 273 cajanifolia, 284 canescens, 282 cavalcantei, 282 cerifera, 282 coriacea, 282 dendrina, 283, 285 falcata, 283 var. falcata, 283, 287 glycinoides, 283 grandifolia, 283 guianensis, 283 var. guianensis, 283, 286 guyanensis, 283 javitensis, 283 var. glabra, 284 var. grandifolia, 283 var. javitensis, 283, 286 laurifolia, 284 parvifolia, 284 pinnata, 263 plumieri, 276 poitaei, 282 polyphylla, 263

810

I NDEX

[Clitoria] rubiginosa, 283 sagotii, 284 var. canaliculata, 284, 285 var. sprucei, 284 sect. Centrosema, 271 simplicifolia, 284, 287 steyermarkii, 284 virginiana, 276 Clusiophyllum sprucei, 178 Cnidoscolus, 107 urens, 107, 108 Cobija, 105 Coco de mono, 763, 765, 766, 770, 771, 772, 776 Coco de mono rebalsero, 770 Cococoroli, 592 Cocojoro, 592 Cocura, 109 Codonanthe, 550 bipartita, 550 calcarata, 550, 551 confusa, 550 crassifolia, 550, 551 dissimulata, 552 ulei, 553 Codonanthopsis, 552 dissimulata, 552, 552 ulei, 553 Coeppertia caudata, 734 Cojón de verraco, 247, 248 Cola de pava, 772 Cola de zorro, 394 Cola de Zorro, 394 Columnea, 553 affinis, 555 aureonitens, 555 calotricha var. austroamericanarum, 545 coccinea, 557 microsepala, 553, 554 sanguinea, 554, 555 steyermarkii, 545 subg. Pentadenia, 553 Comanthera, 41 kegeliana, 45 linderi, 45 Comé-yek, 775

Conami, 191 brasiliensis, 196 Conceveiba, 108 cordatum, 100 guianensis, 109, 109 hostmannii, 109 latifolia, 111 martiana, 111 ptariana, 110, 111 sect. Gavarretia, 147 Conceveibastrum, 108 martianum, 111 ptarianum, 111 Conejita, 122 Congrillo blanco, 188 Congrio, 188, 240, 262, 316, 402 Congrio blanco, 316 Congrio rebalsero, 240 Conori, 177, 641 Conuri, 153 Conuri banero, 177 Conuri de tierra firme, 177 Conuri propio, 177 Copa de tabaca, 765 Coquito, 776 Cordilla, 447 Coreco, 258 Cornoco, 264 Coro-cororu, 592 Coronilla scandens, 277 Corrona, 764 Corytholoma, 571 Corytoplectus, 555 congestus, 556 deltoideus, 555, 556 Cosoiba, 530 Coublandia, 360 frutescens, 361 Coumarouna, 318 magnifica, 319 odorata, 319 oppositifolia, 419 punctata, 320 Couratari, 756 fagifolia, 757 guianensis, 757, 758 lineata, 752 multiflora, 757, 759 oligantha, 757 pulchra, 757 sandwithii, 757

stellata, 757, 758 Couroupita, 761 guianensis, 760, 761 venezuelensis, 761 Coursetia, 287 arborea, 288 ferruginea, 288, 288 Coutoubea, 491 minor, 492, 493 racemosa, 492, ramosa, 492 493 f. racemosa, 492 var. racemosa, 492 reflexa, 492 spicata, 493, 494 Crineja, 433 Crotalaria, 289 anagyroides, 290 var. pauciflora, 290 bracteata, 291 cunifolia, 291 depauperata, 290 guianensis, 283 incana, 290 lectophylla, 290 maypurensis, 290 var. depauperata, 290, 292 var. maypurensis, 290, 292 micans, 290 nitens, 291 pilosa, 291, 293 var. skutchii, 291 pterocaula, 291 retusa, 291 retusifolia, 291 sagittalis, 291 schiedeana, 291 spectabilis, 291 stipularia, 291, 293 stipularis, 291 velutina, 291, 293 Croton, 111 argenteus, 116 argyrophyllus, 116 barlettii, 121 benthamianus, 120 bolivarensis, 117 cajucara, 117 cardonei, 120 castaneifolius, 102 conduplicatus, 117

I NDEX

crenatus, 119 cuneatus, 117, 124 essequiboensis, 117 fragrans, 118 glandulosus var. hirtus, 118 gossypiifolius, 118 guaiquinimae, 118 guianensis, 118 hirtus, 118 icabarui, 118 kavanayensis, 119 lanjouwensis, 119 lobatus, 119 matourensis, 119, 125 var. benthamianus, 119 megalodendron, 119, 126 miquelensis, 122 mollis, 119 monachinoensis, 117 neblinae, 120, 126 nepetifolius, 120, 127 nervosus, 116 nuntians, 120 orinocensis, 120 pakaraimae, 119 palanostigma, 120, 127 polygamus, 143 populifolius var. essequiboensis, 118 potaroensis, 120, 125 pullei, 121 roraimensis, 121, 125 var. subinteger, 121 schiedeanus, 121, 127 scutatus, 121, 128 sessiliflorum, 222 sipaliwinensis, 121, 129 sp. A, 123, 131 sp. B, 123 sp. C, 123 sp. D, 123 sp. E, 132 sp. F, 132 sp. G, 132 sp. H, 133 spiraeifolius, 122 spruceanus, 122 subcoriaceus, 122, 130 subincanus, 122, 130 subserratus, 122, 130 surinamensis, 117 trinitatis, 122, 129

umbratilis, 122 vergarenae, 123, 132 xanthochloros, 119 yavitensis, 123, 129 Cryptocarya canelilla, 706 laevis, 703 Cuai-eneru, 383 Cuervea, 600 kappleriana, 600, 601 mitchellae, 600 ovalifolia, 617 Cujicillo, 242 Cumariche, 341 Cumaroa, 528 Cunuri, 177 Cunuri blanco, 177 Cunuria, 176 casiquiarensis, 212 crassipes, 178 glabra, 177 spruceana, 178 Cupana, 404 Cupana de chamanare, 404 Cupi, 553 Curabayek, 597 Curarina, 511 Curataquilla, 229 Curatari, 756 Curataria, 756 Curawei-yek, 607 Cureo barero, 734 Curtia, 494 conferta, 495, 496 obtusifolia, 495, 496 quadrifolia, 496 tenella, 496 tenuifolia, 495, 496 subsp. tenella, 496 Cururú luiro, 188 Cusapoi-yek, 119 Cuspa, 445 Cuyubi, 656 Cyclolobium amazonicum, 379 hostmannii, 379 Cymbosema, 294 apurense, 295 roseum, 294, 295 Cypella, 661 linearis, 660, 661 Cyrillopsis, 665 micrantha, 665, 667

811

paraensis, 666, 667 Cytisus cajan, 265 sessiliflorus, 420 violaceous, 321 —D— Dactylostemon, 87 schomburgkii, 88 Dalbergaria, 553 aureonitens, 555 sanguinea, 555 Dalbergia, 295 amazonica, 296 aturensis, 298 brownei, 297 ecastophyllum, 297 foliosa, 297, 299 frutescens, 297 var. frutescens, 297, 300 glauca, 297 heptaphylla, 336 hygrophila, 297, 300 intermedia, 297, 299 inundata, 298, 299 monetaria, 298, 301 pachycarpa, 298 pentaphylla, 336 riedelii, 298, 300 sp. A, 298 sp. B, 298 sp. C, 298 sp. D, 298 spruceana, 298, 301 subcymosa, 298 tomentosa, 297 variabilis, 297 Dalechampia, 133 affinis, 135, 137 attenuistylus, 135 brevipes, 139 brownsbergensis, 135 colorata, 139 cynanchoides, 139 dioscoreifolia, 135 dioscoreifolia, 136 heterobractea, 136, 138 heterophylla, 139 latifolia, 139 liesneri, 136 magnoliifolia, 136 megacarpa, 136

812

I NDEX

[Dalechampia] micrantha, 136 mollis, 139 papillistigma, 136 parvibracteata, 136 peruviana, 139 pruriens, 139 roezliana var. amazonica, 136 rubiformis, 139 ruboides, 139 scandens, 138, 139 sidaefolia, 139 tenuiramea, 139 tiliifolia, 138, 139 villosa, 139 Damai, 420 Dau bagibagi, 403 Dawadema, 401 Deguelia scandens, 304 De-ma-ta-gu-tu, 109 Dendrobangia, 649 boliviana, 649, 650 tenuis, 649 Dendrostigma, 465 Dendrostylis, 465 grandifolia, 467 Dendrothrix, 139 multiglandulosa, 141 yutajensis, 140, 141 Dequira-arepillo, 258 Derello, 605 Derris, 302 amazonica, 302, 303 ernestii, 336 floribundus, 335 glabrescens, 335 guyanensis, 304 hedyosma, 335 latifolia, 336 negrensis, 302, 303 nicou, 338 pterocarpus, 304, 304 scandens, 304 urucu, 338 utilis, 338 Desmodium, 304 adscendens, 307 var. orinocense, 308 affine, 307 asperum, 308 axillare, 307

var. acutifolium, 307 var. sintenisii, 307 var. stoloniferum, 307 barbatum, 307, 310 cajanifolium, 307 campyloclados, 308 canum, 308 distortum, 308 dubium, 308 flexuosum, 308 hickenianum, 308 incanum, 308 orinocense, 308, 309 pachyrrhizum, 308 procumbens, 308 var. procumbens, 308 sclerophyllum, 309 scorpiurus, 309 tortuosum, 309 triflorum, 309 wydlerianum, 310 Diastema, 556 racemiferum, 556, 556 Dibrachion brasiliensis, 316 riparium, 316 Dioclea, 310 albiflora, 312 apurensis, 312 broadwayana, 312 elliptica, 315 glabra, 315 guianensis, 312 holtiana, 312 macrantha, 312 macrocarpa, 313 malacocarpa, 313 reflexa, 313, 314 rigida, 313 ruddiae, 313 scabra, 313 var. brownii, 313 var. scabra, 315 steyermarkii, 315 virgata, 315 var. crenata, 315 var. virgata, 315 Diplotropis, 315 brachypetala, 278 brasiliensis, 316 grandiflora, 367 martiusii, 316, 318 nitida, 280

purpurea, 316 var. purpurea, 316, 317 racemosa, 316 var. kaieteurensis, 316 var. racemosa, 316 sp. A, 318 strigulosa, 317 triloba, 318 Dipteryx, 318 applanata, 419 cordata, 416 crassifolia, 416 magnifica, 319 odorata, 319, 319 oppositifolia, 419 var. parviflora, 419 phaeophylla, 392 punctata, 320 reticulata, 419 rigida, 416 rosea, 320 trifoliata, 320 Discocarpus, 141 brasiliensis, 141 essequeboensis, 141 gentryi, 141, 142 mazarunensis, 105 spruceanus, 142, 142 Discophora, 650 froesii, 651 guianensis, 651, 651 panamensis, 651 Ditaxis, 142 polygama, 143, 143 Ditomaga nemorosa, 500 Dolicholus, 385 kuntzei, 386 phaseoloides, 386 pittieri, 388 Dolichos altissmus, 358 cunifolius, 291 erosus, 373 luteolus, 428 maritimus, 269 minimus, 386 pruriens, 357 roseus, 269 scabra, 313 uncinatus, 423 urens, 358 virgatus, 315

