Flora of the Venezuelan Guayana, Volume 9: Rutaceae-Zygophyllaceae [9] 1930723474, 9781930723474

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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 9 RUTACEAE–ZYGOPHYLLACEAE

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

MISSOURI BOTANICAL GARDEN PRESS St. Louis

Copyright © 2005 by the Missouri Botanical Garden Press All rights reserved. ISBN 1-930723-47-4 Printed in U.S.A. Published on XXXXXXXXX 2005 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 Rutaceae (Jacquelyn A. Kallunki) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Sabiaceae (Gerardo A. Aymard C. and Nidia L. Cuello A.) . . . . . . . . . . 39 Santalaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Sapindaceae (Pedro Acevedo-Rodríguez and Hans T. Beck) . . . . . . . . . 46 Sapotaceae (Terrence D. Pennington and K. S. Edwards) . . . . . . . . . . . 89 Sarraceniaceae (Paul E. Berry, Ricarda Riina, and Julian A. Steyermark) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 Scrophulariaceae (Noel H. Holmgren and Karl A. Vincent) . . . . . . . . . 144 Simaroubaceae (W. Wayt Thomas and Edivani V. Franceschinelli) . . 168 Picramniaceae (José Rubens Pirani) . . . . . . . . . . . . . . . . . . . . . . . . . . . 177 Siparunaceae (Susanne S. Renner and Gerlinde Hausner) . . . . . . . . . 180 Smilacaceae (John F. Gaskin and Paul E. Berry) . . . . . . . . . . . . . . . . . 184 Solanaceae (William G. D’Arcy, Carmen Benítez de Rojas, and Michael H. Nee) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194 Sphenocleaceae (Bruce A. Stein) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247 Sterculiaceae (Carmen L. Cristóbal, Janice G. Saunders, and Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248 Strelitziaceae (Lennart L. Andersson) . . . . . . . . . . . . . . . . . . . . . . . . . . 281 Styracaceae (Peter W. Fritsch and Julian A. Steyermark) . . . . . . . . . . 283 Symplocaceae (Julian A. Steyermark and Paul E. Berry) . . . . . . . . . . 290 Taccaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295 Tepuianthaceae (Paul E. Berry and Zachary S. Rogers) . . . . . . . . . . . 297 Ternstroemiaceae (Paul E. Berry and Anna L. Weitzman) . . . . . . . . . 300 Tetrameristaceae (James S. Miller and Paul E. Berry) . . . . . . . . . . . . . . 309 Theaceae (Anna L. Weitzman) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311 Bonnetiaceae (Anna L. Weitzman) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313 Theophrastaceae (James S. Miller and Bertil Ståhl) . . . . . . . . . . . . . . . . 325 Thurniaceae (Ricarda Riina and Paul E. Berry) . . . . . . . . . . . . . . . . . . . . 330 Thymelaeaceae (Zachary S. Rogers, Paul E. Berry, and Julian A. Steyermark) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332 v

vi C ONTENTS

Tiliaceae (Laurence J. Dorr and Willem Meijer) . . . . . . . . . . . . . . . . . . Muntingiaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trigoniaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triuridaceae (Paul J. M. Maas and Hiltje Maas) . . . . . . . . . . . . . . . . . . . Tropaeolaceae (Paul E. Berry and Luther J. Raechal) . . . . . . . . . . . . . . . Turneraceae (María Mercedes Arbo) . . . . . . . . . . . . . . . . . . . . . . . . . . . Typhaceae (Luther J. Raechal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmaceae (James S. Miller and Paul E. Berry) . . . . . . . . . . . . . . . . . . . . Urticaceae (Amy Pool) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valerianaceae (Nereida Xena de Enrich) . . . . . . . . . . . . . . . . . . . . . . . . . . Velloziaceae (James S. Miller and Paul E. Berry) . . . . . . . . . . . . . . . . . . Verbenaceae (Gerardo A. Aymard C.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Violaceae (Ángel Fernández del Valle) . . . . . . . . . . . . . . . . . . . . . . . . . . Viscaceae (Job Kuijt) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vitaceae (Francisco Delascio Chitty and Julian A. Steyermark) . . . . . . . Vochysiaceae (Luis Marcano-Berti) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Winteraceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xyridaceae (Robert Kral) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zamiaceae (Dennis Wm. Stevenson) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zingiberaceae (Hiltje Maas and Paul J. M. Maas) . . . . . . . . . . . . . . . . Zygophyllaceae (Gerardo A. Aymard C. and Nidia L. Cuello A.) . . . . . . .

343 362 364 368 372 373 385 386 391 402 404 407 445 464 492 500 524 526 575 577 580

Addendum: Ceratophyllaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . 585 Appendix A: List of new names published in this volume . . . . . . . . . . 586 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 587

Preface and Acknowledgments This volume concludes the taxonomic treatments of all native and naturalized vascular plant families known to occur in the Venezuelan Guayana. It includes 48 families of flowering plants, from Rutaceae through Zygophyllaceae, and including one family omitted from the alphabetical sequence, as well as several families that have been segregated since the inception of the flora. The Flora of the Venezuelan Guayana project has been strongly supported since its inception by the Missouri Botanical Garden and especially by its director, Peter H. Raven. The Herbario Nacional de Venezuela, now part of the Fundación Instituto Botánico de Venezuela, has supported this project in many ways; it was there that Julian Steyermark conceived the idea of the flora and did most of his background research. The Julian A. Steyermark Fund, established by the late Dr. Steyermark at the Missouri Botanical Garden, provided major funding for the flora. The Wisconsin State Herbarium and the Botany Department at the University of Wisconsin–Madison has also supported this project in its later stages. This volume is based upon work supported by the National Science Foundation under Grants Nos. BSR-8717303, BSR-9045532, and BSR-9201044. The Foundation provides awards for research in the sciences and engineering. The awardee is wholly responsible for the conduct of such research and preparation of the results for publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the authors and do not necessarily reflect the views of the National Science Foundation. Besides the illustrations by Bruno Manara, Christiane Anderson, editor of the series Systematic Botany Monographs, graciously allowed us to use the figure from that series of Phoradendron berryi, which was drawn by Job Kuijt and is Fig. 398 in this volume. Figures 159–161 and the detail of Fig. 155 were drawn by Sarah Gaskin. Numerous institutions and individuals in Venezuela have contributed significantly to different aspects of the flora. These include the Corporación Venezolana de Guayana and its affiliate Electrificación del Caroní, C.A.; the Ministerio del Ambiente y de los Recursos Naturales Renovables; the Instituto Nacional de Parques and the Dirección General de Parques Nacionales; the Servicio Autónomo 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 Otto Huber, Francisco Delascio, Francisco Guánchez, vii

viii P REFACE AND A CKNOWLEDGMENTS

Gilberto Morillo, Zoraida Luces de Febres, and Aníbal Castillo; the Universidad Central de Venezuela, through the Herbario Ovalles at the Facultad de Farmacia and its director emeritus, Stephen Tillett, and the herbarium of the Facultad de Agronomía in Maracay; 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, Lisa Campbell, Germán Carnevali, Kandis Elliot, Shirley Flavin, Ronald Liesner, Luther Raechal, Ivón Ramírez, Erika Rohrbach, and George Yatskievych. A special thanks to Ruth King and George Thornburgh, whose help as volunteers at Missouri Botanical Garden, 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 begin 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 treats the ferns and fern allies, or pteridophytes. It also includes an introduction to the pteridophytes and a key to the pteridophyte families, followed by the corresponding family treatments in alphabetical order. The volume then continues with the first 11 seed plant families, again in alphabetical order, beginning with Acanthaceae and ending with Araceae. Volume 3 includes 20 families from Araliaceae through Cactaceae; Volume 4 includes 35 families from Caesalpiniaceae through Ericaceae; Volume 5 includes 29 families from Eriocaulaceae through Lentibulariaceae; Volume 6 includes 28 families from Liliaceae to Myrsinaceae; Volume 7 includes 18 families from Myrtaceae to Plumbaginaceae; and Volume 8 includes 17 families from Poaceae to Rubiaceae. Family names of flowering plants in the Flora of the Venezuelan Guayana mostly follow those used by A. Cronquist in An Integrated System of Classification of Flowering Plants (Columbia University Press, New York. 1981). Also included are certain families published or recognized after Cronquist’s book, such as Euphroniaceae, Bonnetiaceae, Muntingiaceae, and Picramniaceae. The 48 families that appear in Volume 9 and their numbers of native or naturalized genera and species in the Venezuelan Guayana are given in the following table. ix

x INTRODUCTION

FAMILIES INCLUDED IN THIS VOLUME Genera Species Rutaceae Sabiaceae Santalaceae Sapindaceae Sapotaceae Sarraceniaceae Scrophulariaceae Simaroubaceae Picramniaceae Siparunaceae Smilacaceae Solanaceae Sphenocleaceae Sterculiaceae Strelitziaceae Styracaceae Symplocaceae Taccaceae Tepuianthaceae Ternstroemiaceae Tetrameristaceae Theaceae Bonnetiaceae Theophrastaceae

20 2 1 16 9 1 18 4 1 1 1 16 1 10 1 1 1 1 1 2 1 1 2 2

60 7 1 83 85 10 42 9 7 8 9 79 1 48 1 11 11 1 4 21 1 1 28 3

Genera Species Thurniaceae Thymelaeaceae Tiliaceae Muntingiaceae Trigoniaceae Triuridaceae Tropaeolaceae Turneraceae Typhaceae Ulmaceae Urticaceae Valerianaceae Velloziaceae Verbenaceae Violaceae Viscaceae Vitaceae Vochysiaceae Winteraceae Xyridaceae Zamiaceae Zingiberaceae Zygophyllaceae Ceratophyllaceae

1 4 8 1 1 3 1 2 1 3 6 1 2 15 8 2 1 4 1 5 1 1 3 1

2 10 30 1 7 5 1 21 1 4 13 1 2 74 29 53 10 65 1 95 2 8 4 1

190

971

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 only exceptions to this rule are segregate families such as Muntingiaceae and Picramniaceae that were recognized after their corresponding volumes were published, and Ceratophyllaceae, which was only recognized for the flora area after its volume was published. The gymnosperms, monocotyledons, and dicotyledons are not treated separately within the seed plant category. The authors for each family treatment are cited be-

INTRODUCTION

xi

low the family heading; in multi-authored families, if different parts were contributed by different authors, the respective authors are cited separately under the parts they were responsible for (such as a particular genus or the key to the genera). Each family treatment contains a complete description of the family and the genera that are present in the flora; these descriptions are worldwide in scope. Within each family, genera are arranged in alphabetical order. At the end of each family or genus description, a separate paragraph lists the overall distribution of the family or genus. This is followed by the author’s estimate of the number of subordinate taxa worldwide, as well as the number of taxa present in the Venezuelan Guayana. For each genus, the estimated number of species in all of Venezuela is also given. All keys used in the flora are dichotomous and bracketed. In each couplet, the number in parenthesis refers to the couplet that led to it. The keys to the genera are designed primarily to cover the species that occur in the Venezuelan Guayana and not necessarily species found elsewhere. Slightly more than half the species in the flora are illustrated by black-andwhite line drawings. Volume 9 includes 503 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 “◆” followed by the figure number appears at the end of the paragraph. A separate paragraph is sometimes used to provide taxonomic discussion and/or notes on the uses of the plant in the flora area. Any varieties or subspecies present in the flora area appear below their corresponding species entry, and they contain a similar level of information as regular species entries. A key is included if more than one subspecies or variety is present. Forms found in the flora area do not have separate entries, but they are discussed under the next higher rank.

xii

INTRODUCTION

Throughout the flora, author abbreviations follow Authors of Plant Names (R. K. Brummitt and C. E. Powell, editors. Royal Botanic Gardens, Kew. 1992). The only exception is the use of the traditional abbreviation “H.B.K.” for the authors of the publication Nova Genera et Species Plantarum (A. von Humboldt, A. Bonpland, and K. S. Kunth. Librairie Grecque-Latine-Allemande, Paris. 1815–1825). This series of seven volumes was published in two versions (quarto and folio), each with different page numbers. Page numbers cited in this flora refer to the quarto version, which is more widely available than the folio version, except for volume four, which is the only volume in which the folio edition was distributed before the quarto. Book abbreviations follow Taxonomic Literature: A Selective Guide to Botanical Publications and Collections with Dates, Commentaries and Types, 2nd edition (F. A. Stafleu and R. S. Cowan. Bohn, Scheltema & Holkema, Utrecht, Netherlands. 1976– 1988), and journal abbreviations follow Botanico-Periodicum-Huntianum (G. H. M. Lawrence, A. F. G. Buchheim, G. S. Daniels, and H. Dolezal, editors. Hunt Botanical Library, Pittsburgh, Pennsylvania. 1968) or B-P-H/S: Botanico-Periodicum/ 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), and the website developed from this and previous editions of this publication (http://sciweb.nybg.org/ science2/IndexHerbariorum.asp). 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 Pedro Acevedo-Rodríguez Botany, MRC-166 United States National Herbarium Smithsonian Institution PO Box 37012 Washington, DC 20013 Lennart L. Andersson (deceased) Department of Systematic Botany University of Göteborg P.O. Box 461 SE 405 30 Göteborg SWEDEN María Mercedes Arbo Instituto de Botánica del Nordeste Casilla de Correo 209 3400 Corrientes ARGENTINA Gerardo A. Aymard C. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA Hans T. Beck Department of Biological Sciences Northern Illinois University DeKalb, Illinois 60115 Carmen Benítez de Rojas Universidad Central de Venezuela Facultad de Agronomía Instituto de Botánica Agrícola Herbario “Victor Manuel Badillo” Apartado 4579 Maracay, Aragua 2101 VENEZUELA

Paul E. Berry Department of Ecology and Evolutionary Biology The University of Michigan Natural Science Building (Kraus) 830 North University Ann Arbor, Michigan 48109 Carmen L. Cristóbal Instituto de Botánica del Nordeste Casilla de Correo 209 3400 Corrientes ARGENTINA Nidia L. Cuello A. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA William G. D’Arcy (deceased) Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Francisco Delascio Chitty Jardín Botánico del Orinoco Calle Bolívar Ciudad Bolívar, Edo. Bolívar VENEZUELA Laurence J. Dorr Department of Botany, MRC-166 National Museum of Natural History Smithsonian Institution P.O. Box 37012 Washington, DC 20013

xiii

xiv

CONTRIBUTORS

K. S. Edwards Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB ENGLAND, U.K. Ángel Fernández del Valle IVIC, Centro de Biofísica y Bioquímica Apartado 21827 Caracas 1020 A VENEZUELA Edivani V. Franceschinelli Universidade Federal de Minas Gerais, ICB Departamento de Botânica Caixa Postal 486 30161-970 Belo Horizonte, MG BRAZIL Peter W. Fritsch Department of Botany California Academy of Science 875 Howard St. San Francisco, California 94103 John F. Gaskin USDA-ARS-NPARL PO Box 463 Sidney, Montana 59270 Gerlinde Hausner Werner-Hilpert Strasse 67 D-65197 Wiesbaden GERMANY Noel H. Holmgren New York Botanical Garden Bronx, New York 10458 Bruce K. Holst Marie Selby Botanical Gardens 811 South Palm Ave. Sarasota, Florida 34236 Jacquelyn A. Kallunki New York Botanical Garden Bronx, New York 10458

Robert Kral Botanical Research Institute of Texas 5009 Pecan St. Fort Worth, Texas 76102 Job Kuijt Department of Biology University of Victoria Victoria, British Columbia V8W 3N5 CANADA 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 Luis Marcano-Berti Urb. Sta. Ana, Calle Ejido Edif. Chocanta, Apt. D Mérida 5101 VENEZUELA Willem Meijer (deceased) c/o School of Biological Sciences 101 Morgan Building University of Kentucky Lexington, Kentucky 40506 James S. Miller Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

CONTRIBUTORS

Michael H. Nee The New York Botanical Garden Bronx, New York 10458 Terrence D. Pennington Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB ENGLAND, U.K. José Rubens Pirani Departamento de Botânica, IB/USP Caixa Postal 11461 05422-970 - São Paulo, SP BRAZIL Amy Pool Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Luther J. Raechal c/o Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Susanne S. Renner Botanische Staatssammlung München Menzinger Strasse 67 D-80638 München GERMANY Ricarda Riina Botany Department University of Wisconsin-Madison 430 Lincoln Drive Madison, Wisconsin 53706 Zachary S. Rogers Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Janice G. Saunders Instituto de Botánica Darwinion Casilla de Correo 22 B1642HYD San Isidro Provincia de Buenos Aires ARGENTINA

Bertil Ståhl Botanical Museum Carl Skottsbergs Gata 22 S-413 19 Göteborg SWEDEN Bruce A. Stein NatureServe 1101 Wilson Boulevard, 15th Floor Arlington, Virginia 22209 Dennis Wm. Stevenson The New York Botanical Garden Bronx, New York 10458 Julian A. Steyermark (deceased) c/o Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 W. Wayt Thomas The New York Botanical Garden Bronx, New York 10458 Karl A. Vincent c/o New York Botanical Garden Bronx, New York 10458 Anna L. Weitzman Department of Botany, NHB-166 National Museum of Natural History Smithsonian Institution Washington, DC 20560 Nereida Xena de Enrich Centro de Biología Tropical Universidad Central de Venezuela Apartado 20513 Caracas 1040 VENEZUELA Kay Yatskievych Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

xv

RUTACEAE by Jacquelyn A. Kallunki Aromatic, glandular-punctate trees or shrubs, rarely herbs, sometimes armed. Leaves alternate or opposite, rarely whorled, usually pinnately or palmately compound or 1- or 3-foliolate, rarely pinnately dissected or simple, usually petiolate, exstipulate; glandular punctae usually present but not always pellucid. Inflorescence terminal, lateral, or axillary, sometimes cauline, rarely borne on the lower surface of a leaf, thyrsoid, cymose, racemose, spicate, or coiled. Flowers actinomorphic to somewhat zygomorphic, bisexual or unisexual, hypogynous or rarely perigynous. Calyx of (2–)4 or 5 free or basally connate sepals, usually quincuncial, sometimes imbricate or valvate; corolla of (2–)4 or 5 petals (rarely absent), these alternate with sepals, free or sometimes connate to various degrees, usually imbricate, sometimes valvate, contorted, or quincuncial. Androecium usually of 5 or 10 stamens, sometimes (much) more numerous, sometimes of 2 or 3 fertile stamens and 2–5 staminodes; filaments free or less often connate below and then sometimes adnate to the corolla; anthers introrsely dehiscent by longitudinal slits, sometimes apically apiculate or glandular or basally appendaged. Disk intrastaminal, annular, cupular, or cylindrical, rarely unilateral, sometimes modified into a gynophore, or rarely obsolete. Ovary of (2–)4 or 5(–many) carpels ± connate into a plurilocular ovary with distinct styles or a single common style, or free and connate only in their styles, or rarely reduced to 1 carpel; placentae in a compound ovary usually axile, rarely intruded-parietal when the partitions are not joined in the center, or ventral in an apocarpous ovary; ovules 1–several per locule, often superposed, seldom biseriate. Fruit a (4)5-locular capsule or 1–5 mericarps with ventral, loculicidal dehiscence assisted by a ligneous endocarp of which the two halves separate from each other and forcibly eject the seed, a few- to severalseeded berry or hesperidium, a drupe, or rarely a samara. Seeds 1–several per carpel; endosperm present or absent; embryo straight or curved with rather large, flat or crumpled cotyledons. Worldwide, mostly tropics and subtropics; ca. 157 genera and ca. 1600 species, 20 genera and 60 species in the flora area. Genera cultivated in Venezuela include Citrus, Ruta, Casimiroa, and Triphasia. 1

2

R UTACEAE

Key to the Genera of Rutaceae 1.

Leaves opposite (some individuals may have some alternate or subopposite leaves) ...................................................................................... 2 1. Leaves alternate or appearing basal ........................................................ 5 2(1). Inflorescences cauline; seeds smooth ........................................... 14. Raputia 2. Inflorescences axillary or terminal; seeds tuberculate ........................... 3 3(2). Erect herb, sometimes woody at base; outermost sepal much larger than other sepals, mimicking a bract, and concealing the 6 mm corolla; anthers without basal appendages; disk elongated on one side ...... 6. Ertela 3. Shrub or small tree; sepals sometimes unequal but none of them large enough to conceal the corolla; anthers with basal appendages; disk equilateral .............................................................................................. 4 4(3). Calyx cupular, equally 5-lobed, in fruit glabrous; corolla lobes induplicate-valvate ........................................................ 5. Decagonocarpus 4. Calyx of free (or very shortly connate), unequal sepals, in fruit glabrous or often obscured by dense pubescence; corolla lobes imbricate ............................................................................................. 16. Raveniopsis 5(1). Leaves pinnately compound ...................................................................... 6 5. Leaves simple, 1- or 3-foliolate, or palmately 5-foliolate ........................ 9 6(5). Petals > 1 cm long; style many times longer than ovary; fruit a berry .................................................................................................. 11. Murraya 6. Petals < 1 cm long; style shorter than ovary; fruit samaroid or ventrally dehiscent with separating, bony endocarp .......................................... 7 7(6). Inflorescence an elongate raceme; pedicels 2–2.3 cm long; leaves with 1 or 2 pairs of leaflets, often (1)3-foliolate at base of inflorescence ............................................................................................... 13. Pilocarpus 7. Inflorescence a panicle, less often a short raceme; pedicels much shorter than 2 cm; leaves usually with > 2 pairs of leaflets and not (1)3foliolate at base of inflorescence ........................................................... 8 8(7). Plants polygamous, monocarpic, not spiny; leaves congested at apex of branches; carpels connate, each with 1 apical ovule; fruit a 2(3)winged samara ....................................................................... 18. Spathelia 8. Plants dioecious, not monocarpic, often spiny; leaves generally not congested at apex of branches; carpels free or connate, each with 2 axile or ventral ovules; fruit dehiscent, not winged ................ 20. Zanthoxylum 9(5). Decumbent, pubescent herbs; leaves 3-foliolate, often appearing basal ................................................................................................. 3. Apocaulon 9. Shrubs or trees; leaves various, borne on branches .............................. 10 10(9). Fruit a 5-locular drupe; leaves simple, oblanceolate, 9–96 cm long, erect or ascending, often clustered at branch apex; inflorescence terminal, often robust, many times dichotomous; petals pink or red, barbate within .......................................................................................... 10. Hortia 10. Fruit dehiscent; leaves simple or variously (but not pinnately) compound; plant without the above combination of characters ....... 11 11(10). Specimen with flowers ............................................................................ 12 11. Specimen with fruits ............................................................................... 23 12(11). Calyx 16–46 mm long, red; sepals variously connate, valvate; flowers often borne on short shoots of perennating inflorescence ................. ........................................................................................... 7. Erythrochiton

R UTACEAE 3

12.

13(12). 13.

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

15.

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

17.

18(13). 18. 19(18). 19. 20(19). 20. 21(20). 21. 22(19).

22.

23(11). 23. 24(23). 24. 25(24).

Calyx generally < 8 mm long, green; sepals free or connate at base, with lobes valvate or overlapping in bud; flowers not in perennating inflorescence ........................................................................................ 13 All 5 stamens fertile; petals free (sometimes coherent at very base initially) ............................................................................................... 14 Two stamens fertile; petals coherent at base (forming a tube at least at anthesis, sometimes ultimately falling separately), connate at base (falling as a unit), or free .................................................................... 18 Petals ≥ 1.2 cm long ................................................................................. 15 Petals ≤ 1 cm long .................................................................................... 16 Leaves simple; petals ca. 2 cm long, white-villous within above the base, coherent at base, valvate; disk reduced to a band of tissue adherent to a short, broad gynophore ..................................................... 1. Adiscanthus Leaves 3-foliolate; petals 1.2–6 cm long, closely appressed-pubescent on both surfaces, free, quincuncial or infrequently imbricate; disk cupular or cylindric, embracing the narrowed gynophore ............ 19. Spiranthera Inflorescence unbranched, an elongate raceme ....................... 13. Pilocarpus Inflorescence branched, a thyrse or dichasium ..................................... 17 Leaves 1-foliolate (apex of petiole swollen), usually > 4 cm long, usually not congested; petiole ≥ 4 mm long; inflorescence a thyrse usually with obvious internodes and pedicels ........................................... 8. Esenbeckia Leaves simple (apex of petiole not swollen), ≤ 4 cm long, congested along stem; petiole virtually absent; inflorescence a dichasium, usually appearing subcapitate but sometimes with primary branches to 2.5 cm long ............................................................................... 17. Rutaneblina Leaves palmately 5-foliolate; inflorescence a terminal, long-pedunculate, bifurcate dichasium with secund branches .............. 12. Neoraputia Leaves simple or 1- or 3-foliolate; inflorescence various, if a bifurcate dichasium then cauline and on a peduncle ca. 1 cm long ................. 19 Anthers with basal appendages; petals connate into a tube ................ 20 Anthers without basal appendages; petals free or coherent into a tube .............................................................................................................. 22 Calyx of free, ± unequal sepals .............................................. 16. Raveniopsis Calyx cupular, ± equally 5-lobed or -toothed ......................................... 21 Calyx, pedicel, and vegetative bud with echinoid or shaggy trichomes; corolla tube 10–12 mm long, 0.7–1 times as long as lobes ... 2. Angostura Calyx, pedicel, and vegetative bud with simple trichomes; corolla tube 11–37 mm long, 1.4–3.7 times as long as lobes ......................... 9. Galipea Petals free (or in C. longifolius free at base but adnate at middle to apex of filaments and with lobes only ca. 2 times as long as wide); inflorescence a thyrse or bifurcate and cauline ............... 4. Conchocarpus Petals coherent into a tube 6–10 mm long, lobes (when dry) to 1 mm wide, several times longer than wide, usually recurved at full anthesis; inflorescence a corymboid thyrse or a pseudoraceme .......... 15. Rauia Fruit a (4)5-locular capsule ..................................................................... 24 Fruit of up to 5, ± free mericarps ............................................................ 26 Capsule echinate or muricate .................................................... 8. Esenbeckia Capsule sometimes with raised veins but not echinate or muricate .... 25 Leaves 3-foliolate; calyx not explanate; capsule glabrous; length of peduncle (to lowest pedicel scar) 6–8 cm ................................... 9. Galipea

4

R UTACEAE

25.

26(23). 26. 27(26). 27. 28(27).

28. 29(27). 29. 30(26). 30. 31(30).

31. 32(31). 32. 33(30). 33. 34(33). 34. 35(34). 35. 36(35). 36. 37(36). 37. 38(37).

38.

Leaves 5-foliolate; calyx enlarged and explanate (spread out flat) in fruit; capsule densely tomentulose; length of peduncle (to bifurcation) 21–36 cm .............................................................................. 12. Neoraputia Leaves simple (apex of petiole not swollen) ........................................... 27 Leaves 1-foliolate (apex of petiole swollen) or 3-foliolate ..................... 30 Leaves ≤ 11 cm long ................................................................................. 28 Leaves 30–76 cm long .............................................................................. 29 Leaves obovate, broadly rounded, truncate, or emarginate and often apiculate at apex, glabrous, subsessile, densely reticulate, leathery when dry; seeds smooth ..................................................... 17. Rutaneblina Leaves without the above combination of features; seeds tuberculate ............................................................................................. 16. Raveniopsis Infructescence a terminal dichasium much shorter than peduncle .............................................................................................. 1. Adiscanthus Infructescence a lateral thyrse, ± equal to peduncle ................................. .................................................................. 4. Conchocarpus (C. longifolius) Leaves 1-foliolate ..................................................................................... 31 Leaves 3-foliolate (sometimes 1-foliolate at base of inflorescence) ...... 33 Calyx 16–46 mm, longer than the fruit, initially persistent but ultimately deciduous; fruits often borne on short shoots of perennating, lateral inflorescence ................................... 7. Erythrochiton Calyx < 5 mm, shorter than the fruit, usually persistent; infructescence terminal, not perennating ................................................................... 32 Leaves oblanceolate, often grayish green on upper surface when dry; petiole usually ferruginous .................... 4. Conchocarpus (C. ucayalinus) Leaves elliptic, green on upper surface when dry; petiole not ferruginous .......................................................................... 15. Rauia (R. subtruncata) Infructescence a raceme to 31 cm long with pedicels 2–2.3 cm long ...................................................................... 13. Pilocarpus (P. racemosus) Infructescence a thyrse, dichasium, or bifurcate, if an apparent raceme then pedicels < 1 cm ............................................................................ 34 Infructescence cauline ................................. 4. Conchocarpus (C. ramiflorus) Infructescence terminal or lateral .......................................................... 35 Central leaflet 1.5–7.5 cm long .............................................. 16. Raveniopsis Central leaflet 9–31 cm long ................................................................... 36 Apical buds, calyx, and mericarps with yellow-green pubescence of echinoid or stellate trichomes ................................................ 2. Angostura Above type of pubescence absent ............................................................ 37 Surface of fruiting carpel clearly evident, bearing sparse, short, appressed trichomes (if any) ............................................. 19. Spiranthera Surface of fruiting carpel obscured by dense pubescence (unless surface has been abraded) ............................................................................... 38 Pubescence of fruiting carpel a mixture of mealy trichomes and longer, straight, pointed ones, usually brown or rusty-colored; inflorescence a narrow thyrse; leaf blade with inframarginal vein ........................... ............................................................. 4. Conchocarpus (C. heterophyllus) Pubescence of fruiting carpel matted, usually straw-colored; inflorescence a pseudoraceme; leaf blade lacking an inframarginal vein ........................................................................... 15. Rauia (R. spicata)

Adiscanthus

5

1. ADISCANTHUS Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 186. 1922. Shrubs or trees to 5 m tall. Leaves alternate, simple; petioles swollen at base; blades 30–76 cm long, narrowly oblanceolate, long-cuneate at base, entire, papery when dry. Inflorescence terminal, an erect, long-pedunculate dichasium; epidermis of peduncle usually finely transversely cracked. Flowers actinomorphic. Calyx shallowly cupular, slightly 5-lobed, explanate, undulate, and persistent in fruit, the lobes obtuse; petals 5, free (coherent at base initially), valvate, ca. 2 cm long,

Fig. 1. Adiscanthus fusciflorus

6

R UTACEAE

lanceolate, maroon without, white within, coiled at anthesis, glabrous without, white-villous within above the base. Androecium of 5 fertile stamens; filaments free; anthers lorate, bifid at base, dorsifixed, coiled after anthesis. Disk obsolete, reduced to a band of tissue adherent to a short, broad gynophore between bases of ovary and filaments. Ovary 5-lobed, carpels connate laterally at base and free above except at base of style; ovules 2 per carpel; style 1, columnar; stigma truncate, slightly 5-lobed. Fruit of 1 or 2(–5) free mericarps, each dehiscent ventrally. Seeds 1 per carpel, tear-shaped, tapered to apex, smooth, brown-mottled, dull, glabrous. Amazonian Venezuela, Peru, and Brazil; 1 species. Adiscanthus fusciflorus Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 187. 1922. Treelet ca. 3 m tall, unbranched or with only 2 or 3 branches; dichasia 1 or 2, longpedunculate; flowers maroon with tufts of

white, villous pubescence on inner surface of petals. Evergreen lowland forests, 100–200 m; Amazonas (base of Sierra de la Neblina). Amazonian Peru, northern Brazil. ◆Fig. 1.

2. ANGOSTURA Roem. & Schult., Syst. Veg. 4: 188. 1819. Bonplandia Willd., Mém. Acad. Roy. Sci. Hist. (Berlin) 1802: 26. 1804, non Cav. 1800. Cusparia Humb., Essai Géogr. Pl. 58. 1807, nom. provis. Shrubs or trees to 15 m tall. Leaves alternate, usually 3-foliolate (sometimes 1-foliolate or palmately 4–7-foliolate), petiolate; blades entire, papery when dry. Inflorescence terminal, short- to long-pedunculate, a variously branched thyrse. Flowers sub-actinomorphic. Calyx cupular, ± 5-lobed, persistent in fruit, the lobes quincuncial in bud; corolla white, 5-lobed, densely pubescent with echinoid or stellate trichomes, the tube ± equal to or shorter than lobes, straight, the lobes imbricate, spreading or recurved. Androecium of 2 fertile stamens and 5 linear staminodes; filaments connate into a tube adnate to the corolla for most of its length; anthers oblong, basifixed, free, appendaged at base. Disk cupular, shorter than or equal to ovary, glabrous. Ovary of 5 free carpels; ovules 2 per carpel; style 1; stigma subcapitate. Fruit of 1–5 free mericarps, each ventrally dehiscent. Seeds 1 per carpel, ellipsoid-ovoid, smooth or slightly rugulose, glabrous. Nicaragua, Costa Rica, Venezuela, Ecuador, Peru, Brazil, Bolivia; 8 species, 1 in Venezuela. Angostura trifoliata (Willd.) T.S. Elias, Taxon 19: 575. 1970. —Bonplandia trifoliata Willd., Mém. Acad. Roy. Sci. Hist. (Berlin) 1802: 27. 1804. —Cusparia febrifuga Humb., Essai Géog. Pl. 58. 1807, nom. provis. —Angostura cuspare Roem. & Schult., Syst. Veg. 4: 188. 1819, nom. illeg. —Galipea cusparia A. St.Hil. ex DC., Prodr. 1: 731. 1824, nom. illeg. —Bonplandia angostura Spreng., Syst. Veg. 1: 780. 1825. —Galipea officinalis J. Hancock, Trans. Roy. Med.Bot. Soc. London 1: 25, pl. 11. 1829, nom. illeg. —Sciuris officinalis (J. Hancock) Oken, Allg. Naturgesch. 3(2): 1268. 1841. —Cusparia trifoliata (Willd.)

Engl. in Mart., Fl. Bras. 12(2): 113. 1874. —Galipea febrifuga (Humb.) Baill., Traité Bot. Méd. Phan. 2: 851. 1884. —Bonplandia angostura Rich. ex B.D. Jacks. in Index Kew. 1: 322. 1893. —Cusparia officinalis (J. Hancock) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 167. 1896. —Cusparia angostura (Rich. ex B.D. Jacks.) A. Lyons, Pl. Nam., ed. 2, 153. 1907. —Quina, Quina amarilla. Tree 2–15 m tall; pubescence of echinoid or stellate trichomes; flowers white. Semideciduous forests, common on ferruginous quartzite outcrops, lower montane forests, 50–500 m; Delta Amacuro (Serranía de

Angostura 7

Imataca), Bolívar (Ciudad Bolívar, islands in Lago Guri, Río Caroní, Río Caura, Río Cuyuní, Serranía de Imataca, Upata), Amazonas (upper Río Orinoco). Anzoátegui, Sucre. ◆Fig. 2. The bark of Angostura trifoliata is used as a remedy for malaria, for intermittent fever, and as a blood purifier. It was one of the original ingredients of the famous Angostura bitters now used to flavor cocktails.

Fig. 2. Angostura trifoliata

8

R UTACEAE

3. APOCAULON R.S. Cowan, Mem. New York Bot. Gard. 8: 119. 1953. Decumbent, pubescent, succulent herbs. Leaves alternate, congested and often appearing basal, 3-foliolate, long-petiolate; blades entire, the lower surface pale. Inflorescence axillary, a long-pedunculate dichasium with a central flower and 2 racemose branches. Flowers zygomorphic. Sepals 5, very shortly coherent, of unknown aestivation, very unequal, linear, persistent in fruit; corolla to 7 mm long, white, the tube curved, the (4)5 lobes imbricate, the 2 lobes opposite the lobe flanked by the 2 stamens joined for a slightly longer distance and forming a bilobed lip. Androecium of 2 fertile stamens and 3 awl-shaped staminodes; filaments distinct, adherent to corolla tube; anthers elliptic, basifixed, coherent by their adaxial surfaces, appendaged at base. Disk cupular, enclosing the ovary, minutely puberulent at margin. Ovary of (4)5 free carpels joined at the apex in the style; ovules 2 per carpel; style 1; stigma capitate, 5-lobed. Fruit of 1–5 free mericarps, each ventrally dehiscent with beaked apex. Seeds 1 per carpel, reniform, ± tuberculate, glabrous. Endemic to the Guayana Shield in southern Venezuela; 1 species. Apocaulon carnosum R.S. Cowan, Mem. New York Bot. Gard. 8: 119. 1953. Terrestrial or epiphytic herb. Locally abundant on mossy hummocks, humuscovered boulders, rotten logs, and in trees in moist montane forests; 600–1300 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 3.