I NDEX

Dormidera, 242 Drago, 382 Drepanocarpus, 339 aristulatus, 342 crista-castrensis var. latifolium, 343 dubius, 342 ferox, 342 ß macrophyllum, 342 frondosus, 343 inundatus, 343 lunatus, 343 ovalifolium, 342 paludicola, 298 venezuelensis, 342 Drymonia, 557 coccinea, 557 serrulata, 558, 558 Drypetes, 143 fanshawei, 144 maguireana, 105 spruceana, 105 variabilis, 144, 145 Duara blanca, 171 Dudu, 401, 404 Dujo, 444, 446, 459 —E— Eburu, 298 Ecastaphyllum, 295 amazonicum, 296 foliosum, 297 glaucum, 297 hygrophilum, 297 monetaria var. riedelii, 298 riedeli, 298 tomentosum, 297 Echepeñ, 117 Elachyptera, 601 floribunda, 602, 602 Elaeophora, 207 abutifolia, 209 polyadenia, 209 Elodea, 642 granatensis, 642, 643 orinocensis, 642 Elshotzia, 761 Emmotum, 651 acuminatum, 652 argenteum, 654 celiae, 652

conjunctum, 653, 654 floribundum, 654 fulvum, 653, 654 glabrum, 654 nudum, 652 ptarianum, 654 yapacanum, 654 Endlicheria, 712 anomala, 713 arunciflora, 714 bracteolata, 714 canescens, 714 chalisea, 714 dictifarinosa, 714 directonervia, 714 gracilis, 714 grandis, 749 levelii, 714 macrophylla, 714 maguireana, 703 multiflora, 714, 715 nilssonii, 714 reflectens, 716 rubriflora, 716 szyszylowiczii, 716 vinotincta, 716 wurdackiana, 716 Endopleura, 624 uchi, 625, 625 Enicostema, 496 verticillatum, 497, 498 Episcia, 558 adenosiphon, 563 cordata, 563 cuneata, 573 fimbriata, 559, 560 hirsuta, 563 porphyrotricha, 564 reptans, 559, 560 resioides, 564 sp. A, 559 sp. B, 559 xantha, 559 Epoca-po-yek, 654 Eriocaulaceae, 1 Eriocaulon, 12 aequinoctiale, 13 aquaticum, 13 atabapense, 14 bifidum, 21 bisumbellatum, 42 bonplandianum, 44 brevifolium, 14

813

caulescens, 42 cinereum, 13, 14 dimorphopetalum, 13 fasciculatum, 22, 23 flavescens, 18 gracile, 44 guyanense, 13 humboldtii, 13, 15 jauense, 13 klotzschii, 14 melanocephalum, 13, 14 nitens, 46 sieboldianum, 13 spongiola, 13, 17 steyermarkii, 13, 16 tenue, 47 tenuifolium, 14, 16 tortile, 27 umbellatum, 44, 48 xeranthemoides, 49 Eriope, 679 crassipes, 679 subsp. crassipes, 679, 680 Eriosema, 320 crinitum, 321 var. crinitum, 321 var. stipulare, 321, 322 edule, 386 rufum, 321 var. rufum, 321, 322 simplicifolium, 321 var. micranthum, 321 stipulare, 321 violaceum, 321 volubile, 386 Ero-cuaja, 315 Erythrina, 322 fusca, 323, 324 glauca, 323 mitis, 323 poeppigiana, 324, 324 rubrinervia, 324 Erythroxylaceae, 59 Erythroxylum, 59 acutifolium, 63 aff. subrotundum, 67 albertianum, 66 amazonicum, 63, 68 areolatum, 67 aristigerum, 67 var. bahiense, 67 bahiense, 67

814

I NDEX

[Erythroxylum] cataractarum, 63 christii, 67 citrifolium, 63 comosum, 66 cuatrecasasii, 63 divaricatum, 63, 68 duckei, 63 filipes, 65 fimbriatum, 63 floribundum, 65 foetidum, 63 gomphioides, 63 gracilipes, 63, 69 guanchezii, 64 havanense, 64 var. continentis, 64 hypoleucum, 64 impressum, 64, 69 kapplerianum, 64, 70 kirkianum, 66 laurinum, 65 lenticellosum, 64 ligustrinum, 64 var. grandifolium, 64 lindemanii, 65 lineolatum, 65 lucidum, 65 macrophyllum, 65 var. macrophyllum, 65, 71 var. savannarum, 66 mapuerae, 64 micranthum, 63 mucronatum, 66, 70 novogranatense var. macrophyllum, 63 obtusum, 64 oreophilum, 66, 68 orinocense, 66, 71 ovatum, 64 paraense, 63 recurrens, 63 roraimae, 66 rufum, 66 schomburgkii, 66 spruceanum, 67 squamatum, 67 var. emarginatum, 67 var. microcarpum, 67 var. orinocense, 64 steyermarkii, 67, 69 suberosum, 67, 70

testaceum, 67 trinerve, 67 venezuelense, 66 vernicosum, 67 var. oreophilum, 66 williamsii, 67, 71 Eschweilera, 761 alata, 763 alutacea, 775 bracteosa, 763, 767 chaffanjonii, 765 chartacea, 775 collina, 763 conduplicata, 776 coriacea, 764 corrugata, 776 decolorans, 764, 768 floribunda, 765 gracilipes, 765 grata, 765 laevicarpa, 764 longipes, 766 micrantha, 764 neblinensis, 765 obtecta, 770 odora, 764 paniculata, 765 parviflora, 765, 767 parvifolia, 765, 769 pedicellata, 765 revoluta, 766 rionegrense, 766 roraimensis, 766 subglandulosa, 766, 769 tenuifolia, 768, 770 Escoba, 64 Escobo, 66 Escobo negro, 64 Espuelo de gallo, 444 Etaballia, 325 dubia, 325, 325 guianensis, 325 Euceraea, 455 nitida, 456, 456 rheophytica, 455, 457 sleumeriana, 457 Euphorbia, 145 brasiliensis, 107 caracasana, 147 comosa, 147 cotinifolia, 146, 147 cotinoides, 147 dioeca, 106

globulifera, 106 heterophylla, 147 hirta, 106 humayensis, 147 hypericifolia, 107 hyssopifolia, 107 prostrata, 107 spruceana, 147 tennella, 107 thymifolia, 107 tithymaloides, 186 Euphorbiaceae, 72 Euphronia, 229 acuminatissima, 229, 230 guianensis, 229, 230 hirtelloides, 229, 230 licanioides, 229 Euphroniaceae, 228 Evonymodaphne armeniaca, 718 Exacum guianense, 521 racemosum, 492 spicatum, 494 tenuifolium, 496 —F— Fabaceae, 231 Fairchildia, 394 Falso coco de mono, 771 Fissicalyx, 326 fendleri, 326, 326 Flacourtia benthamii, 472 nitida, 472 Flacourtiaceae, 434 Flor de conejo, 550 Flor de mayo, 212 Fregosillo, 492 Frijol de sapo, 428 Frijole de la lagartilla, 267 Frijolillo, 312 Fruta de paloma, 66, 444 Fruto de burro, 470 Fruto de paloma, 63, 64 —G— Gaedawakka schomburgkianus, 105 Galactia, 327 angustifolia, 327

I NDEX

camporum, 327 gracillima, 327 jussiaeana, 327, 327 var. angustifolia, 327 pinnata, 263 striata, 327 var. tenuiflora, 327 tenuiflora, 327 Galega cathartica, 420 cinerea, 420 sinapou, 420 toxicaria, 422 Garrapato, 496 Gaspadillo, 218 Gaspadillo marrón, 599 Gavarretia, 147 terminalis, 147, 148 Generala, 283, 290, 420 Gengibrillo, 599 Genlisea, 782 anfractuosa, 783 esmeraldae, 783 filiformis, 783, 785 glabra, 783, 785 guianensis, 783, 784 nigrocaulis, 783 pygmaea, 783, 785 repens, 783, 785 roraimensis, 783, 784 sanariapoana, 783, 784 Gentiana aphylla, 536 verticillata, 498 Gentianaceae, 474 Geoffroea inermis, 260 retusa, 262 surinamensis, 262 Gesneria caracasana, 572 elatior, 571 gollmeriana, 572 hirsuta, 561 lindeniana, 572 rubricaulis, 561 sect. Corytholoma, 571 vargasii, 572 Gesneriaceae, 542 Gitara, 86 venezolana, 87 Gliricidia, 328 sepium, 328, 328

Glochidion vacciniifolium, 200 Glycine crinita, 321 phaseoloides, 386 punctata, 380 reflexa, 386 reticulata, 386 rufa, 321 simplicifolia, 321 striata, 327 tenuiflora, 327 Glycydendron, 149 amazonicum, 149, 149 Gnetaceae, 573 Gnetum, 574 amazonicum, 576 camporum, 574 cruzianum, 576 leyboldii, 576 var. woodsonianum, 576 melinonii, 576 microstachyum, 576 nodiflorum, 576 oblongifolium, 576 paniculatum, 576 paraense, 576 schwackeanum, 575, 576 thoa, 576 urens, 576 var. camporum, 574 Goeppertia anomala, 713 multiflora, 714 reflectens, 716 sericea var. bracteolata, 714 Golondrina, 107, 428 Goma, 153 Goma brava, 153 Goma concha blanca, 153 Goma fina, 153 Goma rebalsera, 153 Goma seringa, 153 Guaboa, 423 Guacapú, 280 Guacharaco, 776 Guacharaco amarillo, 776 Guacharaco blanco, 776 Guacharaco rojo, 776 Guacharaco rosado, 776 Guachimaca, 171 Guaco, 356

815

Guada-yek, 724 Guajenera, 383 Guamo, 399, 400 Guamo peludo, 258 Guamo terciopelo, 403 Guanábana silvestre, 437 Guanacoco, 366 Guanak, 43 Guapocoji, 280 Guaquito, 401, 402 Guaraba, 403 Guaraguao, 358 Guarania laurifolia, 213 Guaritoto, 107 Guatero, 771, 772 Guatoso, 771, 772 Guayabo rebalsero, 96 Guayaparu, 401 Guayapu, 401 Guidonia apetala, 460 calophylla, 461 coriacea, 461 cupulata, 461 hirsuta, 445 procera, 461 spinescens, 447 suaveolens, 461 tremula, 448 ulmifolia, 448 Guinda, 448 Gusana, 242 Gustavia, 770 acuminata, 771 acuta, 772 augusta, 771, 773 coriacea, 772 hexapetala, 772, 774 poeppigiana, 772, 774 pulchra, 772 yaracuyensis, 772 Gymnanthes, 149 hypoleuca, 150, 150 Gymnobalanus fendleri, 734 —H— Haematostemon, 151 coriaceus, 151, 151 Haemodoraceae, 576 Haloragaceae, 581

816

I NDEX

Haloragis tetrandra, 582 Hebepetalum, 618 humiriifolium, 619, 619 parviflorum, 620 punctatum, 620 roraimensis, 619 schomburgkii, 623 sp. A, 620 Heburu, 735 Hecastophyllum dubium, 325 Hecatostemon, 457 completus, 457, 458 dasygynus, 458 guazumaefolius, 458 Hedysarum adscendens, 307, 308 asperum, 308 axillare, 307 barbatum, 307 brasilianum, 242 cajanifoliun, 307 canum, 308 diphyllum, 432 distortum, 308 ecastaphyllum, 297 gemellum, 432 hamatum, 394 var. viscosum, 394 incanum, 308 procumbens, 308 scorpiurus, 309 stoloniferum, 307 tortuosum, 309 triflorum, 309 vaginale, 259 Helia acutangula, 485 alata, 485 alba, 485 angustifolia, 485 arborea, 481 bifida, 486 caerulescens, 534 campanuloides, 501 chelonoides, 485 elisabethae, 525 nemorosa, 500 pratensis, 501 pumila, 501 purpurascens, 486 ramosissima, 501

schomburgkii, 486 trifidus, 485 uliginosa, 486 Heliconia, 583 acuminata, 584 subsp. acuminata, 585, 587 subsp. occidentalis, 585 aurea, 585 bihai, 585, 588 cannoidea, 586 caribaea, 585 chartacea, 585, 589 costanensis, 585 densiflora, 585 subsp. angustifolia, 585 subsp. densiflora, 585, 586 episcopalis, 585 glauca, 586 hirsuta, 585, 587 humilis, 585 julianii, 585, 590 lourteigiae, 585 marginata, 586, 589 f. lutea, 586 platystachys, 586, 587 psittacorum, 586, 588 var. rhizomatosa, 586 revoluta, 586 richardiana, 586 schaeferiana, 585 spathocircinada, 586, 590 stricta, 585 Heliconiaceae, 583 Helleria, 640 Hermesia castaneifolia, 91 Hernandia, 592 guianensis, 591, 592 sonora, 592 Hernandiaceae, 592 Hevea, 152 benthamiana, 153, 154 confusa, 153 guianensis, 153 var. guianensis, 153 var. lutea, 153, 155 lutea, 153 microphylla, 153 minor, 153 paludosa, 153 pauciflora, 153

var. coriacea, 153, 156 Hicaquillo, 99 Hippocratea, 602 aspera, 608 comosa, 604 crinita, 604 floribunda, 602 huanucana, 605 kappleriana, 600 mitchellae, 600 multiflora, 613 opacifolia, 615 tenuiflora, 610 verrucosa, 610 volubilis, 602, 603 Hippocrateaceae, 594 Hippomane biglandulosa, 219 glandulosa, 219 Hisingera benthamii, 472 Hoelzelia, 394 Hoja de danta, 100 Hoja de gavilán, 515 Hoja de venado, 617 Hoja del sol, 405 Hoja salado, 472 Holopyxidium, 774 Homalium, 458 anzoateguiense, 459 densiflorum, 459 eleuthrostylum, 459 eurypetalum, 459 guianense, 459, 459 mituense, 459 napimoga, 459 pedicellatum, 459 pittieri, 459 puberulum, 459 racemosum, 459, 459 racoubea, 459 Huertia, 394 Hugoniaceae, 618 Humboldtiella ferruginea, 288 Humiria, 626 balsamifera, 627 var. balsamifera, 628, 630 var. coriacea, 628 var. floribunda, 628 var. guaiquinimana, 628 var. guianensis, 628

I NDEX

var. iluana, 628 var. imbaimadaiensis, 628 var. laurina, 629 var. pilosa, 629 var. savannarum, 629 var. stenocarpa, 629 var. subsessilis, 629 cassiquiari, 628 crassifolia, 629, 630 floribunda, 628 var. guianensis, 628 var. laurina, 629 var. spathulata, 629 var. subsessilis, 629 fruticosa, 629, 630 pilosa, 629 savannarum, 629 wurdackii, 629, 630 Humiriaceae, 623 Humirianthera, 648 ampla, 649 duckei, 649 Humiriastrum, 631 colombianum, 632 cuspidatum, 632 liesneri, 632 obovatum, 632 ottohuberi, 632 piraparanense, 632, 633 Humirium balsamiferum, 627 cuspidatum, 632 floribundum, 628 guianense, 628 oblongifolium, 637 obovatum, 632 surinamense, 628 Hura, 158 crepitans, 157, 158 Hybosema, 328 Hydrocharitaceae, 641 Hydrolea, 646 elatior, 645, 646 minima, 646 multiflora, 646 spinosa, 645, 646 Hydromystria laevigata, 644 Hydrophyllaceae, 644 Hyeronima, 158 alchorneoides, 158, 160 laxiflora, 158

oblonga, 159, 159 Hylenaea, 603 comosa, 604, 604 Hymenolobium, 329 heterocarpum, 329, 330 petraeum, 330 Hypenia, 681 salzmannii, 680, 681 Hypocyrta crassifolia, 550 gibbosa, 546 sect. Codonanthe, 550 Hyptidendron, 681 arboreum, 681, 682 Hyptis, 682 arborea, 681 atrorubens, 685, 690 belizensis, 686 brachiata, 685, 690 brevipes, 685, 691 var. serrata, 685 canescens, 688 capitata, 685 colubrimontis, 686, 688 conferta, 686 var. angustata, 686, 692 constricta, 686 crenata var. angustifolia, 686 dilatata, 686, 695 excelsa, 686 ferruginea, 686 guanchezii, 686 hirsuta, 686, 695 huberi, 687, 695 jurgensenii, 686 var. angustata, 686 laciniata, 687, 689 lacustris, 687 lanceolata, 687, 694 lanicephala, 686 lantanifolia, 687, 689 lundii, 686 luticola, 687, 692 micrantha, 688 microphylla, 687, 692 mutabilis, 687, 694 var. canescens, 688 parkeri, 688, 689 var. verbenifolia, 688 pectinata, 688, 691 polystachya, 688 pyriformis, 688

817

var. pyriformis, 688, 693 var. yutajeensis, 688, 693 recurvata, 688 var. megacephala, 689 salzmannii, 681 sect. Buddleioides, 681 sect. Hypenia, 681 sect. Umbellaria, 681 spicata, 688 suaveolens, 689 verbenifolia, 688 —I— Iburu, 382 Icacinaceae, 646 Icaco, 99 Ihchan, 308 Inchan-pichanemen, 331 Inchapemen, 240 Indigofera, 330 bongardiana, 331 hirsuta, 331 lespedezioides, 331, 331 microcarpa, 331 pascuorum, 331 suffruticosa, 331 tinctoria, 332 Iridaceae, 658 Iris martinicencis, 664 Irlbachia, 498 alata, 485 subsp. alba, 485 subsp. angustifolia, 486 subsp. arborea, 481 subsp. longistyla, 486 bonplandiana, 500 caerulescens, 534 cardonae, 500, 503 nemorosa, 500, 502 paruana, 500 phelpsiana, 500 plantaginifolia, 500, 502 poeppigii, 501 pratensis, 501, 503 pumila, 501, 502 purpurascens, 486 quelchii, 516 ramosissima, 501 recurva, 500 subcordata, 500

818

I NDEX

[Irlbachia] tatei, 501, 503 Iroucana guianensis, 445 Isoloma, 561 pycnosuzygium, 561 Ixia xiphidium, 581 Ixonanthaceae, 664 Iyoynaji, 316 —J— Jabillo, 158 Jablonskia, 161 congesta, 161, 161 Jabón, 456 Jabón de raya, 400 Jadewa-jadakwadiyo, 297 Jaillito, 66, 67 Japarandiba, 770 Jarizo, 343 Jatropha, 162 carthaginensis, 172 curcas, 162, 163 gossypifolia, 162, 163 manihot, 172 urens, 107 Jayo, 64 Jeamo, 358 Jebe, 335, 336 Jedieji-chu, 550 Jibia, 756 Jijije, 358 Jobo macho, 461 Jo-to-roco-anejóro, 172 Jocotouji, 766 Juan Zambrano, 331 Juan Zamora, 327, 331 Juasadi, 448 Jubey de Arau, 99 Jugastrum, 761 christii, 770 obtectum, 770 sifontesii, 770 Julocroton, 111 argenteus, 116 vergarenae, 123 Juncaceae, 672 Juncastrum, 672 Juncus, 672 cyperinus, 674 densiflorus, 673, 674

var. cyperinus, 674 Juvia, 756 —K— Kabaituri, 367 Kadaka, 258 Kaguaitariyek, 367 Kajadi, 280 Kajari, 403 Kamabadek, 364 Kamatata, 296 Kanaguarajodi, 404 Kanau, 732 Kanau-yek, 736 Kapakun, 368 Karamate yo, 240 Karipon, 757 Kasakideimo, 465 Kashimujuru, 171 Katajai, 177 Kawadadek, 66 Kawadi jodedi, 445 Kawudadek, 66 Kayapa, 258 Këmëhkë, 307 Kerosén, 144 Kerosén negro, 144 Kerosenillo, 619 Kippistia cognata, 599 diffusiflora, 599 Kodojijoto grande, 461 Koellikeria, 560 erinoides, 560, 560 Kohleria, 561 hirsuta, 561, 561 longipedunculata, 561 pycnosuzygium, 561 rhodomallos, 561 rubricaulis, 561 Koma koma, 550 Koyamo, 772 Krameria, 675 arida, 676 evolvuloides, 676 ixina, 676 ixine, 675, 676 lanceolata, 676 linifolia, 676 spartioides, 676 Krameriaceae, 674 Kubitzkia, 716

macrantha, 716, 717 Kuhlia, 436 mollis, 437 Kuhpëkwamën, 307 Kummeria, 650 brasiliensis, 651 Kunudi, 177 Kunuri, 177 Kurariyusu, 662 Kure-yek, 258 Kusapui, 119 —L— Lacistema, 676 aggregatum, 676, 677 coriaceum, 676 elongatum, 676 orinocense, 676 Lacistemataceae, 676 Laetia, 460 acuminata, 460 americana, 460 apetala, 460 calophylla, 461 casearioides, 461 completa, 458 coriacea, 461, 461 cupulata, 461, 462 guazumaefolia, 458 guidonia, 473 hirtella, 458 procera, 461, 462 suaveolens, 461, 462 Lagunero, 382 Lamiaceae, 678 Larentia linearis, 661 Lasiolepis aquatica, 13 Lauraceae, 700 Laurel, 719, 725 Laurel amarillo, 709, 724 Laurel blanco, 724, 725, 733, 737, 742 Laurel de aparo, 725 Laurel de babilla, 714 Laurel de rebalse, 713 Laurel de sapo, 188 Laurel espina, 736 Laurel eucalipto, 724 Laurel hediondo, 734, 736 Laurel marrón, 719