Fig. 3. Apocaulon carnosum

Conchocarpus 9

4. CONCHOCARPUS J.C. Mikan, Del. Fl. Faun. Bras. t. 2. 1823 [1820]. See Stearn, William T. 1956. Mikan’s Delectae Florae et Faunae Brasiliensis. J. Soc. Bibl. Nat. Hist. 3: 135–136. Shrubs or small trees to 7 m tall. Leaves alternate (rarely opposite), 1foliolate, 3-foliolate, or palmately 4–7-foliolate, rarely (in C. longifolius) apparently simple (i.e., not swollen at apex or base of petiole), petiolate; blade entire, leathery or papery. Inflorescence terminal or extra-axillary, rarely axillary, short- or long-pedunculate, a variously modified thyrse, borne usually on leafy stems, rarely (e.g., in C. ramiflorus) on leafless stems. Flowers actinomorphic (or zygomorphic). Sepals (4)5, quincuncial, usually connate at base, sometimes free, deciduous or persistent; petals (4)5, imbricate, free, coherent, or connate, often adherent to the staminal filaments, forming a ± tubular corolla, to ca. 2 cm long, white (rarely pink), usually densely sinuate-strigose. Androecium of 2 fertile stamens and 3(5) staminodes (in flora area), of 3 fertile stamens and 2 staminodes, or of 5 fertile stamens; filaments distinct (rarely connate), often variously adherent or adnate to the corolla; anthers ovate, oblong, or lanceolate, basifixed, usually free, rarely connate (e.g., in C. ucayalinus), not appendaged at base, sometimes apiculate. Disk cupular, shorter than or equal to the ovary, glabrous. Ovary of 5 carpels, these usually free (rarely connate axially and basally); ovules 2 per carpel; style usually 1 (5 in C. longifolius), the stigma enlarged or not, sometimes 5-lobed. Fruit of 1–5 mericarps, these usually free (rarely connate at base), dehiscing ventrally. Seed 1 per carpel, ovoid, smooth, subrugulose, or tuberculate, glabrous. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 45 species, 6 (maybe 7) in Venezuela, 4 of these in the flora area. Key to the Species of Conchocarpus 1. 1. 2(1).

2.

3(1).

3.

Leaves simple or 1-foliolate ...................................................................... 2 Leaves 3-foliolate ....................................................................................... 3 Leaves 47–76 × 9–14 cm, long-cuneate at base, not drying grayish green; petiole 6–8 mm long; peduncle flattened; petals coherent and adnate to apex of filaments; anthers of 2 fertile stamens not coherent; styles 5 .............................................................................................. C. longifolius Leaves 13–25(–42) × 3–6.5(–11.7) cm, obtuse at base, often drying grayish green; petiole 19–33(–60) mm long; peduncle not flattened; petals and filaments free; anthers of 2 fertile stamens laterally coherent; style 1 ..................................................................... C. ucayalinus Inflorescence cauline, bifurcate; peduncle < 1 cm long; fruiting mericarp slightly beaked at apex, the surface not obscured by pubescence ................................................................................................ C. ramiflorus Inflorescence a terminal thyrse; peduncle 8–20 cm long; fruiting mericarp rounded at apex, the surface obscured by pubescence ............................................................................................C. heterophyllus

Conchocarpus heterophyllus (A. St.Hil.) Kallunki & Pirani, Kew Bull. 53: 300. 1998. —Galipea heterophylla A. St.-Hil., Bull. Sci. Soc. Philom. Paris, sér. 3, 10: 132. 1823.

Treelet to 3 m tall. Dense evergreen forests, 100–300 m; Bolívar (Reserva Forestal Imataca). Suriname, Brazil (Maranhão to Rio de Janeiro).

10

R UTACEAE

Conchocarpus longifolius (A. St.-Hil.) Kallunki & Pirani, Kew Bull. 53: 308. 1998. —Galipea longifolia A. St.-Hil., Bull. Sci. Soc. Philom. Paris, sér. 3, 10: 131. Sep 1823. —Galipea pentagyna A. St.-Hil., Mém. Mus. Hist. Nat. 10: 281, t. 14A. 1823 [1824]. —Cusparia pentagyna (A. St.-Hil.) Engl. in Mart., Fl. Bras. 12(2): 122. 1874. —Angostura pentagyna (A. St.-Hil.) Albuq., Acta Amazon. 11: 851. 1981. —Cola de pava. Unbranched shrub or treelet 1–7 m tall; leaves large, stiff, clustered at the apex; thyrses long-pedunculate; flowers creamcolored with odor of citronella. Evergreen lowland forests, 100–500 m; Delta Amacuro, Bolívar (Serranía de Imataca). Suriname, French Guiana, eastern Brazil. ◆Fig. 5.

Fig. 4. Conchocarpus ucayalinus

Conchocarpus ramiflorus (Spruce ex Engl.) Kallunki & Pirani, Kew Bull. 53: 319. 1998. —Cusparia ramiflora Spruce ex Engl. in Mart., Fl. Bras. 12(2): 111. 1874. —Angostura ramiflora (Spruce ex Engl.) Albuq., Acta Amazon. 11: 851. 1981.

Fig. 5. Conchocarpus longifolius

Decagonocarpus 11

Treelet to 3 m tall; flowers white. Evergreen lowland forests, 100–200 m; Amazonas (base of Sierra de la Neblina). Brazil (northwestern Amazonas). Conchocarpus ucayalinus (Huber) Kallunki & Pirani, Kew Bull. 53: 323. 1998. —Cusparia ucayalina Huber, Bol. Mus. Paraense Hist. Nat. 4: 573, fig. 4. 1906. —Angostura ucayalina (Huber) Albuq., Acta Amazon. 11: 852. 1981. —Jazmín de monte.

Rauia ulei K. Krause in Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 145. 1914. —Angostura ulei (K. Krause) Gereau, Candollea 45: 386. 1990. Shrub or small tree to 4 m tall; petioles ferruginous. Evergreen lowland to submontane forests, 200–1000 m; Bolívar (Altiplanicie de Nuria, near Cerro Venado), Amazonas (Cerro Huachamacari). Colombia (Santander), Guyana, French Guiana, eastern Peru, Brazil, northern Bolivia. ◆ Fig. 4.

5. DECAGONOCARPUS Engl. in Mart., Fl. Bras. 12(2): 105, t. 21. 1874. Shrubs or trees to 8 m tall. Leaves opposite, simple, petiolate; blades entire, fleshy. Inflorescence axillary, a pedunculate, unilateral, few-flowered, sometimes congested raceme. Flowers slightly zygomorphic. Calyx cupular, ± equally 5-lobed, persistent in fruit, the lobes quincuncial in bud; corolla to 3.5 cm long, orange to red, fleshy, the tube longer than the lobes, slightly curved, ± 5-angled, the 5 lobes induplicate-valvate, recurved to spreading. Androecium of 2 fertile stamens and 3 linear staminodes; filaments connate in a tube adherent in its upper 1/2 to the corolla; anthers elliptic, basifixed, appendaged at base. Disk cupular, enclosing the ovary, glabrous. Ovary of 5 free carpels united only at apex in the style; ovules 2 per carpel; style 1; stigma only slightly enlarged, oblique. Fruit of 5 free mericarps, rarely fewer by abortion, each ventrally dehiscent and with beaked apex. Seeds 1 or 2 per carpel, reniform-elliptic to cuboid, tuberculate, glabrous. Endemic to the Guayana Shield in eastern Colombia, southern Venezuela, and northern Brazil; 2 species, both in the flora area. The two species keyed below may not be distinct; the morphological differences noted in specimens from higher elevations (i.e., Decagonocarpus cornutus) may reflect variation due to environmental factors. Key to the Species of Decagonocarpus 1.

1.

Pedicels 0–2 mm in flower, to 3 mm in fruit; fruiting carpels with 1 dorsal and 2 commissural ridges and a beak to 1.5 mm long; peduncle 1.2–6 cm long .................................................................... D. oppositifolius Pedicels 4–7.5(–19) mm in flower, 7–10 mm in fruit; fruiting carpels with 1 dorsal ridge and a beak 2–6 mm long; peduncle 0.2–3 cm long ................................................................................................... D. cornutus

Decagonocarpus cornutus R.S. Cowan, Mem. New York Bot. Gard. 10(1): 153, fig. 18. 1958. —Decagonocarpus cornutus var. congestus R.S. Cowan, Mem. New York Bot. Gard. 10(1): 155. 1958. Shrub to 3 m tall. Rocky areas in thickets, 1100–1800 m; Amazonas (Cerro Cuao, Cerro Guanay, Cerro Yutajé, Río Coro Coro). Reported from Brazil (northwestern Amazonas).

Published reports of Decagonocarpus cornutus from Colombia were based on misidentification of specimens of D. oppositifolius. Decagonocarpus oppositifolius Spruce ex Engl. in Mart., Fl. Bras. 12(2): 105, t. 21. 1874. Shrub or tree 1–8 m tall; leaves elliptic or obovate; corolla tubular, red or orange (although the inflorescences of specimens

12

R UTACEAE

collected in Venezuela are congested, those of most specimens from Colombia are elongated and more obviously racemose). Forests and open areas on granitic outcrops, 100– 400 m; Amazonas (Cerro Sipapo, basins of

Fig. 6. Decagonocarpus oppositifolius

upper Río Casiquiare, Río Negro, and Río Orinoco). Colombia (Caquetá, Guaínia), Brazil (Amazonas: Cucui on Rio Negro). ◆Fig. 6.

Fig. 7. Ertela trifolia

6. ERTELA Adans., Fam. Pl. 2: 358. 1763. Moniera Loefl., Iter Hispan. 197. 1758, non P. Browne 1756, spelling variants: Monnieria, Monniera, Monneria. Erect herbs, sometimes woody at base. Leaves usually opposite on lower part of stem, sometimes alternate or subopposite after an axis is terminated by an inflorescence and then branches dichotomously, 3-foliolate, petiolate; blades entire, membranous. Inflorescence apparently terminal, a pedunculate dichasium with a central flower and 2 secund, 1–5-flowered branches. Flowers zygomorphic. Sepals 5, free, quincuncial, very unequal in size and shape, with outermost one larger than other sepals and corolla and mimicking a bract, the second ca. 1/2 the size of first, the others minute, persistent in fruit; corolla 2-lipped (1 formed by the single innermost petal and the other 4-lobed), to 6 mm, white, the tube short, straight, the lobes imbricate. Androecium of 2 fertile stamens and 3 linear staminodes; filaments adherent to corolla, those of the fertile stamens connate, those of staminodes free; anthers oblong, basifixed, connate at adjacent edges, not appendaged at base. Disk cupular, shorter than ovary except on 1 side (opposite the 4-lobed lip) elongate, scale-like, and exceeding the ovary. Ovary of 5 free carpels; ovules 2 per carpel; style 1; stigma obliquely capitate (turned to face the fertile stamens). Fruit of 1–5 free mericarps, each dehiscent ventrally and dorsally. Seeds 1 per carpel, ellipsoid-reniform, tuberculate, glabrous. Southwestern Mexico, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela.

Erythrochiton

Ertela trifolia (L.) Kuntze, Revis. Gen. Pl. 1: 100. 1891. —Monnieria trifolia L., Syst. Nat. ed. 10, 2: 1153. 1759. —Albahaca, Mata de hormiga. Erect herb to 1 m tall; flowers white. Semideciduous forests; disturbed river

13

banks, forest edges, and roadsides, 50–700 m; scattered in Bolívar and Amazonas. Barinas, Portuguesa, Sucre, Táchira, Zulia; southwestern Mexico, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 7.

Fig. 8. Erythrochiton brasiliensis

14

R UTACEAE

7. ERYTHROCHITON Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 11: 151, 165. 1823. Shrubs or trees 0.5–12 m tall. Leaves alternate, 1-foliolate, petiolate; blades entire, papery when dry. Inflorescence lateral but not obviously axillary, a pedunculate dichasium, in age often perennating, branched, elongate, cauline, and bearing flowers on short shoots, or (in E. hypophyllanthus Planch. & Linden from Colombia) a sessile, short cyme borne on the lower surface of a leaf. Flowers actinomorphic to zygomorphic. Sepals 5, variously connate at anthesis, valvate, 1.5–4.6 cm long, lanceolate, red, white, or green, usually persistent in fruit; corolla 4.3–7.9 cm long, white, glabrous without (pubescent in E. trichanthus Kallunki from Peru); the petals connate, forming a ± straight tube, the 5 lobes imbricate. Androecium of 5 fertile stamens or of 2 fertile and 3–5 linear staminodes; filaments adnate to the corolla; anthers oblong to lorate, basifixed, when only 2 fertile appendaged at base. Disk tubular, taller than ovary, glabrous. Ovary of 5 free carpels; ovules 2 per carpel; style 1; stigma capitate and 5-lobed. Fruit of 5 free mericarps (rarely 1 aborting) each dehiscing ventrally. Seeds 1 or 2 per carpel, ovoid, reniform, or cuboid, sometimes slightly beaked above the hilum, tuberculate, pubescent. Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 7 species, 1 in Venezuela. Erythrochiton brasiliensis Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11: 166, t. 18C, 22. 1823. —Chiquita blanca, Cola de pava. Shrub or small, often unbranched tree to 12 m tall; flowers large with red calyx and

glabrous, white corolla. Evergreen lowland or lower montane forests, 100–400 m; Bolívar (basins of Río Caroni, Río Paragua, and Río Yuruari in the Río Cuyuní basin), Amazonas (upper Río Orinoco). Colombia, Guyana, Suriname, French Guiana, Ecuador(?), Peru, Brazil, Bolivia. ◆Fig. 8.

8. ESENBECKIA Kunth in H.B.K., Nov. Gen. Sp. (folio ed.) 7: 191. 1825. Shrubs or trees, usually to 12 m tall, occasionally to 25 m. Leaves alternate or opposite, simple or digitately 1–5-foliolate (1-foliolate in the flora area); petiole winged or not; blades entire, papery or ± leathery when dry. Inflorescence terminal or axillary, a broadly or narrowly paniculate thyrse, or much condensed when axillary. Flowers actinomorphic. Calyx 5(7)-lobed, persistent in fruit, the lobes mostly quincuncial; petals 5(7), free, imbricate or valvate, 1.4–6.5 mm long, usually yellow to white, occasionally purplish brown to violet, spreading to reflexed when flowering, persistent or deciduous in fruit. Androecium of 5(7) fertile stamens; filaments free, lying against the grooves of the disk, in some species with basal abaxial swellings; anthers heart-shaped, dorsifixed, versatile, apiculate. Disk annular or cupular, embracing base of ovary, glabrous or papillose. Ovary of 5 basally or completely connate carpels; ovules 2 per carpel, collateral; style 1; stigma capitate or clavate, 5-lobed or entire. Fruit capsular, dorsally smooth, tuberculate, echinate, or muricate, with or without a knob-like projection on each carpel, globose to stellately lobed in cross section, dehiscent septicidally along the abaxial sutures and loculicidally along the ventral sutures to the apex. Seeds 1 or 2 per carpel, obliquely tear-shaped or obliquely subglobose, ± beaked at apex, smooth, shiny or dull, glabrous.

Esenbeckia 15

Mexico, Central America, Jamaica, Martinique(?), Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 26 species, 2 in Venezuela, both in the flora area. Key to the Species of Esenbeckia 1.

1.

Petioles not winged; apex of leaflets rounded or obtuse; petals 3.4–6 mm long; fruits prickly but without knob-like projections on each carpel, globose in cross section ......................................................... E. grandiflora Petioles winged or at least narrowly margined; apex of leaflets mostly distinctly acuminate; petals 2.5–3 mm long; fruits ± muricate and with dorsal knob-like projections on each carpel (more easily discernible as such prior to dehiscence), stellately lobed in cross section ................................................................................ E. pilocarpoides

Esenbeckia grandiflora Mart., Nov. Gen. Sp. Pl. 3: 85. 1831. Venezuela, Trinidad, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina; 2 subspecies, 1 with 2 varieties, 1 subspecies with 1 variety in Venezuela. E. grandiflora subsp. grandiflora var. grandiflora. —Arautamakatadek (Arekuna), Wayarayuyek (Arekuna). Esenbeckia obovalifolia Pittier, Bol. Soc. Venez. Ci. Nat. 9: 122. 1944. Tree 6–20 m tall. Lower montane to montane and gallery forests, 200–1000 m; Delta Amacuro (Altiplanicie de Nuria), Bolívar (upper Río Caroni basin in Gran Sabana, Serranía de Imataca). Aragua, Falcón, Nueva Esparta; Trinidad-Tobago, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina. The second variety of this subspecies, var. intermedia (Mart. ex Engl.) Kaastra, occurs in Brazil (Rio de Janeiro). Esenbeckia pilocarpoides Kunth in H.B.K., Nov. Gen. Sp. (folio ed.) 7: 192, t. 655. 1825. Martinique(?), Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, Brazil, Bolivia; 2 subspecies, 1 in Venezuela. E. pilocarpoides subsp. pilocarpoides. —Gaspadillo grande. Esenbeckia venezuelensis Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 159. 1896. Shrub or small tree to 4 m tall. Semideciduous to evergreen lowland or lower

Fig. 9. Esenbeckia pilocarpoides subsp. pilocarpoides

montane forests, sometimes on rocky soil or on granitic outcrops or near river rapids, 50–700 m; Bolívar (Altiplanicie de Nuria, Río Caroni, Río Cuyuní, west of Río Maniapure), Amazonas (El Motín and Bobadilla on upper Río Orinoco). Northern Venezuela; Martinique(?), Colombia, Trinidad-Tobago, Guyana, Suriname, Amazonian Brazil, Bolivia. ◆Fig. 9.

16

R UTACEAE

9. GALIPEA Aubl., Hist. Pl. Guiane 662, t. 269. 1775. Trees or shrubs to 15 m tall. Leaves alternate, 1- or 3-foliolate, petiolate; blades subsessile or petiolulate, entire, often bidentate at apex, papery when dry. Inflorescence terminal or lateral, pedunculate, an elongate, elliptic, or corymboid thyrse or a capitate dichasium. Flowers slightly zygomorphic. Calyx cupular, 5lobed or -denticulate, persistent in fruit; corolla to 5 cm long, white, appressedpubescent, the tube straight or curved, the 5 lobes imbricate, subequal. Androecium of 2 fertile stamens and 3–6 linear or ligulate staminodes often tipped with a globose gland; filaments connate into a tube tightly adherent or adnate to the corolla tube; anthers narrowly oblong, basifixed, appendaged at base. Disk cupular, shorter than or equal to ovary, glabrous. Ovary of 5 carpels connate laterally and ventrally or less often free; ovules 2 per carpel; style 1; stigma capitate, ± 5-lobed. Fruit a capsule with 5 mericarps connate ventrally and dorsally at maturity, each dehiscing ventrally but remaining connate in basal 1/2 dorsally, or of 5 free mericarps, each dehiscing ventrally. Seeds 1 per carpel, ellipsoid, smooth or slightly rugulose, glabrous or pubescent. Guatemala(?), Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; ca. 15 species, 2 in Venezuela, 1 of these in the flora area. Galipea davisii Sandwith, Bull. Misc. Inform. 1935: 118. 1935. —Ehparayo’ (Panare), Quino blanco. Tree to 15 m tall; leaves 3-foliolate, rather shiny; corymboid thyrses of fragrant flowers with slightly curved, funnelform corollas to 2.5 cm long; capsules pale, glandular-punctate, ca. 2 cm long. Semideciduous

forests on granitic slopes, 100–400 m; western Bolívar, northern Amazonas. Guyana, Brazil (northern Pará and Roraima). ◆Fig. 10.

Fig. 10. Galipea davisii

Hortia 17

10. HORTIA Vand., Fl. Lusit. Bras. Spec. 14. 1788. Shrubs or trees to 38 m tall. Leaves alternate, simple, petiolate, erect or ascending, often clustered at branch ends; petiole often swollen at base; blades 9– 96 cm long, oblanceolate, acute, decurrent, or often long-cuneate at base, entire, leathery when dry. Inflorescence terminal, a robust, many times dichotomous, many-flowered, flat- or round-topped dichasium, often hidden from below by large, ascending leaves; epidermis of branches and peduncle often transversely cracked. Flowers actinomorphic. Calyx cupular, 5-lobed to subentire, persistent in fruit, the lobes quincuncial in bud; petals 5, free, valvate, to 8 mm long, lanceolate, various shades of pink or red, with inflexed tip, barbate within, finally recurved. Androecium of 5 fertile stamens; filaments free, at base immersed in disk; anthers oblong, bifid at base, dorsifixed. Disk 5-lobed in cross section, much shorter than ovary, glabrous. Ovary of 5 connate carpels, ± tapering to style; ovules 2 per carpel; style 1, longitudinally 5-grooved, shorter than or equal to ovary. Fruit a 5-locular, ± globose drupe; pericarp glandular, fleshy; each locule with bony endocarp. Seeds 1 or 2 per locule, ellipsoid or tear-shaped, smooth, glabrous. Panama, Colombia, Venezuela, Guyana, Brazil; 11 species, 4 in Venezuela, all of these in the flora area. Key to the Species of Hortia 1. 1.

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

Leaves 9–26 cm long, narrowly oblong, oblanceolate, or obovate, acute or decurrent to an obvious petiole 1–4.5 cm long .................................... 2 Leaves mostly 50–96 cm long (those immediately below the inflorescence as short as 24 cm), oblanceolate or narrowly oblanceolate, longcuneate and decurrent to within 0.5 cm of swollen petiolar base ...... 3 Leaves narrowly oblong or oblanceolate ....................................... H. coccinea Leaves obovate ......................................................................... H. vandelliana Pedicels 2–3 mm wide, glandular-crusty as are sepals; leaves 60–96 × 12– 20 cm, broadly rounded and retuse or apiculate at apex ...... H. neblinensis Pedicels ca. 1–1.2 mm wide, not glandular-crusty; leaves (24–)50–78 × (5.3–)8.5–13.5 cm, obtuse, short-acuminate, or rounded at apex ......................................................................................................... H. regia

Hortia coccinea Spruce ex Engl. in Mart., Fl. Bras. 12(2): 183. 1874. Shrub or small tree; petals red. Woodland or savanna margins on granitic outcrops near rivers, 100–200 m; Amazonas (Budare in upper Río Temi basin, Caño San Miguel, base of Cerro Yapacana, lower Río Atacavi, Río Casiquiare, Río Pasimoni). Brazil (Amazonas). ◆Fig. 11. Hortia neblinensis Maguire & B. Boom, Mem. New York Bot. Gard. 51: 121. 1989. Tree to 25 m tall and 40 cm diam.; flower buds large for genus. Montane forests, 1200–1300 m; Amazonas (likely on Sierra de

la Neblina). Brazil (Amazonas: Serra da Neblina, Serra Pirapucú). Hortia regia Sandwith, Bull. Misc. Inform. 1931: 184. 1931. Tree ca. 20 m tall. Lower montane forests, 500–600 m; Bolívar (La Escalera at km 109 south of El Dorado). Guyana. Hortia vandelliana Groppo, Novon 15: 141. 2005. Tree or shrub to 20 m tall; fruits green becoming light yellow. Evergreen lowland forest, 100–200 m; Amazonas (near Yavita). Brazil (Amazonas, Roraima).

18

R UTACEAE

Fig. 11. Hortia coccinea

Fig. 12. Murraya paniculata

11. MURRAYA J. König ex L., Mant. Pl. 2: 554. 1771, nom. cons. Chalcas L., Mant. Pl. 1: 68. 1767. Trees or shrubs. Leaves alternate, odd-pinnate, petiolate; leaflets alternate, entire or crenulate, papery when dry. Inflorescence terminal or upper-axillary, a variably branched, ± flat-topped dichasium. Flowers actinomorphic. Sepals 5, free

Neoraputia 19

or united, ovate or triangular, persistent in fruit; petals 5, free, imbricate, 1–2 cm long, lanceolate to oblanceolate, white. Androecium of 10 fertile stamens; filaments free, linear-awl-shaped, subequal or alternately long and short; anthers small, subglobose to elliptic. Disk annular or short-cylindric, shorter than ovary. Ovary of 2–5 connate carpels; ovules 1 or 2 per carpel; style 1, slender, longer than the ovary, deciduous; stigma capitate. Fruit an ovoid or globose berry; pericarp thin, glandular, red or black; pulp mucilaginous. Seeds 1 or few, glabrous or tomentellous. Tropical Asia, Pacific Islands, 2 species widely cultivated and sometimes naturalized in most tropical areas; ca. 12 species, 1 in Venezuela. Murraya paniculata (L.) Jack, Malayan Misc. 1: 31. 1820. —Chalcas paniculata L., Mant. Pl. 1: 68. 1767. Murraya exotica L., Mant. Pl. 2: 563. 1771. —Chalcas exotica (L.) Millsp., Publ. Field Columbian Mus., Bot. Ser. 1: 25. 1895. Tree to 10 m tall; flowers very fragrant, white; fruits red or black. Gallery forests

and adjacent savannas, ca. 50 m; Bolívar (east of Ciudad Bolívar near Río Marhuanta). Widespread in cultivation or locally escaped in Venezuela; native to tropical Asia, but widely cultivated and often naturalized in warm parts of the world. ◆Fig. 12.

12. NEORAPUTIA Emmerich, Rodriguésia 30(45): 234. 1978. Raputia Aubl. pro parte, sensu Engl. in Mart., Fl. Bras. 12(2): 102, t. 20, 21. 1874. Trees 3–15 m tall. Leaves alternate or in 1 species sometimes subopposite, 1– 7-foliolate, petiolate; blades entire, papery when dry. Inflorescence terminal, pedunculate, usually a dichasium with 2 racemose branches, or in 1 species irregularly 1–3 times di- or trichotomous, with secund branches. Flowers ± actinomorphic. Calyx cyathiform, ± 5-dentate or -lobed, sometimes irregularly so and then sometimes appearing bilabiate, slightly enlarged, explanate, and persistent in fruit; corolla 1.5–2.5 cm long, white, salmon-pink, or brick-red, densely appressed-pubescent, the petals 5, coherent at base forming a straight tube shorter than or equal to lobes, the lobes imbricate, obtuse, spreading. Androecium of 2 fertile stamens and 3 linear staminodes awl-shaped at apex; filaments distinct, adherent to corolla tube; anthers oblong, basifixed, with small rim-like appendages at base. Disk cupular, taller than ovary, glabrous. Ovary of 5 carpels connate ventrally and basally; ovules 2 per carpel; style 1; stigma capitate, 5-sulcate. Fruit a 5-parted capsule, the mericarps connate ventrally and basally at maturity, each dehiscing ventrally and then sometimes remaining connate only at base, densely tomentulose. Seeds (1)2 per carpel, subglobose to ellipsoid, ± angled, smooth, brown-mottled, glabrous. Venezuela, Peru, Brazil; ca. 5 species, 1 in Venezuela. Neoraputia paraënsis (Ducke) Emmerich, Rodriguésia 30(45): 243. 1978. —Raputia paraënsis Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 184. 1922. —Guachimacán. Neoraputia cowanii Emmerich, Rodriguésia 30(45): 259, fig. 9. 1978.

Tree to 10 m tall; leaves 5-foliolate; flowers salmon-pink. Evergreen lowland forests, 100–300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, road from Casa Blanca to Anacoco). French Guiana, Brazil (Acre, Pará, Maranhão, Bahía). ◆Fig. 13.

20

R UTACEAE

Fig. 13. Neoraputia paraënsis

13. PILOCARPUS Vahl, Eclog. Amer. 1: 29. 1796 [1797]. Shrubs or trees to 12(–20) m tall. Leaves alternate to subverticillate by crowding at tips of branches, simple, digitately 1- or 3-foliolate, or odd-pinnate; petioles if present sometimes narrowly winged; blades entire, usually emarginate, papery to leathery when dry. Inflorescence terminal or axillary, racemose. Flowers actinomorphic. Calyx 4- or 5-lobed or -toothed, persistent in fruit; petals (4)5, free, induplicate-valvate, coherent in bud by uncinnately inflexed tips, ovate, 1.9–5 mm long, brownish, red, purple, pink, or yellow-green, reflexed at anthesis, deciduous. Androecium of (4)5 fertile stamens; filaments free, accumbent in grooves of disk; anthers heart-shaped with dorsal gland near apex, dorsifixed, versatile or not. Disk annular to cupular, completely adnate to the ovary, glabrous or rarely pubescent. Ovary of 5 basally connate and distally free carpels; ovules 1 or 2 per carpel, collateral but if both develop seeds then becoming superposed; style 1,

Pilocarpus 21

shorter than ovary; stigma 5-lobed. Fruit of 1–5 free (united at very base) mericarps, each dehiscent ventrally, arcuately veined or tuberculate. Seeds 1 or 2 per carpel, usually reniform, keeled abaxially, smooth, shiny, glabrous. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Paraguay, Argentina; 16 species, 2 in Venezuela, 1 of these in the flora area. Pilocarpus racemosus Vahl, Eclog. Amer. 1: 29, t. 10. 1796 [1797]. Mexico, Central America, West Indies, Venezuela, French Guiana; 2 subspecies, 1 with 2 varieties, 1 subspecies with 1 variety in Venezuela. P. racemosus subsp. racemosus var. racemosus Tree 3–20 m tall, leaves odd-pinnate grading to 1- or 3-foliolate toward base of

inflorescence; pedicels to 2 cm long. Semideciduous to evergreen lower montane forests, sometimes on exposed granitic outcrops, 200–600 m; Bolívar (islands in Lago Guri, basin of Río Caroni, Río Yuruari in the Río Cuyuní basin). Lara; western Mexico, West Indies, French Guiana. ◆Fig. 14.

The second variety of this subspecies, var. yucatanus Kaastra, occurs in Mexico (Yucatán).

Fig. 14. Pilocarpus racemosus subsp. racemosus var. racemosus

22

R UTACEAE

14. RAPUTIA Aubl., Hist. Pl. Guiane 670, t. 272. 1775. Myllanthus R.S. Cowan, Mem. New York Bot. Gard. 10(2): 34. 1960. Shrubs or trees to 13 m tall. Leaves opposite, 1- or 3-foliolate, petiolate; blades entire, papery to leathery when dry. Inflorescence axillary, cauline (rarely in axil of a leaf), pedunculate, a coiled, unilateral raceme, 1–4 at a node. Flowers zygomorphic. Sepals 5, connate at very base, quincuncial, broadly overlapping, persistent in fruit; corolla curved, 2-lipped, to 1.5 cm long, white or pale green, the tube short, the 5 lobes imbricate, the innermost 1 connate to a lesser extent and the others forming a 4-lobed, recurved lip. Androecium of 2 fertile stamens and (2)3 awl-shaped staminodes; filaments distinct but adherent to corolla by pubescence; anthers oblong or narrow-ovate, basifixed, connate, with obvious asymmetric or falcate appendages at base. Disk cupular, shorter to taller than ovary, glabrous. Ovary of 5 free carpels; ovules 2 per carpel; style 1; stigma obliquely capitate. Fruit of 1–5 free mericarps, each ventrally dehiscent. Seeds 1 per carpel, oblong, smooth, glabrous. Amazonian Colombia, Venezuela, French Guiana, Amazonian Peru, northern Brazil; 10 species, 3 in Venezuela, all in the flora area. Key to the Species of Raputia 1.

1. 2(1).

2.

Leaves 1-foliolate, subcoriaceous, the upper surface shiny; secondary veins 27–36 per side, straight, forming a ± straight intramarginal vein ........................................................................................ R. szczerbanii Leaves 3-foliolate, chartaceous, the upper surface dull: secondary veins 14–32 per side, straight or curved-ascending, looping at margin ...... 2 Leaflets sessile or with petiolules ca. 0.4 cm long, oblanceolate to oblong, central leaflet 25–54 × 8–14 cm; inflorescence 2.5–4 cm long; fruiting mericarps glabrous ................................................................... R. maroana Leaflets with petiolules 0.5–2 cm long, elliptic, central leaflet 16–23 × 5.5–8 cm; inflorescence 8.5–14 cm long; fruiting mericarps densely strigulose ............................................................................... R. neblinensis

Raputia maroana (R.S. Cowan) Kallunki, Brittonia 42: 176. 1990. —Myllanthus maroana R.S. Cowan, Mem. New York Bot. Gard. 10(2): 35, fig. 22. 1960. Shrub or tree 2–6 m tall; leaves glabrous, drying yellow-brown abaxially. Evergreen lowland forests, 100–200 m; Amazonas (near Maroa, Río Autana). Brazil (northwestern Ama-zonas). ◆Fig. 15. Raputia neblinensis (R.S. Cowan) Kallunki, Brittonia 42: 177. 1990. —Myllanthus neblinensis R.S. Cowan, Mem. New York Bot. Gard. 10(2): 37, fig. 22. 1960.

Tree to 5 m tall; known only by the type collection (in bud). Talus forests, 900–1000 m; Amazonas (slopes of Sierra de la Neblina). Endemic. ◆Fig. 16. Raputia szczerbanii (Steyerm.) Kallunki, Brittonia 46: 282. 1994. —Lubaria szczerbanii Steyerm., Brittonia 32: 17, fig. 1. 1980. Tree to 5 m tall. Rocky outcrops along rivers on tepui summit, ca. 700–800 m; Bolívar (Cerro Guaiquinima). Endemic. ◆Fig. 17.

Raputia 23

Fig. 15. Raputia maroana

Fig. 16. Raputia neblinensis

24

R UTACEAE

Fig. 17. Raputia szczerbanii

15. RAUIA Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11: 151, 167. 1823. Shrubs or trees to 10 m tall. Leaves alternate, 1- or 3-foliolate, petiolate; blades entire, papery when dry. Inflorescence terminal, pedunculate, a corymboid thyrse or rarely a pseudoraceme. Flowers actinomorphic. Calyx cupular, 5denticulate or -dentate, persistent in fruit; corolla to 20 mm long, white or cream, densely appressed-pubescent, the petals free at very base and ± coherent and

Rauia 25

adherent to filaments above the base forming a straight tube ≤ the lobes, the 5 lobes very narrow, imbricate in bud, spreading or recurved. Androecium of 2 fertile stamens and 3 or 5 linear, awl-shaped staminodes; filaments distinct, adherent by abaxial pubescence to the petals above the base; anthers lorate, basifixed, not appendaged at base, apiculate. Disk cupular, equal to or taller than ovary, denticulate, glabrous. Ovary of 5 carpels connate ventrally especially at apex and sometimes also laterally at base; ovules 2 per carpel; style 1, apparently remaining within the corolla tube; stigma clavate. Fruit a capsule of 4 or 5 mericarps connate ventrally and/or basally or free. Seeds 1(2?) per carpel, subglobose, ellipsoid, or reniform, smooth, glabrous or pubescent. Venezuela, Guyana, Suriname, Brazil; ca. 8 species, 2 in Venezuela, both in the flora area. Rauia is similar to some Conchocarpus, and their relationship requires further study.

Fig. 18. Rauia subtruncata

26

R UTACEAE

Key to the Species of Rauia 1.

1.

Leaves 3-foliolate; inflorescence a pseudoraceme; staminodes 5 (2 epipetalous); fruiting mericarps obovate, conchiform, obtusely carinate dorsally, densely appressed-pubescent; seeds pubescent with short, erect trichomes............................................................................. R. spicata Leaves 1-foliolate; inflorescence a corymboid thyrse; staminodes 3; fruiting mericarps subglobose; transversely wrinkled; seeds unknown .............................................................................................. R. subtruncata

Rauia spicata Haye in Pulle, Recueil Trav. Bot. Néerl. 30: 182. 1933. Tree 10 m tall. Granitic outcrops and ledges along rapids, 200–300 m; Bolívar (Río Toro). Suriname.

Rauia subtruncata Steyerm., Ann. Missouri Bot. Gard. 75: 315, fig. 3. 1988. —Jasmín grande, Nicolás. Tree to 6 m tall. Semideciduous lowland forests, 100–300 m; Bolívar (between El Manteco and San Pedro de las Bocas). Guyana. ◆Fig. 18.

16. RAVENIOPSIS Gleason, Brittonia 3: 166. 1939. Shrubs or small trees, with simple or branched trichomes. Leaves usually opposite, occasionally some leaves (in R. steyermarkii) alternate, simple or 1–3foliolate; blades entire, papery to leathery when dry. Inflorescence axillary or terminal, a simple spike or raceme (rarely a panicle of racemes), sometimes a congested, unilateral, and ± capitate dichasium, a dichasium with 2 coiled, racemose branches and a terminal flower at apex of peduncle, or rarely a solitary flower. Flowers slightly zygomorphic to markedly 2-lipped. Sepals 5, free or shortly connate, quincuncial, usually unequal (sometimes markedly so), persistent in fruit; corolla to 2.5 cm long, white, pink, orange, or red, often densely pubescent, the tube cylindric, ± curved, the 5 lobes imbricate, spreading, subequal to markedly unequal. Androecium of 2 fertile stamens and 3 linear staminodes; filaments ± connate, tightly adherent or adnate to the corolla; anthers narrowly oblong, basifixed, sometimes laterally coherent, sometimes pubescent, with minute saccate to larger falcate or flabellate appendages at base. Disk cupular, truncate to ± lobate, shorter than or equal to ovary. Ovary of 5 carpels, free laterally and united at base of style or sometimes at base ventrally; ovules 2 per carpel; style 1; stigma ± 5-lobed. Fruit of 1–5 free mericarps, each ventrally dehiscent and with beaked apex. Seed 1 per carpel, ellipsoid, tuberculate, glabrous. Endemic to the Guyana Shield in southern Venezuela and northern Brazil; 19 species, 16 in Venezuela, all in the flora area. At least one of the two reports from Colombia was based on misidentification of specimens of Decagonocarpus oppositifolius. Key to the Species of Raveniopsis 1. 1. 2(1).

Leaves 3-foliolate (occasionally simple leaves also present) .................. 2 Leaves simple ............................................................................................. 4 Pubescence of branched hairs; leaves papery when dry; inflorescence pedunculate, dichasial, with 1 terminal flower and 2 coiled, racemose branches; mature corollas pubescent with branched hairs without and ca. 16–25 mm long ................................................................... R. stelligera

Raveniopsis 27

2.

Pubescence of simple hairs; leaves leathery or papery when dry; inflorescence pedunculate and racemose (rarely branched with racemose branches) or subsessile and congested; mature corollas ± glabrous and ca. 15–20 mm long or densely pubescent and ca. 10 mm long ......................................................................................................... 3 3(2). Leaves leathery when dry, with dense, buff-colored, woolly hairs covering the lower surface; petiole 1.5–4 mm long; inflorescence subsessile, congested, densely pubescent, appearing almost capitate initially; mature corollas densely pubescent without, ca. 10 mm long ................................................................................................ R. cowaniana 3. Leaves papery when dry, with appressed straight hairs especially on veins beneath; petiole 12–35 mm long; inflorescence pedunculate, unilaterally racemose, rarely branched (fasciated?) and with branches racemose; mature corollas almost glabrous without, ca. 15–20 mm long ......................................................................................... R. trifoliolata 4(1). Pubescence of branched hairs ................................................................... 5 4. Pubescence of simple hairs ....................................................................... 9 5(4). Hairs on lower surface of leaf dendritic (with elongate rachis and many irregular branches); inflorescence spicate, secund, congested; corolla white ....................................................................................................... 6 5. Hairs on lower surface of leaf echinoid (with many branches originating at ± one point, either stipitate or sessile), leaf surface appearing lepidote; inflorescence racemose or dichasial with spicate branches; corolla white or red ............................................................................... 7 6(5). Leaf blade 1.4–1.8 times as long as wide; inflorescence ferruginous, appearing bristly due to the straight, relatively long branches of the hairs; fruiting mericarps with ± straight, simple hairs ...... R. steyermarkii 6. Leaf blade 2–3.8 times as long as wide; inflorescence buff, turning gray, appearing soft due to the sinuous, fine, relatively shorter branches of the hairs; fruiting mericarps with sinuous, sometimes dendritic hairs ................................................................................................. R. tomentosa 7(5). Leaf blade 1.5–4.1 cm long; hairs on lower surface of leaf sessile, most somewhat peltate; inflorescence racemose, secund; corolla white ..................................................................................................... R. breweri 7. Leaf blade 3–11 cm long; hairs on lower surface of leaf of 2 kinds, darker ones stipitate and somewhat peltate, paler ones sessile and echinoid; inflorescence dichasial; corolla red ....................................................... 8 8(7). Leaf blade widest near middle, acute or cuneate at base; corolla 1.2– 1.4 cm long ............................................................................... R. jauaensis 8. Leaf blade widest below middle, usually obtuse, rounded, or subcordate (rarely acute) at base; corolla 1.6–2.2 cm long ................. R. peduncularis 9(4). Leaves narrowly linear, 2–3 mm wide, strongly revolute; corolla 6–7 mm long .............................................................................................. R. linearis 9. Leaves elliptic, ovate, or obovate, > 3 mm wide, sometimes revolute; corolla > 10 mm long ........................................................................... 10 10(9). Inflorescence a single flower; peduncle ca. 2 mm long .................R. paruana 10. Inflorescence usually > 1 flower, or if 1 flower then peduncle at least 7 mm long (R. ruellioides) ................................................................... 11

28

R UTACEAE

11(10). Inflorescence a dense, secund spike; pubescence obscuring lower surface of leaf; veins on lower surface of leaf 15–20, equally prominent ................................................................................................... R. nubicola 11. Inflorescence dichasial, racemose, or subcapitate; pubescence, if present, not obscuring lower surface of leaf, or if so then major veins on lower surface of leaf 5–11, often alternating with less prominent ones .... 12 12(11). Inflorescence dichasial with a central flower and 2 racemose branches, without bracteal leaves at base of bifurcation; peduncle 3–5.8 cm; corolla white ............................................................................ R. abyssicola 12. Inflorescence subcapitate, racemose, or if dichasial with 2 bracteal leaves at base of bifurcation, these deciduous before fruiting; peduncle 0.8–4 cm; corolla white (sometimes with pink lobes) or red ............. 13 13(12). Corolla red ................................................................................................ 14 13. Corolla white ............................................................................................ 15 14(13). Lower surface of leaves hirsute with erect or ascending hairs; inflorescence subcapitate; sepals with spreading hairs 0.7–1.6 mm long, the margin appearing bristly-ciliate to naked eye .......... R. capitata 14. Lower surface of leaves with appressed pubescence on midrib and usually also on secondary veins; inflorescence a 2–4-flowered raceme (may appear capitate in bud); sepals with appressed hairs 0.3–0.5 mm long, the margin not appearing bristly-ciliate to naked eye ................................................................................................ R. ruellioides 15(13). Inflorescence secund and racemose, with flowers often few and congested and then appearing ± capitate; peduncle 0.3–1.5 cm long; sepals 5.2–10 mm long, with coarse pubescence of appressed to spreading hairs not obscuring both surfaces; lower surface of leaf tomentose, with curling or sinuous hairs, rarely glabrescent between veins ........................................................................................... R. fraterna 15. Inflorescence dichasial, with 2 reduced bracteal leaves at base of bifurcation, the lateral branches sometimes few-flowered and not elongated so that inflorescence appears ± capitate; peduncle 1.8–4 cm long; sepals 2.5–6 mm long, with dense sericeous pubescence completely covering both surfaces; lower surface of leaf sericeous, with ± straight, appressed hairs, sometimes glabrescent between secondary veins ............................................................................ R. sericea Raveniopsis abyssicola R.S. Cowan, Mem. New York Bot. Gard. 10(2): 32, fig. 11. 1960. Shrub 2–3 m tall with white flowers. Along canyons on tepui slopes, 1000–1100 m; Amazonas (Sierra de la Neblina). Endemic. Raveniopsis breweri Steyerm., Brittonia 32: 48. 1980. Erect shrub to 1.5 m tall; lower surface of leaf densely pubescent, appearing rustylepidote to the naked eye; flowers fragrant.

Rare to frequent in understory of shrubby thickets, in open vegetation on boggy ground, moist meadows, rocky river beds on sandstone, 1700–2000 m; Bolívar (Auyántepui). Endemic. Raveniopsis capitata R.S. Cowan, Mem. New York Bot. Gard. 23: 863. 1972. Shrub to 1.5 m tall; racemes capitate in bud, racemose when flowering. Open rocky sandstone areas, along streams in thickets near meadows, 1900–2100 m; Bolívar (Cerro Jaua), Amazonas (Cerro Parú). Endemic.

Raveniopsis 29

Raveniopsis cowaniana Steyerm. & Luteyn, Ann. Missouri Bot. Gard. 71: 316. 1984. Wiry shrub to 1.2 m tall; corollas red. Heliamphora savannas and scrub on ridges, 1700–1900 m; Amazonas (northwestern plateau of Sierra de la Neblina). Endemic. Raveniopsis fraterna R.S. Cowan, Mem. New York Bot. Gard. 10(2): 31, fig. 13. 1960. Shrub to 2.5 m tall with elongate branches. In montane thickets, on rocks, or in riparian woods, 2100–2500 m; Bolívar (Macizo del Chimantá). Endemic. Raveniopsis fraterna is closely related to R. sericea, also of Macizo del Chimantá, and may hybridize with it. Raveniopsis jauaensis Steyerm., Bol. Soc. Venez. Ci. Nat. 132: 338. 1976. Shrub to 3 m tall. Tepui summit forest and meadows, 1400–1500 m; Bolívar (Cerro Guanacoco, Cerro Sarisariñama). Endemic. Raveniopsis linearis (Gleason) R.S. Cowan, Mem. New York Bot. Gard. 10(2): 33. 1960. —Ravenia linearis Gleason, Bull. Torrey Bot. Club 58: 375. 1931. Shrub 0.3–1.5 m tall with small, white flowers. Montane thickets along streams; 1000–2000 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 23. Raveniopsis nubicola R.S. Cowan, Mem. New York Bot. Gard. 10(2): 34, fig. 17. 1960. Shrub 0.7–1.5 m tall; corollas pale orange-red; calyx, vegetative bud, and lower surface of leaves with very dense, yellowish indument. Montane scrub forests, ca. 2000 m; Amazonas (Sierra de la Neblina). Endemic. Raveniopsis paruana (R.S. Cowan) R.S. Cowan, Mem. New York Bot. Gard. 10(2): 32, fig. 15. 1960. —Ravenia paruana R.S. Cowan, Mem. New York Bot. Gard. 8: 122. 1953. Sprawling shrub 0.1–1.5 m tall; corollas orange-red. Locally frequent in rolling tepui meadows, ca. 2000 m; Amazonas (Cerro Parú). Endemic.

Raveniopsis peduncularis Pittier & Lasser, Bol. Soc. Venez. Ci. Nat. 9: 121. 1944. Shrub to 3 m tall; lower surface of leaf with dense pubescence, appearing lepidote. Frequent at edges and in understory of montane thickets, open rocky places in tepui meadows, 700–1800 m; Bolívar (Cerro Guaiquinima). Endemic. ◆Fig. 19. Raveniopsis ruellioides (Oliv.) R.S. Cowan, Mem. New York Bot. Gard. 10(2): 30, fig. 21. 1960. —Ravenia ruellioides Oliv. in Thurn, Timehri 5: 191. 1886, “Revenia.” Ravenia tatei Gleason, Bull. Torrey Bot. Club 58: 376. 1931. Ravenia ruellioides var. ptariana Steyerm., Fieldiana, Bot. 28: 273. 1952. Raveniopsis ruellioides var. angusta R.S. Cowan, Mem. New York Bot. Gard. 10(2): 30, fig. 21. 1960. Shrub 0.2–3 m tall; corolla tubular, to 2.5 cm long. Lower to upper montane thickets, commonly along streams or on sandstone, 300–2600 m; widespread in Bolívar and Amazonas. Guyana. ◆Fig. 20. A collection from Sierra de la Neblina, Nee 31149, has the tubular, red corollas typical of Raveniopsis ruellioides, but has much larger leaves and a spreading, bristly pubescence. Raveniopsis sericea R.S. Cowan, Mem. New York Bot. Gard. 10(2): 30, fig. 12. 1960. Shrub to 2 m tall with elongate, wiry branches. Locally common in low shrubby woods, often on rocks on tepui summit, 1800–2500 m; Bolívar (Macizo del Chimantá). Guyana. ◆Fig. 21. Raveniopsis stelligera (R.S. Cowan) R.S. Cowan, Mem. New York Bot. Gard. 10(2): 27, fig. 20a–c. 1960. —Galipea stelligera R.S. Cowan, Mem. New York Bot. Gard. 8: 121. 1953. Raveniopsis stelligera var. plicata R.S. Cowan, Mem. New York Bot. Gard. 10(2): 29, fig. 20d–g. 1960. Raveniopsis liesneri Steyerm., Ann. Missouri Bot. Gard. 75: 317, fig. 4. 1988.

30

R UTACEAE

Fig. 19. Raveniopsis peduncularis

Fig. 20. Raveniopsis ruellioides

Fig. 22. Raveniopsis tomentosa

Fig. 21. Raveniopsis sericea

Fig. 23. Raveniopsis linearis Fig. 24. Raveniopsis stelligera

Shrub or treelet 1–5 m tall; corollas tubular, red-orange. Montane forests, savannas, among rocks on slopes, often along streams, 500–1500 m; Amazonas (Cerro Duida, Cerro Yutajé, Sierra de la Neblina). Endemic. ◆Fig. 24. Raveniopsis steyermarkii R.S. Cowan, Mem. New York Bot. Gard. 14(3): 12. 1967. Shrub to 2.5 m tall with axillary, many-

flowered spikes that persist and become much elongated. Tepui scrub, 1700–1900 m; Bolívar (Auyán-tepui). Endemic. Raveniopsis tomentosa Gleason, Brittonia 3: 167. 1939. Shrub, sometimes monopodial, to 3 m tall, with very dense, dendritic pubescence. Locally common in tepui scrub on sandstone, 2100–2300 m; Bolívar (Auyán-tepui). Endemic. ◆Fig. 22.

Rutaneblina

Raveniopsis trifoliolata R.S. Cowan, Mem. New York Bot. Gard. 10(2): 25, fig. 19. 1960.

31

Shrub 1–2 m tall; flowers red. Locally abundant, tepui slope scrub, ca. 1100 m; Amazonas (Sierra de la Neblina). Endemic.

17. RUTANEBLINA Steyerm. & Luteyn, Ann. Missouri Bot. Gard. 71: 314. 1984. Shrubs 0.3–1.5 m tall. Leaves alternate, often crowded, simple, subsessile; blades obovate, broadly rounded, truncate, or emarginate at apex, entire or scarcely crenulate, leathery when dry. Inflorescence terminal, a ± sessile, condensed dichasium, often appearing subcapitate and not surpassing the leaves, or with primary branches to 2.5 cm long. Flowers actinomorphic. Sepals 5, free, quincuncial, slightly unequal, triangular, green (sometimes with pink margins) or dull lavender, persistent in fruit; petals 5, free, contorted in bud, 3.5 mm long, lanceolate, carinate, acute, white to pink, glabrous. Androecium of 5 fertile stamens; filaments free; anthers deltoid-ovoid, apiculate, basifixed. Disk obsolete. Ovary of 5 free, sessile carpels united only ventrally in the style; ovules 2 per carpel; style 1; stigma capitate, 5-lobed. Fruit of 5 free mericarps, occasionally fewer through abortion, each dehiscent ventrally. Seed 1 per carpel, tear-shaped, beaked at apex, smooth, shiny, glabrous. Endemic to the Guyana Shield in southern Venezuela; 1 species. Rutaneblina pusilla Steyerm. & Luteyn, Ann. Missouri Bot. Gard. 71: 314, fig. 6. 1984. Low shrub with curving branches; leaves often clustered at tips of branches, leaf surface venation densely reticulate. Along streams in tepui meadows with Bonnetia maguireorum, 1600–1900 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 25.

Fig. 25. Rutaneblina pusilla

32

R UTACEAE

18. SPATHELIA L., Sp. Pl. ed. 2, 386. 1762, nom. cons. Spathe P. Browne, Civ. Nat. Hist. Jamaica 187. 1756, nom. rejic. Sohnreyia K. Krause, Notizbl. Königl. Bot. Gart. Berlin 6: 147. 1914. Diomma Engl. ex Harms in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 19a: 460. 1931. Trees to 12 m tall, monocarpic, unbranched with leaves congested at apex. Leaves alternate, pinnate, 2–100-jugate, petiolate; leaflets opposite to alternate, crenate or entire, papery to somewhat leathery and often brittle when dry. Inflorescence terminal or rarely axillary (Huber 13063, NY), a panicle to 3 m long. Flowers actinomorphic, bisexual and staminate on same plant. Sepals 5, free or slightly connate at base, valvate or imbricate, elliptic, sometimes glandular at apex, ultimately deciduous from fruit; petals 5, free, quincuncial, to 8 mm, elliptic, rounded and sometimes glandular at apex, white. Androecium of 5 stamens, these slightly larger in staminate than in bisexual flowers; filaments free, winged or not, the wing when present bidentate or entire, in some free at apex from filament, in some connate slightly at base; anthers elliptic, bifid, dorsifixed. Disk subtending ovary. Ovary of 2 or 3 connate carpels, laterally compressed when 2; ovule 1 per carpel, appearing apical; style 0 or 1 and very short; stigma 2- or 3-lobed, deciduous. Fruit a 2- or 3-winged samara, wings narrower to broader than seedbearing portion. Seeds 1 per locule, subglobose to ellipsoid, smooth, dull, glabrous. West Indies, Venezuela, Peru, northern Brazil; ca. 17 species, 1 in Venezuela. Spathelia ulei (Engl. ex Harms) R.S. Cowan & Brizicky, Mem. New York Bot. Gard. 10(2): 62. 1960. —Diomma ulei Engl. ex Harms in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 19a: 460. 1931. —Marua-yek. Diomma fruticosa Steyerm., Fieldiana, Bot. 28: 272. 1952. —Spathelia fruticosa (Steyerm.) R.S. Cowan & Brizicky, Mem. New York Bot. Gard. 10(2): 61. 1960. Spathelia chimantaensis R.S. Cowan & Brizicky, Mem. New York Bot. Gard. 10(2): 63, fig. 1g–i. 1960. Spathelia neblinaensis R.S. Cowan & Brizicky, Mem. New York Bot. Gard. 10(2): 63, fig. 1e–f. 1960. Spathelia jauaensis R.S. Cowan, Mem. New York Bot. Gard. 23: 863. 1972. Monopodial palm-like treelet to 12 m tall; flowers fragrant, in large panicles; leaves pinnate, clustered at apex; samaras often corky and reddish at maturity. Dense or open, shrubby, forested, or savanna habitats, often along watercourses, on sandstone, in Bonnetia thickets, sometimes in ecotone between riparian forests and savannas, 100–2500 m; Bolívar (Gran Sabana, Macizo del Chimantá to Cerro Jaua and Cerro Guanay and Sierra de Mai-

Fig. 26. Spathelia ulei

Spiranthera 33

gualida, Roraima-tepui), Amazonas (Cerro Aracamuni, Cerro Sipapo, Cerro Yutajé, Sierra de la Neblina, Sierra de Maigualida). Endemic. ◆Fig. 26. Five species of Spathelia have been described from the flora area. These were separated on differences in the distribution of punctae on the leaflets and of pubescence on the fruits, the size of fruits, the outline of fruit wings and ovaries, the size of the leaflets, and the length of the rachises, but consistent differences cannot be discerned when all collections are examined. Two collections (Huber 12719, NY, and Huber 13063, NY) from Sierra de Maigualida

(Amazonas-Bolívar border) differ from the other collections by the abundant ferruginous indument on the leaves and fruit and by the fewer (2–5 versus 7–24) pairs of rather rugose leaflets. According to the collector, the Maigualida plants are shorter than the others and grow in colonies rather than as scattered individuals. These populations may deserve taxonomic recognition, but until the genus is better known, Huber’s collections are being included as an extreme of S. ulei. Plants in an area flower simultaneously and flowers are pollinated by butterflies and bees.

19. SPIRANTHERA A. St.-Hil., Bull. Sci. Soc. Philom. Paris 1823: 130. 1823. Shrubs or trees to 18 m tall. Leaves alternate, 3-foliolate, petiolate; blades entire, papery to somewhat leathery when dry, often pale beneath. Inflorescence terminal, a pedunculate thyrse. Flowers actinomorphic. Calyx cupular, 5-lobed, deciduous, the lobes triangular, acute; petals 5, free, usually quincuncial, less often imbricate, 12–60 mm long, narrowly oblanceolate, white or cream, densely appressed-pubescent. Androecium of 5 fertile stamens; filaments free, filiform; anthers lorate, slightly bifid at base, basifixed, recurved or coiled after anthesis. Disk fleshy or membranous, cupular or cylindric, embracing the gynophore, entire

Fig. 27. Spiranthera guianensis

34

R UTACEAE

or unequally laciniate, the lobes often variously pubescent. Ovary of 5 carpels connate ventrally at base and 5-lobed above, borne on a gynophore; ovules 2 per carpel; style 1, elongate; stigma clavate to slightly capitate, slightly 5-lobed. Fruit of 1 or 2(–5) free mericarps, each beaked, ventrally dehiscent. Seeds 1 per carpel, ellipsoid, smooth, dull, glabrous. Venezuela, Guyana, Peru, Brazil, Bolivia; 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Spiranthera 1.

1.

Calyx 2.5–3 mm long; petals 5.3–6 cm long; combined length of ovary and gynophore 6.5–9 mm; terminal leaflet narrowly obovate, 10– 21 cm long .............................................................................. S. guianensis Calyx 0.8–1.2 mm long; petals 1.2–1.5 cm long; combined length of ovary and gynophore 1.7–3.5 mm; terminal leaflet elliptic, narrowly elliptic, or narrowly obovate, 14–27 cm long ...................................... S. parviflora

Spiranthera guianensis Sandwith, Bull. Misc. Inform. 1928: 368. 1928. —Quina amarilla, Quina negra. Tree to 18 m tall; flowers fragrant. Lower montane forests, ca. 600 m; eastern Bolívar. Guyana, Brazil (northern Amazonas). ◆Fig. 27.

Spiranthera parviflora Sandwith, Bull. Misc. Inform. 1939: 548. 1940. Shrub or tree to 15 m tall. Seasonally flooded forests along black-water rivers, upland forests, 100–900 m; Bolívar (near Icabarú), Amazonas (Río Guainía, upper Río Negro, Río Temi). Guyana, Peru, Brazil (Pará, Maranhão).

20. ZANTHOXYLUM L., Sp. Pl. 270. 1753. Fagara L., Syst. Nat. ed. 10, 2: 897, 1362. 1759. Shrubs or trees often with spines or prickles on trunk, branches, petioles, leaf rachises, and peduncles. Leaves alternate, pinnate or 1- or 3-foliolate, sometimes with winged rachises, petiolate; leaflets opposite to alternate, entire or crenate, with glands between the teeth, papery to leathery when dry. Inflorescence terminal, axillary, or otherwise lateral, usually a panicle, sometimes a raceme. Flowers actinomorphic, unisexual (plants dioecious). Sepals 3–5, free, quincuncial, persistent in fruit; petals 3–5, free, quincuncial or imbricate, to ca. 5 mm long, often with a gland at apex, white or yellowish. Androecium in staminate flowers of 3–5 stamens; filaments free, awl-shaped to filiform; anthers ovate or oblong, bifid at base, connective often with apical gland, in pistillate flowers absent or much reduced. Disk subtending ovary or rudimentary ovary, often inconspicuous. Ovary in pistillate flowers of 1–5 free or connate, sessile or stipitate carpels; ovules 2 per carpel; styles free or connate, shorter than ovary; stigmas free or connate, capitate, peltate, or 2–5-lobed; ovary in staminate flowers reduced usually to a ± conical projection, subtended by the disk and tapered to the apex, without stigma. Fruit of 1–5 free or connate, sessile or stipitate mericarps, each ventrally dehiscent. Seeds 1 per carpel, subglobose, black, smooth, shiny, glabrous. Tropical and temperate regions of the Old and New Worlds; ca. 225 species, ca. 23 in Venezuela, 14 of these in the flora area. Engler (Nat. Pflanzenfam. ed. 2, 19a. 1931) presented abbreviated keys to groups of species, but a monograph of the genus has not been published. Names could not be assigned to 2 of the taxa represented by fertile collections from the

Zanthoxylum 35

flora area. Pending further study, they have been assigned letters and included in the key. The genus is also represented in the flora area by 4 sterile specimens, which remain unnamed and are not included in the key. Gentry & Stein 46748 (MO) and 46749 (MO, NY) are from the Sierra de la Neblina base camp on the Río Mawarinuma, Amazonas state. Two other undetermined collections, Williams 11752 (VEN) and Steyermark 86419 (VEN), are from Bolívar.

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

2.

3(1).

3.

4(3).

4.

5(4).

5.

6(5).

6.

Pubescence of branched trichomes ........................................................... 2 Pubescence absent or of simple trichomes ............................................... 3 Leaflets in (3–)7–10 pairs, the largest ones 8.2–14.3 × 3.8–6 cm, when mature paler below with persistent, dense, stellate, echinoid, and dendroid trichomes; margin indistinctly crenulate, appearing ± entire to naked eye .................................................................................... Z. ekmanii Leaflets in 3–17 pairs, the largest ones 4.5–10.7 × 0.9–2.3 cm, when mature with scattered, minute, stellate trichomes; margin distinctly crenate to naked eye .............................................................. Z. rhoifolium Leaf rachis distinctly winged; spines recurved at nodes of ± zigzag stem; terminal leaflet 0.8–2.8 × 0.4–1.7 cm; fruit a solitary free, stipitate mericarp ........................................................................................ Z. fagara Leaf rachis slightly margined to terete; spines when present straight or not confined to nodes; terminal leaflet or one of upper pair 2–14 × 1– 6 cm; fruit (unknown in sp. B) of 1–5 free, sessile or stipitate mericarps or a 4–6-parted capsule ....................................................... 4 Fruit a 4–6-parted, echinate capsule; leaves odd-pinnate; leaflets in 2 or 3 pairs, lowermost ones 0.6–2.9 × 0.5–2 cm, usually < 1/2 as long as terminal leaflet ...................................................................... Z. syncarpum Fruit of 1–6 free, non-echinate mericarps; leaves odd- or even-pinnate; leaflets in 2–8 pairs, but if 2 or 3 then lowermost ones 2.6–12.5 × 1.3– 5.5 cm and > 1/2 as long as terminal or one of uppermost pair ........... 5 Pubescence of shoot, leaf rachis, and inflorescence axes usually fulvous; fruiting mericarps 4–6, pubescent, sessile; leaflets pubescent on both sides with ascending to erect trichomes, glabrescent on upper surface except for the midrib .......................................................... Z. martinicense Pubescence of shoot, rachis, and inflorescence axes if present not fulvous; fruiting mericarps 1–5, glabrous, sessile or stipitate; leaflets glabrous (sometimes with minute trichomes on lower surface of midrib visible at 30×) ............................................................................ 6 Fruiting mericarps 1–5, stipitate; leaflets usually distinctly and regularly crenate to the naked eye, glabrous even on midrib below, often pale-verrucose; inflorescence axes usually transversely lenticellate, glabrous ................................................................... Z. caribaeum Fruiting mericarps 1–3, sessile (unknown in sp. B); leaflets entire to crenate, glabrous except in some cases with minute trichomes on lower surface of midrib (visible at 30×), not pale-verrucose; inflorescence axes not lenticellate, ± pubescent .................................. 7

36

R UTACEAE

7(6). 7.

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

9.

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

11.

12(11).

12. 13(12).

13.

Inflorescence racemose; leaflets in 2 pairs, subcrenate, obtuse to rounded to retuse at apex, obovate, 1.5–6 cm long ....................................... Z. sp. B Inflorescence paniculate; leaflets in 2–7 pairs, crenate to entire, acuminate at apex, elliptic, oblanceolate, or obovate, 5–16 cm long ................................................................................................................ 8 Sepals 3; carpel 1; leaflets with easily detachable, lepidote scales (= glands or hardened exudate?) on both surfaces ............ Z. juniperinum Sepals 5; carpels 3 (1–3 maturing); leaflets without detachable lepidote scales ...................................................................................................... 9 Leaflets in 7 pairs, obviously crenate especially in upper 1/2, obovate, broadly rounded to truncate with obtuse apiculum, leathery when dry .......................................................................................................... Z. sp. A Leaflets in (1)2–7 pairs, entire or subcrenate, elliptic, narrow-ovate, or oblanceolate, acuminate, obtuse, or rounded and apiculate, leathery or papery when dry ............................................................................. 10 Leaflets rigid-leathery when dry, shiny on both surfaces, the upper surface gray-green when dry, the lower one brown ............. Z. amapaense Leaflets papery when dry, shiny or not, not colored as above when dry .............................................................................................................. 11 Inflorescences narrow panicles, in axils of leaves; carpel and developing mericarp 1 per flower; leaflets 1–4 pairs, acuminate; rachis and lower surface of midrib glabrous ............................................................ Z. huberi Inflorescences broader panicles, terminal; carpels 3 per flower (unknown in Z. compactum) and developing mericarps 1–3 per flower; leaflets in 3–7 pairs, acuminate, obtuse, or rounded and with a blunt apiculum at apex; rachis and lower surface of midrib glabrous or with minute, erect trichomes .................................................................................... 12 Sepals of staminate flowers connate at base, with lobes not obviously imbricate at base, 1–1.1 mm long; anthers 1.2–1.4 mm long ........... ............................................................................................... Z. compactum Sepals of staminate flowers imbricate at base, 0.5–0.6 mm long; anthers 0.4–0.5 mm long .................................................................................. 13 Leaflets shiny on upper surface, seemingly entire, acuminate, the acumen 8–14 mm long; leaf rachis and lower surface of midrib glabrous, the rachis flattened or grooved and often narrowly margined above; carpels 3 per flower, 1–3 developing ....... Z. pentandrum Leaflets dull on upper surface, subcrenate, rounded with blunt apiculum to 3 mm long; leaf rachis and lower surface of midrib with minute, erect trichomes, the rachis subterete; carpels 3 per flower, only 1 developing ................................................................................. Z. apiculatum

Zanthoxylum amapaense (Albuq.) P.G. Waterman, Taxon 24: 363. 1975. —Fagara amapaensis Albuq., Publ. Inst. Nac. Pesq. Amaz. Bot. 27: 11. 1968. —Bocsúo negro, Bosú, Bosuo, Mapurite. Tree to 25 m tall with spines to 2 mm long on branches and on peduncle of terminal inflorescence; leaflets rigid-leathery when

dry, 6–12.2 × 3–5 cm (24 × 10 in sterile shoot); sepals 5; carpels 3(4), of which only 1 develops; mature mericarp 8–9 mm long, rather carinate along line of dehiscence. Evergreen lowland to lower montane forests, 200–700 m; Delta Amacuro (Serranía de Imataca), northeastern Bolívar. French Guiana.

Zanthoxylum 37

Zanthoxylum apiculatum (Sandwith) P.G. Waterman, Taxon 24: 363. 1975. —Fagara apiculata Sandwith, Bull. Misc. Inform. 1935: 120. 1935. —Bosí, Erizo. Tree to 14 m tall with spines to 10 mm long on branches and to 3 mm on rachises. Evergreen lowland to lower montane forests, 100–600 m; Bolívar (near El Dorado, Río Suapure and Amaruay-tepui). Guyana. Zanthoxylum caribaeum Lam., Encycl. 2: 39. 1786. —Fagara caribaea (Lam.) Krug & Urb. in Urb., Bot. Jahrb. Syst. 21: 562. 1896. —Mapurite morado, Mapurito. Tree to 10 m tall with spiny trunk. Semideciduous forests bordering savannas, 50–400 m; Bolívar (Altiplanicie de Nuria, islands in Lago Guri, lower Río Caroni from La Paragua to Puerto Ordaz). Scattered in northern Venezuela; Mexico, Central America, West Indies, Colombia, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 28. Zanthoxylum compactum (Huber ex Albuq.) P.G. Waterman, Taxon 24: 363. 1975. —Fagara compacta Huber ex Albuq., Publ. Inst. Nac. Pesq. Amazonia Bot. 28: 12, t. 11. 1968. —Bosúa. Tree to 6 m tall with short broad spines on trunk; leaflets alternate, 8–14 × 3.5–5.1 cm; inflorescence terminal with globose buds; petals of staminate flowers ca. 2 mm long; anthers drying red-brown. Habitat not indicated, ca. 50 m; Delta Amacuro (Caño Jota-Sabuca). Zanthoxylum ekmanii (Urb.) Alain, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 9: 24. 1950. —Fagara ekmanii Urb., Repert. Spec. Nov. Regni Veg. 20: 302. 1924. —Bocsuo amarillo, Bojúa, Boxúo amarillo, Samán montañero. Tree to 30 m tall with spiny trunk; flowers yellow, fragrant. Evergreen lowland forests, 50–300 m; Delta Amacuro (near Río Toro), northeastern Bolívar (El Dorado to Upata). Southern Mexico, Central America, West Indies, Suriname, French Guiana, Ecuador, Peru, Brazil.

Zanthoxylum fagara (L.) Sarg., Gard. & Forest 3: 186. 1890. —Schinus fagara L., Sp. Pl. 389. 1753. —Jalapatrás, Uña de gavilán. Fagara pterota L., Syst. Nat. ed. 10: 897. 1759. —Zanthoxylum pterota (L.) Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 3. 1823. Spiny shrub or tree to 5 m tall. Savannas and thickets, 50–200 m; northeastern Bolívar (Puerto Ordaz to El Manteco). Scattered through rest of Venezuela; widespread in Neotropics to Argentina. ◆Fig. 31. Zanthoxylum huberi P.G. Waterman, Taxon 24: 366. 1975. Tree 10–22 m tall. Evergreen lowland to lower montane forests, 200–1000 m; Bolívar (55 km south of Jabillal on Río Caura, Macizo del Chimantá [Toronótepui]). Brazil. Zanthoxylum juniperinum Poepp. in Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 77. 1845. —Naranjuelo, Paneco. Fagara ocumarensis Pittier, Bol. Minist. R.R. E.E. 8–12: pp. 276. 1929. (Arb. Arbust. Venez. 9/19: 120. 1929.) —Zanthoxylum ocumarense (Pittier) Steyerm., Fieldiana, Bot. 28: 273. 1952. Tree to 18 m tall with spiny trunk. Evergreen lowland to lower montane forests, 200–500 m; Delta Amacuro (Serranía de Imataca). Aragua, Carabobo, Lara, Mérida, Monagas, Sucre; Mexico, Central America, Colombia, Suriname, Ecuador, Peru, Brazil. Zanthoxylum martinicense (Lam.) DC., Prodr. 1: 726. 1824. —Fagara martinicensis Lam., Encycl. suppl. 2: 627. 1812. —Bosú, Bosúo, Mapurite blanca, Mapurito. Tree to 40 m tall with large spines on trunk. Semideciduous forests, edges of savannas, 100–400 m; Delta Amacuro (Sacupana), Bolívar (near La Paragua and El Palmar). Falcón, Yaracuy; West Indies. Zanthoxylum pentandrum (Aubl.) R.A. Howard, J. Arnold Arbor. 64: 269. 1983. —Fagara pentandra Aubl., Hist. Pl.