I NDEX

Laurel negro, 724 Laurel orillero, 713 Laurel paraguito, 719, 737 Laurel rastrojero, 724 Laurel rebalsero, 714 Laurel tigrito, 737 Laurel verde, 725 Laurelillo, 733 Laurelito, 724 Laurembergia, 582 tetrandra, 582, 582 Laurus caerulea, 744 canaliculata, 733 floribunda, 735 globosa, 724 hihua, 724 javitensis, 735 leucoxylon, 736 puberula, 737 reticulata, 725 triplinervis, 712 Lawadema-eje-de-deu, 401 Leche cochino, 403 Leche de cochino, 258 Lechero, 221 Lechozo, 473 Lecointea, 332 amazonica, 332, 333 Lecticula, 793 spruceana, 793 Lecythidaceae, 750 Lecythis, 774 alutacea, 775 bracteosa, 763 chaffanjonii, 765 chartacea, 775, 777 coriacea, 764 corrugata, 776 subsp. corrugata, 776, 777 subsp. rosea, 776, 778 davisii, 776 var. gracilipes, 776 fagifolia, 757 gracilipes, 765 karuaiensis, 775 longipes, 766 micrantha, 764 multiflora, 757 odora, 764 paniculata, 765 parviflora, 765

pedicellata, 766 rosea, 776 subglandulosa, 766 tenuifolia, 770 zabucajo, 776, 778 Lecythopsis, 756 Leiothamnus, 524 elisabethae, 525 Leiothrix, 18 celiae, 18, 18 echinulata, 47 flavescens, 18 var. alpina, 20 var. chimantensis, 18 var. flavescens, 19, 20 marahuacensis, 43 steyermarkii, 46 turbinata, 27 umbratilis, 18 var. brevipes, 20 Leiphaimos, 534 alba, 537 aphylla, 536 aurantiaca, 536 chionea, 536 flavescens, 537 pittieri, 537 spruceana, 537 tenella, 537 tenuiflora, 537 Lemna, 780 aequinoctialis, 780, 780 Lemnaceae, 779 Lemniscia, 640 floribunda, 640 guianensis, 640 Lengua de tigre, 576 Lengua pia poco, 492 Lentibulariaceae, 782 Leonotis, 696 nepetifolia, 696, 697 Leonurus, 696 heterophyllus, 696 japonicus, 696, 697 sibiricus, 696 Leptolobium costulatum, 367 nitens, 240 Leretia, 654 ampla, 649 cordata, 654, 655 nitida, 654 poeppigiana, 655

vellozii, 654 Licaria, 717 armeniaca, 718 brasiliensis, 718 cannella, 718, 720 chrysophylla, 719 cymbarum, 734 debilis, 719 dolichantha, 719 macrophylla, 719 martiniana, 719 oppositifolia, 719, 720 polyphylla, 719, 720 schultesii, 719 simulans, 719 tomentosa, 719 trinervis, 719 vernicosa, 719 Lightia guianensis, 229 licanioides, 229 Limnobium, 643 laevigatum, 644, 644 Limoncillo, 444 Lindackeria, 463 latifolia, 463, 464 maynensis var. laxiflora, 465 paludosa, 464, 465 Lisianthus, 483 acutangulus, 485 acutilobus, 500 albus, 485 angustifolius, 485 arboreus, 481 bifidus, 486 campanuloides, 501 cardonae, 500 chelonoides, 485 chimantensis, 483 elisabethae, 525 imthurnianus, 483 jauaensis, 501 pratensis, 501 pumilus, 501 pyriformis, 486 quelchii, 516 ramosissimus, 501 recurvus, 500 scabridulus, 501 schomburgkii, 486 sect. Brachycodon, 498 sect. Megacarpaea, 504

819

820

I NDEX

[Lisianthus] sect. Symbolanthus, 524 subcordatus, 500 tatei, 501 trifidus, 485 virgatus, 485 Lisyanthus, 483 alatus, 485 angustifolius, 485 bifidus, 486 caerulescens, 534 purpurascens, 486 sect. Chelonanthus, 483 sect. Macrocarpaea, 504 trifidus, 485 uliginosus, 486 Lonchocarpus, 333 benthamianus, 336 boliviensis, 335 crassispermus, 335 crucisrubierae, 335 densiflorus, 335 dipteroneurus, 335 discolor, 336 ernestii, 336 fendleri, 335 subsp. pubescens, 335 floribundus, 335 glabrescens, 335 guaricensis, 374 hedyosmus, 335 heptaphyllus, 336 imatacensis, 336, 338 killipii, 302 latifolius, 336 margaritensis, 336 martynii, 336 negrensis, 302 nicou, 338 nitidulus, 335 paniculatus, 336 pentaphyllus, 336 pictus, 336, 337 pterocarpus, 304 punctatus, 336 sanctae-marthae, 335 sericeus, 336, 337 stenopteris, 335 stenurus, 335 tubicalyx, 338 urucu, 338 utilis, 338 violaceus, 336

Lunania, 465 parviflora, 465, 466 Lute, 459 —M— Ma’nour’uruwe’dasu, 280 Mabea, 163 anomala, 166 arenicola, 166, 168 argutissima, 171 atroviridis, 167 eximia, 167 floribunda, 167 frutescens, 166, 169 linearifolia, 166, 169 longepedicellata, 167 longibracteata, 166 lucida, 167 maynensis, 167 montana, 167, 168 muricata, 167 nitida, 167, 168 var. albiflora, 167 occidentalis, 167 orbiculata, 166 pallida, 167 parguazae, 167 parvifolia, 171 piriri, 167, 169 pulcherrima, 167 schomburgkii, 171 sp. A, 171 sp. B, 171 subsessilis, 170, 171 taquari, 170, 171 var. angustifolia, 171 trianae, 171 verrucosa, 167 Maca, 404 Machaerium, 339 acuminatum, 341, 346 acutifolium, 341 var. pseudacutifolium, 341 affine, 342 altiscandens, 342 amazonense, 342 aristulatum, 342 bracteatum, 343 caicarense, 342 compressicaule, 344 var. manoense, 344

decorticans, 342 dubium, 342 eggersii, 345 ferox, 342, 346 ferrugineum, 344 ß parviflorum, 344 floribundum, 342 var. floribundum, 342, 347 frondosum, 343 guaremalense, 343 inundatum, 343, 348 kegelii, 343, 349 latifolium var. manoense, 344 leiophyllum, 343 var. latifolium, 343, 348 longistipitatum, 342 lunatum, 343, 350 macrophyllum, 344 var. macrophyllum, 344 madeirense, 344, 351 multifoliolatum, 344, 349 myrianthum, 344, 350 nervosum, 344 pachyphyllum, 343 pseudacutifolium, 341 quinatum, 344 var. parviflorum, 344, 352 var. quinatum, 344, 352 rectipes, 342 robiniifolium, 344, 345 rosescens, 342 sieberi, 344 sp. A, 345 steinbachianum, 343 tovarense, 345 venezuelense, 342 woodworthii, 342 Macho, 405 Macrocarpaea, 504 arborea, 516 autanae, 507 cerronis, 516 marahuacae, 505, 507 neblinae, 506, 507 piresii, 507 quelchii, 516 rugosa, 506, 507 salicifolia, 516 sp. A, 507 tepuiensis, 516

I NDEX

Macroptilium, 353 gracile, 353, 354 lathyroides, 353 var. lathyroides, 353, 354 longepedunculatum, 353 monophyllum, 354, 355 Macuca, 404 Macuca grande, 400 Macuca montañero, 400 Macure, 367 Madi, 658 Maguey, 388 Mahomo chino, 335 Maikanwakuyek, 90 Maikonaji, 319 Majagua sabanero, 444 Majagua verde, 188 Majaguillo, 764 Majagüillo, 766 Majaguillo negro, 764 Majako shodo, 401 Majei, 123 Majomo, 336 Majomo rebalsero, 335 Majomo sabanero, 288 Mamey hediono, 761 Mana-choro-yek, 750 Mangle, 91, 186 Mangle de río, 117 Manglillo, 186 Manihot, 171 anomala, 172 subsp. pubescens, 172 brachyloba, 172 carthaginensis, 172 diffusa, 172 esculenta, 172 maguireana, 173 orinocensis, 174 rusbyi, 172 surinamensis, 173 tristis, 173 subsp. saxicola, 173, 173 subsp. tristis, 174 utilissima, 172 Manteca rebalsera, 167 Manteco, 116 Manteco de agua, 117 Mañu, 330 Mapadaro, 423 Mapagarro, 100

Mappia ampla, 649 cordata, 654 nitida, 654 poeppigiana, 654 Maprounea, 174 amazonica, 175 guianensis, 174, 175 Maraca de cascabel, 291 Margaritaria, 175 nobilis, 175, 176 Maripa-yen-ya-pupen-yek, 344 Marsypianthes, 698 chamaedrys, 698, 698 hyptoides, 698 Martia, 655 Masiri, 431 Maspara, 447 Mata ratón, 328 Matamatá de altura, 756 Mato zabandu, 331 Mayna, 465 grandiflora, 440 grandifolia, 467, 467 latifolia, 463 laxiflora, 465 paludosa, 465 zuliana, 467 Megalostylis poeppigii, 136 Meibomia, 304 axillaris, 307 var. acutifolia, 307 campyloclada, 308 Melocotoncito, 599, 613 Melosa, 120 Mentolito, 661 Menudito, 335 Merecure de ardita, 637 Mesosphaerum, 682 atrorubens, 685 brevipes, 685 capitatum, 685 confertum, 686 dilatatum, 686 excelsum, 686 hirsutum, 687 laciniatum, 687 lacustre, 687 lanceolatum, 687 lantanifolium, 687 microphyllum, 687

mutabile, 687 parkeri, 688 pectinatum, 688 recurvatum, 688 suaveolens, 689 Mespilodaphne ceanothifolia, 733 myriantha, 736 oblonga, 737 Metari-yek, 368 Metsajabaja, 600 Mezilaurus, 722 anacardioides, 722 itauba, 722 lindaviana, 721, 722 maguireana, 722 sprucei, 721, 722 wurdackiana, 722 Micrandra, 176 cunuri, 177 glabra, 177, 179 minor, 177, 180 rossiana, 178, 179 siphonioides, 178, 180 var. minor, 177 sp. A, 178, 180 spruceana, 178 sprucei, 178, 180 Micropteryx poeppigiana, 324 Microstachys, 182 corniculata, 182, 183 Midi, 629 Miel de pajarito, 66, 552 Miniatuia, 521 Miriyon, 401 Moeroris stipulata, 199 Molenillo, 167 Momi, 120 Momishi, 100, 123 Momisi, 119 Momo, 177 Monopteryx, 355 angustifolia, 355, 356 inpae, 355 jahnii, 326 sp. A, 356 uaucu, 356 Moradita, 100 Moraea linearis, 661 Mucanana, 382