38

R UTACEAE

Fig. 28. Zanthoxylum caribaeum

Fig. 29. Zanthoxylum syncarpum Fig. 30. Zanthoxylum sp. B

S A B I A C E A E 39

Guiane 78, t. 30. 1775. —Boxúo negro, Mapurito amarillo. Tree to 20 m tall with spiny trunk. Evergreen lowland forests, 100–400 m; Delta Amacuro (near El Palmar), Bolívar (Altiplanicie de Nuria). Miranda; West Indies, Trinidad, Guyana, Suriname, French Guiana, Brazil. Zanthoxylum rhoifolium Lam., Encycl. 2: 39. 1786. —Fagara rhoifolia (Lam.) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 118. 1896. —Bosnó, Mapurite, Mapurite blanco. Tree to 20 m tall with spiny trunk. Deciduous to semideciduous lowland and lower montane forests and disturbed evergreen lowland forests, 100–400 m; eastern Bolívar, Amazonas (Caño Jayuwapuey a tributary of the Río Ocamo). Distrito Federal, Lara, Mérida, Zulia; widespread throughout Neotropics. Zanthoxylum syncarpum Tul., Ann. Sci. Nat. Bot. sér. 3, 7: 279. 1847. —Fagara syncarpa (Tul.) Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(4): 116. 1896. —Bosúa, Mapurite. Tree to 10 m tall with spiny trunk. Dry areas, savannas, near sea level to 200 m; Bolívar (Caicara, El Palmar, Puerto Ordaz). Scattered in northern Venezuela. ◆Fig. 29.

Fig. 31. Zanthoxylum fagara

Zanthoxylum sp. A. —Yachiyek (Are-kuna). Tree 13 m tall with spines to 8 mm long on branches and to 5 mm on peduncle; leaflets 3.8–6 × 6.6–12.5 cm. Montane forests, ca. 1000 m; Bolívar (Gran Sabana). Zanthoxylum sp. B Spiny tree. Habitat not indicated, 100– 200 m; Amazonas (Salto Yureba on tributary of lower Río Ventuari). ◆Fig. 30.

SABIACEAE by Gerardo A. Aymard C. and Nidia L. Cuello A. Trees, shrubs, or rarely lianas. Leaves alternate to subopposite, simple or pinnately compound, exstipulate, leaflets sometimes heteromorphic; base of petioles often swollen and ligneous (Meliosma); petiolules often pulvinate. Inflorescences terminal or axillary cymes or elongate panicles (Ophiocaryon), ramiflorous or cauliflorous. Flowers small, usually bisexual. Sepals 3–5, unequal, imbricate; petals (4)5, opposite the sepals, unequal, imbricate. Stamens 5, opposite or sometimes alternate with the petals, all functional (Sabia) or else the outer 3 reduced to staminodes and the inner 2 functional (Meliosma, Ophiocaryon); filaments or connective forming a swollen or collar-like extension; anther opening transversely or by valves. Ovary superior, ovoid to conical, 2(3)-carpellate and -locular; ovules 1–3 per locule, placentation axile; styles simple or bifid, usually

40

S ABIACEAE

connate, stigmas or stigmatic surfaces 1 or 2; nectary disk usually present, annular, surrounding the ovary. Fruits drupaceous or dry and indehiscent, 1locular, the endocarp bony or crustaceous. Seeds 1 per fruit, with a large embryo; cotyledons folded or coiled (Ophiocaryon); endosperm reduced or absent. Pantropics (most diverse in the tropics and subtropics of southeastern Asia); 3 genera and ca. 80 species, 2 genera and 7 species in the flora area. Key to the Genera of Sabiaceae 1. 1.

Leaves simple; stigma 1; anthers transversely dehiscent ......... 1. Meliosma Leaves pinnately compound; stigmas 2; anthers with valvate dehiscence ............................................................................................. 2. Ophiocaryon

1. MELIOSMA Blume, Catalogus 10. 1823. Trees or shrubs. Leaves alternate, rarely subopposite, frequently clustered at the twig apices, simple (Venezuelan species) or compound, entire or frequently dentate; petiole usually thickened at base. Inflorescence terminal to cauliflorous or ramiflorous, paniculate, usually pyramidal; bracts reduced, inconspicuous. Flowers small, zygomorphic. Sepals 4 or 5; petals 4 or 5, the outer ones larger and usually ± suborbicular, the inner ones much-reduced, thin and strap-shaped, opposite the fertile stamens and ± fused to the filament bases. Stamens 5, 3 sterile and 2 fertile; filaments short, flat, incurved at tip; anthers bilocular, the thecae short and thick, transversely dehiscent, often separated by the thickened connective; staminodes 3, opposite and ± fused with the bases of the larger petals, sometimes forming a cap over the pistil. Ovary 2(3)-locular, with 2 ± superimposed ovules per locule; style minute to elongate; stigma 1, minute; disk usually present, 5-toothed. Fruit a globose to obovoid drupe, with a single large seed. Pantropics (most diverse in the Neotropics); 50 species, 10 in Venezuela, 4 of these in the flora area. Key to the Species of Meliosma 1. 1. 2(1).

2. 3(2).

3.

Branchlets with dense, ferruginous, appressed trichomes; leaves pilose with gland-tipped trichomes.. .................................................... M. gentryi Branchlets glabrous, with strigose trichomes or finely puberulous; leaves glabrous or puberulous along veins on lower surface .............. 2 Leaves 7–15 cm wide, puberulous along veins on lower surface, the base cuneate; petioles 3.5–7 cm long; inflorescence 20–40 cm long .................................................................................................. M. vasquezii Leaves 2–8 cm wide, glabrous on both surfaces, the base acute, obtuse or subacute; petioles 0.5–3 cm long; inflorescence 7–12 cm long ........... 3 Leaves subcoriaceous to coriaceous, the apex acute or acuminate, the base obtuse or subacute; petioles 0.5–3 cm long; floral bracts ovate or lanceolate-oblong, subacute, densely pubescent; flowers sessile ................................................................................................... M. herbertii Leaves chartaceous to subcoriaceous, the apex rounded or obtuse, the base acute; petioles 0.5–1.7 cm long; floral bracts suborbicular, obtuse, glabrescent; flowers pedicellate ........................ M. aff. meridensis

Meliosma 41

Meliosma gentryi Aymard & Cuello, Biollania 10: 1. 1994. Tree to 10 m tall; fruits green. Moist montane forests, ca. 1100 m; Amazonas (eastern side of Cerro Cuao at Caño Piedra ca. 75 km southeast of Puerto Ayacucho). Endemic.

The only specimen (A. Fernández 5479, MO, PORT) of this taxon from the flora area is in fruit. It is closely related to Meliosma meridensis from the Andes and the Coastal Cordillera. Additional collections are needed to establish whether it represents a new species instead of a geographic disjunction.

Meliosma herbertii Rolfe, Kew Bull. 81: 244. 1893. West Indies, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; 2 varieties, both in the flora area.

Meliosma vasquezii Gentry, Ann. Missouri Bot. Gard. 73: 823. 1986. —Pi’tzejécute (Bale) Tree 5–10 m tall; moist nonflooded forests, 100–200 m; Amazonas (middle Río Casiquiare). Amazonian Colombia, Ecuador, Peru, Bolivia.

Key to the Varieties of M. herbertii 1. Inner minor petals bicuspidate at the apex; petioles 1–3 cm long, glabrous ....................................... var. herbertii 1. Inner minor petals entire at the apex; petioles 0.5–1 cm long, sparsely strigose ..................................... var. tepuiensis M. herbertii var. herbertii. —Caruache, Töpishécute (Bale). Tree 10–15 m tall; fruit purple-black. Evergreen lowland and lower montane forests, 100–400 m; Delta Amacuro (northeast of El Palmar, Río Toro), Bolívar (Altiplanicie de Nuria), Amazonas (middle Río Casiquiare). Anzoátegui, Monagas, Sucre; West Indies, Guyana, Colombia, Ecuador, Peru, Brazil (Amazonas, Rondônia), Bolivia. ◆Fig. 32. M. herbertii var. tepuiensis (Steyerm. & Maguire) Steyerm. & Gentry, Fl. Venez. 5(1): 210. 1992. —Meliosma tepuiensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 448. 1967. Tree to 15 m tall. Moist montane forests, 1900–2000 m; Bolívar (Macizo del Chimantá [southeast of Apacará-tepui]). Endemic. Meliosma aff. meridensis Lasser, Bol. Soc. Venez. Ci. Nat. 8(55): 233. 1943. Meliosma uberrima Idrobo & Cuatrec., Caldasia 7(33): 199. 1955. Meliosma nubicola Steyerm. & Lasser, Bol. Soc. Venez. Ci. Nat. 26: 463. 1966. Tree to 30 m tall; fruits green. Moist montane forests, 1200–1300 m; Bolívar (4 km north of Cerro Impacto, Sierra de Maigualida). Aragua, Distrito Federal, Lara, Mérida, Trujillo; Colombia (Cundinamarca).

Fig. 32. Meliosma herbertii var. herbertii

42

S ABIACEAE

2. OPHIOCARYON R. Schomb. ex Endl., Gen. Pl. suppl. 1: 1425. 1841. Phoxanthus Benth., Trans. Linn. Soc. London 22: 127. 1857. Shrubs to small trees to 20 m tall. Leaves alternate, compound, heteromorphic, even-pinnate or odd-pinnate, sometimes trifoliolate or simple on flowering branches; leaflets entire, opposite, subopposite, or alternate, subchartaceous, coriaceous, to rigid-coriaceous, elliptic, ovate, or oblong-oblanceolate. Inflorescence axillary or terminal, elongate-paniculate, 15–170 cm long; bracts present. Flowers small, actinomorphic, bisexual. Sepals 4 or 5, unequal; petals 4 or 5, unequal, white to reddish or pink, the outer ones ovate-obtuse to triangularacuminate or caudate, the inner ones usually linear or oblong. Stamens 2 or 3, opposite the petals; staminodes 2 or 3; anthers valvately dehiscent. Ovary 2carpellate and -locular, globose, with 1 or 2 ovules per locule; styles 2, minute; stigmas 2; disk present, entire or 5-toothed. Fruit a globose drupe. Seed 1, without endosperm; embryo spirally twisted or coiled. Colombia, Venezuela, Guyana, French Guiana(?), Ecuador, Peru, Brazil; 9 species, 3 in Venezuela, all in the flora area. Key to the Species of Ophiocaryon 1. 1. 2(1). 2.

Leaflets thin-textured; petals white to reddish, apically acuminate to caudate ................................................................................ O. manausense Leaflets thick-chartaceous or rigid-coriaceous; petals pink, apically obtuse ..................................................................................................... 2 Leaves 35–55 cm long; leaflets elliptic, thickly chartaceous, 6–24 × 2.5– 7 cm; margins of sepals eciliate ................................................... O. duckei Leaves 70–80 cm long; leaflets elliptic to elliptic-ovate, rigid-coriaceous, 28–30 × 8–15 cm; margins of sepals ciliate ................................... O. sp. A

Ophiocaryon duckei Barneby, Mem. New York Bot. Gard. 23: 115. 1972. Small tree to 10 m tall; corolla pink. Expected in upper Río Negro area of Estado Amazonas. Ophiocaryon duckei is known presently only from seasonally flooded black-water (igapó) forests at low elevations in northern Amazonas, Brazil. Ophiocaryon manausense (W.A. Rodrigues) Barneby, Mem. New York Bot. Gard. 23: 118. 1972. —Phoxanthus manausensis W.A. Rodrigues, Publ. Inst.

Nac. Pesq. Amazônia Bot. 17: 3–7. 1964. Tree 5–10 m tall. Evergreen lowland nonflooded forests, 100–200 m; Amazonas (lower Río Casiquiare). Amazonian Colombia, Ecuador, Peru, and Brazil. Ophiocaryon sp. A Tree 10–15 m tall; petals pink. Montane forests, 1200–1300 m; expected in Amazonas. Brazil (Amazonas: upper Río Matucará, Serra Pirapucú, southwest of Sierra de la Neblina). ◆Fig. 33. This taxon is based on Silva & Brazão 60883 (MO, NY).

Ophiocaryon 43

1 mm

3 cm 1 mm 10 cm Fig. 33. Ophiocaryon sp. A

44

S ANTALACEAE

SANTALACEAE by James S. Miller Small trees, shrubs, or perennial or annual herbs, hemiparisitic on roots of other plants or rarely epiphytic and hemiparasitic on branches of trees (Dendrotrophe), green and photosynthetic, the stems occasionally thorny. Leaves opposite or occasionally alternate, simple, entire, exstipulate, sometimes reduced to scales. Inflorescences terminal or axillary, racemose, spicate, paniculate, dichasial, capitate, or the flowers solitary and axillary. Flowers bisexual or unisexual, the plants then monoecious or dioecious, actinomorphic, epigynous or less often perigynous or hypogynous, the perianth monochlamydeous. Tepals (3)4 or 5(–8), connate in a tube with the apices free, often green and fleshy, basally adnate to the ovary. Stamens (3)4 or 5(–8), opposite the tepals; filaments usually basally adnate to the tepals; anthers bithecal, dehiscing longitudinally or rarely by a single apical pore; pollen 3-aperturate. Ovary inferior or half-inferior, usually surrounded by an intrastaminal disk, (2)3(–5) carpellate, unilocular, the placentation free central; style terminal; stigma capitate; ovules 1–4, pendulous from the free central placenta, anatropous or less commonly hemitropous. Fruits indehiscent, nut-like or drupaceous; seeds lacking testa, with abundant, carnose, oily, or starchy endosperm. Nearly cosmopolitan (mainly Old World); 30–35 genera and ca. 400 species, 1 species in the flora area. 1. THESIUM L., Sp. Pl. 207. 1753. Austroamericum Hendrych, Bol. Soc. Argentina Bot. 10: 126. 1963. Perennial or annual herbs or subshrubs, hemiparasitic on roots of other plants. Leaves usually alternate, linear, lanceolate, or reduced and scale-like, only rarely with an expanded blade, exstipulate. Inflorescences terminal or axillary, racemose, spicate, paniculate, cymose, or capitate. Flowers bisexual, actinomorphic, epigynous or perigynous, monochlamydeous. Tepals (4)5, connate in a tube with the apices free, the lobes often pubescent on the inner surface. Stamens (4)5; filaments basally adnate to the perianth tube; anthers bithecal, dehiscing longitudinally; staminal disk usually indistinct. Ovary unilocular with 2–4 ovules pendent from the free central placenta; style terminal, simple, or almost absent; stigma capitate or obscurely 3-lobed. Fruits dry, indehiscent, nut-like, often 10ribbed, the perianth often persistent; seeds with carnose endosperm. Neotropics, temperate or subtropical regions or mountains of tropical areas in the Old World; ca. 240–300 species, 1 in Venezuela. Thesium tepuiense Steyerm., Fieldiana, Bot. 28. 1: 225. 1951. —Austroamericum tepuiense (Steyerm.) Hendrych, Bol. Soc. Argentina Bot. 10: 128. 1963. Annual, root hemiparasite to 30 cm tall. Flowers greenish. Fruits yellow-green. Wet

and swampy savannas, moist depressions of dry savannas, dry, rocky savannas, moist banks of pools and creeks, 900–2200 m; Bolívar (Gran Sabana and summits of tepuis). Adjacent Guyana. ◆Fig. 34.

Thesium 45

Fig. 34. Thesium tepuiense

46

S APINDACEAE

SAPINDACEAE by Pedro Acevedo-Rodríguez and Hans T. Beck Trees, shrubs, or woody or herbaceous climbers. Stems of climbing genera usually with multiple vascular cylinders. Leaves alternate, pinnately or ternately compound, rarely opposite or simple; stipules minute to large, present only in climbing genera; distal leaflet in most arboreal and shrubby genera rudimentary. Inflorescence thyrses, racemes, or panicles. Flowers mainly 4- or 5-merous, actinomorphic or zygomorphic, bisexual or unisexual by reduction (plants functionally monoecious, or dioecious or polygamous). Calyx of (3)4 or 5(–7) equal or unequal, distinct or basally connate sepals; petals 4 or 5, distinct, ornamented by an adaxial appendage, usually white or light yellow, rarely absent, the appendages variously shaped, mostly petaloid, simple, bifurcate, or hood-shaped, basally adnate to the petal or just a prolongation of petal margins; disk extrastaminal, annular or unilateral, often modified into 2 or 4 glandular lobes. Androecium actinomorphic or zygomorphic; stamens (4–7)8(–10); filaments free or basally connate, equal or unequal in length; anthers dorsifixed or basifixed, introrse, opening by longitudinal slits. Gynoecium syncarpous, (2)3(–5)-carpellate, carpels with 1–many ovules. Fruits septifragal or loculicidal capsules, schizocarps with winged or rounded mericarps, or berries. Mostly pantropics or subtropics, extending also to subtemperate zones; ca. 140 genera and 1800 species, 16 genera and 83 species in the flora area. Key to the Genera of Sapindaceae by Pedro Acevedo-Rodríguez 1. 1. 2(1).

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

5(2).

5.

Climbing plants often bearing tendrils and usually producing milky sap ................................................................................................................ 2 Erect shrubs or trees; not producing milky sap ....................................... 7 Fruit (schizocarp) splitting into 3 winged, 1-seeded mericarps, the mericarps indehiscent (seeds not exposed), dispersed as individual units ....................................................................................................... 3 Fruit a septifragal capsule (exposing the seeds), variously shaped, sometimes bearing dorsal wings .......................................................... 5 Wing extending to both ends of mericarp, surrounding the locule; locule in central position ........................................................... 6. Houssayanthus Wing placed to either end of mericarp, not surrounding the locule ....... 4 Mericarp wing distal; flowers actinomorphic, disk annular; leaves trifoliolate; thyrse umbel-like ................................................ 13. Thinouia Mericarp wing proximal; flowers zygomorphic, disk unilateral, 2–4lobed; leaves trifoliolate or biternately compound; thyrse with elongate rachis, not umbel-like ............................................... 11. Serjania Capsule fleshy or woody, thick-walled, red, less often green or yellow; seed with a fleshy coat covering most of its surface; leaves variously compound, but usually pinnately 5-foliolate ........................... 9. Paullinia Capsule thin, papery, turning from light green to straw-colored; seeds with heart-shaped or reniform whitish arillode surrounding the hilum, leaves trifoliolate or biternate .................................................. 6

S A P I N D A C E A E 47

6(5).

6.

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

12.

13(11). 13. 14(13).

14.

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

Leaves trifoliolate; stems terete and lenticellate, slightly woody, usually producing milky sap; disk 4-lobed; capsules semi-inflated, the edges of the locules compressed around the septae margin into a wing-like projection ................................................................................... 15. Urvillea Leaves biternate; stems sulcate and herbaceous, not producing milky sap; disk 2-lobed (the lobes sometimes elongated); capsules completely inflated, without marginal wing ....................... 2. Cardiospermum Leaves simple ............................................................................................. 8 Leaves compound ....................................................................................... 9 Plant usually viscid; flowers apetalous; stamens 10; anthers nearly sessile; fruit a membranous capsule ....................................... 5. Dodonaea Plant not viscid; flowers with 4 petals; stamens not sessile; fruit of 1 or 2 fleshy monocarps ..................................... 1. Allophylus (A. amazonicus) Leaves trifoliolate, the terminal leaflet well developed ............ 1. Allophylus Leaves pinnately compound, the terminal leaflet rudimentary (sometimes slightly developed in Dilodendron) .......................................... 10 Leaves bipinnate, with numerous small (1–2 cm long) leaflets .............................................................................................. 4. Dilodendron Leaves once-pinnate, with larger (> 5 cm long) leaflets ........................ 11 Petiole and/or leaf rachis winged ............................................................ 12 Petiole and leaf rachis not winged .......................................................... 13 Inflorescence a panicle; flowers 4-merous, petals projecting beyond calyx, lacking inner appendages; style obsolete; stigma subcapitate; leaflets mostly opposite, chartaceous, elliptic, with undulate margins; fruit ellipsoid to globose, green at maturity; pericarp leathery; seeds covered with fleshy tan-colored pulp ..................................... 8. Melicoccus Inflorescence thyrse-like; flowers 5-merous, petals not projecting beyond the calyx, bearing a pair of reduced marginal appendages (sometimes lacking); style elongate; stigma apiculate; leaflets opposite or alternate, coriaceous, with entire margin; fruit brown at maturity, 2lobed, one of the lobes smaller or rudimentary, with a thin, smooth pericarp; seeds black, shiny, surrounded by sticky pulp ....... 10. Sapindus Flower disk unilateral, semiannular; fruit splitting into 3 inflated samaroid mericarps .................................................................. 14. Toulicia Flower disk annular; fruit a capsule or a berry ..................................... 14 Petals 4–7 mm long, with a single, entire appendage (adnate to the inner base of petal), the appendage elongated, conspicuously erect, densely sericeous or villous within; fruit a berry ................................... 12. Talisia Petals 1–3 mm long, with a pair of marginal appendages, the appendages sparsely to densely ferruginous within; fruit a berry or a capsule ................................................................................................. 15 Flowers 4-merous, ovary 2-locular; lower surface of leaflets foveate at vein axils beneath .................................................................. 16. Vouarana Flowers 5-merous, ovary 3-locular; lower surface of leaflets foveate or not at vein axils beneath ..................................................................... 16 Fruit a berry, 2-lobed, one lobe smaller or rudimentary, the pericarp thin and smooth .............................................................................. 10. Sapindus Fruit a dehiscent capsule, 2- or 3-lobed, the pericarp woody or coriaceous .............................................................................................................. 17

48

S APINDACEAE

17(16). Calyx divided to the base into distinct sepals, the sepals > 2 mm long; leaflet margin often toothed ...................................................... 3. Cupania 17. Calyx with sepals united at base to form a short cupular tube, the sepals 1–2 mm long (except M. macrostylis and M. macrolepis); leaflet margins entire (except M. scrobiculata, which has undulate to remotely serrate margins) ......................................................... 7. Matayba 1. ALLOPHYLUS L., Sp. Pl. 348. 1753. by Pedro Acevedo-Rodríguez Shrubs or trees. Leaves trifoliolate (in the flora area) or unifoliolate; stipules absent; petioles elongate. Inflorescences racemes, panicles, or thyrses. Flowers zygomorphic, functionally pistillate or staminate (plants apparently dioecious). Calyx 4(5)-merous, with the 2 external sepals smaller than the internal ones; petals 4(5), usually white and spatulate, with a basal appendage adnate to the margins of petal; disk annular, semiannular or 4-lobed. Stamens 8; filaments unequal, connate at base into a short tube; anthers dorsifixed. Ovary 2(3)carpellate, each carpel with a single ovule; style terminal, with 2 stigmatic branches. Fruit drupaceous, fleshy or leathery, globose, yellowish to red, of 1 or 2 ellipsoid or obovoid monocarps, 1 usually rudimentary. Seeds subglobose; embryo with abaxial cotyledon bent over the biplicate adaxial cotyledon. Pantropics and subtropics; ca. 255 species, 6 in Venezuela, 4 of these in the flora area. Key to the Species of Allophylus 1. 1. 2(1). 2. 3(2).

3.

Leaves unifoliolate (seemingly simple) ................................... A. amazonicus Leaves trifoliolate ...................................................................................... 2 Lower surface of leaflets canescent-tomentose ......................... A. racemosus Lower surface of leaflets glabrous or finely appressed-pubescent ......... 3 Leaflets oblong-elliptic, to 5 cm long, the upper surface foveolate, lower surface with appressed whitish trichomes, especially on veins; margins revolute, undulate or remotely serrate near the longacuminate apex; base acute or attenuate; petioles to 3 cm long, ferruginous-tomentose ........................................................... A. parimensis Leaflets elliptic, 10–14 cm long, lower surface glabrous, except for a few appressed trichomes along the midvein; margins entire, slightly revolute; base attenuate, slightly unequal; petioles 2–10 cm long, puberulent ............................................................................... A. punctatus

Allophylus amazonicus (Mart.) Radlk. in Engl. & Prantl., Nat. Pflanzenfam. III. 5: 312. 1895. —Schmidelia amazonica Mart., Flora 22(Beibl. 1): 6. 1839. Shrub or small tree; leaves unifoliolate; leaflets chartaceous, obovate to elliptic, crenate-serrate, glabrous; monocarps obovoid, puberulent to glabrous. Amazonas (Isla Ratón). Widespread throughout the Amazon basin.

Allophylus parimensis Steyerm., Ann. Missouri Bot. Gard. 75: 333. 1988. Tree to 10 m tall; leaves subcoriaceous. Montane forest, ca. 800 m; Amazonas (Simarawochi). Endemic. Allophylus punctatus (Poepp. & Endl.) Radlk. in Engl. & Prantl, Nat. Pflanzenfam. III. 5: 312. 1895. —Schmidelia punctata Poepp. & Endl.,

Cardiospermum 49

Fig. 35. Allophylus racemosus

Nov. Gen. Sp. Pl. 3: 38, no. 3, t. 244. 1844. Shrub or small tree, (3–)5(–12) m tall; leaf with the lower surface with dark vein network; thyrses simple, 5–9 cm long. Lowland forests, 50–300 m; widespread in Bolívar. Widely distributed throughout Neotropics. Allophylus racemosus Sw., Prodr. 62. 1788. —Comida de guacharaca, Fruta de paloma. Schmidelia occidentalis Sw., Fl. Ind.

Occid. 2: 665. 1798. —Allophylus occidentalis (Sw.) Radlk., Sitzungber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 20: 230. 1891 [1890]. Shrub to 4 m tall; leaves softly pubescent, the lower surface with prominent veins; berries subglobose, orange. Semideciduous forests and adjoining savannas, 100–300 m; Bolívar (2–8 km east of La Paragua, Reserva Forestal El Caura, Río Asa, 18 km north of Upata). Widespread elsewhere in Venezuela; Mexico, Central America, West Indies, Guyana. ◆Fig. 35.

2. CARDIOSPERMUM L., Sp. Pl. 366. 1753. by Pedro Acevedo-Rodríguez Herbaceous tendrilled vines, not producing milky sap. Stems furrowed, terete, or slightly angled; cross section of stems with a single vascular cylinder. Leaves ternately or biternately compound, membranous or chartaceous; stipules minute, early deciduous; petioles and rachis not winged; leaflets deeply serrate or lobed. Inflorescences thyrses, bearing tendrils at base of rachis, the flowers in cincinni distally along main rachis. Flowers 4–8 mm long, zygomorphic, functionally staminate or pistillate; calyx 4- or 5-merous; petals 4, distinct, spatulate, white, with basally adnate, hood-shaped appendage on adaxial surface; appendage yellow and fleshy at apex; disk unilateral, 2-lobed, the lobes sometimes elongate; receptacle enlarged into a short androgynophore. Stamens 8; filaments unequal, usually pubescent, connate at base; anthers dorsifixed. Fruits septifragal, marginicidal (dehiscent by the disjunction of the united margins of the carpels),

50

S APINDACEAE

inflated capsules, with thin-membranous walls, the septae persistent after dehiscence, with seeds persistently attached. Seeds spherical, black, with a small, heart-shaped or reniform, white arillode around hilum; embryo with abaxial cotyledon bent over the biplicate adaxial cotyledon. Central America, South America, a few paleotropic and subtemperate species; ca. 12 species, 3 in Venezuela, all in the flora area. Key to the Species of Cardiospermum 1. 1. 2(1).

2.

Petals > 5 mm long; disk glands elongate, horn-like; capsules ellipsoid, to 6 cm long; leaflets 5–10 cm long .................................. C. grandiflorum Petals < 5 mm long; disk glands short, ovoid; capsules subglobose or turbinate-trigonate, to 3 cm long; leaflets 2–4 cm long ...................... 2 Plants thin, herbaceous, usually glabrous; seeds to 5 mm diameter; leaflets deeply lobed to sinuate; inflorescence with a whorl of 3(4) cincinni ............................................................................... C. halicacabum Plants robust, semiwoody, variously pubescent; seeds to 4 mm diameter; inflorescence with several whorls of cincinni along main axis .................................................................................................. C. corindum

Cardiospermum corindum L., Sp. Pl. 526. 1763. Semiwoody vine to 3 m long; capsules subglobose. In early successional vegetation, dry forests, along roadsides, 100–200 m; Bolívar (Guasipati). Widespread elsewhere in Venezuela; widespread in Central America, the West Indies, and the rest of

Fig. 36. Cardiospermum corindum

South America, apparently introduced in Africa and tropical Asia. ◆Fig. 36. Cardiospermum grandiflorum Sw., Prod. 64. 1788. Robust vine to 5 m long, variously pubescent; capsules ellipsoid. Lower slope forests, open disturbed areas, 300–500 m; Bolívar (Cerro Bolívar). Widespread elsewhere in

Cupania 51

Venezuela; widespread in tropical and subtropical areas of the world. Cardiospermum halicacabum L., Sp. Pl. 366. 1753. Herbaceous vine with perennial rootstock, many-branched from base; capsules turbinate, ovoid, or subglobose. Probably native to the Neotropics, now nearly cosmopolitan; 2 varieties, both in the flora area. Key to the Varieties of C. halicacabum 1. Capsules 2–3 cm long, subglobose ................................ var. halicacabum 1. Capsules to 1.5 cm long, turbinatetrigonate, truncate or depressed at apex ............................... var. microcarpum

C. halicacabum var. halicacabum Leaflets usually deeply lobed. Deciduous woodlands, ca. 200 m; Bolívar (Los Aceites northwest of El Manteco). Widespread elsewhere in Venezuela; elsewhere in tropical and subtropical areas of the world, introduced into temperate areas. C.

halicacabum var. microcarpum (Kunth) Blume, Rumphia 3: 185. 1847. —Cardiospermum microcarpum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 104. 1821. —Once horas. Leaflets sinuate. Secondary growth, riversides, roadsides, near sea level to 200 m; Delta Amacuro (Tucupita), Amazonas (Río Atabapo, Río Negro, Río Orinoco, Río Sipapo). Widespread throughout the tropics.

3. CUPANIA L., Sp. Pl. 200. 1753. by Pedro Acevedo-Rodríguez Small to large trees. Leaves pinnately compound with rudimentary distal leaflet; stipules absent; rachis cylindrical or angular; leaflets mostly serrate, alternate or opposite, the petiolules usually short and not enlarged. Inflorescences axillary or terminal panicles or thyrses, with flowers produced in dichasia, monochasia, cincinni, or congested cymes. Flowers 5-merous, actinomorphic, bisexual or unisexual (plants polygamous, dioecious). Sepals short (2–4 mm long), imbricate, usually tomentose; petals as long as the sepals, with a pair of marginal tomentose appendages; disk annular and usually lobed. Stamens (4, 6)8, exserted; filaments of equal length, tomentose and filiform; anthers dorsifixed, ovoid or oval with retuse apex. Ovary 3-carpellate, each carpel with a single ovule. Fruits 2- or 3-locular, woody or leathery capsules. Seeds with fleshy arillode at base; embryo with straight cotyledons. Neotropics and subtropics; ca. 50 species, 11 in Venezuela, 7 of these in the flora area. Cupania is distinguished from Matayba by its imbricate, larger sepals (not valvate and short) and usually serrate leaflet margins (not mostly entire). Both characters intergrade, however, making generic delimitation unclear. A revision of the group may show them to be congeneric. Key to the Species of Cupania 1. 1. 2(1).

2.

Leaves with 2 leaflets; margins entire ..................................... C. kukenanica Leaves with > 4 leaflets; margins serrate or crenate .............................. 2 Leaves usually with > 10 leaflets; lower surface of leaflets cinereoustomentose; margins finely serrate (3 or 4 serrations per cm), the serrations sharp ........................................................................... C. cinerea Leaves with 10 or fewer leaflets; lower surface of leaflets variously pubescent but not cinereous; margins remotely serrate (1 or 2 serrations per cm), the serrations usually obtuse .................................... 3

52

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

S APINDACEAE

Plant pubescent, tomentose, or puberulous, especially on lower surface of leaflets, petioles, rachises, and young stems ................................... 4 Plant hirsute or hispid, especially on lower surface of leaflets, petioles, rachises, and young stems .................................................................... 5 Leaflets acute or acuminate at apex; capsules with 2 or 3 divaricate lobes ...................................................................................... C. scrobiculata Leaflets retuse or rounded at apex; capsules pyriform ................. C. latifolia Plant hispid; leaflets oblong or oblong-elliptic, with subentire margins ..................................................................................................... C. hispida Plant hirsute; leaflets elliptic, obovate, or oblanceolate, with repanddentate margins .................................................................................... 5 Leaflets coriaceous to chartaceous, with acute to apiculate apex and revolute margins .......................................................................... C. hirsuta Leaflets strongly coriaceous (subrigid), rugose, with retuse apex and strongly revolute margins ..................................................... C. rubiginosa

Cupania cinerea Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 39. 1844. —Chaparrillo, Chaparro de agua. Small to large tree, with short buttress roots; leaflets obovate or oblanceolate, obtuse at apex, the lower surface grayish or canescent. Seasonally flooded forests, riparian forests, or bordering savannas, 50– 400 m; Bolívar (Río Asa above Raudal Cotúa, middle and lower Río Caura, Río Nichare), Amazonas (mouth of Caño Yapacana, Río Atabapo, Río Casiquiare, near Río Mavaca, Río Orinoco near Tamatama). Widely distributed elsewhere in Venezuela; Panama, Colombia, Ecuador, Peru, Brazil (Acre), Bolivia. ◆Fig. 40. Cupania hirsuta Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 565. 1879 [1880]. —Chaparrillo, Fruta de paloma. Tree to 10 m tall; leaflets elliptic to obovate, with sharply serrate margins, the lower surface hirsute; capsules pyriform, reddish, hirsute; seeds black with yellowish arillode. Lowland forests, 100–400 m; Delta Amacuro (Río Toro, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Ciudad Bolívar, 62 km south of El Dorado, lower Río Orinoco). Guyana, Suriname, French Guiana. ◆Fig. 39. Cupania hispida Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 564. 1879 [1880]. Small tree; leaflets to 20 cm long, with strongly revolute margins, the lower surface

hispidulous; flowers congested in inflorescence. Evergreen lowland forests, 100– 300 m; Amazonas (Río Cuao). Brazil (Amazonas). Cupania kukenanica Steyerm., Fieldiana, Bot. 28: 344. 1952. Cupania roraimae Steyerm., Fieldiana, Bot. 28: 346. 1952. Tree ca. 8 m tall; leaflets coriaceous, the lower surface of leaflet lighter than upper one, the margins entire; inflorescence spiciform, densely flowered. Rocky stream banks, 1000– 1100 m; Bolívar (Gran Sabana, Río Kukenán, south of Roraima-tepui). Endemic. Cupania latifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 126. 1821. Cupania papillosa Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 557. 1879 [1880]. Cupania semidentata Radlk., Bull. Torrey Bot. Club 25: 337. 1898. Tree to 15 m tall; leaflets coriaceous and usually rugose, the lower surface puberulent; capsules pyriform, ferruginous-tomentose; seeds shiny, black. Evergreen to semideciduous lowland to lower montane forests, 50–500 m; Bolívar (Cerro Arimagua east of El Piñal, El Palmar, lower Río Orinoco). Widespread elsewhere in Venezuela; Panama, Colombia, Ecuador, Peru, Amazonian Brazil. ◆Fig. 38. Cupania rubiginosa (Poir.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 566.

Cupania 53

Fig. 37. Cupania scrobiculata

Fig. 38. Cupania latifolia

54

S APINDACEAE

Fig. 39. Cupania hirsuta

1879 [1880]. —Robinia rubiginosa Poir. in Lam., Encycl. 6: 227. 1804. Cupania tepuiensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 445. 1952. Tree to 10 m tall; leaflets rigidly coriaceous, rugose, the lower surface with prominent veins. Lowlands to high montane cloud forests, 100–2000 m; Bolívar (Aparamán-tepui, Cerro Carmene, Macizo del Chimantá, Roraima-tepui), Amazonas (near Raudal Arata, Río Atabapo, Río Ocamo). Lesser Antilles, Trinidad, Guyana, Peru, Brazil (Amazonas). Cupania scrobiculata Rich., Actes Soc. Hist. Nat. Paris 1: 109. 1792. —Zapatero. Tree to 9 m tall; leaves ca. 25 cm long; leaflets 9–12 cm long, entire, crenate, or remotely serrate, obtuse at base; petiolules swollen; capsules ca. 2 cm long, tomentose.

Fig. 40. Cupania cinerea

Semideciduous forests to evergreen montane forests, 50–900 m; Bolívar (El Abismo, El Manteco to San Pedro de las Dos Bocas, El Paují, near Las Claritas, south of Maniapure, lower Río Botanamo, lower Río Cuchivero basin, Río Parguaza, Río Uairén in Río Kukenán basin, Túriba), Amazonas (western base of Cerro Sipapo, road from Puerto Ayacucho to Gavilán, Río Casiquiare, between Samariapo and Puerto Ayacucho, San Carlos de Río Negro, Sierra Parima). Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 37. Three forms were recognized by Radlkofer (in Engl., Pflanzenr. IV. 165 (Heft 983): 1052. 1934) based on leaflet characters. Recognition of forms, however, for the Venezuelan material is not possible because of intergradation.

Dilodendron 55

4. DILODENDRON Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 355. 1878 [1879]. by Pedro Acevedo-Rodríguez Trees. Leaves bipinnate or subtripinnate; terminal leaflets on rachises fully developed or rudimentary; secondary rachises marginate and alternate; leaflets alternate or opposite, unequal at base and deeply serrate on margins. Inflorescences terminal panicles, with flowers in simple or compound dichasia. Flowers 5-merous, functionally staminate or pistillate (plants apparently dioecious). Calyx slightly zygomorphic with one sepal larger than the others; petals much shorter than sepals, with narrow, marginal appendages; disk annular and flattened. Stamens 6 or 8, exserted; filaments of equal length, subulate, free to base; anthers sagittate, dorsifixed. Ovary with 3 1-ovular carpels. Fruits loculicidal, 2- or 3-locular, woody capsules. Embryo with abaxial cotyledon bent over the biplicate adaxial cotyledon. Costa Rica, Panama, Colombia, Venezuela, Peru, Brazil, Bolivia, Paraguay; 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Dilodendron 1.

1.

Lower surface of leaflets glabrous, tertiary venation evidently reticulate; capsules trigonous-globose, the valves densely pilose within, not reflexed or splitting to the base .......................................... D. costaricense Lower surface of leaflets densely strigillose, tertiary venation not evident; capsules laterally compressed, the valves glabrous or sparsely pilose within, reflexed and splitting to the base ........ D. elegans Fig. 41. Dilodendron elegans

56

S APINDACEAE

Dilodendron costaricense (Radlk.) Gentry & Steyerm., Ann. Missouri Bot. Gard. 74: 536. 1987. —Dipterodendron costaricense Radlk., Smithsonian Misc. Collect. 61(24): 7. 1914. Tree (3–)10–20 m tall; leaves to 0.5 m long; leaflets lanceolate, rachis narrowly winged in younger portions. Sandy riversides, 200–500 m; Bolívar (Río Caura). Mérida, Zulia; Costa Rica, Panama, Colombia. Dilodendron elegans (Radlk.) Gentry & Steyerm., Ann. Missouri Bot. Gard. 74:

537. 1987. —Dilodendron bipinnatum var. elegans Radlk. in Mart., Fl. Bras. 13(3): 597. 1900. —Dipterodendron elegans (Radlk.) Radlk., Smithsonian Misc. Collect. 61(24): 7. 1914. —Caro Montañero. Dipterodendron venezuelense Steyerm., Fieldiana, Bot. 28: 346. 1952. Tree 8–25 m tall; leaflets alternate or subopposite, oblong-elliptic. Sandy riversides; 200–500 m; Bolívar (near El Dorado, Río Asa, Río Hacha). Barinas, Carabobo, Zulia; Costa Rica, Peru. ◆Fig. 41.

5. DODONAEA Mill., Gard. Dict. abr. ed. 4. 1754. by Pedro Acevedo-Rodríguez Shrubs. Young branches slightly ribbed. Leaves alternate, simple, petiolate or sessile; stipules absent. Inflorescences axillary or terminal with determinate growth, the flowers variously clustered along the rachis. Flowers unisexual or bisexual (plants dioecious or polygamous-dioecious). Calyx of (3–)5(–7) free sepals; petals absent; disk absent. Stamens usually twice as many as the sepals; filaments short; anthers sagittate or oblong. Ovary of (2)3–5(6) carpels, each with 2 ovules. Fruits winged, thin-walled capsules. Seeds usually black, lenticular or subspheric; embryo with both cotyledons spirally coiled. Mostly Pantropics, with a few species extending into southern temperate zones, primarily Australia (with 59 endemics); 68 species, 1 in Venezuela.

Fig. 42. Dodonaea viscosa

Houssayanthus

Dodonaea viscosa Jacq., Enum. Syst. Pl. 19. 1760. Viscid shrub 1–2 m tall; flowers yellowish, ovary 3-carpellate; mature capsules usually with reddish tinge. Shrubby savannas, roadsides, 1100–1200 m; Bolívar

57

(Sabanita near San Ignacio, San Ignacio de Yuruaní). Widespread elsewhere in Venezuela; tropics and subtropics worldwide, temperate South America and Africa. ◆Fig. 42. Dodonaea viscosa is a very widespread and polymorphic species.

6. HOUSSAYANTHUS Hunz., Kurtziana 11: 17. 1978. by Pedro Acevedo-Rodríguez Woody vines. Stems terete; cross section of stem with a central and 3 peripheral vascular cylinders. Leaves trifoliolate or pinnately 5-foliolate; stipules minute, early deciduous. Inflorescence a thyrse with a pair of tendrils at base of rachis, the flowers produced in lateral cincinni along the rachis. Flowers zygomorphic, functionally unisexual. Calyx of 5 unequal sepals; petals 4, distinct, with a basally adnate hood-shaped appendage on adaxial surface; disk unilateral, 4-lobed. Stamens 8, slightly exserted; filaments unequal; anthers dorsifixed, sometimes with an apical appendage. Ovary of 3 uniovular carpels; style terminal with 3 stigmatic branches. Fruits schizocarps, splitting into 3 mericarps containing a central locule with a narrow wing around it. Seeds lenticular; embryo with straight cotyledons bent at apex. Venezuela, Brazil, Bolivia, Paraguay, northeastern Argentina; ca. 3 species, 1 in Venezuela. Houssayanthus macrolophus (Radlk.) Hunz., Kurtziana 11: 17. 1978. —Cardiospermum macrolophum Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 261. 1878 [1879]. —Urvillea macrolopha (Radlk.) Radlk. in Mart., Fl. Bras. 13(3): 428. 1897. Woody, tomentose vine 2–3 m long; leaves trifoliolate; leaflets ovate, oblong, or subtrilobate, the lower surface canescent, the margins crenate-dentate; mericarps to 2 cm long. Seasonally flooded river margins, 100– 300 m; Delta Amacuro (Santa Catalina). Apure. ◆Fig. 43.

Fig. 43. Houssayanthus macrolophus

58

S APINDACEAE

7. MATAYBA Aubl., Hist. Pl. Guiane 349. 1775. by Pedro Acevedo-Rodríguez Large or small trees. Leaves pinnately compound with a rudimentary distal leaflet; stipules absent; rachis cylindrical or angular; leaflets entire, alternate or opposite; petiolules usually short and not enlarged. Inflorescences axillary or terminal panicles or thyrses, with flowers produced in dichasia, monochasia, cincinni, or congested cymes. Flowers 5-merous, actinomorphic, bisexual or unisexual (plant polygamous-dioecious). Sepals short (< 2 mm long), valvate, usually tomentose; petals as long as the sepals or longer, with a pair of marginal tomentose appendages; disk annular, usually lobed. Stamens (4–)8, exserted; filaments of equal length, tomentose and filiform; anthers dorsifixed, ovoid or oval with retuse apex. Ovary 3-carpellate, each carpel with a single ovule. Fruits a 2- or 3-locular, woody or leathery, loculicidal capsule. Seed nearly globose or ellipsoid; embryo with straight cotyledons. Tropical and subtropical areas of the New World; ca. 50 species, 18 in Venezuela, 16 of these in the flora area. Key to the Species of Matayba 1. 1. 2(1).

Leaves with only 1 pair of opposite leaflets ............................................. 2 Leaves with 2 or more pairs of opposite or alternate leaflets ................. 3 Leaflets obtuse or acute at apex, coriaceous to subcoriaceous; tertiary veins finely reticulate .............................................................. M. ptariana 2. Leaflets retuse at apex, chartaceous; tertiary veins loosely reticulate ............................................................................................. M. kavanayena 3(1). Leaflets coriaceous to rigidly coriaceous .................................................. 4 3. Leaflets chartaceous to subcoriaceous ..................................................... 7 4(3). Leaves with 2 or 4 opposite or subopposite leaflets; tertiary veins as prominent as the secondary .................................................................. 5 4. Leaves with 4 or more, alternate or subopposite leaflets; tertiary veins less prominent than the secondary ...................................................... 6 5(4). Leaflets elliptic or lanceolate, acute or acuminate at apex; vein axils not foveolate ..................................................................................... M. robusta 5. Leaflets spatulate or elliptic, rounded and retuse at apex; vein axils foveolate ................................................................................. M. yutajensis 6(4). Leaflets elliptic, oblong, or lanceolate, acuminate at apex, oblique at base, usually > 15 cm long ............................................... M. camptoneura 6. Leaflets elliptic or obovate, rounded or obtuse at apex, cuneate at base, to 17 cm long ................................................................................. M. opaca 7(3). Leaves with 4 (sometimes 2) opposite or subopposite leaflets ..... M. inelegans 7. Leaves with > 4 (sometimes 4 leaflets also present) alternate or opposite leaflets .................................................................................................... 8 8(7). Leaflets obtuse or rounded at apex .......................................................... 9 8. Leaflets acuminate or acute at apex ...................................................... 10 9(8). Leaflets elliptic, usually with a retuse apex ........................ M. atropurpurea 9. Leaflets obovate, not retuse at apex ................................................ M. opaca 10(8). Leaflets shortly acuminate at apex, the acumen obtuse ....................... 11

Matayba 59

10. 11(10). 11. 12(10). 12. 13(12). 13. 14(13). 14. 15(14).

15. 16(14). 16. 17(16). 17. 18(13). 18. 19(18). 19. 20(18). 20.

Leaflets acute, acuminate, or long-acuminate at apex, the acumen tapering ................................................................................................ 12 Leaflets oblong, elliptic, or oblanceolate, the margins usually wavy; capsule slightly flattened, 2-coccate, stipitate ................... M. arborescens Leaflets elliptic, the margins entire, weakly revolute; capsule subglobose, sessile ...................................................................... M. guianensis Leaflets acute at apex; capsule subglobose, sessile ................. M. guianensis Leaflets acuminate or long-acuminate at apex; capsule slightly flattened and coccate or trigonous ...................................................................... 13 Leaflets lanceolate, long-acuminate at apex .......................................... 14 Leaflets oblong or elliptic, acuminate at apex ....................................... 18 Leaflets with prominent primary and secondary veins on lower surface; capsule 2- or 3-coccate, shortly stipitate ............................................ 15 Leaflets with inconspicuous or slightly prominent secondary veins on lower surface; capsule globose or pyriform ........................................ 16 Style as long as or longer than the ovary; capsules < 1 cm long, usually 2-coccate, deeply emarginate at apex, with smooth pericarp .............................................................................................. M. macrostylis Style shorter than the ovary; capsules ca. 2 cm long, 3-coccate, emarginate at apex, with warty pericarp .................................. M. elegans Capsule globose, sessile ............................................................ M. guianensis Capsule pyriform, long-stipitate (> 7 mm long) ..................................... 17 Capsule slightly flattened, depressed at apex, 2-locular ............ M. longipes Capsule pyriform, apiculate at apex, 3-locular ........................ M. spruceana Leaflets with secondary veins prominent on lower surface; tertiary veins subclathrate; capsules short-stipitate (2 mm long) ........................... 19 Leaflets without prominent secondary veins; tertiary veins reticulate; capsule long-stipitate (5 mm long) ..................................................... 20 Leaflets with secondary veins sunken on upper surface ..... M. adenanthera Leaflets with secondary veins not sunken on upper surface ................ ........................................................................................... M. camptoneura Leaves with < 10 leaflets; leaflets acute at apex ...................... M. spruceana Leaves with > 10 leaflets; leaflets caudate at apex .................. M. peruviana

Matayba adenanthera Radlk. in Mart., Fl. Bras. 13(3): 615. 1900. —Zapatero. Tree to 12 m tall; leaflets 7–20 cm long, the lower surface with prominent venation; capsules reddish, pubescent or glabrous, smooth. Forest margins, 100–200 m; Bolívar (lower Río Caura, upper Río Parguaza), Amazonas (base of Cerro Sipapo, Río Atabapo, Yavita). Brazil (Amazonas). Matayba arborescens (Aubl.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 625. 1879 [1880]. —Sapindus arborescens Aubl., Hist. Pl. Guiane 357. 1775.

Small tree; leaflets 6–8, alternate, usually with retuse apex; capsules 2-locular, reddish, glabrous, slightly flattened. Savanna tree islands or savanna edges, gallery forests, flooded forests, humid roadsides, 100–800 m; Delta Amacuro (14 km southeast of Piacoa), Bolívar (middle Río Caura, Río Oris in middle Paragua basin, Río Parguaza, San Francisco de Yuruaní), Amazonas (Caño Yagua, La Esmeralda ridge, Río Coro Coro, near Santa Bárbara del Orinoco, base of Sierra de la Neblina, Sierra Parima). Widely distributed elsewhere in Venezuela; Trinidad, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 46.

60

S APINDACEAE

Matayba atropurpurea Radlk. in Mart., Fl. Bras. 13(3): 617. 1900. —Matayba discolor var. atropurpurea Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 630. 1879 [1880]. Small tree or shrub to 10 m tall; leaflets oblong or elliptic, lower surface lighter than upper surface, tertiary venation finely reticulate; flowers minute; capsules longstipitate. In white sandy soil among shrubs, 100–1100 m; Amazonas (Río Casiquiare, Río Orinoco 8 km below mouth of Río Atabapo, San Carlos de Río Negro, Sierra de la Neblina). Colombia.

ing brown, with upper surface darker; capsules globose, usually verrucose. Savannas, secondary and gallery forests, tree islands in savannas, 100–500 m; Bolívar (Canaima, Caño Asisa, Río Caura, Río Cucurital southeast of San Salvador de Paúl, Río Paragua, Río Parguaza, Serranía de los Pijiguaos, Urimán to Río Ambutuir), Amazonas (Cerro Yutajé, Culebra to Huachamacari, Isla Ratón, in and around Puerto Ayacucho, Río Coro Coro west of Cerro Yutajé, Río Pará, Río Ventuari, San Fernando de Atabapo, Serranía Pará). Widespread elsewhere in Venezuela; Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

Matayba camptoneura Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 629. 1879 [1880]. Matayba reducta Steyerm., Fieldiana, Bot. 28: 351. 1952. Tree to 20 m tall; leaflets 6, chartaceous to coriaceous, 10–35 cm long, with prominent veins on lower surface and oblique base; inflorescences short panicles. Gallery and humid forests, 100–300 m; Bolívar (upper Río Caura, near Las Pavas, Río Cuyuní, Río Ichún, upper Río Paragua), Amazonas (below La Esmeralda, San Carlos de Río Negro). Guyana, Suriname, French Guiana.

Matayba inelegans Spruce ex Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 627. 1879 [1880]. Matayba sororopaniana Steyerm., Fieldiana, Bot. 28: 348. 1952. Small tree; leaflets 4, glabrous, 8–14 cm long; rachis cylindric; capsules green, pyriform, stipitate (the stipe 5–8 mm long). Evergreen lowland and lower montane forests, 100–2100 m; Delta Amacuro (Caño Jotajana off Caño Guiniquina), Bolívar (Sororopán-tepui), Amazonas (base of Cerro Sipapo, Río Cunucunuma, Río Pacimoni, Río Sipapo, between San Carlos de Río Negro and Solano, San Fernando de Atabapo, base of Sierra de la Neblina, Yavita to Maroa road). Guyana, French Guiana, Peru, Brazil.

Matayba elegans Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 629. 1879 [1880]. —Carapillo, Fruto de paloma, Mulato, Zapatero. Tree 5–8(–15) m tall; leaves 30–45 cm long; leaflets 6–8, lanceolate, seldom elliptic, the base slightly unequal; capsules (2)3lobed or subpyriform, reddish or pink. Riparian or moist forests, 50–400 m; Bolívar (Cerro Marimarota, Reserva Forestal La Paragua, Reserva Forestal Río Caura, Río Aro, middle Río Caura, Río Parguaza), Amazonas (south of Isla Ratón, Río Coro Coro). Zulia; Colombia. ◆Fig. 49. Matayba guianensis Aubl., Hist. Pl. Guiane 331. 1775. Shrub or small tree; leaflets 8 or more, chartaceous to subcoriaceous, usually dry-

Matayba kavanayena (Steyerm.) Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 426. 1966. —Cupania kavanayena Steyerm., Fieldiana, Bot. 28: 343. 1952. Tree to 8 m tall; leaflets glabrous, the lower surface lighter than upper one, the tertiary venation whitish. Dense riparian forests, 100–1200 m; Bolívar (Quebrada Kavanayén, Río Caruay), Amazonas (Atabapo). Endemic. ◆Fig. 48. Matayba longipes Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 626. 1879 [1880 ]. Matayba tovarensis Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 626. 1879 [1880].

Matayba 61

Matayba venezuelana Steyerm., Fieldiana, Bot. 28: 353. 1952. Matayba longipes subsp. tepuiensis Steyerm., Ann. Missouri Bot. Gard. 75: 1069. 1988. Tree to 20 m tall; leaflets 4, oblong or lanceolate, 6–20 cm long; capsule flattened, 2-locular, long-stipitate (the stipe ca. 1 cm long). Lower montane to montane forests, 400–1400 m; Amazonas (Caño Asisa, slopes of Cerro Coro Coro, base of Sierra de la Neblina, Serranía Parú). Zulia. Matayba macrostylis Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 625. 1879 [1880]. Matayba macrolepis Radlk. in Mart., Fl. Bras. 13(3): 605. 1900. Trichilia caudata Killip & Cuatrec., Revista Acad. Colomb. Ci. Exact. 5: 36. 1942. Small tree; leaflets usually drying brown, the upper surface darker; capsules reddish, short-stipitate. Lowland evergreen forests, gallery forests, 100–200 m; Amazonas (Isla Ratón, Río Casiquiare, Río Cataniapo, Río Manipitare, Río Siapa). Brazil (Amazonas, Pará). Matayba opaca Radlk., Sitzungsber. Math.Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 628. 1879 [1880]. Tree 4–12 m tall; leaflets 4–8, chartaceous to coriaceous, elliptic or obovate, with obtuse or rounded apex; stems lenticellate. Venezuela, Guyana, Suriname, Brazil; 2 varieties, both in the flora area. Key to the Varieties of M. opaca 1. Leaflets coriaceous, shiny (especially the upper surface), with strongly revolute margins and obtuse apex .. var. fallax 1. Leaflets chartaceous to subcoriaceous, dull on both surfaces, margin sometimes revolute, and apex rounded or slightly retuse .................... var. opaca M. opaca var. fallax (Radlk.) Uittien in Pulle, Fl. Suriname 2(1): 384. 1937. —Matayba fallax Radlk. in Mart., Fl. Bras. 13(3): 612. 1900. Open savannas on white sand, 100–900 m;

Bolívar (Cerro Pitón, Río Acanán, Río Chicanán, Río Yuruaní), Amazonas (Río Casiquiare, Río Yatúa, south of Samariapo toward Río Sipapo). Suriname, Brazil. ◆Fig. 47. M. opaca var. opaca. —Panucoa. Matayba affinis Steyerm., Ann. Missouri Bot. Gard. 75: 1067. 1988. Gallery forests, savannas, ca. 100 m; Bolívar (Cerro Akurimá along middle Río Caroní, Icabarú, Río Aponguao, Río Asa, Río Karaurín, upper Río Trueno, Sierra de Lema), Amazonas (Caño Caname, Río Asisa, Río Autana, upper Río Orinoco, Río Siapa, Sabana de Moyo on south bank of lower Río Ventuari, Santa Bárbara del Orinoco). Guyana, Suriname. Matayba peruviana Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 627. 1879 [1880]. Matayba oligandra Sandwith, Bull. Misc. Inform. 1935: 123. 1935. Trichilia ptariana Steyerm., Fieldiana, Bot. 28: 278. 1952. —Matayba oligandra var. ptariana (Steyerm.) Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 426. 1966. Matayba chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17: 447. 1967. Matayba jauaensis Steyerm., Bol. Soc. Venez. Ci. Nat. 33: 347. 1976. Matayba oligandra var. occidentalis Steyerm., Ann. Missouri Bot. Gard. 75: 1069. 1988. Small tree; leaves with 5–7 pairs of opposite or alternate leaflets, these with caudate, acuminate apex; stamens 4–8; capsule reddish, long-stipitate (stipe 1 cm long). Evergreen lowland to montane forests, dwarf forests, 100–1800 m; Bolívar (Cerro Sarisariñama, Cerro Venamo, Río Apacará, Río Tírica), Amazonas (Cerro Duida, Cerro Sipapo, Río Arauicaua, Río Orinoco, Río Yatúa). Guyana. ◆Fig. 50. Matayba ptariana Steyerm., Fieldiana, Bot. 28: 348. 1952. Small tree to 20 m tall; petiolules swollen; capsules slightly flattened, 2- or 3locular, depressed at apex, glabrous and stipitate (the stipe ca. 7 mm long). Endemic; 2 subspecies, both in the flora area.

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Fig. 44. Matayba ptariana subsp. ptariana

Fig. 45. Matayba spruceana

Fig. 46. Matayba arborescens

Matayba 63

Fig. 47. Matayba opaca var. fallax

Fig. 48. Matayba kavanayena

Fig. 49. Matayba elegans

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Fig. 50. Matayba peruviana

Key to the Subspecies of M. ptariana 1. Petioles < 1 cm long; leaflets rounded at apex .................. subsp. guaiquinimae 1. Petioles 1–2 cm long; leaflets acute or acuminate at apex .... subsp. ptariana M.

ptariana subsp. guaiquinimae Steyerm., Ann. Missouri Bot. Gard. 74: 635. 1987. Shrub 2–3 m tall. Shrublands on tepui summit, montane forests, 1000–1400 m; Bolívar (Cerro Guaiquinima, Cerro Uroi in Sierra de Lema). Endemic. M. ptariana subsp. ptariana Low forests, shrublands along rivers, 400–1600 m; widespread in Bolívar (Cerro Guaiquinima, Cerro Marutaní, Cerro Sarisariñama, 35 km west of Chiguao, El Paují, Ptari-tepui, Río Canaracuni, Río Carapo, Río Caroní, Río Ichún, headwaters of Río Paragua and Río Uaiparú, Sierra Pakaraima), Amazonas (Sierra Parima). Adjacent Brazil. ◆Fig. 44.

Matayba robusta Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 627. 1879 [1880]. Tree to 6 m tall; leaflets 2–4, coriaceous, dull, strongly reticulate; inflorescences short, aggregate racemes; capsules pyriform. Blackwater river banks, ca. 100 m; Amazonas (Río Pasimoni). Amazonian Brazil. Matayba spruceana (Benth.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 9: 627. 1879 [1880]. —Cupania spruceana Benth., Hooker’s J. Bot. Kew Gard. Misc. 3: 199. 1851. Small tree; leaves with 4–6 leaflets; inflorescence short, aggregate racemes. Wooded ridges, 1600–2000 m; Bolívar (Sororopán-tepuí). Brazil (Pará). ◆Fig. 45. Matayba yutajensis Steyerm., Ann. Missouri Bot. Gard. 75: 1070. 1988. Tree to 4 m tall; leaflets strongly reticulate with revolute margins and retuse apex; capsules red, slightly flattened, with 1

Melicoccus 65

or 2 developed cocci, shortly stipitate. Moist canyons, sandstone outcrops, 500–1600 m; Amazonas (Cerro Duida in headwaters of Caño Culebra, Cerro Marahuaka, Cerro

Yutajé, Río Coro Coro). Endemic. Matayba yutajensis is vegetatively very similar to M. juglandifolia; further studies may show them to be conspecific.

8. MELICOCCUS P. Browne, Civ. Nat. Hist. Jamaica 210. 1756. by Pedro Acevedo-Rodríguez Trees. Leaves pinnately compound, the terminal leaflet rudimentary or lacking; stipules lacking; rachis winged; leaflets 2 or 4, opposite. Inflorescence terminal panicles, on lateral branches, the flowers solitary along the secondary rachis. Flowers 5-merous, actinomorphic, functionally pistillate or staminate (plants apparently dioecious). Calyx cup-shaped, of equal sepals; petals reflexed, appendages lacking; disk prominent, annular, and slightly lobed. Stamens 8, exserted; filaments of equal length; anthers dorsifixed. Ovary 2-carpellate, each carpel with a single ovule; style obsolete; stigmatic surface lobed, subcapitate. Fruits 1(2)-seeded, subglobose or ellipsoid berries, with leathery foveolate pericarp. Seeds with fleshy outer coat. Embryo with fleshy and straight cotyledons. Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, northeastern Argentina; 10 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Melicoccus 1.

1.

Leaf rachis usually winged; inflorescence a raceme (flowers not grouped in dichasia); calyx and corolla 4-merous; gynoecia bicarpellate ................................................................................................. M. bijugatus Leaf rachis not winged; inflorescence a thyrse (flowers in simple or compound dichasia); calyx and corolla 5-merous; gynoecia tricarpellate .............................................................................. M. oliviformis

Melicoccus bijugatus Jacq., Syst. Enum. Pl. 19. 1760. —Melicocca bijuga (Jacq.) L., Sp. Pl. ed. 2, 495. 1762. —Mamón, Mamoncillo. Tree to 20 m tall; bark smooth and grayish; flowers greenish or yellowish, fragrant; berry green; seed coat tan. Deciduous forests, also commonly cultivated or spontaneous, near sea level to 200 m; Bolívar (Ciudad Bolívar, El Dorado, Puerto Ordaz). Venezuelan Coastal Cordillera, Andes; Colombia, Guyana, Suriname, French Guiana, Brazil, introduced to Central America and West Indies. ◆Fig. 51. The outer coat of the seed of Melicoccus bijugatus is juicy and edible and is often sold in markets.

Melicoccus oliviformis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 100. 1821. —Talisia oliviformis (Kunth) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 342. 1878, “olivaeformis.” Tree 5–35 m tall; stems sulcate, lenticellate, ash-colored; leaves paripinnate; leaflets 2 or 4, opposite; calyx ferruginoustomentose; petals rhombate, with 2 marginal appendages. Dry forests, scrublands, riparian forests, savannas, 100–500 m; Amazonas (Río Atabapo, Río Ocamo, near Raudal Arata). Anzoátegui, Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Miranda, Nueva Esparta, Zulia; Colombia, Trinidad, Ecuador, Peru, introduced in Central America and the West Indies.

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Fig. 51. Melicoccus bijugatus

9. PAULLINIA L., Sp. Pl. 365. 1753; emend. Schum., Skr. Naturhist.-Selsk. 3(2): 119. 1794. by Hans T. Beck Lianas or scandent shrubs. Stems terete or sulcate; cross section of stems simple (a single vascular cylinder) or with many vascular cylinders (a central cylinder surrounded by 3 peripheral cylinders). Leaves compound, typically pinnate and 5-foliolate, sometimes ternate, biternate, or multijugate; stipules small to large, persistent or caducous; petiole and rachis marginate, winged or not winged; leaflets membranous to coriaceous; margins subentire to serrate-dentate, glands often occurring on teeth; venation reticulate or clathrate. Inflorescence thyrsoid, terminal, axillary or cauliflorous; occasionally when axillary inflorescences presenting a pair of curved tendrils at the base of the rachis; bracts and bracteoles usually subulate and minute. Flowers zygomorphic, functionally staminate or pistillate (plants polygamo-dioecious), whitish. Sepals 5 or 4 (by fusion of lowermost), the outer sepals smaller, quincuncial or imbricate; petals 4, free, with adnate petaloid scales on adaxial surfaces, scales of central petals longer than those of lateral petals; disk unilateral, modified into 4 lobes. Stamens 8, slightly unequal, sometimes connivent and clustered. Pistils of 3 uniovular carpels; style 1; stigmas 3. Fruits loculicidal, sessile or stipitate capsules, dorsally 3-winged (rarely 9-winged), spiny or wingless. Seeds usually black and lustrous, partially enclosed by a white fleshy coat (aril). Neotropics south to Argentina, tropical and subtropical Africa and Madagascar; ca. 125 species, 32 in Venezuela, 20 of these in the flora area.

Paullinia 67

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

2.

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

5.

6(4).

6.

7(3). 7. 8(7). 8. 9(8).

9.

Leaves ternate ........................................................................................... 2 Leaves not ternate ..................................................................................... 3 Petioles marginate or narrowly winged; cross section of stem usually with > 1 vascular cylinder (a central cylinder surrounded by 3, 2, or 1 peripheral cylinders), rarely simple; stipules deltoid, caducous or sometimes persistent, stipular scars on mature stems visible; leaflet margins serrate-dentate, teeth rounded and glandular, apices acute or rounded; vein axils on lower surface conspicuously barbate; tertiary venation reticulate; bracts subulate, 1–2 mm long; capsules clavate, without spiny projections ............................................... P. cururu Petioles not winged; cross section of stem simple; stipules rounded, caducous, stipular scars not conspicuous; leaflet margins subentire, glands occurring on basal margins or on petiolules of lower leaflets, apices long-acuminate or aristate; vein axils on lower surface of leaves sparsely barbate; tertiary venation reticulate to subclathrate; bracts deltoid, ca. 0.5 mm long; capsules ellipsoid, with spiny projections ............................................................................................. P. paullinioides Leaves biternate, trijugate, or multijugate .............................................. 4 Leaves 5-foliolate, pinnate ........................................................................ 7 Leaves trijugate ......................................................................................... 5 Leaves biternate or multijugate ............................................................... 6 Tertiary venation clathrate; petioles marginate or winged, rachises winged or not, or marginate; stipules foliaceous, 1–4 cm long, caducous, stipule scars visible; stems terete; inflorescence rachises robust, 2–4 mm diameter; capsules globose, 6-ribbed, stipitate ...................... P. ingaefolia Tertiary venation reticulate; petioles not winged to marginate, rachises marginate or winged; stipules deltoid, ca. 1 mm long, stipule scars not visible; stems subterete; inflorescence rachises usually thin, ca. 1 mm diameter; capsules pear-shaped and 3-winged, becoming wingless at maturity (wings evanescent), substipitate ............................ P. micrantha Leaves biternate; rachises not winged or marginate; stipules deltoid, ca. 1 mm long; inflorescences axillary, solitary, with elongate rachises; capsules 3-winged, the wings 5–7 mm wide at middle .......... P. fuscescens Leaves 4- or 5-jugate, sometimes 3- or 6-jugate; rachises marginate; stipules linear-lanceolate, 3–7 mm long; inflorescences axillary, solitary or fasciculate, with short condensed rachises; capsules 3winged, wings 5–12 mm wide at middle ............................. P. serjaniifolia Cross section of stems with many vascular cylinders (a central cylinder with 3 peripheral cylinders) ................................................................. 8 Cross section of stem simple (1 central vascular cylinder) ................... 10 Petioles and rachises not winged ................................................. P. alsmithii Petioles and rachises winged .................................................................... 9 Stipules linear-lanceolate with entire margins; lower surface of leaves glabrous, barbate at vein axils; bracts needle-like, 1–2 mm long; capsules pear-shaped or clavate, ± stipitate, mostly glabrous ................ P. pinnata Stipules oblong to lanceolate with scariose margins; lower surface of leaves pubescent, barbate at vein axils; bracts linear-lanceolate, 4–

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10(7). 10. 11(10). 11. 12(11). 12. 13(12).

13.

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

15. 16(11).

16.

17(10). 17. 18(17). 18. 19(18).

19.

7 mm long, conspicuously protruding beyond cincinni; capsules ovoidellipsoid, sessile, usually pubescent ........................................ P. leiocarpa Flowering, fruiting, and older stems sulcate or costate ........................ 11 Flowering, fruiting, and older stems terete or subterete, never deeply sulcate .................................................................................................. 17 Petioles not winged .................................................................................. 12 Petioles winged ........................................................................................ 16 Rachises winged ....................................................................................... 13 Rachises not winged ................................................................................ 14 Young stems densely pubescent; stipules incised; leaflet margins with mucronate glandular teeth, bracts conspicuously longer than sessile cincinni; capsules triquetrous, sessile, densely pubescent ................................................................................................. P. rubiginosa Young stems glabrous, glabrescent, or sparsely pubescent; stipules lanceolate; leaflet margins with glandular teeth, not mucronate; bracts not exceeding pedunculate cincinni; capsules 3-winged, stipitate, glabrous .......................................................................................... P. caloptera Tertiary venation reticulate; capsules ellipsoidal, 3-winged ...... P. nuriensis Tertiary venation clathrate; capsules globose, not winged ................... 15 Stipules ovate, incised, often persistent and conspicuous on older stems; bracts scarious; plant densely pubescent; capsules densely pubescent ....................................................................................................... P. rugosa Stipules oblong, entire, often absent on older stems; bracts inconspicuous; plant sparsely pubescent; capsules glabrous ...................... P. cupana Bracts oblong, 5–15 mm long, conspicuously protruding beyond sessile cincinni; stipules elongate, to 7 cm long, oblong; capsules pear-shaped or turbinate, the valves moderately thick, 2–4 mm ................ P. bracteosa Bracts subulate, 1–3 mm long; stipules as in P. bracteosa but usually smaller, typically 5–10 mm long; capsules pear-shaped, valves thin, ca. 1 mm thick ............................................................................ P. imberbis Lower surface of leaflets glabrous except for barbate vein axils; capsules with 9 wings, wing margins verrucose .................................... P. verrucosa Lower surface of leaflets densely pubescent, puberulent, or glabrescent; capsules unwinged or with 3 wings .................................................... 18 Lower surface of leaflets glabrescent; margins remotely dentate, the glands not prominent; capsules globose, not winged ............ P. capreolata Lower surface of leaflets puberulent or pubescent; margins dentate, the teeth glandular; capsules ellipsoidal, winged .................................... 19 Lower surface of leaflets puberulent, occasionally glabrescent; leaflet margins with glandular teeth, not mucronate, rachises not winged, varying to marginate; inflorescence rachises usually thin (ca. 1 mm diameter); cincinni closely spaced on inflorescence rachises; stipules oblong, persistent; stems subterete, occasionally shallowly sulcate; capsules equally 3-winged, glabrous, purple-red, not bicolorous, wing veins widely spaced .................................................................. P. rufescens Lower surface of leaflets densely golden brown-pubescent; leaflet margins with glandular, mucronate teeth, rachises not winged; inflores-

Paullinia 69

cence rachises usually thick (2–3 mm diameter); cincinni sparsely spaced on inflorescence rachises; stipules deltoid, early to late deciduous, stipule scars visible; stems terete; capsules unequally 3-winged, markedly pubescent, the wings commonly red, contrasting with the golden yellow fruit, wing veins narrowly spaced .................. P. anisoptera Paullinia alsmithii Macbr., Field Mus. Nat. Hist., Bot. Ser. 13 3A(2): 332. 1956. Liana. Wet forests, frequent along river banks, 100–400 m; Bolívar (Río Parguaza), Amazonas (Río Cunucunuma, Río Orinoco, San Fernando de Atabapo). Táchira; Ecuador, Peru, Brazil (Acre). Paullinia anisoptera Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(2): 397. 1858. —Bejuco cresta de gallo. Liana. Wet forests, 100–400 m; occasional in Delta Amacuro and Bolívar. Distrito Federal; Guyana, French Guiana, Brazil (Pará). ◆Fig. 54. Pending systematic revision may show Paullinia livescens Radlk. to be conspecific with P. anisoptera. Paullinia bracteosa Radlk., Bull. Herb. Boissier sér. 2, 5: 321. 1905. Liana. Wet forests, 100–500 m; Amazonas (Sierra Parima, Ugueto). Venezuelan northern Coastal Range; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil (Acre, Amazonas, Pará, Roraima), Bolivia. Paullinia caloptera Radlk. in Engl. & Prantl, Nat. Pflanzenfam. III. 5: 304. 1895. —Rabo de iguana, Tar seeho waina he héhéchë, To’oóto. Liana. Riparian and evergreen forests, 50–200 m; Bolívar (Túriba), Amazonas (widespread). Apure, Falcón, Sucre; Costa Rica, Panama, Colombia, Brazil (Amazon basin). ◆Fig. 55. Paullinia caloptera is widespread and polymorphic. However, in Venezuela it is rather stable in its morphology. Paullinia caloptera is used medicinally as a vomitive and for dry coughs. Paullinia capreolata (Aubl.) Radlk., Monogr. Serjania 70, no. 17. 1875. —Enourea capreolata Aubl., Hist. Pl. Guiane 587, t. 235. 1775. —Bejuco marrón, Manjar.