821

822

I NDEX

Muco, 761 Mucuna, 357 altissima, 358 hassjoo, 358 pruriens, 357 var. utilis, 357, 358 rostrata, 358, 359 urens, 358, 359 utilis, 357 Mucunana, 382 Mucutena, 382 Mudujai, 400 Muellera, 360 frutescens, 360, 361 moniliformis, 361 Muerto, 771 Mulato, 597 Muneu-yek, 736 Mureyenu-yek, 368 Musa bihai, 585 Mutare-yek, 368 Myriadenus, 431 Myrocarpus, 361 venezuelensis, 361, 361 Myrodendrum amplexicaule, 627 balsamiferum, 627 Myrospermum, 362 frutescens, 362, 363 sect. Myroxylon, 363 Myroxylon, 363 balsamum, 363, 364 benthamii, 472 —N— Nama spinosa, 646 Napimoga guyannensis, 459 Nasinaba, 735 Nautilocalyx, 561 adenosiphon, 563, 565 arenarius, 563 cataractarum, 563, 566 chimantensis, 563 cordatus, 563 fasciculatus, 563 maguirei, 563 pallidus, 563, 567 pemphidius, 563 porphyrotrichus, 564, 565

punctatus, 564 resioides, 564, 567 sp. A, 564, 566 sp. B, 564 sp. C, 564 sp. D, 564, 569 sp. E, 564 sp. F, 564 sp. G, 564 Neblinantha, 507 neblinae, 508, 509 parvifolia, 509 Nectandra, 722 aurea, 724 barcellensis, 734 bredemeyeriana, 724 cuspidata, 724 cymbarum, 734 fulva, 724 glandulifolia, 725 globosa, 724, 726 grandis, 749 guianensis, 725 hihua, 724 kaburiensis, 725 kunthiana, 749 leucantha var. attenuata, 724 var. guianensis, 724 lineata, 725 martinicensis, 725 meyeriana, 749 neesiana, 736 olivacea, 724 pearcei, 725, 726 pichurim, 725 pisi, 724 pittieri, 725 polyphylla, 719 reticulata, 725 rubra, 750 ruforamula, 725 sanguinea, 725 spumea, 734 vaga, 724 willdenoviana, 724 Nematanthus calcaratus, 550 Neohuberia, 761 Nepeta mutabilis, 687 pectinata, 688 Nescafé, 357

Neurocarpum, 280 cajanifolium, 284 javitense, 283, 284 simplicifolium, 284 Neurotheca, 509 loeselioides, 510, 510 Nicandra, 510 Nicolás, 109 Nicolsonia orinocensis, 308 Nigua, 445, 446, 459 Nigüito, 444 Niña, 629 Nirgua, 448 Níspero, 333, 404 Nissolia acuminata, 341 ferruginea, 344 leiophylla, 343 quinata, 344 robiniifolia, 344 Nosamo de casa, 361 —O— O’ mochahcho, 373 Ochthocosmus, 667 attenuatus, 668, 670 berryi, 668 floribundus, 668, 670 grandifolius, 669 longipedicellatus, 668, 670 micranthus, 667 multiflorus, 668 var. angustifolius, 669 var. canaripoensis, 669 var. multiflorus, 669, 670 roraimae, 669 var. grandifolius, 669, 671 var. roraimae, 669 Ocimum, 698 campechianum, 699, 699 micranthum, 699 Ocotea, 727 aciphylla, 732, 738 var. chimantaensis, 732 adenotrachelium, 732 amazonica, 733 atrata, 733 barcellensis, 734

I NDEX

basirecurva, 733 bofo, 733 bracteosa, 733 budowskiana, 734 canaliculata, 733 castanea, 737 caudata, 733 ceanothifolia, 733 celiana, 749 cernua, 733 costulata, 732 cowaniana, 733 crassifolia, 733 cujumary, 734 cuprea, 737 cymbarum, 734, 739 debilis, 734 depauperata, 734 directoramea, 734 dissimilis, 742 duidensis, 734 duotincta, 736 erectifolia, 734, 740 esmeraldana, 734, 739 fasciculata, 734 fendleri, 734 ferruginea, 749 finium, 732 flavantha, 735 floribunda, 735 florulenta, 736 fulvifolia, 732 glabra, 735 glandulosa, 734 glaucophylla, 735 globifera, 737 glomerata, 735 gracilis, 735, 741 guianensis, 735, 741 huberi, 735 javitensis, 735 julianii, 735, 740 lasseriana, 736 laticostata, 735 leucoxylon, 736, 738 liesneri, 736 lineata, 725 longifolia, 736 maguireana, 732 martiniana, 737 megacarpa, 736 myriantha, 736 neblinae, 736

neesiana, 736 nilssonii, 737 oblonga, 737 opifera, 736 pauciflora, 737 perrobusta, 737 perseiphylla, 745 persulcata, 736 pichurim, 725 puberula, 737 punctulata, 733 revoluta, 737 roraimae, 719, 732 roseopedunculata, 737, 741 rubra, 750 sanariapensis, 737 schomburgkiana, 737 sericiflora, 732 sp. A, 742 splendens, 737 steyermarkiana, 749 subalveolata, 737 tillettsiana, 742 tomentella, 742 tomentosa, 742 venosa, 742 vernicosa, 719 wachenheimii, 735 wurdackiana, 740, 742 yutajensis, 742 Octopleura, 509 loeselioides, 510 Ojito de pescado, 447 Ojo de burro, 358 Ojo de cari cari, 313 Ojo de grulla, 175 Ojo de zamuro, 313, 358 Okiyeu, 405 Okoi-yek, 403 Oleiocarpus, 318 Oliverio, 404 Ollita, 766 Olvidanovia, 343 Omphalea, 183 diandra, 183, 184 elaeophoroides, 183 megacarpa, 183 Oncoba latifolia, 463 maynensis var. laxiflora, 465 Onotillo, 100

823

Oparu, 323 Orchyllium alpinum, 789 humboldtii, 791 quelchii, 793 schimperi, 791 Oregano canaotero, 394 Oregano montañero, 394 Oreja de tigre, 283 Oreodaphne aciphylla, 732 adenotrachelium, 732 amazonica, 733 bracteosa, 733 caudata, 733 cernua, 733 costulata, 732 crassifolia, 733 diospyrifolia var. incompta, 737 fasciculata, 734 florulenta, 736 glomerata, 735 gracilis, 735 martiniana, 737 pauciflora, 737 schomburgkiana, 737 semecarpifolia, 712 splendens, 737 Oripopo, 766 Ormosia, 364 amazonica, 366 bolivarensis, 366, 369 chlorocalyx, 367 coarctata, 366 coccinea, 367 var. subsimplex, 368 costulata, 367, 370 crassicarpa, 368 cuneata, 367 discolor, 367 euneura, 366 grandiflora, 367 heterophylla, 368 lignivalvis, 367 macrocalyx, 367 macrophylla, 367, 369 maguireorum, 368 micrantha, 367 microsperma, 368 nobilis, 368 var. bolivarensis, 366 var. nobilis, 368

824

I NDEX

[Ormosia] pacimonensis, 423 paraensis, 368, 371 sp. A, 372 sp. B, 372 steyermarkii, 368 subsimplex, 368, 370 toledana, 367 williamsii, 368 Oroma, 764 OruKaiyet, 316 Oteratana, 298 —P— Pachyrhizus, 372 erosus, 372, 373 Paci-ra-ru-yek, 750 Paepalanthus, 20 albopulvinatus, 41 anomalus, 42 apacarensis, 20 var. humilis, 22 aristatus, 21, 35 auyantepuiensis, 21 bifidus, 21, 33 biformis, 42 brunneus, 24 bulbifer, 48 capillaceus, 37 f. proliferus, 37 var. proliferus, 37 cardonae, 21 caulescens, 43 chimantensis, 21, 31 convexus, 21 var. major, 23 var. parvicephalus, 24 var. strigosus, 23 cristatus, 21, 34 cumbricola, 21, 28 dichotomus, 21 var. dichotomus, 22, 35 var. pumilus, 22 duidae, 37 var. parvifolius, 37 eriophyllus, 47 fasciculatus, 22, 33 fasciculoides, 22, 34 fertilis, 44 filipes, 47 flavescens, 18 formosus, 22, 28

fraternus, 22 var. chimantensis, 26 var. marahuacensis, 18 var. spathulatus, 26 fulgidus, 23, 32 var. zuloagensis, 23 gleasonii, 23 gracilis, 44 griseus, 24 guyanensis, 21 heteropeplus, 43 holstii, 23 humboldtii, 44 jauensis, 37 var. caulescens, 37 jenmanii, 45 kegelianus, 45 killipii, 27 kunhardtii, 23 lamarckii 23, 24 leucocyaneus, 24 lilliputianus, 26 macrocaulon var. venamensis, 25 maguirei, 27 major, 23 meseticola, 18 minutus, 45 moldenkeanus, 22 nitens, 46 oblongus, 46 oyapockensis, 24, 33 parvicephalus, 24 var. parvicephalus, 24 var. wurdackii, 24, 33 pauperrimus, 26 pendulus, 25 perplexans, 25 var. steyermarkii, 23 var. wurdackii, 24 perpusillus, 24 phelpsiae, 24, 30 polytrichoides, 24, 34 robustus, 23 roraimae, 38 roraimensis, 25 salticola, 27 savannarum, 22 var. glabrescens, 22 saxicola var. conicus, 48 schomburgkii, 25, 31 scopulorum, 25, 29

var. auyantepuiensis, 25 sessiliflorus, 26 var. venezuelensis, 26, 35 simplex, 47 squamuliferus, 26 stegolepoides, 26, 32 var. acutalis, 26 steudelianus, 42 steyermarkii, 22 subcaulescens, 26 subtilis, 26, 34 sulcatus, 26 tafelbergensis, 24 tatei, 27 tenuis, 47 tortilis, 27, 29 turbinatus, 27 umbellatus, 48 venetifolius, 46 venustoides, 27 venustus, 27, 33 vigiensis, 24 villipes, 24 viscosus, 24 williamsii, 48 yapacanensis, 27 var. hirsutus, 27, 35 var. yapacanensis, 27 Pagaea, 498 nemorosa, 500 poeppigii, 501 pumila, 501 ramosissima, 501 recurva, 500 subcordata, 500 Paja vinera, 13 Palmito, 446 Palo amarillo, 736 Palo blanco, 262 Palo cachicamo, 440 Palo cachimbo, 771 Palo candela, 188 Palo de agua dulce, 109, 147, 212 Palo de arco, 333 Palo de boya blanco, 178 Palo de cachicamo, 467 Palo de cachimbo de rebalse, 772 Palo de camarón, 440, 473 Palo de chamanare, 400 Palo de conejo, 446