Liana. Humid and riparian forests, secondary vegetation, 50–700 m; Delta Amacuro (Río Amacuro, east of Río Grande, San Victor, Serranía de Imataca), Bolívar (El Dorado), Amazonas (Isla Macará on Río Negro, La Esmeralda, Río Casiquiare, Río Cunucunuma, Río Ocamo, Río Mawarinuma, San Carlos de Río Negro). Apure, Aragua, Distrito Federal, Lara, Miranda, Portuguesa, Sucre, Yaracuy; Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 52. Paullinia capreolata is a widespread, highly polymorphic species complex. The fruit of P. capreolata is reported as edible. Paullinia cupana Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 117. 1821. —Cupana, Ojo (Baniba). Paullinia sorbilis Mart., Fl. Bras. 7: 266. 1826. Liana or scandent shrub. Found in cultivation, 100–200 m; Amazonas (San Carlos de Río Negro, San Fernando de Atabapo, Victorino). Not reported from elsewhere in Venezuela, but commonly found in cultivation in Brazil (Amazonas, Pará). Paullinia cupana is hypothesized to be native to the upper Río Negro, Río Casiquiare, and upper Río Orinoco region. Seeds of Paullinia cupana are the source of guaraná, which is used both indigenously and industrially in Brazil for the manufacture of a stimulating, caffeine-rich beverage. However, in Venezuela, the consumption of these stimulant beverages made from the seeds occurs only locally. The three collections recorded from Amazonas represent collections through time: 1804, Humboldt & Bonpland; 1887, Gaillard; 1942, Williams. Each specimen provides data about the uses of the species, indicating their cultivated status. These specimens appear identical with material from central Amazonia. Ducke’s claim that a wild typical variety, Paullinia cupana var.

70

S APINDACEAE

cupana, known from the Río Orinoco, is different from the cultivated variety, P. cupana var. sorbilis (Mart.) Ducke, known from central Amazonia, is not supported by systematic collections. Paullinia cururu L., Sp. Pl. 365. 1753. —Bejuco de loro, Bejuco tres filos, Tonina. Scandent shrub. Riparian forests, 200– 300 m; Bolívar (Río Parágua). Widespread in Venezuelan states north of the Río Orinoco; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil. Paullinia fuscescens Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 120. 1821. —Bejuco Santa Ana, Berga de toro, Cacho de toro. Scandent shrub. Forests, forest savannas, 200–300 m; Delta Amacuro (Caño Guinipa, Sacupana), Bolívar (El Palmar, Represa Guri). Widespread in Venezuelan states north of the Río Orinoco; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador. Paullinia imberbis Radlk., Mongr. Paullinia 177. 1896. Liana. Clearings in riparian and premontane forests, 100–300 m; Amazonas (Gavilán, Río Cuao, Tamatama). Mérida, Portuguesa; Panama, French Guiana, Peru, Brazil (Amapá, Amazonas, Maranhão, Mato Grosso, Pará), Bolivia. Paullinia ingaefolia Rich. ex Juss., Ann. Mus. Natl. Hist. Nat. 4: 349. 1804. —Cuna baruk. Liana; fruits edible. Lowland humid riparian and wet montane forests, 100–1300 m; Bolívar (Quebrada O-paru-má near Kavanayén, Ptari-tepui, Radaul Mavaca on Río Parguaza), Amazonas (from Culebra to Huachamacari, Río Cunucunuma, Río Matacuni, Sierra Parima, Simarawochi). Apure, Miranda, Táchira; Costa Rica, Panama, Guyana, French Guiana, Ecuador, Peru, northern Brazil. ◆Fig. 59. Paullinia leiocarpa Griseb., Fl. Brit. W. Ind. 124. 1859. —Bejuco tres filos, Cuehiemuni (Yekwana).

Liana; fruits edible. Wet forests, commonly riparian, 50–300 m; common in Bolívar (La Paragua, Macarapa on Río Paragua, middle Río Caura), Amazonas (Puerto Ayacucho). Widespread in northern Venezuela; Colombia, Trinidad-Tobago, Guyana, Brazil (Goiás, Pará, Roraima). ◆Fig. 60. Paullinia micrantha Cambess. in A. St.Hil., Fl. Bras. Merid. 1: 373. 1825 [1828]. Scandent shrub. Wet and riparian forests, 100–900 m; Delta Amacuro (east-northeast of El Palmar, east of Río Grande), Bolívar (La Escalera, Santa Elena de Uairén). Guyana, Suriname, French Guiana, northern and southeastern Brazil. ◆Fig. 56. Paullinia nuriensis Steyerm., Acta Bot. Venez. 3: 179. 1968. Scandent shrub; leaflets elliptic, the margins entire, lower pairs with small glands at base. Forested summits, ca. 600 m; Bolívar (Cerro El Picacho north of Las Chicharras). Endemic. Paullinia paullinioides Spruce ex Radlk., Monogr. Paullinia 239. 1896. —Bejuco barba de mono. Liana. Low forests and river banks, ca. 100–200 m; Amazonas (Río Mawarinuma). Apure, Mérida; Costa Rica, Colombia, Ecuador, Peru, northern Brazil. Paullinia pinnata L., Sp. Pl. 366. 1753. —Aruhuhu, Arujujuj, Arusi-moroconi, Aru-simu, Bejuco tres filos rebalsero, Casabe colorado, Pico de paloma. Liana; fruits edible. Humid mixed evergreen forests, gallery forests, 50–100 m; Delta Amacuro (Caño Araguabisi between Caño Araguao and Caño Güiniquina, Misión San Francisco de Guayo, Río Acure, Río Cuyubini, Sacupana), Bolívar (Tumeremo). Falcón, Lara, Monagas, Sucre; widespread from Mexico through Central America and the West Indies, South America, tropical and subtropical Africa, including Madagascar. As the only species of Paullinia in Africa, P. pinnata has many common names and uses. However, it is not known if it was introduced from South America. ◆Fig. 61.

Paullinia 71

Paullinia rubiginosa Cambess. in A. St.Hil., Fl. Bras. Merid. 1: 371. 1825 [1828]. Liana. Wet forests, 300–900 m; Bolívar (Sierra de Lema), Amazonas (Simarawochi). Mérida, Táchira; Panama, Colombia, Peru, Brazil. Paullinia rufescens Rich. ex Juss., Ann. Mus. Natl. Hist. Nat. 4: 349. 1804. —Kun-a-wa-lu-yek, Tupukuiden. Liana. Wet evergreen and riparian forests, 100–1100 m; Delta Amacuro (town of Sierra Imataca east-southeast of Los Castillos, Río Acure), Bolívar (El Dorado, El Paují, Macizo del Chimantá [Toronó-tepui], Río Karún, upper Río Paragua, Río Tirica, Santa Elena de Uairén), Amazonas (northeast of San Carlos de Río Negro). Yaracuy; Guyana, Suriname, Brazil (Pará). ◆Fig. 57. Paullinia rugosa Benth. ex Radlk., Monogr. Paullinia 219. 1896. Liana. Wet evergreen forests, sandy savannas, steep rocky savannas, savanna forests, 100–400 m; Amazonas (Caño Yapacana, Capihuara, upper Río Casiquiare, Río Matacuni, Río Orinoco, Río Siapa, Sierra Parima, Simarawochi, common in Puerto Ayacucho area along roads to Samariapo,

Paso El Diablo, and Gavilán). Apure, Aragua, Distrito Federal, Táchira; Costa Rica, Panama, Colombia, Peru, northern Brazil. ◆Fig. 53. Paullinia serjaniifolia Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 356. 1862, “serjaniaefolia.” Liana. Disturbed mesophytic forests, savannas, 100–200 m; Amazonas (Puerto Ayacucho). Carabobo; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil (Amazonas). Paullinia verrucosa Radlk., Abh. Math.Phys. Cl. Königl Bayer Akad. Wiss. 19(1): 247. 1896. Paullinia novemalata Radlk., Monogr. Paullinia 181. 1896. Liana. Wet riparian forests, 100–200 m; Bolívar (upper Río Caroní, Santa Elena de Uairén). Guyana, Suriname, Brazil (Amazonas, Roraima). ◆Fig. 58. Paullinia verrucosa is poorly represented in herbaria. Leaflets on fruiting stems vary tremendously in size from 2 to 12 cm, and leaflet number varies outside the flora area from 3-foliolate (ternate) to 5-foliolate. However, the specimens from the flora area show the 5-foliolate condition.

Fig. 52. Paullinia capreolata

72

S APINDACEAE

Fig. 53. Paullinia rugosa

Fig. 54. Paullinia anisoptera

Paullinia 73

Fig. 55. Paullinia caloptera

Fig. 56. Paullinia micrantha

Fig. 58. Paullinia verrucosa

Fig. 57. Paullinia rufescens

74

S APINDACEAE

Fig. 59. Paullinia ingaefolia

Fig. 60. Paullinia leiocarpa

Fig. 61. Paullinia pinnata

Sapindus 75

10. SAPINDUS Plum. ex L., Sp. Pl. 367. 1753. by Pedro Acevedo-Rodríguez Small to medium-sized trees. Leaves pinnately compound, the terminal leaflet rudimentary; stipules absent; rachis and petioles winged, margined, or naked; leaflets alternate or opposite, coriaceous, falcate, lanceolate, with long-acute apex and oblique base. Inflorescence a thyrse with cincinni as secondary units. Flowers 5-merous, slightly zygomorphic, functionally pistillate or staminate (plants polygamous-monoecious). Calyx with 2 smaller outer sepals; petals with reduced marginal appendages; disk annular and cupular. Stamens 8, exserted; filaments of equal length; anthers dorsifixed. Ovary 3-carpellate, each carpel with a single ovule. Fruits mostly globose berries, usually with only 1 well-developed locule, the others reduced or rudimentary (sometimes 2 globose lobes are developed); pericarp chartaceous, smooth and shiny. Seeds globose, black, shiny, ca. 8 mm diameter; embryo with straight cotyledons. Pantropics; ca. 14 species, 1 in Venezuela. Sapindus saponaria L., Sp. Pl. 367. 1753. —Jaboncillo, Para-para. Tree to 5 m tall; berry green to yellowish, with 1 or 2 globose lobes, usually one of them partially developed; seed surrounded by sticky pulp. Dry or open habitats, near sea level to 200 m; Delta Amacuro (Río Orinoco),

Bolívar (Río Orinoco), Amazonas (San Carlos de Río Negro). Widespread and often cultivated elsewhere in Venezuela and Neotropics, introduced into Old World tropics. ◆Fig. 62. Sapindus saponaria is widely used for soap and fish poison, and is often cultivated along streets.

Fig. 62. Sapindus saponaria

76

S APINDACEAE

11. SERJANIA Plum. ex Mill., Gard. Dict. abr. ed. 4. 1754. by Pedro Acevedo-Rodríguez Woody or herbaceous vines, often producing milky sap. Cross section of stem with a single or multiple vascular cylinders, usually 3, but sometimes 8 or 10. Leaves ternately compound (ternate, biternate, or triternate), or pinnately 5foliolate; stipules small and early deciduous. Inflorescences axillary thyrses or terminal thyrsoids with widely spaced cincinni as secondary branches. Flowers zygomorphic (plants polygamous-dioecious). Calyx of 4 or 5 unequal sepals; petals 4, mostly spatulate, with basally adnate hood-shaped appendage; disk unilateral, 2or 4-lobed; receptacle sometimes enlarged into a short androgynophore. Stamens 8, slightly exserted; filaments unequal; anthers dorsifixed, without an apical appendage. Ovary of 3 uniovular carpels; style terminal, with 3 stigmatic branches. Fruits schizocarps, splitting into 3 samaroid mericarps, with a proximal wing. Seeds lenticular to globose; embryo with straight or biplicate cotyledons. Tropical and subtropical areas of the New World; ca. 230 species; 16 in Venezuela, 9 of these in the flora area. Key to the Species of Serjania 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

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

6. 7(6).

7.

Cross section of stem with numerous vascular cylinders ....................... 2 Cross section of stem with a single vascular cylinder ............................. 9 Cross section of stem with 8 or 10 peripheral cylinders, disposed in a circular way around the central cylinder ................................................. 3 Cross section of stem with 3 or 5(6) peripheral cylinders, disposed in a triangular way around the central cylinder ........................................ 5 Plant densely pubescent; lower surface of leaflets canescent ......... S. adusta Plant glabrous or sparsely pubescent; lower surface of leaflets glabrous or with few trichomes ............................................................................ 4 Leaflets long-acuminate at apex, lineate (with conspicuous dark lines or vein pattern) on lower surface; cincinni usually elongate (1 cm or more) and scorpioid; bracts and bracteoles with glandular trichomes on margins; sepals lineate ......................................................... S. pyramidata Leaflets acute or obtuse, not lineate; cincinni usually < 1 cm, and not scorpioid; bracts and bracteoles variously pubescent but not glandular at margin; sepals not lineate ................................................ S. caracasana Stems with 5 or 6 peripheral cylinders; leaflets > 9 ................. S. clematidea Stems with 3 peripheral cylinders; leaflets 9 (the leaves biternate) ...... 6 Plants scarcely woody (not robust), not producing milky sap; calyx with third and fifth sepals connate 3/4 to whole length; receptacle enlarged into a short androgynophore; disk 2-lobed ............................ S. communis Plants robust, usually producing milky sap; calyx with third and fifth sepals distinct; receptacle not enlarged as above; disk 4-lobed ......... 7 Leaflets glabrous, coriaceous, 5–10 cm long, serrate only on upper part of blade, apiculate at apex, the teeth glandular; leaf rachis usually winged; stems obtusely angled .......................................... S. paucidentata Leaflets pubescent on both sides, chartaceous, 2–6 cm long, serrate on upper 1/2 of blade; leaf rachis not winged; stems sharply angled ....... 8

Serjania 77

8(7). 8.

9(1). 9.

Lower surface of leaflets ferruginous-tomentose; terminal leaflet rhombic; inflorescence axis ferruginous-tomentose .......................... S. rhombea Lower surface of leaflets sparsely pubescent, especially along veins; terminal leaflet lanceolate or ovate (rarely rhombic); inflorescence axis canescent ................................................................................ S. clematidea Stems terete; bracts and bracteoles with glandular trichomes on margins ........................................................................................... S. membranacea Stems 5-angled; bracts variously pubescent, but not as above ............................................................................................... S. grandifolia

Serjania adusta Radlk., Monogr. Serjania 156. 1875. Woody vine producing milky sap; leaves biternate or 5-pinnately compound; leaflet margins with rounded glandular teeth; inflorescence cincinni scorpioid, ca. 1 cm long. Forest slopes, open areas, 100–400 m; Bolívar (Serranía Baraguán). Widespread elsewhere in Venezuela; Cuba. Serjania caracasana (Jacq.) Willd., Sp. Pl. 2: 465. 1799. —Paullinia caracasana Jacq., Pl. Hort. Schoenbr. 1: 52. 1797. Serjania mariquitensis Triana & Planch., Ann. Sci. Nat. Bot. ser. 4, 18: 347. 1862. Serjania ierensis Britton, Bull. Torrey Bot. Club 50: 52. 1923. Woody vine to 10 m long, producing abundant milky sap, glabrous or pubescent; cross section of stems with 6–8 peripheral vascular cylinders; leaves trifoliolate, pinnately 5foliolate, or biternate. Moist open areas, woods, cloud forests, near sea level to 600 m; scattered in northern Bolívar. Widespread elsewhere in Venezuela; Neotropics. ◆Fig. 63. Serjania caracasana is a polymorphic species. Serjania clematidea Triana & Planch. Ann. Sci. Nat. Bot. ser. 4, 18: 349. 1862. Woody vine to 6–7 m long, glabrescent; stems with 6 sharp ribs; sepals canescent; petals cream or yellowish; mericarps reddish when mature, with ferruginous-tomentose locules. Scrub forests, 50–300 m; Bolívar (Cerro San Borja along lower Río Suapure, Río Villacoa, Serranía de los Pijiguaos), Amazonas (Caño Yutajé, Isla Carestia in Río Orinoco, near Puerto Ayacucho, Río Orinoco, northwest of Samariapo). Widespread elsewhere in Venezuela; Colombia, Brazil (Amazonas). ◆Fig. 64.

Serjania communis Cambess. in A. St.Hil. et al., Fl. Bras. Merid. 1: 362. 1827 [1828]. Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; 3 varieties, 1 in Venezuela. S. communis var. communis Herbaceous or slightly woody vine, producing watery sap (not milky), to 5 m long; mericarps with flattened cocci. Riversides, 100–300 m; Amazonas (along rapids on upper Río Orinoco). Widespread elsewhere in Venezuela; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 65. Serjania grandifolia Sagot ex Radlk., Monogr. Serjania 166. 1875. Woody vine to 6 m long, with abundant milky sap; anthers usually pink or light violet; mericarps reddish when mature, with globose locules. Moist forests, open areas, 800–1300 m; Bolívar (El Dorado, Río Cuyuní). Guyana, Suriname, French Guiana, Peru. Serjania membranacea Splitg., Tijdschr. Naturl. Gesch. Physiol. 9: 105. 1842. Woody vine to 20 m long; stems usually with reddish tinge and hollow pith; mericarps reddish, with subglobose locules, wing very narrow below the locule. Open areas, thickets, along rivers or roadsides, 50– 1100 m; Delta Amacuro (Río Amacuro, Sierra Imataca), Bolívar (between El Dorado and La Gran Sabana, Río Venamo, Ueitepui), Amazonas (San Simón de Cocuy). Widespread elsewhere in Venezuela; Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Amazonian Brazil. Serjania paucidentata DC., Prodr. 1: 603. 1824. —Rabo de iguana, Tres filos.

78

S APINDACEAE

Fig. 63. Serjania caracasana

Fig. 64. Serjania clematidea

Serjania 79

Fig. 65. Serjania communis var. communis

Fig. 66. Serjania pyramidata

80

S APINDACEAE

Stems triangular, with 5 sharp ribs; cross section of stems with 3 triangular peripheral cylinders; mericarps pinkish tinged, with subglobose ferruginous-tomentose locules. Roadsides, forest margins, lowland forests, 50–900 m; Delta Amacuro (near Los Castillos de Guayana, Río San José), Bolívar (road between Anacoco and Las Trincheras, Río Toro). Widespread elsewhere in Venezuela; Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, French Guiana, Brazil. Serjania pyramidata Radlk., Monogr. Serjania 155. 1875. —Bejuco moreno. Stems producing milky sap; cross section of stems with 10 peripheral steles, surrounding the central one; cincinni elongate,

scorpioid; mericarps reddish tinged, with subglobose glabrous locules. Open areas, moist lowland forests, 50–400 m; Delta Amacuro (lower Río Orinoco), Bolívar (Reserva Forestal Imataca, Río Cuyuní, Río Grande, road to San Pedro, road between Upata and San Félix). Widespread elsewhere in Venezuela; Nicaragua, Costa Rica, Panama Colombia, Peru, Brazil. ◆Fig. 66. Serjania rhombea Radlk., Monogr. Serjania 324. 1874. Woody vine with sharply angled, hirsute stems; leaves biternate, terminal leaflets rhombic. Savannas, forest edges, 50–200 m; Amazonas (Caño Yureba). Widespread elsewhere in Venezuela; Honduras to Panama, Colombia, Ecuador, Peru.

12. TALISIA Aubl., Hist. Pl. Guiane 349. 1775. by Pedro Acevedo-Rodríguez Unbranched shrubs or small to large trees. Leaves pinnately compound, with a rudimentary terminal leaflet; stipules absent; leaflets opposite or alternate, usually with swollen petiolules. Inflorescences axillary or terminal thyrses, with flowers in monochasia or dichasia. Flowers 5-merous, actinomorphic, staminate or pistillate (plants monoecious). Calyx usually cup-shaped; petals reflexed or ascending, with adnate appendage; appendages petaloid, sericeous, and basally adnate or marginal; disk annular, 5- or 8-lobed. Stamens 5–8, slightly exserted; filaments of equal or unequal length; anthers lanceolate or ellipsoid, basifixed. Ovary 3-carpellate, each carpel with a single ovule; style subulate, distally crowned by a capitate to cylindrical stigmatic surface. Fruits 1(–3)-seeded, ellipsoid to globose, berries, with leathery, fleshy, or woody pericarp. Seeds with fleshy outer coat; embryo with fleshy, transversely straight cotyledons. Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 52 species; 16 in Venezuela, 13 of these in the flora area. The outer seed coat or pericarp of Talisia is edible and tasty. A few species have been used as a fish poison. Key to the Species of Talisia 1. 1. 2(1). 2. 3(2).

3.

Leaves with 4–8(–10) leaflets ................................................................... 2 Leaves with 8–18 leaflets .......................................................................... 6 Leaflets obovate or oblanceolate, abruptly acuminate ............................ 3 Leaflets lanceolate, oblong-lanceolate, or oblong-elliptic, acuminate or obtuse at apex ........................................................................................ 4 Leaflets 4–6, obovate, abruptly acuminate; stems smooth to striate; inflorescences terminal, > 25 cm long; fruit glabrous; mesocarp ca. 1 mm thick ............................................................................................. T. obovata Leaflets 4–8, oblanceolate or seldom elliptic, with blunt acumen at apex;

Talisia 81

stems grayish, with many rounded lenticels; inflorescences axillary, usually cauliflorous, < 10 cm long; fruit appressed-pubescent; mesocarp ca. 2 mm thick ............................................................... T. hexaphylla 4(2). Leaflets chartaceous .................................................................... T. chartacea 4. Leaflets coriaceous .................................................................................... 5 5(4). Leaflets 4–10, obtuse at apex; secondary and tertiary veins not prominent; fruit glabrous ................................................................... T. laevigata 5. Leaflets 4 or 5, acuminate at apex; secondary and tertiary veins prominent on lower surface; fruit appressed-pubescent ........................ T. firma 6(1). Inflorescence rachis, sepals, and lower side of leaflets bearing glandularstipitate trichomes ................................................................. T. hemidasya 6. Plant glabrous, inflorescence rachis tomentose or glabrous ................... 7 7(6). Leaflets caudate at apex ................................................................. T. caudata 7. Leaflets acuminate, acute, obtuse, or rounded at apex ........................... 8 8(7). Leaflets obtuse or rounded and retuse, < 15 cm long ....................... T. retusa 8. Leaflets acuminate or acute, at least some > 15 cm long ........................ 9 9(8). Stamens 8 ...................................................................................... T. acutifolia 9. Stamens 5 ................................................................................................. 10 10(9). Secondary veins slightly prominent on both surfaces; disk glabrous; filaments glabrous or puberulent ............................................. T. macrophylla 10. Secondary veins sunken on upper surface, prominent on lower surface; disk hirsute; filaments pilose ...................................................... T. nervosa Talisia acutifolia Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 349. 1878. Talisia medrii Guarim, Acta Amazon. 9: 235. 1979. Tree 2–3 m tall; leaves 15–50 cm long; leaflets 6–12, coriaceous, glabrous; leaf rachis terete or slightly angular, puberulent; inflorescence ca. 25 cm long, tomentose; calyx tomentose; disk hirsute. Seasonally flooded or disturbed forests, 100–200 m; Amazonas (Río Casiquiare, San Carlos de Río Negro). Amazonian Brazil (Río Negro). Talisia caudata Steyerm., Ann. Missouri Bot. Gard. 75: 1072. 1988. Tree ca. 2 m tall; leaves to 60 cm long; leaflets 16–24, alternate; leaf rachis puberulous; fruit 2.2–2.4 cm long, ellipsoid, puberulous. Lowland forests, 100–200 m; Amazonas (Cerro Sipapo, Río Cuao). Endemic. Talisia chartacea Acev.-Rodr., BioLlania Edición Especial No. 6: 144. 1997. Tree to 30 m tall, glabrous throughout; leaflets 4–6, oblong-lanceolate, chartaceous; berry fleshy(?), purplish green, with thickened mesocarp. Lowland evergreen forests

on consolidated sandy grounds, 200–300 m; Bolívar (southwest of Cerro Guaiquinima), Amazonas (between Atabapo and La Esmeralda). Endemic. Talisia firma Radlk., Sitzungsber. Math.Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 346. 1878. Tree 4–20 m tall; leaves < 20 cm long; leaflets 4 or 5, opposite (if 5, one subterminal), lanceolate, coriaceous, discolor; sepals greenish yellow, covered with appressed rusty trichomes; stamens 8, unequal; fruit ellipsoid, ca. 2 cm long, densely covered with rusty trichomes. Gallery or seasonally flooded forests, flooded black-water swamps, 50–200 m; frequent in Amazonas. Endemic. Talisia hemidasya Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 349. 1878. —Cotoperí montañero, Cotoperíz montañero, Cotupería. Talisia glandulifera Steyerm., Ann. Missouri Bot. Gard. 75: 1072. 1988. Tree to 20 m tall; leaves to 50 cm long; leaflets (5–)8–16, opposite or alternate, falcate, subcoriaceous; leaf rachis terete or angular; inflorescence hirsutulose, with glan-

82

S APINDACEAE

dular trichomes; fruits ca. 2.5 cm long, nearly globose, ferruginous-tomentose. Lowland evergreen forests, near sea level to 100 m; Delta Amacuro (Los Castillos, Río Grande), Bolívar (Caserío Los Rosos northeast of Upata, El Palmar, El Tigre to La Soledad road). Guyana, Suriname, French Guiana. ◆Fig. 69. Talisia hexaphylla Vahl, Eclog. Amer. 2: 29. 1798. Panama, Colombia, Venezuela, Trinidad, Guyana, French Guiana, Peru, Amazonian Brazil, Bolivia; 3 subspecies, 1 in Venezuela. T. hexaphylla subsp. hexaphylla. —Cotoperí, Cotoperíz. Talisia panamensis Pittier, Contr. U.S. Natl. Herb. 20: 129. 1918. Talisia cararensis Cuatrec., Rev. Acad. Colomb. Cienc. 8: 306. 1951. Cauliflorous tree 10–30 m tall; leaves to 35 cm long; leaflets 4–8, coriaceous, lighter on lower surface; fruits globose, ca. 2.5 cm long. Gallery forests, forested slopes, disturbed lowland forests, wooded areas bordering savannas, 100–600 m; Delta Amacuro (Río Cuyubini), Bolívar (Altiplanicie de Nuria, Cerro Cotorra, south of El Dorado, Represa Guri, Río Paragua). Panama, Colombia, Trinidad, Guyana, Peru, Amazonian Brazil, Bolivia. Talisia laevigata Acev.-Rodr., BioLlania Edición Especial No. 6: 145. 1997. —Mamoncillo, Palo azul. Tree ca. 16 m tall; leaves < 15 cm long; leaflets 6–8, oblong-elliptic, involute, coriaceous, glabrous, glossy; inflorescence terminal, ca. 30 cm long; fruit glabrous. Habitat and elevation not given; Bolívar (Reserva Forestal La Paragua). Endemic. Talisia macrophylla (Mart.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 347. 1878 [1879]. —Cupania macrophylla Mart., Flora 21(2): 74. 1838. Talisia allenii Croat, Ann. Missouri Bot. Gard. 63: 527. 1976. Talisia pentantha Steyerm., Ann. Missouri Bot. Gard. 75: 1075. 1988. Talisia sancarlosiana Steyerm., Ann. Missouri Bot. Gard. 75: 1075. 1988. Treelet 3–5 m tall; stems terete and fur-

rowed; leaves 45–60 cm long; leaflets 8–16, opposite or alternate, 11–30 cm long, oblong or elliptic, glabrous or puberulent, chartaceous, with veins prominent on lower side; fruit globose, smooth, 2–2.5 cm long. Rain forests, gallery forests, 100–500 m; Bolívar (Canaima, 16 km northwest of Kilómetro 88, 3 km north of San Carlos de Río Negro), Amazonas (Raudal de los Guaharibos on upper Río Orinoco, between San Carlos and Solano, near Yutajé). Costa Rica, Panama, Colombia, the Guianas, Ecuador, Peru, Brazil. ◆Fig. 68. Talisia nervosa Radlk., Smithsonian Misc. Collect. 61(24): 4. 1914. Talisia grandifolia Cuatrec., Rev. Acad. Colomb. Cienc. 8: 306. 1951. Talisia tirirensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 448. 1967. Talisia dwyeri Croat, Ann. Missouri Bot. Gard. 63: 528. 1976. Talisia amaruyana Steyerm., Ann. Missouri Bot. Gard. 75: 1070. 1988. Talisia amaruayana Steyerm., Ann. Missouri Bot. Gard. 75: 1070. 1988. Tree to 8 m tall; leaves to 60(–150) cm long; leaflets 8–14, elliptic to oblong, shortacuminate at apex, usually drying dark brown; stamens 5; fruit globose to ellipsoid, smooth, ca. 2 cm long. Lowland and submontane evergreen forests, 100–800 m; Bolívar (Amaruay-tepui, Macizo del Chimantá [Chimantá-tepui], Río Tírica), Amazonas (Brazo Casiquiare, Cerro Huachamacari, upper Río Cuao, Río Yatua, San Carlos de Río Negro, Yavita to Maroa road). Costa Rica, Panama, Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 67. Talisia obovata A.C. Sm., Brittonia 2: 154. 1936. Tree 5–8 m tall; leaves to 40 cm long; leaflets obovate, 4–6, abruptly acuminate, finely reticulate-veined; sepals acute; fruits green, ellipsoid. Nonflooded lowland evergreen forests, 100–200 m; Amazonas (upper Río Orinoco, Yavita to Maroa road, 4.5 km northeast of San Carlos de Río Negro). Peru, Brazil. Talisia retusa R.S. Cowan, Brittonia 7: 403. 1952. —Tiestigo. Talisia heterodoxa Steyerm., Ann. Missouri Bot. Gard. 75: 1073. 1988.

Talisia 83

Fig. 67. Talisia nervosa

Fig. 68. Talisia macrophylla

84

S APINDACEAE

Fig. 69. Talisia hemidasya

Tree 10–23 m tall and to 40 cm diameter; leaflets 8–12, opposite or alternate, chartaceous or subcoriaceous, elliptic or obovate, glabrous; fruits globose, ca. 3 cm long, glabrous, with leathery pericarp. Nonflooded forests, 100–300 m; Bolívar (Represa Guri), Amazonas (Río Guayapo). Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia.

13. THINOUIA Triana & Planchon, Ann. Sci. Nat. Bot. ser. 4, 18: 368. 1862. by Pedro Acevedo-Rodríguez Lianas. Stems terete, lenticellate, becoming warty with age, cross section of young stem with a single vascular cylinder, older stems with many cortical steles. Leaves trifoliolate; petioles not winged; leaflets coriaceous. Inflorescences umbellike thyrses (congested cincinni along a shortened rachis) seldom bearing tendrils, axillary or aggregate into terminal thyrsoid inflorescences. Flowers 5-merous, small (3–4 mm), actinomorphic, bisexual. Calyx cup-shaped; petals obovate to spatulate, with a pair of short, marginal appendages; disk annular. Stamens 6–8, exserted; filaments of equal length; anthers ellipsoid, dorsifixed. Fruit splitting into 3 mericarpic units, leaving a common carpophore attached to the pedicel. Seeds nearly spherical; embryo with abaxial cotyledon bent over the biplicate adaxial one. Neotropics; 12 species, 2 in Venezuela, 1 of these in the flora area.

Toulicia 85

Thinouia myriantha Triana & Planch., Ann. Sci. Nat. Bot. ser. 4, 18: 369. 1862. Leaflets coriaceous, remotely serrate toward distal portion, lower pair of secondary veins starting at base of blade, base rounded or truncate, apex apiculate. Evergreen lowland forests, 50–400 m; Delta Amacuro (Río Grande). Yaracuy; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana. ◆Fig. 70.

Fig. 70. Thinouia myriantha

14. TOULICIA Aubl., Hist. Pl. Guiane 359. 1775. by Pedro Acevedo-Rodríguez Small unbranched trees. Leaves pinnately compound, with rudimentary terminal leaflet; leaflets usually falcate, opposite or alternate, with prominent petiolules. Inflorescences terminal panicles or axillary thyrses, with flowers in lateral cincinni. Flowers zygomorphic, staminate or pistillate (plant polygamous-dioecious). Calyx of 5 distinct, unequal sepals; petals 4, distinct, with a petaloid appendage on adaxial surface, the appendages bifid or marginal; disk unilateral, semiannular. Stamens 8; filaments of equal length; anthers ellipsoid, dorsifixed. Ovary 3-carpellate, each carpel with a single ovule; style distal, with 3 stigmatic branches. Fruits splitting into 3 samaroid mericarps, with a basal wing and a papery, inflated locule. Seeds dark brown, subglobose; embryo with abaxial cotyledon bent over biplicate adaxial one. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 12 species, 3 in Venezuela, all in the flora area.

86

S APINDACEAE

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

Leaflets long-petiolulate (> 1 cm long) ...................................... T. petiolulata Leaflets short-petiolulate (< 5 mm long) .................................................. 2 Leaflets oblong or oblong-elliptic, to 25 cm long; petals 5–5.5 mm long; fruits to 5 cm long ................................................................... T. guianensis Leaflets ovate, ovate-lanceolate, or elliptic, to 10 cm long; petals to 3.5 mm long; fruits to 3 cm long .............................................. T. pulvinata

Toulicia guianensis Aubl., Hist. Pl. Guiane 359. 1775. —Carapo blanco. Small to medium-sized tree; wood yellowish; leaves 0.5–1 m long; lower surface of leaflets usually brown. Seasonally flooded lowland forests, riversides, 50–200 m; Delta Amacuro (Río Grande), Bolívar (Río Caura, Río Orinoco, Río Suapure), Amazonas (Río Negro). Guyana, Suriname, French Guiana. ◆ Fig. 71.

Toulicia petiolulata Radlk., Notizbl. Königl. Bot. Gart. Berlin 6: 155. 1915. Tree to 20 m tall; leaflets 15–25 cm long, long-petiolulate (0.8–2.5 cm); schizocarps green. Forested river banks, 200–300 m; Amazonas (Río Ocamo). Amazonian Brazil. Toulicia pulvinata Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 371. 1878. —Carapo blanco, Sarrapia.

Fig. 71. Toulicia guianensis

Urvillea 87

Tree to 25 m tall, with gray bark and whitish wood; leaves 0.3–0.6 m long. Riversides, secondary forests, 100–200 m; Bolívar

(Reserva Forestal La Paragua), Amazonas (Río Orinoco). Guyana, Suriname, French Guiana.