I NDEX

Palo de cuca, 224 Palo de cuchara, 224 Palo de cují, 444 Palo de escoba, 444 Palo de gallineta, 446 Palo de golondrona, 401 Palo de lagarto mato, 515 Palo de máguari, 64 Palo de mata, 109 Palo de mataguaro, 111 Palo de mato, 515 Palo de morrocoy, 400 Palo de perro de agua, 224 Palo de rabipelado, 772 Palo de rana, 515 Palo de ratón, 445 Palo de raya, 403 Palo de sardina, 183 Palo de temblador, 515 Palo gallineta, 64, 459 Palo macure, 367 Pan de acure, 144, 437, 448 Pandara-yek, 171 Panillo, 401 Para-cutaco, 404 Paradrymonia, 568 metamorphophylla, 568 sp. A, 568 sp. B, 568 sp. C, 569 sp. D, 569 sp. E, 569 Para-para, 383 Parature negro, 722 Pardillo, 447 Paretaura, 404 Pa-ronta-yek, 313 Pasita, 444 Pata de gallina, 167, 206 Pata de grulla, 167 Pata de paloma, 92, 122 Pata de paují, 167 Patrisia acuminata, 469 affinis, 469 bicolor, 469 dentata, 469 parviflora, 470 spruceana, 472 tomentosa, 470 Pausandra, 185 flagellorachis, 186 martinii, 185, 186

Pedilanthus, 186 tithymaloides, 186 subsp. tithymaloides, 186, 186 Pega pega rosada, 331 Pega-pega, 307, 308, 309, 433 Peine del diablo, 267 Peinesito, 444 Pekea, 761 Pelote, 330 Peludita, 394 Peniolilla, 323 Pentadenia, 553 microsepala, 553 Peonía, 367, 368 Peonía abrus, 239 Peonía rebalsera, 368 Peonilla, 368 Peonío, 367, 368 Peonío de cerro, 283 Pepa de zamuro, 358 Pera, 187 bicolor, 188, 189 citriodora, 188, 190 decipiens, 188, 190 distichophylla, 188, 189 ferruginea, 191 glabrata, 188, 189 nitida, 188 schomburgkiana, 188 tomentosa, 190, 191 Pericoca, 366, 461 Pericoco, 324 Peridium bicolor, 188 var. nitidum, 188 var tomentosum, 191 decipiens, 188 ferrugineum, 191 glabratum, 188 Peritassa, 604 compta, 605 granulata, 607 huanucana, 605, 606 laevigata, 605, 606 subsp. obtusa, 605 peruviana, 607 pruinosa, 607 Perro de agua banero, 188 Persea, 742 americana, 744 areolatocostae, 744, 745

825

caerulea, 744 cinnamonifolia, 712 croatii, 744 croizatii, 744 fastigiata, 744, 746 var. pilosa, 744 fluviatilis, 744, 746 grandiflora, 745, 747 jenmanii, 745, 746 lignitepala, 744 maguirei, 745, 747 perseiphylla, 745 pseudofasciculata, 745 sprucei, 712 steyermarkii, 745 Perumatá, 66 Peru-yek, 636 Phaseolus, 373 adenanthus, 427 appendiculatus, 427 campestris, 353 candidus, 427 diversifolius, 428 gracilis, 353 juruanus, 427 lasiocarpus, 428 lathyroides, 353 linearis, 428 longepedunculatus, 353 longifolius, 428 lunatus, 373, 374 monophyllus, 355 peduncularis, 428 sect. Macroptilium, 353 truxillensis, 427 vexillatus, 430 vulgaris, 374 var. vulgaris, 374 Phellocarpus amazonum, 382 Philodice, 35 hoffmannseggii, 36, 36 Philotria granatensis, 642 orinocensis, 643 Phlomis nepetaefolia, 696 Phoebe areolatacostae, 744 cinnamonifolia, 712 erectifolia, 734 filamentosa, 712 semecarpifolia, 712

826

I NDEX

[Phoebe] steyermarkiana, 735 Phyllanthus, 191 adenophyllus, 198 amarus, 195 anadenus, 200 atabapoensis, 195, 201 attenuatus, 196, 202 bolivarensis, 196 borjaensis, 196 brachycladus, 199 brasiliensis, 196 caribaeus, 196 caroliniensis, 196 subsp. caroliniensis, 196 subsp. guianensis, 196, 201 carrenoi, 196 chimantae, 196 conami, 196 congestus, 161 corcovadensis, 200 cornifolius, 197 croizatii, 200 delicatissimus, 199 diffusus, 199 dinizii, 198 duidae, 197, 204 elsiae, 197, 202 fluitans, 197 francavillanus, 198 gallinetae, 198 glaucoviridis, 198 guianensis, 196 hyssopifolioides, 197, 204 ichthyomethius, 199 jablonskianus, 197 jauaensis, 197 juglandifolius, 197 subsp. cornifolius, 197, 203 lediformis, 197 lindbergii, 197 longistylus, 197 maguirei, 198 major, 198 microphyllus, 198 minutifolius, 198, 201 minutulus, 198, 203 monocladus, 197 myrsinites, 198

subsp. francavillanus, 198, 205 subsp. myrsinites, 198 neblinae, 198 nobilis, 175 obfalcatus, 199, 205 orbiculatus, 199, 203 orinocensis, 199 paezensis, 199 paraqueensis, 199 pimichinianus, 198 piscatorum, 199, 201 pseudoconami, 199 var. pubescens, 199 pycnophyllus, 199 rupestris, 199, 204 schomburgkianus, 196 sequoiifolius, 200 stipulatus, 199, 204 strobilaceus, 200 subapicalis, 200 subsp. sequoiifolius, 200 subsp. subapicalis, 200, 205 subsp. A, 200 tenellus, 200 tepuicola, 200 urinaria, 200 vacciniifolius, 200, 205 subsp. vinillaensis, 200 ventuarii, 200 Pica Pica, 357 Pico rojo, 147 Pilón, 260, 262, 333, 423 Pilón negro, 66 Pilón rebalsero, 262 Pilón rosado, 613 Pinito blanco, 437 Piñón montañero, 119 Pionía, 323 Pionilla, 367 Pionina, 367 Pionío, 368 Piper aggregatum, 676 Piranhea, 206 longepedunculata, 206, 206 trifoliata, 207, 207 Piricoca, 461 Pirigara, 770

hexapetala, 772 Pirigaria, 770 Piscidia, 374 carthaginensis, 374, 375 guaricensis, 374 Pitopitosiji, 258 Pitumba guianensis, 446 Platymiscium, 375 duckei, 377 var. nigrum, 377 nigrum, 377 pinnatum, 376, 377 polystachyum, 377 var. fendleri, 377 trinitatis, 377 Platypodium, 377 elegans, 377 subsp. maxonianum, 377, 378 maxonianum, 377 Platysema triquetra, 276 Pleurisanthes, 655 emarginata, 655 sp. A, 656, 656 Pleurothyrium, 748 amapáense, 748 cowanianum, 749 Plukenetia, 207 abutifolia, 209 loretensis, 208 macrostyla, 209 multiglandulosa, 208 penninervia, 209 polyadenia, 209, 210 volubilis, 209 Poatoru, 109 Pocay yek, 764 Podocalyx, 212 loranthoides, 211, 212 Poecilanthe, 378 amazonica, 379, 379 hostmannii, 379 Pogonophora, 212 schomburgkiana, 212, 213 f. elliptica, 212 var. longifolia, 212 Pogopetalum, 651 acuminatum, 652 Poinsettia heterophylla, 147

I NDEX

Poiretia, 380 punctata, 380, 380 Polypompholyx schomburgkii, 792 Poma-taca, 43 Ponsigué montañero, 636 Pontopidana, 761 Poraqueiba, 656 acuminata, 658 paraensis, 656, 657 sericea, 657, 658 Poraresia anomala, 212 Pore-taurai, 404 Possira, 394 arborescens, 399 triphylla, 399 Potalia, 510 elegans, 511, 512 maguireorum, 513, 515 resinifera, 514, 515 Pretaurayese, 404 Prionostemma, 607 aspera, 607, 608 Pristimera, 608 nervosa, 609, 610 tenuiflora, 609, 610 verrucosa, 610 Prockia orinocensis, 438 Pronilla, 368 Psychotria sessiliflora, 533 Pterocarpus, 381 acapulcensis, 382, 384 amazonicus, 383 amazonum, 382 draco, 382 frutescens, 297 grandis, 383 lunatus, 343 magnicarpa, 383 officinalis, 382, 383 podocarpus, 382 rohrii, 383, 384 var. rubiginosus, 383 rupestris, 383 santalinoides, 383 ulei, 382 vernalis, 382 Punteral, 67, 447 Pyrrorhiza, 577

neblinae, 577, 578 —Q— Querosén, 144 Querotín, 144 Quinchoncho, 265 —R— Rabo de zamuro, 119 Rabo pelado, 771 Racoubea guyannensis, 459 Raddia impressifolia, 613 Raíz de zamuro, 331 Rastrojera, 121 Rastrojero, 437, 445 Rastrojero blanco, 437 Raya, 465 Realito, 298 Rebentillo, 166 Rebentillo amarillo, 166 Rechsteineria, 571 caracasana, 572 gollmeriana, 572 incarnata, 571 Redetu koru, 186 Reichertia, 519 Reventillo, 91, 96, 100, 117, 166, 171, 212 Reventillo de bocón, 99 Rhodostemonodaphne, 748 celiana, 749 grandis, 749 kunthiana, 749, 749 steyermarkiana, 749 Rhoogeton, 570 leeuwenbergianus, 571 viviparus, 570, 571 Rhopalostylis buettnerioides, 136 Rhynchosia, 385 caliensis, 388 crinita, 321 edulis, 386 kuntzei, 386 melanocarpa, 386, 387 melanosticta, 386 minima, 386, 387 phaseoloides, 386, 387

827

var. erecta, 386 reticulata, 386 var. kuntzei, 386 rufa, 321 schomburgkii, 387, 388 sect. Eriosema, 320 simplicifolia, 321 violacea, 321 Richeria, 213 grandis, 213, 214 var. divaricata, 213 var. genuina, 213 var. obovata, 213 var. racemosa, 214 laurifolia, 213 loranthoides, 212 obovata, 213 racemosa, 214 Ricinella, 89 triloba, 89 Ricinus, 214 communis, 215, 215 Rittera, 394 dodecandra, 399 pinnata, 403 Riveria, 394 Robasesina, 342 Robinia coccinea, 367 ferruginea, 288 panacoco, 402 sepium, 328 sericea, 336 tomentosa, 402 violacea, 336 Roble, 377 Roble blanco, 377 Roble colorado, 377 Rogersonanthus, 515 arboreus, 516, 517 coccineus, 516, 517 quelchii, 516, 517 tepuiensis, 516 Romero de sabana, 331 Rondonanthus, 36 acopanensis, 36 var. acopanensis, 37, 39 var. obtusifolius, 37 capillaceus, 37, 38 caulescens, 37 duidae, 37, 39 flabelliformis, 37, 40