15. URVILLEA Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 105. 1821. by Pedro Acevedo-Rodríguez Herbaceous to woody vines. Stems terete and lenticellate, becoming 3-lobed with age, producing milky sap; cross section of stem with a single vascular cylinder. Leaves ternately (trifoliolate) or biternately compound; stipules minute, deciduous. Inflorescence a thyrse with closely spaced cincinni. Flowers zygomorphic, bisexual. Calyx of 5 unequal sepals; petals 4, distinct, usually < 4 mm long, spatulate, with an adnate, hood-shaped appendage on adaxial surface; disk unilateral, 4lobed, receptacle enlarged into a short androgynophore. Stamens 8; filaments unequal; anthers sometimes with an apical appendage. Ovary of 3 uniovular carpels; style terminal with 3 stigmatic branches. Fruits thin, papery, semi-inflated capsules, with narrow marginal wing. Seeds subglobose, black, with a heartshaped or reniform white arillode around the hilum; embryo with the abaxial cotyledon bent over the biplicate adaxial cotyledon. Central America, South America except Chile; ca. 12 species, 1 in Venezuela. Urvillea ulmacea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 106. 1821. Leaves trifoliolate, the lower surface ferruginous-tomentose; inflorescences densely flowered. Open areas along roads and riversides, 50–200 m; Bolívar (west of Río Aro, Río Suapure), Amazonas (Río Cataniapo). Widespread elsewhere in Venezuela; Central America, South America except Chile. ◆Fig. 72.

Fig. 72. Urvillea ulmacea

88

S APINDACEAE

16. VOUARANA Aubl., Pl. Guiane. 2 (suppl.): 12, fig. 374. 1775. by Pedro Acevedo-Rodríguez Medium-sized trees. Leaves pinnately compound, with a rudimentary terminal leaflet; leaflets opposite or subopposite, the lower surface usually foveate at vein axils. Inflorescence axillary or terminal thyrsoids with flowers in lateral dichasia. Flowers 4- or 5-merous, actinomorphic or zygomorphic, bisexual or unisexual (plants polygamous-monoecious). Calyx of distinct, concave sepals; petals rhombic, shorter than the sepals, with 2 marginal appendages; disk annular and

Fig. 73. Vouarana anomala

S APOTACEAE 89

lobed. Stamens 6–8; filaments of nearly equal length, subulate and pubescent; anthers ellipsoid, dorsifixed. Ovary obclavate, 2-carpellate, each carpel with a single, basal ovule; stigma subulate and papillose. Fruit a 1- or 2-seeded, pyriform, woody capsule. Seeds ellipsoid with a basal aril. Venezuela, Guyana, Suriname, French Guiana, northern Brazil; 2 species, 1 in Venezuela. Vouarana anomala (Steyerm.) Acev.-Rodr., BioLlania Edición Especial No. 6: 146. 1997. —Toulicia anomala Steyerm., Ann. Missouri Bot. Gard. 75: 1076. 1988. Shrub 2–3 m tall; leaflets membranous,

lanceolate, falcate, foveate at veins axils; margins entire or crenate; flowers 4-merous, actinomorphic; stamens 6–8. Riversides, 100–200 m; Bolívar (Río Suapure 70–80 km above its mouth). Endemic. ◆Fig. 73.

SAPOTACEAE by Terrence D. Pennington and K. S. Edwards Trees or shrubs, rarely geoxylic (with large underground stems), suffrutices or lianas, sometimes spiny. Branches usually sympodial. Latex nearly always present in trunk, branches, and fruit, usually white, rarely yellow or blue. Indumentum nearly always of malpighiaceous trichomes (simple in Delypdora). Leaves spirally arranged or alternate and distichous, less frequently verticillate, simple, entire, and very rarely spinous; stipules present or absent; venation craspedodromous (secondary veins terminate at the leaf margin), eucamptodromous (secondary veins gradually decreasing apically inside the margin, or brochidodromous (secondary veins joined together by a series of submarginal loops, this sometimes ± straight and forming a submarginal vein). Petiole rarely bearing a pair of minute stipules. Inflorescences fasciculate or flowers occasionally solitary, axillary, ramiflorous, or cauliflorous, fascicles occasionally arranged along short leafless axillary shoots, forming a panicle, fascicle base sometimes developing into short densely scaly shoots (brachyblasts). Flowers bisexual or unisexual (plant monoecious or dioecious), actinomorphic. Calyx a simple whorl of 4–6 free or partly fused, imbricate or quincuncial sepals, or 6–11 sepals in a closely imbricate spiral, or with 2 whorls of 2–4 sepals and then the outer whorl valvate or only slightly imbricate; corolla rotate, cyathiform, or tubular, gamopetalous; tube shorter than, equal to, or exceeding the lobes; lobes 4–18, entire, lobed or partly divided, or divided to the base into 3 segments, median segment entire, 2 lateral or dorsal segments entire, laciniate, or shallowly or deeply divided. Stamens 4–35(–43), fixed in the lower or upper half of the corolla tube or at the base of the lobes, rarely free; in a single whorl opposite the corolla lobes, or (when more numerous than the corolla lobes) some opposite and some alternate with the corolla lobes, or sometimes several stamens clustered opposite each lobe, or arranged in 2 or 3 alternating whorls within the corolla tube; exserted or included; filaments often geniculate in bud, free or rarely fused into a staminal tube, or partially fused to the

90

S APOTACEAE

staminodes; anthers often extrorse; staminodes 0–8(–12) in a single whorl alternating with the stamens or fixed in the corolla lobe sinus, simple or variously lobed, toothed, or divided, sometimes petaloid. Disk annular or patelliform, surrounding the ovary base and sometimes fused with it, or absent. Ovary superior, 1– 15(–30)-locular, loculi usually uniovulate, rarely 2(–5)-ovulate, placentation axile, basi-ventral, or basal; style simple, included or exserted, style head simple or minutely lobed. Fruit a berry or rarely a drupe, pericarp fleshy or less frequently leathery or woody. Seeds 1–many, globose, ellipsoid, oblong, often strongly laterally compressed, testa usually smooth, shining, free from the pericarp, less frequently roughened, wrinkled, or pitted, and then often adherent to the pericarp; scar adaxial, basi-ventral, or basal, narrow or broad, sometimes extending to cover most or all of the seed. Embryo vertical, oblique, or horizontal, with thin foliaceous or thick flat or plano-convex, usually free, cotyledons; radicle included or exserted; endosperm present or absent. Pantropics; 53 genera and ca. 1100 species, 9 genera and 85 species in the flora area. The genus Sarcaulus may eventually be found in the flora area, particularly S. brasiliensis (A. DC.) Eyma, which is known from Central America and the Guianas. The family Sapotaceae provides many economically important products: latex often used in dentistry and electrical insulation from balata and gutta percha (see discussion under Manilkara bidentata subsp. bidentata) and chewing gum; a heavy, hard, durable timber; and many edible fruits, such as sapote [Pouteria sapota (Jacq.) H.E. Moore & Stearn], caimito [Pouteria caimito (Ruiz & Pav.) Radlk.], and star apple, also called caimito [Chrysophyllum cainito L.]. Key to the Genera of Sapotaceae 1. 1. 2(1). 2. 3(2). 3. 4(2). 4. 5(4).

5.

Calyx of 2 whorls of (2)3(4) sepals, outer whorl usually valvate; corolla lobes divided into 3 segments ................................................ 6. Manilkara Calyx a single whorl of 4–6 imbricate or quincuncial sepals; corolla lobes undivided ............................................................................................... 2 Stipules present, well developed, leaving a conspicuous scar ................ 3 Stipules absent .......................................................................................... 4 Flowers pedicellate; staminodes present; seed with dull rough testa; scar broad, covering to 2/3 of seed surface .............................. 1. Chromolucuma Flowers sessile; staminodes absent; seed smooth, shining; scar usually narrow ...................................................................................... 4. Ecclinusa Staminodes present, as many as the corolla lobes .................................. 5 Staminodes fewer than corolla lobes or absent ....................................... 6 Leaves usually alternate and distichous; venation craspedodromous or brochidodromous, often with secondary and tertiary veins closely parallel and leaf appearing finely striate; seed laterally compressed, with endosperm .................................................................... 7. Micropholis Leaves spirally arranged; venation variable, but not closely parallel and leaves never finely striate; seed shape variable, usually without endosperm .................................................................................. 8. Pouteria

Chrysophyllum 91

6(4). 6. 7(6). 7. 8(6). 8. 9(8).

9.

Corolla tubular, campanulate, or globose, stamens included ................. 7 Corolla rotate or broadly cyathiform, stamens exserted ......................... 8 Ovary usually 2-locular; embryo with plano-convex cotyledons and included radicle, endosperm absent ......................................... 8. Pouteria Ovary usually 5-locular; embryo with thin foliaceous cotyledons, radicle exserted, copious endosperm .......................................... 2. Chrysophyllum Ovary 1-locular, locule with 2 basal ovules; seed scar basal or basiventral ......................................................................................... 3. Diploon Ovary 2–6-locular, locules 1-ovulate, ovules axile; seed scar adaxial .... 9 Leaves spirally arranged, usually minutely punctate on lower surface; inflorescence mostly axillary; filaments not strongly narrowed below anther insertion; ovary 2- or 3(4)-locular; fruit a berry ....... 5. Elaeoluma Leaves often opposite or verticillate, not punctate; inflorescence usually cauliflorous or ramiflorous; filaments strongly narrowed below anther insertion; ovary usually 5-locular; fruit a drupe ..................... 9. Pradosia

1. CHROMOLUCUMA Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 160. 1925. Trees. Leaves spirally arranged; stipules present, large; venation eucamptodromous. Flowers unisexual (plant monoecious or dioecious), pedicellate. Corolla cyathiform or shortly tubular, tube usually equaling or slightly longer than lobes (slightly shorter in staminate flowers of C. rubriflora). Stamens fixed in the upper half or at the top of the corolla tube, included; filaments short; staminodes present. Disk absent. Ovary broadly truncate, 2–5-locular. Seed with dull rough testa, and broad adaxial scar covering up to 2/3 of seed; embryo with plano-convex, free cotyledons, radicle slightly exserted; endosperm absent. Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 2 species, 1 in Venezuela. Chromolucuma rubriflora Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 160, t. 18. 1925. —Mayabo. Tree 10–30 m tall, 30 cm diameter; latex scarce, yellow; flowers with orange-red calyx

and whitish corolla; fruit tawny. Evergreen lowland forests, swampy areas, 100–200 m; Amazonas (lower Río Baría, Río Cunucunuma, Río Negro). Central Amazonian Brazil. ◆Fig. 74.

2. CHRYSOPHYLLUM L., Sp. Pl. 192. 1753. Trees or shrubs. Leaves alternate and distichous or spirally arranged; stipules absent; venation brochidodromous or eucamptodromous, tertiary veins often parallel to the secondaries and descending from the margin, or oblique and closely parallel, or reticulate. Inflorescence axillary, ramiflorous, or cauliflorous. Flowers unisexual or bisexual, fasciculate, rarely solitary. Calyx a single whorl of (4)5(6) imbricate or quincuncial sepals, sometimes accrescent in fruit, frequently ciliate; corolla globose, campanulate or cylindrical, tube shorter than, equaling, or exceeding the lobes, lobes (4)5(–8), simple. Stamens (4)5(–8), fixed in the lower or upper part of the corolla tube, included; anthers extrorse in bud, hairy or glabrous; staminodes usually absent, rarely present as small lanceolate or subulate structures in the corolla lobe sinuses, alternating with the stamens. Ovary 4- or 5(–12)locular, placentation axile; style included. Fruit l–many-seeded. Seed laterally compressed, with a narrow adaxial scar, sometimes extending around the base of

92

S APOTACEAE

Fig. 74. Chromolucuma rubriflora

Chrysophyllum 93

the seed, or not laterally compressed and then the scar broader, basi-ventral or adaxial; testa smooth and shining, or rough and then adherent to the pericarp. Embryo vertical, with thin foliaceous or thick flat cotyledons and exserted radicle, endosperm abundant or about equaling the thickness of the cotyledons. Pantropics; ca. 70 species, 11 in Venezuela, 8 of these in the flora area. Key to the Species of Chrysophyllum 1. 1. 2(1). 2. 3(1). 3. 4(3).

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

7(5).

7.

Leaves not clustered, alternate and distichous; seed scar broad ........... 2 Leaves clustered, spirally arranged; seed scar narrow ........................... 3 Stamens fixed at the apex of the corolla tube; anthers glabrous ................................................................................................ C. argenteum Stamens fixed in the lower half of the corolla tube; anthers sparsely villose ....................................................................................... C. sparsiflorum Seed coat rough, not shining, adherent to the pericarp .......................... 4 Seed coat smooth, shining, free from the pericarp .................................. 5 Leaves broadly oblanceolate or obovate, usually persistent, the lower surface rufous-brown-sericeous; fruit sometimes verrucose or echinate ............................................................................................ C. prieurii Leaves narrower, the lower surface not rufous-brown-sericeous; fruit never verrucose or echinate ............................................... C. amazonicum Sepals usually accrescent in fruit; seed scar adaxial and extending around the base ..................................................................................... 6 Sepals not accrescent in fruit; seed scar adaxial, not extending around the base .................................................................................................. 7 Indumentum of lower surface of leaf closely appressed or leaves glabrous; fruit subglabrous ............................................... C. sanguinolentum Indumentum of lower surface of leaf ferruginous-tomentose, with short, strongly crisped trichomes; fruit densely rufous-villose ........ .................................................................................. C. ucuquirana-branca Leaves usually broadly oblanceolate or obovate, apex obtuse, rounded, or emarginate; venation usually brochidodromous; secondary veins 7– 10 pairs; corolla tube exceeding the lobes ............................ C. pomiferum Leaves usually elliptic, apex usually narrowly attenuate or acute; venation usually eucamptodromous; secondary veins 13–20 pairs; corolla tube shorter than the lobes ............................................... C. lucentifolium

Chrysophyllum amazonicum T.D. Penn., Fl. Neotrop. Monogr. 52: 595. 1990. —Waremba-dek. Tree to 20 m tall; latex copious, white; flowers greenish yellow or green; fruit ripening through yellow to orange. Lower montane forests, 300–800 m; Bolívar (upper Río Paragua). Amazonian Colombia, Guyana, Peru, Amazonian Brazil. Chrysophyllum argenteum Jacq., Enum. Syst. Pl. 15. 1760. Costa Rica, Panama, Greater and Lesser

Antilles, Colombia, Venezuela, north and central Amazonian Brazil; 5 varieties, 3 in Venezuela, 1 of these in the flora area. C. argenteum subsp. auratum (Miq.) T.D. Penn., Fl. Neotrop. Monogr. 52: 545. 1990. —Chrysophyllum auratum Miq. in Mart., Fl. Bras. 7: 97. 1863. —Caimito, Caimito largo, Caimito morado, Veru caimito. Tree 4–20 m tall, 10–30 cm diameter; bark gray-brown, shallowly fissured; latex abundant, milky; wood soft; lower surface of

94

S APOTACEAE

leaves usually sericeous with golden trichomes; flowers pale green-yellow; corolla tube 3–4 times as long as lobes; fruit usually ellipsoid, 1–2.5 cm long; 1-seeded; shiny black-purple on maturing. Evergreen lowland forests, semi-humid forests and dry seasonal forests along streams, disturbed forests, near sea level to 400 m; Delta Amacuro (Río Cuyuní, Serranía de Imataca), northern Bolívar, Amazonas (Salto Tencua). Widespread elsewhere in Venezuela; Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil. The fruits of Chrysophyllum argenteum are edible and sweet, and the timber is used for construction. Chrysophyllum lucentifolium Cronquist, Bull. Torrey Bot. Club 73: 304. 1946. Panama, Colombia, Venezuela, Ecuador, Peru, eastern coastal and Amazonian Brazil; 2 subspecies, 1 of these in Venezuela. C. lucentifolium subsp. pachycarpum Pires & T.D. Penn., Fl. Neotrop. Monogr. 52: 607. 1990. —Capure, Capurillo, Chaparro manteco, Palo manteco. Tree 15–41 m tall, 20–61 cm diameter; trunk fluted, buttresses 0.6–3.4 m high, outer bark rough, gray, hard; inner bark light orange or brown; latex sticky, white; flowers pale green-yellow, aromatic; fruit rough, ovoid, orange-yellow, fleshy, edible, 3.5–7.5 cm long, 5-seeded. Evergreen lowland to lower montane forests, savannas, 50–300 m; Amazonas (Guara near San Juan Manapiare), northern Bolívar. Northern Venezuelan Coastal Cordillera, Mérida; Panama, Colombia, Ecuador, French Guiana, Amazonian Peru, Brazil, Bolivia. Chrysophyllum pomiferum (Eyma) T.D. Penn., Fl. Neotrop. Monogr. 52: 602. 1990. —Achrouteria pomifera Eyma, Recueil Trav. Bot. Néerl. 33: 193, fig. 3. 1936. —Capurillo. Tree 6–35 m tall, 15–33 cm diameter; outer bark brown, soft, and flaky, inner bark cream-colored; latex thin, sticky, white; flowers greenish yellow, calyx and corolla light green; fruit yellow; sap white. Evergreen to semideciduous lowland to lower montane forests, 100–900 m; Bolívar (northern portions, northwestern Gran Sabana), Ama-

zonas (Río Mawarinuma). Apure; Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, and Brazil. ◆Fig. 76. The fruits of Chrysophyllum pomiferum are edible, the latex has been used for adulterating chicle, and the timber is hard and resistant to decay. Chrysophyllum prieurii A. DC., Prodr. 8: 161. 1844. —Cuyuri, Palo de cuyure, Palo triste. Tree 15–25 m tall, 40 cm diameter; bark brown, scaly, sometimes fissured; latex scarce, yellow-white; flowers with agreeable scent; fruit globose, fleshy, 4- or 5-seeded, yellow to orange. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro, Trapichote). Panama, Colombia, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil. ◆Fig. 77. The very hard wood of Chrysophyllum prieurii is used locally in construction. Chrysophyllum sanguinolentum (Pierre) Baehni, Boissiera 11: 74. 1965. —Ragala sanguinolenta Pierre, Not. Bot. Sapot. 60. 1891. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil; 3 subspecies, all in the flora area. Key to the Subspecies of C. sanguinolentum 1. Lower leaf surface closely and finely appressed-puberulous with ferruginous or whitish trichomes; fruiting calyx usually strongly accrescent (sepals strongly thickened, ca. 1 × 1.5 cm) ........................ ...................................... subsp. balata 1. Lower leaf surface glabrous; fruiting calyx weakly accrescent (sepals not or only weakly thickened, < 1 cm long) ... 2 2. Petiole usually < 2 cm long; leaves obovate, with rounded or tapered base ...................... subsp. sanguinolentum 2. Petiole usually 2–4 cm long; leaves usually broadly elliptic or broadly ellipticoblong with rounded or truncate base ................................... subsp. spurium C. sanguinolentum subsp. balata (Ducke) T.D. Penn., Fl. Neotrop. Monogr. 52: 587. 1990. —Ecclinusa balata Ducke,

Chrysophyllum 95

Rev. Int. Bot. Appl. Agric. Trop. 10: 850. 1930. —Saunan-yek (Arekuna), Tu-kuda-mo, Ucuquirana, Ucuquirana de selva baja. Tree 15–30 m tall, 17.5–50 cm diameter; narrow buttresses; outer bark gray to dark brown, slightly rough with shallow longitudinal cracks; latex copious, sticky, milky white; flower buds light green-coppery and pubescent; mature fruit yellow. Riverine forests, seasonally dry forests, wet mossy forests, seasonally flooded forests, palm forests, (100–)1600–2000 m; Bolívar (Macizo del Chimantá, Ptari-tepui, headwaters of Río Pacairao, Sororopán-tepui), Amazonas (Caño Yagua, Cerro Coro Coro, Río Matacuni, Río Mawarinuma, near San Carlos de Río Negro). Amazonian Colombia, Guyana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 78. The latex of Chrysophyllum sanguinolentum subsp. balata is used for balata, and the fruit pulp is edible.

C. sanguinolentum subsp. sanguinolentum. —Moyeja. Tree to 10 m tall; latex white; flowers greenish; fruits rough-skinned, pale brown, edible. Forests on lateritic soil, 100–200 m; Amazonas (San Carlos de Río Negro). Guyana, Suriname, French Guiana, Amazonian Peru and Brazil (Amapá). C.

sanguinolentum subsp. spurium (Ducke) T.D. Penn., Fl. Neotrop. Monogr. 52: 585. 1990. —Ecclinusa spuria Ducke, Bull. Mus. Hist. Nat. (Paris) sér. 2, 4: 743. 1932. —Pouteria spuria (Ducke) Pittier et al., Cat. Fl. Venez. 2: 283. 1947. —Moyeja, Temare, Temare montañero. Tree 15–22 m tall; latex abundant, yellowish white; yellow- or green-brown, edible fruit. Evergreen lowland, nonflooded forests, 100–200 m; Amazonas (around San Carlos de Río Negro). Colombia, Ecuador, Peru, Amazonian Brazil. Chrysophyllum sparsiflorum Klotzsch ex Miq. in Mart., Fl. Bras. 7: 90. 1863. —Pascualito. Chrysophyllum steyermarkii Monach., Fieldiana, Bot. 28: 480. 1953. Tree to 15 m tall; latex milky white; flowers pale green; fruit maturing yellowish.

Fig. 75. Chrysophyllum sparsiflorum

Fig. 76. Chrysophyllum pomiferum

96

S APOTACEAE

Fig. 77. Chrysophyllum prieurii

Fig. 78. Chrysophyllum sanguinolentum subsp. balata

Diploon 97

Semideciduous to evergreen lowland and riparian forests, 50–200 m; Bolívar (La Camilera 40 km west of El Manteco, Puerto El Jobal, Represa Guri, Upata). Anzoátegui; Guyana, Amazonian Brazil and Bolivia. ◆Fig. 75. Chrysophyllum ucuquirana-branca (Aubrév. & Pellegr.) T.D. Penn., Fl.

Neotrop. Monogr. 52: 589. 1990. —Ecclinusa ucuquirana-branca Aubrév. & Pellegr., Adansonia n.s. 1: 23. 1961. Tree to 20 m tall, 30 cm diameter; latex present; leaves discolorous. Evergreen lowland forests, 100–200 m; Amazonas (road from San Carlos de Río Negro to Solano). Central to northern Amazonian Brazil.

3. DIPLOON Cronquist, Bull. Torrey Bot. Club 73: 466. 1946. Trees. Leaves alternate and distichous, or weakly spirally arranged; stipules absent; venation brochidodromous, the secondaries joining below the margin to form a submarginal vein, intersecondaries long, usually extending to the margin, giving the leaves a slightly striate appearance. Corolla rotate, with very short tube greatly exceeded by the lobes. Stamens fixed at the top of the tube, exserted; filaments thickened basally; staminodes absent. Disk absent. Ovary glabrous, 1-locular with 2 basal ovules. Seed with small, broad, basal or basi-ventral scar; embryo with plano-convex, free cotyledons, radicle extending to the surface; endosperm absent. Southern Venezuela, Guyana, Amazonian Peru, Brazil (Amazonas, coastal regions from Alagoas to Paraná), Bolivia; 1 species.

Fig. 79. Diploon cuspidatum

98

S APOTACEAE

Diploon cuspidatum (Hoehne) Cronquist, Bull. Torrey Bot. Club 73: 466. 1946. —Chrysophyllum cuspidatum Hoehne, Ostenia 302, t. 8. 1933. —Cacho. Diploon venezuelana Aubrév., Mem. New York Bot. Gard. 23: 225. 1972.

Tree to 20 m tall, 40 cm diameter; wood hard, used for timber; flowers white. Evergreen lower montane forests, 200–500 m; Delta Amacuro (Serranía de Imataca). Other distribution as in genus. ◆Fig. 79.

4. ECCLINUSA Mart., Flora 22(Beibl. 1): 2. 1839. Trees or rarely shrubs. Leaves spirally arranged, usually loosely clustered at the shoot apex; stipules present, caducous, leaving a conspicuous scar; venation usually eucamptodromous, rarely brochidodromous, intersecondaries usually absent, tertiaries usually oblique, numerous, close, parallel, rarely reticulate or areolate. Inflorescence axillary or in the axils of fallen leaves. Flowers sessile, subtended by small ovate persistent bracts, usually unisexual (monoecious or dioecious). Calyx a single whorl of (4)5 free, quincuncial sepals; corolla small (usually < 5 mm long), campanulate or shortly tubular, the lobes usually exceeding the tube or rarely equaling it; lobes 5(–7), simple. Stamens 5(–7) included, usually fixed near halfway or in the upper half of the corolla tube, rarely in the lower half; filaments well-developed, free; anthers extrorse, glabrous; staminodes absent. Ovary (3–)5(–9)-locular, placentation axile or basi-ventral; style included. Fruit 1–several-seeded, often thin-walled and constricted between the seeds. Seed globose, ellipsoid, sometimes slightly laterally compressed or shaped like the segment of an orange, testa smooth, thin, shining; scar adaxial and nearly always extending around the base of the seed, usually narrow. Embryo vertical, with thick plano-convex cotyledons, radicle not exserted, extending to the surface; endosperm absent. Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 11 species, 8 in Venezuela, all in the flora area. Key to the Species of Ecclinusa 1. 1. 2(1). 2. 3(2).

3.

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

Tertiary venation reticulate or areolate; leaves often < 5 cm long ............................................................................................ E. atabapoensis Tertiary veins parallel, oblique, usually numerous; leaves nearly always > 5 cm long ............................................................................................. 2 Indumentum of young shoots and leaves fine, closely appressed ........... 3 Indumentum or young shoots, and sometimes leaves, of crisped or spreading trichomes .............................................................................. 4 Stipules 4–9 mm long; leaves 4.8–15(–20) cm long, usually elliptic or oblong-elliptic; corolla lobes and ovary loculi 5; fruit thin-walled, 1.5– 3 cm long ................................................................................ E. guianensis Stipules 1–2 cm long; leaves 16–42 cm long, usually oblanceolate; corolla lobes 5–7 and ovary loculi 7–9; fruit thick-walled, 5–6 cm long ................................................................................................. E. lanceolata Leaves strongly bullate ................................................................... E. bullata Leaves not bullate ..................................................................................... 5 Lower leaf blade densely and shortly ferruginous-tomentose, leaves sulcate ................................................................................................ E. ulei Lower leaf blade not ferruginous-tomentose, leaves usually not sulcate ................................................................................................................ 6

Ecclinusa 99

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

7.

Leaves 2.5–4 × as long as broad, oblanceolate, base acute to narrowly attenuate or cuneate; corolla ca. 3.5 mm long ........................ E. ramiflora Leaves about twice as long as broad, usually broadly elliptic, base obtuse to rounded; corolla 2–2.3 mm long. ...................................................... 7 Young shoots densely brown-lanate; leaves 7–14 cm long, lower blade densely brown-tomentose; fruit 1–1.5 cm long, thin-walled and constricted between the seeds, seed scar adaxial and extending around the base of the seed .............................................................. E. orinocoensis Young shoots finely golden-puberulous; leaves 15–27 cm long, lower blade sparsely brown-pubescent; fruit ca. 4 cm long, thick-walled, not constricted between the seeds, seed scar adaxial ................. E. parviflora

Ecclinusa atabapoensis (Aubrév.) T.D. Penn., Fl. Neotrop. Monogr. 52: 627. 1990. —Neoxythece atabapoensis Aubrév., Mem. New York Bot. Gard. 23: 221. 1972. Shrub 0.5–2 m tall, 5–10 cm diameter; leaves green-blue, the lower surface sometimes gray-green and glaucous; latex sticky, white; flower buds cream-colored to pale brown; flowers bright green, calyx red-ferruginous, corolla greenish, stamens yellow; fruit subglobose-globose, ca. 0.6 cm diameter and to 2 cm long, yellow, orange, redferruginous, or green. White-sand savannas, shrub islands in savannas, 100–200 m, Amazonas (Caño Yagua, Cerro Yapacana, Río Atabapo basin, Río Guainía, Río Guayapo, San Antonio). Colombia (Guainía). ◆Fig. 81. Ecclinusa bullata T.D. Penn., Fl. Neotrop. Monogr. 52: 634. 1990. —Cuquirana, Irana, Pendare, Pendare de sapo. Tree 7–25 m tall, ca. 25 cm diameter, sparsely branched with erect to ascending branches; leaves corrugate; thick, latex sticky, white; fruit velvety, orange-tawny, 5– 6 cm long when mature and containing latex. Evergreen lowland to montane forests, 100–1200 m; Amazonas (Cerro Marahuaka, Río Puruname, San Carlos de Río Negro, Yavita to Maroa). Brazil (Amazonas: Rio Negro). The latex of Ecclinusa bullata has been used for mixing with chicle. Ecclinusa guianensis Eyma, Recueil Trav. Bot. Néerl. 33: 203. 1936. —Chicle, Chicle de capure, Patillo-chicle, Pulguillo. Tree 6–30 m tall, 20–50 cm diameter;

trunk cylindrical, bark brown, rough with shallow fissures; latex milky; flowers fragrant, yellow-white; fruit yellow with small pellucid spots. Evergreen to semideciduous lowland to lower montane forests, 50–600 m; Delta Amacuro (Serranía de Imataca), northeastern Bolívar, Amazonas (Río Ventuari, Tencua). Distrito Federal, Miranda, Zulia; Guyana, Suriname, French Guiana, Ecuador, Amazonian Brazil, a few doubtful records from Panama and Peru. ◆Fig. 83. The latex of Ecclinusa guianensis is often mixed with balata. Ecclinusa lanceolata (Mart. & Eich.) Pierre, Not. Bot. Sapot. 57. 1891. —Passaveria lanceolata Mart. & Eich. in Mart., Fl. Bras. 7: 86, t. 38. 1863. Tree to 20 m tall with hard wood; latex present; fruit matures to orange-yellow. Montane forests, ca. 1300 m; Amazonas (Sierra Parima). Apure, Mérida, Táchira; Panama, Colombia, Guyana, French Guiana, Ecuador, Amazonian Peru and Brazil. ◆Fig. 82. Ecclinusa orinocoensis Aubrév., Mem. New York Bot. Gard. 23: 216. 1972. Slender monopodial treelet 1.5–5 m tall; latex abundant, sticky, white; buds cream, flowers yellow-green in sessile clusters; fruit round, yellow-green with brown pubescence, 5-lobed, ca. 2 cm diameter. Savannas, flooded sabanetas, periodically flooded sandy savannas, 100–200 m; Amazonas (basins of Cano Cotúa and Cano Yapacana). Endemic. ◆Fig. 80. Ecclinusa parviflora T.D. Penn., Fl. Neotrop. Monogr. 52: 628. 1990. —Temare de montaña.

100

S APOTACEAE

Tree 3–8 m tall, with a dense crown; latex sticky, white; flowers green; fruit brown, ca. 4 cm long. Evergreen lowland forests, flooded savannas, 50–200 m. Amazonas (near Puerto Ayacucho). Endemic. Ecclinusa ramiflora Mart., Flora 22(Beibl. 1): 2. 1839. Tree to 20 m tall, unbuttressed; latex abundant, white or cream-colored; flowers greenish white; mature fruit yellow or orange. Montane forests, ca. 1200 m; Amazonas (Cerro Marahuaka). Sucre, Táchira; Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Ecclinusa ulei (Krause) Gilly ex Cronquist, Bull. Torrey Bot. Club 73: 311. 1946. —Chrysophyllum ulei Krause, Notizbl. Konigl. Bot. Gart. Berlin 6: 171. 1914. Chrysophyllum ficoides Steyerm., Acta Bot. Venez. 2: 279. 1967. Shrub 3–5 m or tree 3–20 m tall, ca. 25 cm diameter; leaves sulcate, the lower surface castaneous-brown; latex copious, thick, white; flower buds castaneous-brown; flowers olive green; anthers yellow; fruit castaneous-brown. Forested slopes, gallery for-

Fig. 80. Ecclinusa orinocoensis

ests, mossy dwarf forests, low scrub, savannas, 1400–2100 m; Bolívar (Acopán-tepui, Auyán-tepui, Cerro Jaua, Macizo del Chimantá, Ptari-tepui, Roraima-tepui, Sierra de Maigualida). Endemic.

Ecclinusa 101

Fig. 81. Ecclinusa atabapoensis

Fig. 82. Ecclinusa lanceolata

102

S APOTACEAE

Fig. 83. Ecclinusa guianensis

5. ELAEOLUMA Baill., Hist. Pl. 11: 293. 1891. Trees or shrubs. Leaves spirally arranged, usually minutely punctate on lower surface; stipules absent; venation eucamptodromous or brochidodromous, higher order veins often obscure, forming a lax reticulum. Inflorescence mostly axillary. Flowers unisexual. Calyx a single whorl of (4)5 imbricate or quincuncial sepals; corolla broadly cyathiform to rotate, tube shorter than the lobes or rarely equaling the lobes in pistillate, lobes 5(6), spreading. Stamens 5(6), fixed at the top of the corolla tube and exserted; filaments well developed; anthers glabrous; staminodes usually absent, rarely 1 or 2. Disk absent. Ovary 2- or 3(4)-locular, placentation axile; style short. Fruit a 1-seeded berry. Seed broadly ellipsoid, not or sometimes laterally compressed, testa smooth to slightly wrinkled, shining; scar adaxial, full length, narrow or broad; embryo vertical, cotyledons plano-convex, radicle slightly exserted; thin sheath of endosperm present. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil; 4 species, all in the flora area. Key to the Species of Elaeoluma 1. 1. 2(1). 2.

Young shoots, lower surface of young leaves, and inflorescence densely crisped-ferruginous-tomentose ..................................................... E. crispa Young shoots, lower leaf surface of young leaves, and inflorescence subglabrous, or with some fine, closely appressed indumentum ....... 2 Intersecondary veins short or absent; seed laterally compressed, scar 1– 3 mm wide ....................................................................................... E. nuda Intersecondary veins long, well developed; seed not laterally compressed, scar 0.5–1.1 cm wide ............................................................... 3

Manilkara 103

3(2).

3.

Leaves 8–19.5 cm long, usually elliptic or oblong, apex frequently acute or attenuate, secondary veins usually 9–12 pairs; fruit 2–3 cm long ............................................................................................... E. glabrescens Leaves usually 2–6 cm long, oblanceolate, apex usually obtuse, rounded or emarginate, secondary veins usually 5–7 pairs; fruit 1.2–1.4 cm long ............................................................................... E. schomburgkiana

Elaeoluma crispa T.D. Penn., Fl. Neotrop. Monogr. 52: 246. 1990. —Dallama, Tetoncillo de rebalse. Tree 10–12 m tall, 13–30 cm diameter; trunk scaly, bark dull gray-brown; latex abundant, white; young shoots ferruginoustomentose; flowers pale green; fruit yellow, shiny, and hard. Seasonally flooded blackwater riparian forests, 100–200 m; Amazonas (Caño San Miguel, Río Atacavi, Río Baría, Río Pasimoni, San Carlos de Río Negro). Endemic. Elaeoluma glabrescens (Mart. & Eich.) Aubrév., Adansonia n.s. 1: 26. 1961. —Lucuma glabrescens Mart. & Eich. in Mart., Fl. Bras. 7: 72, t. 46, fig. 1. 1863. —Boya de mono, Palo rayao, Punwoyo, Wene iye, Yuquito banero. Tree or treelet 6–20 m tall, 7–50 cm diameter; trunk curved, outer bark dark brown, soft and furrowed, with small scales, inner bark bright orange and soft; latex scant, sticky, white-cream; buds green; flowers greenish white; fruit ovoid, bright green, edible, sweet. Evergreen lowland forests, semideciduous forests on granitic slopes, igapo and periodically flooded riparian forests, 50–400 m; Bolívar (Río Parguaza), southwestern Amazonas. Miranda; Panama, Amazonian Colombia, Guyana, Amazonian Peru, Brazil (Amazonian to Mato Grosso and Goiás). ◆Fig. 85. Elaeoluma nuda (Baehni) Aubrév., Mem. New York Bot. Gard. 23: 224. 1972.