828

I NDEX

[Rondonanthus] micropetalus, 25 roraimae, 38, 38 Rosa de muerto, 771 Roucheria, 620 angulata, 619 calophylla, 620, 621 columbianum, 620 humiriifolia, 619 laxiflora, 621, 623 parviflora, 620 punctata, 620 schomburgkii, 623 sp. A, 622, 623 Rusbyanthus, 504 Ryania, 468 acuminata, 469 angustifolia, 469, 471 bicolor, 469 candollei, 469 casiquiarensis, 470 dentata, 469 var. dentata, 469, 471 kunthii, 469 mansoana, 470 parviflora, 470 pyrifera, 470 var. subuliflora, 470 var. tomentosa, 470 sagotina, 469 speciosa, 469 var. bicolor, 469 var. minor, 469 var. stipularis, 470 var. subuliflora, 470 var. tomentosa, 470 spruceana, 470, 472 stipularis, 470, 472 tomentosa, 470 —S— Sacaraiyek, 229 Saccifolium, 519 bandeirae, 519 Sacoglottis, 633 amazonica, 634, 635, 636 ceratocarpa, 636 cuspidata, 632 cydonioides, 636 excelsa var. colombiana, 632

guianensis, 635, 636 f. dolichocarpa, 636 f. sphaerocarpa, 636 var. hispidula, 636 var. major, 636 maguirei, 635, 636 mattogrossensis, 636 oblongifolia, 637 obovata, 632 retusa, 638 sect. Eusaccoglottis, 633 sect. Humiriastrum, 631 sect. Schistostemon, 636 subg. Eusaccoglottis, 633 subg. Humiriastrum, 631 subg. Schistostemon, 636 uchi, 625 Sagotia, 215 brachysepala, 216 racemosa, 216, 217 var. brachysepala, 216 var. genuina, 217 var. ligularis, 217 var. microsepala, 217 tafelbergii, 217 Sajoko-nudu, 461 Salacia, 610 acreana, 613 amplectens, 613 anomalum, 599 attenuata, 617 belizensis, 599 brevistaminea, 597 catalinensis, 608 cognata, 599 cordata, 613, 614 cylindrocarpa, 617 diffusiflora, 599 elliptica, 613 fluminensis, 617 gigantea, 613 gleasoniana, 599 gracilis, 615 grandiflora, 613 granulata, 607 hippocrateoides, 599 impressifolia, 613, 614 juruana, 613 kanukuensis, 613 krukovii, 599 laxiflora, 617 lineolata, 599

macrantha, 613 megistophylla, 613 multiflora, 613 subsp. mucronata, 614, 615 subsp. multiflora, 615 obovata, 615 opacifolia, 615 pittieriana, 615 podostemma, 599 pruinosa, 607 sphaerocarpa, 599 Salvajito biuri, 550 Salvinia laevigata, 644 Samyda arborea, 444 hirsuta, 445 parviflora, 444 petiolaris, 461 pitumba, 446 procera, 461 silvestris, 447 spinescens, 447 suaveolens, 461 Sánamo, 404 Sandwithia, 217 guianensis, 217, 218 heterocalyx, 218 Sangre de drago, 382 Sangre de toro, 342 Sangre drago, 118 Sangredrago, 382, 383 Sangrito, 118, 120, 297, 325, 382, 383, 608 Sangrito alado, 383 Sapium, 218 albomarginatum, 221 aubletianum, 221 aucuparium, 219 biglandulosum, 221 var. aubletianum, 221 var. klotzschianum, 221 var. lanceolatum, 221 glandulosum, 219, 220 guaricense, 221 hemsleyanum, 221 hippomane, 219 jenmanii, 221 klotzschianum, 221 lanceolatum, 221 microdentatum, 221

I NDEX

moritzianum, 221 naiguatense, 221 obtusilobum, 221 paucinervium, 221 paucistamineum, 221 prunifolium, 221 yutajense, 141 Sapucaya, 774 Sardina, 501 Sarrapia, 319, 320 Sarrapia mona, 319, 320 Sarrapia real, 320 Sarusa, 485 Sasafrás, 732, 734 Sasafrás de loma, 485, 486 Savia, 221 sessiliflora, 222, 222 Sawana, 757 Sawe, 440 Schiekia, 579 congesta, 579 orinocensis, 579 subsp. orinocensis, 578, 579 subsp. savannarum, 579 subsp. silvestris, 579 Schistostemon, 636 auyantepuiensis, 637 fernandezii, 637, 639 oblongifolius, 637, 639 retusus, 638, 638 Schuebleria, 494 conferta, 496 obtusifolia, 496 tenella, 496 tenuifolia, 496 Schultesia, 519 benthamiana, 520, 521 brachyptera, 520, 521 f. heterophylla, 521 guianensis, 521 heterophylla, 521 pohliana, 522 schomburgkiana, 521 subcrenata, 522 Scutellaria, 699 purpurascens, 700, 700 Sebastiana corniculata, 183 guyanensis, 183 sect. Microstachys, 182 Senefeldera, 222

chiribiquetensis, 225 inclinata, 224 karsteniana, 224 multiflora var. acutifolia, 224 yutajensis, 141 Senefelderopsis, 224 chiribiquetensis, 225 croizatii, 225, 225 sipapoensis, 225 venamoensis, 225 Seringa, 153 Seringa de mono, 153 Serpicula, 582 Sesbania, 388 emerus, 388 exasperata, 389, 390 Setiscapella pusilla, 793 subulata, 793 Sextonia, 750 rubra, 750 Shakuru, 171 Shi-ña-teu, 400 Shodocona, 404 Sibara-koni, 399 Sicyomorpha nervosa, 610 peruviana, 607 pruinosa, 607 Siete conchas, 343 Simallo, 620 Siñate, 330 Sina-widiqui, 470 Sinningia, 571 elatior, 571, 572 incarnata, 571 Sipapoantha, 522 ostrina, 522, 523 Siphonia lutea, 153 pauciflora, 153 Sisyrinchium, 661 alatum, 662 marchio, 662 tinctorium, 662 vaginatum, 662, 662 Sobó, 262 Socoroco negro, 66 Soemmeringia, 390 semperflorens, 390, 390 Souza

829

marchio, 662 Sowi, 401 So-wo, 426 Spallanzania, 770 Sparattanthelium, 592 botucudorum var. uncigerum, 593 tupiniquinorum, 593, 593 uncigerum, 593 Spirodela, 780 intermedia, 780, 780 Spirotropis, 391 longifolia, 391, 392 sp. A, 392 Spixia distichophylla, 188 Stahelia, 530 spennerioides, 533 surinamensis, 533 Stenolobium, 266 coeruleum, 267 Stilaginella laxiflora, 158 oblonga, 159 Stizolobium, 357 altissimum, 358 aterrimun, 357 hassjo, 357 pruriens, 357 Stomoisia juncea, 791 spicata, 790 Stylosanthes, 392 angustifolia, 393 capitata, 393 gracilis, 394 guianensis, 393 var. gracilis, 393, 394 var. guianensis, 394 hamata, 394 humilis, 394 viscosa, 394 Suruauray, 99 Surúba, 177 Suru-wai-yek, 177 Swartzia, 394 acuminata, 403 var. puberula, 404 var. tridynamia, 404 alata, 401 alato-sericea, 399 angustifoliola, 399, 406

830

I NDEX

[Swartzia] arborescens, 399, 407 argentea, 399 var. argentea, 400, 408 aymardii, 400 benthamiana, 400 var. benthamiana, 400 var. yatuensis, 400 bifida, 399 brachyrachis, 400 var. glabrata, 400 buntingii, 400, 406 cardiosperma, 400 caudata, 400 conferta, 400 var. conferta, 401, 406 cowanii, 401, 408 cupavenensis, 401 cuspidata, 401 dicarpa, 401 dipetala, 401 discolor, 402 dodecandra, 399 dolichopoda, 405 floribunda, 401 fugax, 402 grandifolia, 401, 409 var. leiogyne, 402 guianensis, 401 jenmanii, 401 laevicarpa, 401 laxiflora, 402 leiogyne, 402 leptopetala, 402, 410 longifolia, 392 macrocarpa, 402 maguirei, 402 melanoxylon, 402 microcarpa, 402, 411 microstylis, 401 oedipus, 402 opacifolia, 404 pachyphylla, 402 palustris, 402 panacoco, 402 var. cardonae, 403 var. tepuiensis, 403, 412 parviflora, 399 parvifolia, 403, 411 piarensis, 403 picta, 403 var. bolivarensis, 403, 413

var. picta, 403, 413 pinnata, 403 pittieri, 403, 412 platygyne, 404 polycarpa, 402 polyphylla, 403, 407 rariflora, 399 roraimae, 404 rosea, 400 rotundata, 402 schomburgkii, 404 var. guayanensis, 404 sericea, 404 var. sericea, 404, 414 similis, 402 sp. A, 405, 410 sp. B, 405 sp. C, 405 sp. D, 405 sp. E, 405 sp. F, 405 sprucei, 404 var. sprucei, 404 var. tessellata, 405 steyermarkii, 405, 415 stipellata, 405 tepuiensis, 403 tessmannii, 405, 414 tomentosa, 402 var. cardonae, 403 triphyllata, 399 triptera, 405 urubuensis, 404 vaupesiana, 405, 415 var. glauca, 405 wurdackii, 405 Sweetia nitens, 240 Symbolanthus, 524 aracamuniensis, 542 aureus, 525, 526 camanensis, 525 elisabethae, 525, 526 subsp. occidentalis, 527 huachamacariensis, 525 quelchii, 516 rosmarinifolius, 526, 527 sessilis, 527 yaviensis, 527 yutajensis, 527 Syngonanthus, 41 acephalus, 41, 50 acopanensis, 36

akurimensis, 45 albopulvinatus, 41, 50 allenii var. parvus, 45 amapensis, 42, 51 amazonicus, 42, 53 anomalus, 42, 57 baldwinii, 42 biformis, 42, 53 bisumbellatus, 42 brevifolius, 42 bulbifer, 48 caulescens, 42, 57 var. hirsutus, 43 cowanii, 43, 56 var. involucratus, 43 var. longipedunculatus, 43 var. simplex, 43 var. tabulatus, 43 densifolius var. venezuelensis, 46 densus var. pumilus, 42 drouetii, 48 var. parviceps, 47 duidae, 43 var. duidae, 43, 50 var. humilis, 43 var. longifolius, 46 eriophyllus, 47 fenestratus, 44, 56 fertilis, 44 var. glandulosus, 44 var. orinocensis, 44 flavipes, 46 glandulosus, 43 f. epapillosus, 43 var. epapillosus, 43 gracilis, 44, 53 guianensis, 24 heteropeplus, 43 huberi, 42 humboldtii, 44 var. elongatus, 44 var. glabrescens, 44 var. glandulosus, 44 var. humboldtii, 44, 57 var. macrocephalus, 45 var. orinocensis, 44 var. parvus, 45 var. simplex, 48 jenmanii, 45, 52

I NDEX

kegelianus, 45, 53 lanatus var. alpinus, 46 longipes, 45, 56 macrocaulon, 42 macrocephalus, 45 minutus, 45 nitens, 45, 51 oblongus, 46 var. aequinoctialis, 46, 54 obtusifolius, 37 ottohuberi, 46, 55 pakaraimensis, 46 var. pakaraimensis, 46, 49 var. rivularis, 46 paraensis, 42 phelpsiae, 43 var. elongatus, 43 var. pilosus, 37 var. viridis, 37 reflexus, 47, 51 var. longifolius, 47 rivularis, 46 savannarum, 22 var. glabrescens, 22 setifolius, 47, 53 similis var. venezuelensis, 46 simplex, 47 var. appendiculifera, 42 tenuis, 47 var. bulbifer, 48, 52 var. minor, 47 var. tenuis, 48 tiricensis, 48, 50 trichophyllus, 48 tricostatus, 49 umbellatus, 48, 54 f. proliferens, 48 vareschii, 47 vaupesanus, 48 venezuelensis, 22 williamsii, 48, 55 xeranthemoides, 49, 52 f. brevifolius, 49 var. alpinus, 49 var. angustifolius, 49 var. tricostatus, 49 yapacanensis, 27 var. hirsutus, 27 Systemonodaphne, 716

macrantha, 717 —T— Tabaco de morrocoy, 485, 500 Tabaco de venado, 485, 500 Tabaco picure, 485 Tabaliz morado, 100 Tabaquilla, 485, 500 Tabaquillo, 188 Tabarí, 752, 766, 776 Tabau-yek, 724 Tachia, 527 gracilis, 528 grandifolia, 528 var. grandifolia, 530 var. orientalis, 530 schomburgkiana, 529, 530 Tachigali purpurea, 316 Tahuari rosado, 776 Tampipio, 757, 766 Tangranta-moi, 313 Tanyeechemen, 330 Tapa de tabaca, 766 Tapa tabaco, 757 Tapaculo, 445, 446 Taparo de monte, 761 Tapeinostemon, 530 borrerioides, 533 breweri, 531, 531 capitatum, 533 jauaensis, 531 longiflorum, 531 var. australe, 531 var. longiflorum, 532, 533 ptariense, 533 rugosum, 533 sessiliflorum, 533 spenneroides, 532, 533 Taralea, 416 casiquiarensis, 319 cordata, 416 var. cordata, 416, 417 var. rigida, 416 crassifolia, 416, 417 oppositifolia, 418, 419 reticulata, 418, 419 rigida, 416 steyermarkii, 419 Tarapaurai-yek, 118

831

Tarawuina, 426 Tártago, 215 Tasajo, 403 Tau-wan-yek, 709 Teichmeyeria, 770 Temare de arrendajo, 147 Temare montañero, 178 Temblador, 515 Temora-urai, 99 Tenahkwa-men, 13 Tento, 367, 368 Tepa pa meh, 64 Tephrosia, 419 adunca, 420 cathartica, 420 cinerea, 420, 421 var. littoralis, 420 littoralis, 420 senna, 420 sessiliflora, 420, 421 sinapou, 420 toxicaria, 422 Teramnus, 422 uncinatus, 423 subsp. uncinatus, 422, 423 Terciopelo, 403 Ternatea, 280 arborescens, 282 javitensis, 283 simplicifolia, 284 Tetracoryne angustifolia, 469 Tetrapollinia, 533 caerulescens, 534, 534 Tetrorchidium, 226 rubrivenium, 226, 227 Thermophila cordata, 613 rugulosa, 615 Thoa urens, 576 Tibure, 92, 100 Tiburi, 92 Timbo, 338 Tinajito, 258, 776 Tiu, 92 Tiu-yek, 120 Tocorito, 335, 336 Tocorito blanco, 336 Toddy, 357 Toluifera, 363 balsamum, 364

832

I NDEX

Tonina, 58 fluviatilis, 58, 58 Tonsella laevigata, 605 Tontelea, 615 attenuata, 617, 617 coriacea, 617 cylindrocarpa, 617 fluminensis, 617 laxiflora, 617 mauritioides, 617 ovalifolia, 617 subsp. ovalifolia, 617 parviflora, 599 Too’o, 367 Tortolito, 448 Totuma de mono, 770 Tounatea, 394 acuminata, 403 var. puberula, 403 guianensis, 401 laxiflora, 402 Tragia, 227 corniculata, 183 fendleri, 227, 228 grandifolia, 90 guayanensis, 228 japurensis, 227 Tragiopsis fruticulosa, 183 Trifolium guianense, 393 Trimezia, 662 chimantensis, 663, 664 fosteriana, 663, 664 martinicensis, 664 Trompilla, 465 Tu-basabosa, 298 Tucana guarana, 404 Tucurito, 341 Tunadi, 258 Tunari, 216 Tuneene, 258 Turä-yek, 392 Turi, 656 Turí, 64 Tussacia, 548 pulchella, 549 Tutuma de mono, 776 Tylopsacas, 572 cuneatum, 573, 573 Tylosperma, 572 cuneatum, 573

—U— Uairatombepe, 654 Uanden, 188 Uaypeu-roy-yek, 297 Udora granatensis, 642 Uña de gato, 298 Uña de gavilán, 342 Uña de murciélago, 344, 656 Uoki-yek, 403 Urapo, 343 Uraurai-yek, 629 Uruate, 444 Usibe, 167 Utricularia, 785 adpressa, 789, 803 alpina, 789, 801 alutacea, 789 amazonasana, 791 amethystina, 789, 800 f. alutacea, 789 angulosa, 791 arenicola, 792 var. kavanayena, 792 aureolimba, 789 aureomaculata, 789, 798 ayacuchae, 790 benjaminiana, 789 bolivarana, 789 breviscapa, 790, 795 calycifida, 790, 800 campbelliana, 790, 801 var. minor, 790 chiribiquitensis, 790, 803 coccinea, 791 concinna, 791 congesta, 793 f. deminutiva, 793 connellii, 792 costata, 790, 802 cucullata, 790, 794 cuspidata, 790 dubia, 792 erectiflora, 790, 803 fimbriata, 790, 796 flaccida, 792 fockeana, 791 foliosa, 790, 795 gibba, 790 guianensis, 790 guyanensis, 791, 802 heterochroma, 791, 798

hispida, 791, 800 humboldtii, 791, 802 f. cuneata, 791 hydrocarpa, 791 jamesoniana, 791, 801 juncea, 791, 803 kaieturensis, 789 leptantha, 793 lloydii, 791, 803 longeciliata, 791, 796 magnifica, 792 maguirei, 790 mirabilis, 792, 797 modesta, 789 montana, 789 myriocista, 792, 794 nana, 792, 803 naviculata, 792, 795 neottioides, 792, 797 nervosa, 792, 799 obtusa, 790 olivacea, 792, 795 oliveriana, 792, 797 orinocensis, 793 pectinata, 792 pubescens, 792, 794 punctifolia, 789 pusilla, 793, 798 quelchii, 793, 801 resupinata, 793 roraimensis, 789 rubricaulis, 791 sandwithii, 793, 796 schimperi, 791 schultesii, 793, 800 sciaphila, 792 simulans, 793, 796 spatulifolia, 789 spicata, 790 spruceana, 793 steyermarkii, 793, 797 stricta, 791 subpeltata, 792 subulata, 793, 799 tenuifolia, 790 tenuissima, 793, 802 tepuiana, 789 trichophylla, 793, 798 tricolor, 794, 799 triloba, 794, 798 turumiquirensis, 789 venezuelana, 792 versicolor, 789

I NDEX

viscosa, 794, 797 —V— Vainepá, 342 Valentonia, 655 Vantanea, 640 cupularis, 641 guianensis, 640 minor, 640, 641 parviflora, 641 Vara blanca, 437, 444, 448 Vara de muerto, 772 Vatairea, 423 guianensis, 423, 424 paraensis, 423, 424 surinamensis, 423 Vataireopsis, 426 speciosa, 425, 426 surinamensis, 426 Veconcibea latifolia, 111 Vela, 771 Vela de barco, 473 Vela de muerto, 772 Verdolaga, 500 Vesiculina cucullata, 790 Ve-yek, 403 Vicia littoralis, 420 Vigna, 426 adenantha, 427, 429 candida, 427 juruana, 427 lasiocarpa, 428 linearis, 428 var. linearis, 428, 428 longifolia, 428 luteola, 428, 429 peduncularis, 428 var. peduncularis, 428, 429 vexillata, 430 var. vexillata, 429, 430 Vohiria, 537, 539 Voyria, 534 acuminata, 535, 538 aphylla, 536, 539 araguensis, 536 aurantiaca, 536, 538 caerulea, 536, 538 chionea, 536

clavata, 536 corymbosa, 536 subsp. alba, 537 subsp. corymbosa, 537 flavescens, 537, 539 parviflora, 540 pittieri, 537, 539 rosea, 537 sect. Voyriella, 540 spruceana, 537, 539 tenella, 537, 539 tenuiflora, 537 Voyriella, 540 oxycarpha, 540 parviflora, 540, 540 —W— Wachendorfia orinocensis, 579 Wadi, 405 Wa-hú, 402 Wakauyek, 486 Wanaca-có, 366 Wanaekoko, 367 Warama-ana-chiyek, 90 Warapa, 764 Wareka marakae-yó, 291 Warim-bay-yek, 111 Wenderothia, 267 Winka, 262 Wolffiopsis, 781 Wolffia, 781 brasiliensis, 781, 781 welwitschii, 781 Wolffiella welwitschii, 781, 781 Wonka, 262 Wontai, 667 Wurdackanthus, 541 argyreus, 541, 542 Wurdackia, 36 flabelliformis, 37 Wvaraba, 404 —X— Xiphidium, 581 caeruleum, 580, 581 floribundum, 581 fockeanum, 581 giganteum, 581 Xyladenius, 436

833

gladulosus, 437 Xylosma, 472 benthamii, 472, 473 pallidifolium, 472 —Y— Ya’-ra paya yo’, 437 Yakui dan, 147 Yará, 440 Yara yara, 473 Yaraiyara, 467 Ya-raya-yek, 105 Yariyala, 440 Yawademo, 401 Yawademo de rebalse, 402 Yuca, 172 Yuca amarga, 172 Yuca dulce, 172 Yumari, 658 Yuquilla, 183 Yurí, 658 Yurí de caracol, 637 Yuwiji, 501 —Z— Zapatero, 419 Zollernia, 430 grandifolia, 430, 431 paraënsis, 431 ulei, 431 Zornia, 431 brasiliensis, 432 curvata, 432 diphylla, 432 gemella, 432 guanipensis, 433 herbacea, 433 lasiocarpa, 433 latifolia, 433 var. latifolia, 433 marajoara, 433 myriadena, 433 pubescens, 433 reticulata, 433 sericea, 433, 433 surinamensis, 433 tenuifolia var. latifolia, 433 Zuelania, 473 guidonia, 473, 474 tremula, 448