—Pouteria nuda Baehni, Candollea 14: 72. 1952. Pouteria auyantepuiensis Steyerm., Acta Bot. Venez. 2: 278. 1967. —Obo-ye-muyek (Arekuna). Treelet or tree 3–25 m tall; bark with long cracks 10–15 mm apart, 1–3 mm deep; latex copious, milky white; flowers pale green to deep red, corolla light green, calyx dull pink; fruit yellow, blue-green, or brown-purple, ca. 2.5 cm long. Seasonally dry riparian forests, evergreen forested slopes, often steep, sandy stream banks, 1100–2200 m; Bolívar (Auyán-tepui, Cerro Jaua, Ptari-tepui), Amazonas (Cerro Coro Coro, Cerro Marahuaka, Cerro Yutajé, Sierra de la Neblina). Suriname, Brazil (Amazonas: Serra da Neblina, Serra Aracá). ◆Fig. 86. Elaeoluma schomburgkiana (Miq.) Baill., Hist. Pl. 11: 294. 1891. —Myrsine schomburgkiana Miq. in Mart., Fl. Bras. 10: 315. 1856. Oxythece steyermarkiana Monach., Fieldiana, Bot. 28: 485. 1953. Shrub or tree to 6 m tall, with a dense crown; leaves silvery below; latex sticky, white; flowers small and greenish white to yellow; fruit ripening purple or black. Seasonally flooded forests, savannas, especially shrub islands, 50–1300 m; Bolívar (Cerro Guaiquinima, Gran Sabana), Amazonas (Río Atabapo basin, Río Guainía, Río Pasimoni, Río Sipapo). Guyana, northern and central Amazonian Brazil. ◆Fig. 84.

6. MANILKARA Adanson, Fam. Pl. 2: 166. 1763. Unarmed trees or rarely shrubs with sympodial branching. Shoot apex often covered with transparent varnish-like wax. Small caducous stipules present or absent. Leaves spirally arranged, very rarely opposite or whorled, often densely clustered at the stem apex. Venation brochidodromous with a marginal vein, secondaries usually straight, parallel, ascending, looping below the margin and sometimes forming a submarginal vein; intersecondaries and tertiaries often prominent, parallel to the secondaries and descending from the margin; quaternary veins often prominently reticulate or areolate. Inflorescence axillary or in the axils of leaf

104

S APOTACEAE

Fig. 84. Elaeoluma schomburgkiana

Fig. 85. Elaeoluma glabrescens

Fig. 86. Elaeoluma nuda

Manilkara 105

scars; flowers solitary or fasciculate, bisexual. Calyx of 2 whorls or (2)3(4) free or slightly united sepals, the outer whorl valvate or only slightly imbricate; corolla nearly always glabrous, tube usually much shorter than the lobes, rarely equaling or exceeding them, lobes 6(–9), usually spreading, usually divided to the base into 3 segments, median segment usually erect, narrow, clawed, clasping the stamen, 2 lateral segments spreading, shorter than, equaling, or exceeding the median segment, entire, deeply divided or laciniate, or less frequently corolla lobe only partly divided or 2- or 3-lobed at the apex, or entire; corolla occasionally carnose. Stamens 6(–12) in a single whorl inserted at the top of the corolla tube, very rarely fixed within the tube; filaments free or partially fused to the staminodes; anthers extrorse, nearly always glabrous; staminodes (0–)6(–9), alternating with the stamens, truncate or variously lobed and divided, often ending in 2 filiform teeth, nearly always glabrous. Ovary 6–12-locular, hairy or glabrous, placentation axile or basi-ventral; style exserted. Fruit 1–several-seeded, smooth or scaly, glabrous, indehiscent, fleshy; seed ellipsoid to obovoid, strongly laterally compressed, with a hard shining woody testa; scar narrowly elongate, basi-ventral or extending along most of the adaxial surface. Embryo vertical with foliaceous cotyledons and exserted radicle; endosperm copious. Neotropics, Africa, Asia, and the Pacific, approx. 62 species, 3 in Venezuela, all in the flora area.

Fig. 87. Manilkara bidentata subsp. bidentata

106

S APOTACEAE

Key to the Species of Manilkara 1.

1. 2(1).

2.

Lower surface of leaves with fine, closely appressed, yellowish indumentum, often forming a pellicle; secondary veins 30–35 pairs; ovary puberulous ......................................................................... M. huberi Lower surface of leaves without obvious appressed indumentum; secondary veins 12–25 pairs; ovary glabrous ................................................. 2 Leaves usually oblanceolate (if elliptic or oblong-elliptic, then stipules present), glabrous; submarginal veins absent; calyx narrow and tapering gradually into the pedicel; lateral corolla lobe segments divided or not ............................................................................. M. bidentata Leaves elliptic or oblong-elliptic, stipules absent, microscopic brown trichomes (< 0.1 mm long) present on lower leaf surface; secondary veins join to form a conspicuous submarginal vein; calyx broad at base and abruptly contracted into the pedicel; lateral corolla lobe segments divided nearly to base ............................................................ M. bolivarensis

Manilkara bidentata (A. DC.) Chev., Rev. Int. Bot. Appl. Agric. Trop. 12: 270. 1932. —Mimusops bidentata A. DC., Prodr. 8: 204. 1844. Panama, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of M. bidentata 1. Stipules present, 2.5–4.5 mm long, leaving an obvious scar; leaves 10–27 cm long (mean length 15.6 cm), often elliptic, apex frequently obtuse or acute ................................. subsp. bidentata 1. Stipules absent or rarely present, but then < 1 mm long; leaves 7–21 cm long (mean length 12.2 cm) usually oblong or oblanceolate, apex usually rounded ........................... subsp. surinamensis M. bidentata subsp. bidentata. —Cuberu, Perguo, Perguo balata, Perguo blanco, Perguo morado, Pulguillo. Mimusops balata var. schomburgkii Pierre, Bull. Mens. Soc. Linn. Paris 1: 507. 1885. Manilkara williamsii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22(3): 165. 1940. Tree 15–35 m tall, 19–59 cm diameter; trunk straight and cylindrical, buttresses to 1.7 m high; bark fissured, gray-brown to red-brown; wood very hard, red; latex vary-

ing in amount from scarce to abundant, sticky, milky white; flowers creamy-white, fragrant; fruit brown-green or green-yellow, edible, ovoid, 1- or 2-seeded, seeds black. Evergreen lowland to lower montane forests, savanna islands, on granitic boulders, occasionally on dry forested slopes, 50–400 (–600) m; Delta Amacuro (Caño Capurito, Serranía de Imataca), northern Bolívar. Distrito Federal, Miranda, Monagas, Sucre; Panama, Guadeloupe, Martinique, Colombia, Trinidad, Guyana, French Guiana, Peru, Brazil (Pará, Roraima). ◆Fig. 87. Manilkara bidentata subsp. bidentata was heavily exploited in the 1900s as the source of commercial balata (American gutta percha, Suriname gutta percha), which is produced by cleaning and chemically purifying the latex. Its chemical inertness and property of becoming plastic, but not elastic, at high temperatures and then becoming hard again when cooled made it very suitable for electrical and marine cable insulation. Balata is chemically identical to, but less pure than, true gutta (produced from several species of the Malayan genus Palaquium) and both have now been largely replaced by synthetic materials. However, it is still extensively used in dentistry for root canal fillings where it remains superior to synthetics. The trees are periodically tapped for balata, a process which does little damage to the forest, but unfortunately, they are also frequently felled for their durable timber.

Micropholis 107

M. bidentata subsp. surinamensis (Miq.) T.D. Penn., Fl. Neotrop. Monogr. 52: 61. 1990. —Mimusops surinamensis Miq. in Mart., Fl. Bras. 7: 43. 1863. —Balata, Chicle, Masarandu, Masaranduba, Pendare, Pendare purue (Arekuna), Purgo. Shrub 1–2 m tall, treelet 3–4 m tall, or tree 5–30 m tall, 30–35 cm diameter; trunk unbuttressed or with small buttresses; bark dark, rough, scaly, and deeply fissured; latex viscous, varying from scarce to copious in amount, white; flowers white or yellowgreen, scented; fruit green when young, changing to red and shiny black on maturing, ca. 1.2 cm diameter, with white latex. Evergreen and semi-evergreen forests, river sides, low periodically flooded caatinga, among shrubby vegetation on white sand, 50–400 m; Delta Amacuro (Serranía de Imataca), widespread in Bolívar and Amazonas. Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amapá, Amazonas, Pará, Roraima). The fruit of Manilkara bidentata subsp. surinamensis is edible and the timber durable. The latex has been exploited in Colombia, Venezuela, and Brazil as a source of chicle (chewing gum).

Manilkara bolivarensis T.D. Penn., Fl. Neotrop. Monogr. 52: 77. 1990. Tree to 25 m tall. Montane gallery forests, ca. 1100 m; Bolívar (drainage of the Río Pacairao in the Gran Sabana). Endemic. Manilkara huberi (Ducke) Chev, Rev. Int. Bot. Appl. Agric. Trop. 12: 276. 1932. —Mimusops huberi Ducke, Arch. Jard. Bot. Rio de Janeiro 2: 14. 1918. —Chupón, Purguo, Pulgo negro. Tree 30–40 m tall, diameter 2–3 m; trunk buttressed; bark gray-brown, deeply fissured; latex copious, white; leaves pale yellowish on lower surface; flowers greenish white to yellowish; fruit green or greenish yellow. Evergreen lowland forests, 100–200 m; Amazonas (Capibara, Río Ventuari). Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, Maranhão, Mato Grosso, Pará, Rondonia, Roraima). The timber of Manilkara huberi is durable and the fruit edible. The latex was described by Richard Spruce as with the consistency of good cream, and its taste perfectly creamy and agreeable. However, due to its extreme viscosity, it has also been used as a substitute for glue and may cause serious constipation.

7. MICROPHOLIS Pierre, Not. Bot. Sapot. 37. 1891. Trees or shrubs. Leaves alternate and distichous or spirally arranged; stipules absent; venation brochidodromous (with a submarginal vein), or craspedodromous, secondary veins closely parallel, often not differentiated from the lower order venation, and then the leaf appearing finely striate. Inflorescence axillary, ramiflorous, or cauliflorous, fasciculate, sometimes developing into short scaly persistent shoots. Flowers often unisexual. Calyx a single whorl of (4)5 free, imbricate or quincuncial sepals; corolla campanulate to short- or long-cylindrical, the tube nearly always exceeding the lobes, rarely equaling them, the lobes (4)5, erect to reflexed, simple. Stamens (4)5, fixed near the top of the corolla tube, included or exserted; filaments short and straight or long and geniculate (at least in bud); anthers extrorse in bud, glabrous; staminodes (4)5, in the corolla lobe sinuses, alternating with the stamens, usually lanceolate or subulate, rarely petaloid. Disk present or absent. Ovary (4)5-locular, placentation axile; style included or exserted. Fruit 1–several-seeded. Seed laterally compressed, testa smooth or often minutely transversely wrinkled, shining or dull; scar adaxial, extending the length of the seed, usually narrow; embryo vertical, with thin foliaceous cotyledons and exserted radicle, surrounded by thick endosperm. Mexico, Central America, West Indies, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 38 species, 14 in Venezuela, 13 of these in the flora area.

108

S APOTACEAE

Key to the Species of Micropholis 1.

Stamens exserted; corolla > 10 mm long (except M. obscura); corolla lobes widely spreading or reflexed ....................................................... 2 1. Stamens included; corolla < 10 mm long; corolla lobes erect or only slightly spreading .................................................................................. 5 2(1). Corolla < 10 mm long; tree with convex buttresses ..................... M. obscura 2. Corolla > 10 mm long; buttresses if present not convex ......................... 3 3(2). Mature leaves persistently rufous-brown appressed-puberulous on lower surface ..................................................................................... M. splendens 3. Mature leaves glabrous or essentially so (indumentum if present very fine and visible only with lens) ............................................................. 4 4(3). Young shoots and inflorescence ferruginous-tomentose............. M. maguirei 4. Young shoots finely appressed-puberulous ............................... M. spectabilis 5(1). Leaves deeply sulcate along the midrib, margin strongly revolute (to 0.5 cm of the leaf margin reflexed) ................................... M. venamoensis 5. Leaves not as above ................................................................................... 6 6(5). Secondary venation spaced, not finely striate; higher order venation (usually reticulate), visible to the naked eye ....................................... 7 6. Secondary venation close, finely striate, higher order venation indistinguishable from the secondary venation, or visible only with a lens ................................................................................................................ 8 7(6). Leaves > twice as long as broad; apex usually acute or acuminate ............................................................................................... M. guyanensis 7. Leaves < twice as long as broad, apex usually rounded or emarginate ......................................................................................... M. suborbicularis 8(6). Lower surface of leaf finely and persistently sericeous .... M. humboldtiana 8. Lower surface of leaf glabrous .................................................................. 9 9(8). Secondary venation very close, finely striate, indistinguishable from the higher order venation .......................................................................... 10 9. Secondary venation striate, but higher order venation can be distinguished with a lens .............................................................................. 11 10(9). Secondary venation steeply ascending; corolla glabrous, lobes truncate; style exserted; fruit 4–7 cm long ....................................... M. melinoniana 10. Secondary venation shallowly ascending; corolla tube often appressedpubescent, lobes obtuse or rounded; style included; fruit 2.5–3 cm long ..................................................................................................... M. egensis 11(9). Corolla 5–6.5 mm long ................................................................. M. mensalis 11. Corolla 2–3 mm long ................................................................................ 12 12(11). Leaves 7–12 cm long; apex obtusely acuminate; pedicel 0.8–1.4 cm long; ovary flattened or winged, puberulous .......................... M. casiquiarensis 12. Leaves 4–9 cm long; apex caudate or narrowly attenuate; pedicel 1.5– 7 mm long; ovary ovoid, not winged, densely pubescent ....... M. venulosa Micropholis casiquiarensis Aubrév., Mem. New York Bot. Gard. 23: 211. 1972. Tree to 30 m tall; flowers cream-colored; fruit maturing yellow to reddish black. Ev-

ergreen lowland forests, 100–200 m; Amazonas (Río Yatúa). Amazonian Brazil. Micropholis egensis (A. DC.) Pierre in Urb., Symb. Antill. 5: 127. 1904.

Micropholis 109

—Sideroxylon egense A. DC., Prodr. 8: 182. 1844. —Cajou. Micropholis wurdackii Aubrév., Mem. New York Bot. Gard. 23: 212. 1972. Treelet or tree 3–40 m tall or shrub to 5 m; latex white; flowers creamy-white, green at the apex; fruit pinkish red to red, shiny, 3 cm × 1.5 cm, edible. Low, periodically flooded riparian forests, high nonflooded forests, rocks on open bank, 50–100 m; Bolívar (Cano Coroso of the Río Caripo, Río Orinoco near El Jobal, Temblador), Amazonas (widespread). Guárico; Panama, Colombia, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 91.

face of leaves brown or gray-green; latex sticky, white; flowers yellow-green; fruit globose, brown-green to purple. Evergreen lowland to montane forested slopes, gallery forests, 100–1700 m; widespread in Bolívar and Amazonas. Widespread elsewhere in Venezuela; Costa Rica, Panama, Antilles, Venezuela, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian and coastal Brazil, Bolivia. The wood of Micropholis guyanensis subsp. guyanensis is used in construction, and the latex was formerly used to adulterate balata. The fruit is edible.

Micropholis guyanensis (A. DC.) Pierre, Not. Bot. Sapot. 40. 1891. —Sideroxylon guyanensis A. DC., Prodr. 8: 182. 1844. Costa Rica, Panama, Puerto Rico, Lesser Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian and coastal Brazil, Bolivia; 2 subspecies, both in the flora area.

Micropholis humboldtiana (Roem. & Schult.) T.D. Penn., Fl. Neotrop. Monogr. 52: 212. 1990. —Chrysophyllum humbodltianum Roem. & Schult., Syst. Veg. 4: 813. 1819. Chrysophyllum obtusatum Willd. ex Roem. & Schult., Syst. Veg. 4: 813. 1819. Chrysophyllum aquaticum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 236. 1818 [1819]. Shrub 3–5 m tall; lower surface of leaves ferruginous; latex scarce; flowers whitish green; fruit dark wine-red. Seasonally flooded riparian forests, flooded savannas, 100–200 m; Amazonas (Caño Cotua, Cerro Yapacana, lower Río Baria). Amazonian Brazil.

Key to the Subspecies of M. guyanensis 1. Leaves usually > 15 cm long, broadly elliptic or broadly oblong-elliptic, often about twice as long as broad, base obtuse or rounded; petiole usually > 2 cm long .......................... subsp. duckeana 1. Leaves often < 15 cm long, usually narrowly elliptic or oblanceolate, 2–4 times as long as broad, base narrowly attenuate, cuneate or acute; petiole usually < 2 cm long ............ subsp. guyanensis M. guyanensis subsp. duckeana (Baehni) T.D. Penn., Fl. Neotrop. Monogr. 52: 180. 1990. —Pouteria duckeana Baehni, Candollea 18: 161, fig. 53. 1962. Tree 30 m tall, 20–140 cm diameter; flowers cream-colored. Evergreen lowland forest on sandy soils, 100–700 m; Amazonas (Cerro Marahuaka slopes, Río Mawarinuma). Colombia, Guyana, Suriname, French Guiana, Peru. Amazonian Brazil. M. guyanensis subsp. guyanensis. —Chicle, Chipitudek (Arekuna), Wacadu. Tree 12–80 m tall, 14–50 cm diameter; outer bark brown, hard, shallowly longitudinally cracked, inner bark orange; lower sur-

Micropholis maguirei Aubrév., Mem. New York Bot. Gard. 23: 210. 1972. —Cachito, Huevo frito, Temare. Treelet or tree 10–25 m tall, 20–25 cm diameter; buttresses narrow, 1 m tall; lower surface of leaves at first with ferruginous indumentum, soon glabrous; latex scarce, white; flowers white, fragrant. Evergreen lowland forests, 100–200 m; Amazonas (Río Guainía, near San Carlos de Río Negro, Yavita). Brazil (Amazonas: upper Rio Negro). ◆Fig. 88. Micropholis melinoniana Pierre, Not. Bot. Sapot. 40. 1891. —Cajón, Pico de lombriz. Tree ca. 28 m tall, 26 cm diameter; flowers scented, greenish white or cream; fruit 4–7 cm long, yellow, orange, reddish, or purple, sometimes slightly sulcate. Ever-

110

S APOTACEAE

green lowland forests, 100–200 m; Bolívar (Río Caura at Salto Pará), Amazonas (Río Manapiare). Mexico, Guatemala, Belize, Costa Rica, Panama, French Guiana, Amazonian Ecuador, Peru and Brazil. ◆Fig. 89. Micropholis melinoniana provides a durable and resistant timber that has been used locally for construction. The fruit is edible. Micropholis mensalis (Baehni) Aubrév., Adansonia n.s. 3: 21. 1963. —Pouteria mensalis Baehni, Candollea 14: 64. 1952. Tree 5 m tall, 8 cm diameter; outer bark gray-brown and smooth, inner bark cream; latex sticky, white; fruit green, rugose. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma). Guyana, Suriname, French Guiana, Amazonian Brazil. Micropholis obscura T.D. Penn., Fl. Neotrop. Monogr. 52: 227. 1990. Tree to 30 m tall, 50 cm diameter; buttresses thin, convex; outer bark dark brown, soft, scaly, inner bark orange, hard; leaves glossy; latex sticky, white; flowers pale yellow. Seasonally flooded forests along blackwater rivers, evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma, near San Carlos de Río Negro). Guyana, French Guiana, Peru, Brazil (Amazonas, Pará). Micropholis spectabilis (Steyerm.) T.D. Penn., Fl. Neotrop. Monogr. 52: 221. 1990. —Pouteria spectabilis Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 436, fig. 8. 1966. Tree to 25 m tall; lower surface of leaves yellow-green; flowers greenish yellow. Montane forests, ca. 1200 m; Bolívar (Gran Sabana northeast of Luepa). Endemic. Micropholis splendens Gilly ex Aubrév., Mem. New York Bot. Gard. 23: 210. 1972. —Palo de sol, Primo chipón, Yavaraji (Yanomami). Tree 15–30 m tall, 20–50 cm diameter; latex sticky, very slow to exude, white; outer bark brown, soft, scaling in small pieces, inner bark orange, hard; leaves ferruginous, the lower surface rufous; flowers caulescent, corolla white, filaments white, anthers brown, stigma and style green; fruit coppery-brown. Evergreen lowland forests, open areas,

along dry ridges and river banks, 100–800 m; Amazonas (Cerro Huachamacari, Culebra, Río Mawarinuma, San Carlos de Río Negro). Brazil (Amazonas: Rio Negro basin), possibly French Guiana. Micropholis suborbicularis Aubrév., Mem. New York Bot. Gard. 23: 209. 1972. Shrub, treelet, or tree 1–15 m tall, treelet 10 cm diameter; leaves ferruginous, sericeous, the lower surface chocolate-brown; latex scarce, white; flower buds ferruginous; flowers white, cream, pale green, or brownish; fruit green becoming ferruginous-brown, ovoid, 1–1.5 cm × 0.5 cm. Elfin tepui forests, forests at savanna edges, low scrub, shrub islands in savannas, along water courses, 1300–2000 m; Bolívar (Cerro Guaiquinima, Cerro Jaua, Gran Sabana, Macizo del Chimantá, Sierra Pakaraima), Amazonas (Cerro Aracamuni, Laguna Asisa). Endemic. Micropholis venamoensis (Steyerm.) T.D. Penn., Fl. Neotrop. Monogr. 52: 205. 1990. —Pouteria venamoensis Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 434, fig. 7. 1966. Tree 15–30 m tall; leaf margin revolute, midvein deeply sulcate; flowers yellowgreen. Humid, mossy, montane forests, 800– 1100 m; Bolívar (Cerro Venamo, La Escalera). Endemic. ◆Fig. 90. Micropholis venulosa (Mart. & Eich.) Pierre, Not. Bot. Sapot. 40. 1891. —Sideroxylon venulosum Mart. & Eich. in Mart., Fl. Bras. 7: 52, t. 20, fig. 2, t. 37, fig. 4. 1863. —Capurillo, Dewaka (Arekuna), Palo de chamanare, Purguillo felix, Yuquito. Treelet or tree 5–33 m tall, 22–35 cm diameter; bark reddish, shallowly fissured; latex abundant, sticky, milky white; flowers white or pale green; fruit pale yellow-green, tinged with red becoming maroon, ovoid-globose. Evergreen lowland forests, montane forests, flooded river margins, (50–)200(–1000) m; Delta Amacuro (Serranía de Imataca), Bolívar (Cerro Venamo, Río Tirica, Serranía de Imataca), Amazonas (widespread). Panama, Colombia, Ecuador, Peru, Guyana, Suriname, French Guiana, Amazonian and coastal Brazil (Bahia). The fruit of Micropholis venulosa is sweet and edible, and the bark is reputed to have medicinal properties.

Micropholis 111

Fig. 88. Micropholis maguirei

Fig. 89. Micropholis melinoniana

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S APOTACEAE

Fig. 90. Micropholis venamoensis

Fig. 91. Micropholis egensis

8. POUTERIA Aubl., Pl. Guiane 85, pl. 33 (excl. fruct.) 1775. Podoluma Baill., Hist. Pl. 11: 290. 1891. Trees or shrubs, rarely a geoxylic suffrutex. Leaves usually spirally arranged, rarely opposite; stipules absent; venation eucamptodromous or brochidodromous, usually without a submarginal vein, never finely striate. Inflorescence axillary or ramiflorous, fasciculate, fascicles single or occasionally arranged along short leaf-

Pouteria 113

less shoots. Flowers often unisexual. Calyx a single whorl or 4–6 free, imbricate or quincuncial sepals, or 6–11 in a closely imbricate spiral; corolla cyathiform to tubular, rarely rotate, tube shorter than, equal to, or exceeding the lobes, lobes 4–6(–9), usually erect, rarely spreading, simple, sometimes fringed-ciliate or papillose. Stamens 4–6(–9), fixed in lower or upper half of corolla tube, or rarely at base of lobes, rarely free, usually included, rarely exserted; filaments generally short; anthers usually extrorse or laterally dehiscent, usually glabrous, rarely hairy; staminodes usually same number as corolla lobes, less frequently partially or completely lacking, inserted in the corolla sinus or inside the tube, sometimes fringed-ciliate or papillose. Disk rarely present. Ovary 1–6(–15)-locular, placentation axile; style included or exserted. Fruit a 1–several-seeded berry. Seed broadly ellipsoid, planoconvex, shaped like the segment of an orange or laterally compressed, testa smooth, wrinkled or pitted; scar adaxial, usually full-length, narrow, broad or sometimes covering almost all the seed surface. Embryo vertical, usually with plano-convex cotyledons and included radicle, less frequently with thin foliaceous cotyledons and exserted radicle; endosperm usually absent, less frequently present. Pantropics; ca. 325 species, 51 in Venezuela, 42 of these (including 1 Podoluma treated here that may belong in Pouteria) in the flora area. Key to the Species of Pouteria Three taxa whose affinities are not clear are not included in the key. These are Pouteria aff. engleri, Pouteria aff. minima, and Podoluma pacimoniensis and are listed at end of the species entries. 1. 1.

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

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

5.

6(5).

Flowers 4-merous (sepals, corolla lobes, stamens, staminodes, and ovary loculi 4); corolla lobes and staminodes usually fringed-ciliate ........... 2 Flowers not 4-merous; sepals 5, or if 4, then ovary 2-locular or corolla lobes 6–10; corolla lobes and staminodes not usually fringed-ciliate .............................................................................................................. 20 Lower leaf surface with indumentum of spreading or erect trichomes ...................................................................................................... P. hispida Lower leaf surface with closely appressed indumentum or glabrous .... 3 Venation brochidodromous with submarginal vein, secondary veins 20– 35 pairs; intersecondaries numerous, extending to the submarginal vein; leaves drying dark brown .......................................... P. gomphiifolia Venation eucamptodromous, or if brochidodromous, then without a submarginal vein ......................................................................................... 4 Venation brochidodromous ........................................................................ 5 Venation eucamptodromous ...................................................................... 8 Higher order venation lax, not finely reticulate; secondary veins usually 13–23 pairs; corolla ca. 3 mm long; fruit soft-skinned, shrinking around the seeds on drying; glabrous .......................................... P. plicata Higher order venation finely reticulate, secondary veins often fewer; corolla 2–14 mm long; fruit tough-skinned, not shrinking on drying; velutinous or glabrous ........................................................................... 6 Leaf apex rounded; corolla 2–5.5 mm long; seed scar 2–3.5 mm wide ...................................................................................................... P. hispida

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6. 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(14). 15. 16(14). 16. 17(16).

17.

18(13). 18. 19(18).

19.

Leaf apex usually acute or attenuate; corolla 4–14 mm long; seed scar usually broader ...................................................................................... 7 Leaf usually 5–10 × 1.7–5 cm; base often narrowly attenuate; secondary veins 8–13 pairs; leaf glabrous; petiole not channeled .............. P. caimito Leaf usually 13–25 × 5–10 cm; base acute; secondary veins usually 13– 22 pairs, lower surface of leaves often finely sericeous; petiole margins strongly infolded ............................................................. P. guianensis Leaf base rounded or truncate .................................................... P. glomerata Leaf base acute to attenuate ..................................................................... 9 Lower leaf surface finely sericeous ......................................................... 10 Lower leaf surface glabrous, or sometimes with close minute indumentum (visible only with lens), sometimes forming a pellicle ....... 11 Petiole 2–5 cm long; corolla 0.5–1.4 cm long; seed scar 0.6–1.1 cm broad ................................................................................................. P. guianensis Petiole 0.3–1 cm long; corolla 2–4.5 mm long; seed scar covering most of the seed, leaving only a narrow free abaxial strip ................. P. glomerata Seed scar covering nearly all the seed surface ........................... P. glomerata Seed scar adaxial, broad or narrow, but not covering > half the seed surface ....................................................................................................... 12 Leaves about twice as long as broad; secondary veins 6–8 pairs; petiole 2–4.5 cm long; fruit 7–10 cm long ............................................. P. maguirei Leaves narrower, secondary veins more numerous, fruit usually smaller .............................................................................................................. 13 Higher order venation finely reticulate ................................................. 14 Higher order venation oblique or horizontal, or if reticulate then rather lax and often obscure, not finely reticulate ....................................... 18 Leaf apex rounded or obtuse; corolla 2–6.5 mm long ............................ 15 Leaf apex acute to attenuate; corolla 5–14 mm long ............................. 16 Petiole not channeled; seed often slightly laterally compressed ... P. caimito Petiole channeled; seed not laterally compressed .......................... P. hispida Secondary veins usually 8–13 pairs; petiole not channeled; seed often slightly laterally compressed ...................................................... P. caimito Secondary veins usually 13–25 pairs, petiole channeled or margins strongly infolded; seed not laterally compressed .............................. 17 Young shoots usually with stiff pubescence; higher order vein reticulum fine, sharp, and conspicuous; leaf dries green; petiole channeled .......................................................................................................... P. torta Young shoots appressed-puberulous; higher order venation coarser and less conspicuous, leaf often dries reddish brown; petiole margins strongly infolded ..................................................................... P. guianensis Tertiary veins rather numerous, oblique to horizontal; anthers with tuft of trichomes at apex .................................................................. P. fimbriata Tertiary veins reticulate or obscure, if oblique then few and widely spaced; anthers glabrous .................................................................... 19 Leaves not glandular-striate, at least 3 times as long as broad, elliptic, oblong, or oblong-elliptic; secondary veins usually 13–23 pairs; petiole margins strongly infolded ............................................................ P. plicata Leaves usually glandular-striate, oblanceolate, or if oblong or elliptic

Pouteria 115

20(1).

20.

21(20). 21. 22(21). 22. 23(21). 23. 24(23). 24. 25(24). 25. 26(25). 26. 27(23).

27.

28(27).

28.

29(28). 29. 30(29). 30. 31(30).

then only ca. 2 times as long as broad, secondary veins 6–13 pairs; petiole flat or only slightly channeled ...................................... P. stipitata Sepals 4; corolla 5.5–22 mm long; corolla lobes, stamens, and staminodes usually 6; ovary loculi usually 6–10; style long, often exserted ....................................................................................................... P. venosa Sepals 4 or 5; corolla usually < 6 mm long; corolla lobes, stamens, and staminodes usually 4 or 5; ovary loculi 1–5; style exserted or not .............................................................................................................. 21 Corolla rotate, tube shorter than lobes; stamens fixed at top of tube or base of lobes, always exserted ............................................................ 22 Corolla cyathiform or tubular; stamens inserted in upper or lower part of corolla tube, rarely free, always included .......................................... 23 Leaves broadly oblanceolate or obovate, apex usually rounded; secondary veins 9 or 10 pairs ....................................................... P. canaimaensis Leaves oblong-elliptic, oblong-lanceolate, or elliptic, apex obtuse to caudate, secondary veins 15–22 pairs ....................................... P. eugeniifolia Seed endospermous; embryo with thin, foliaceous cotyledons ............. 24 Seed without endosperm; embryo with thick plano-convex cotyledons 27 Stamens free ................................................................................. P. laevigata Stamens fixed in the upper half or at the top of the corolla tube ......... 25 Leaves opposite, with numerous fine horizontal tertiary veins .... P. arcuata Leaves spirally arranged, tertiary veins few, oblique, or obscure ........ 26 Mature leaves densely short-tomentose below, with crisped reddish brown to ferruginous trichomes ............................................ P. sipapoensis Mature leaves glabrous ............................................................ P. cayennensis Flowers 5-merous (sepals, corolla lobes, stamens, staminodes, and ovary loculi 5); corolla tubular, 0.8–1.1 cm long, tube exceeding the lobes ............................................................................................. P. surumuensis Sepals, corolla lobes and stamens 4–6, ovary loculi usually 1 or 2; corolla 1.5–6.5 mm long, usually cyathiform; tube shorter or longer than the lobes ...................................................................................................... 28 Venation usually brochidodromous; higher order venation obscure; lower surface of leaves usually glaucous; staminodes 0–5, usually vestigial .............................................................................................................. 29 Venation eucamptodromous or brochidodromous; higher order venation usually conspicuous; lower surface of leaves not usually glaucous; staminodes usually same number as corolla lobes, well developed .............................................................................................................. 35 Corolla 5.5–6.5 mm long, tube exceeding the lobes ............ P. ambelaniifolia Corolla not exceeding 4 mm long, tube nearly always only equaling or shorter than lobes ................................................................................ 30 Lower surface of leaves persistently golden-ferruginous-sericeous; secondary veins 20–25 pairs ..................................................... P. gabrielensis Lower surface of leaves not persistently sericeous; secondary veins fewer .............................................................................................................. 31 Young shoots and inflorescence densely pubescent to tomentose with spreading indumentum; leaves about twice as long as broad, apex rounded, truncate, or emarginate ................................................. P. rigida

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31. 32(31).

32.

33(32).

33. 34(33).

34.

35(28). 35. 36(35). 36. 37(36).

37.

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

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

41.

Indumentum of young shoots and inflorescence minute, closely appressed; leaves narrow ........................................................................ 32 Leaf margin strongly revolute; lower blade persistently puberulous with sparse minute appressed-ferruginous indumentum (lens); leaf base acute to rounded ................................................................ P. pimichinensis Leaf margin not strongly revolute; lower blade subglabrous or rarely with pale appressed indumentum; leaf base attenuate or cuneate .............................................................................................................. 33 Young shoots and inflorescence golden-sericeous; leaves thin-textured, with pustular or pitted upper blade; secondary veins 6–13 pairs .............................................................................................. P. scrobiculata Young shoots and inflorescence not sericeous; leaves coriaceous, not pustular or pitted; secondary veins usually more numerous ................. 34 Midrib sunken on upper surface of leaves; stamens inserted in lower half of corolla tube, with well developed filaments; fruit slender, apex acute or narrowly attenuate ........................................................ P. elegans Midrib raised on upper surface of leaves; stamens fixed near apex of corolla tube, with very short filaments; fruit broader, with rounded apex .................................................................................................. P. cuspidata Higher order venation (tertiary or above) finely areolate; ovary 1-locular .................................................................................................. P. reticulata Higher order venation not finely areolate, or if so, then ovary 2(–5)-locular ......................................................................................................... 36 Stamens free or fixed at base of corolla tube; corolla lobes 4 ............... 37 Stamens fixed in the upper half of the corolla tube, or if in lower half then corolla lobes 5 .............................................................................. 40 Venation brochidodromous; tertiary veins parallel to secondaries and descending from margin; lower surface of leaves glaucous; seed coat smooth ........................................................................................... P. egregia Venation eucamptodromous or brochidodromous; tertiary veins neither parallel to the secondaries nor descending from the margin; leaves not glaucous; seed coat usually wrinkled or fissured .............................. 38 Venation mostly brochidodromous; fruit apex rounded; seed coat wrinkled and finely transversely fissured ........................... P. lucumifolia Venation eucamptodromous; fruit apex acute to rostrate; seed coat usually slightly or strongly wrinkled, not fissured ................................. 39 Leaves glabrous; secondary veins 7–10 pairs, widely spaced, convergent and strongly arcuate ................................................................... P. coriacea Leaves appressed-puberulous on lower surface; secondary veins 7– 14 pairs, closer, parallel or only slightly convergent, weakly arcuate ........................................................................................................ P. bangii Stamens fixed in the lower half of the corolla tube; leaf with a submarginal vein .................................................................................... P. anomala Stamens fixed near the top of the corolla tube; submarginal vein absent .............................................................................................................. 41 Staminodes absent or fewer than the corolla lobes; lower surface of leaves thinly appressed-puberulous with minute brown trichomes; fruit 10–11 cm long ....................................................................... P. ucuqui Staminodes as many as corolla lobes ..................................................... 42

Pouteria 117

42(41). Lower surface of leaves with persistent indumentum .......................... 43 42. Lower surface of leaves essentially glabrous ......................................... 44 43(42). Indumentum of lower surface of leaves erect or spreading; leaves 25– 70 cm long ............................................................................... P. crassiflora 43. Indumentum of lower surface of leaves of fine closely appressed trichomes; leaves smaller ................................................................. P. ucuqui 44(42). Ovary 4- or 5-locular .................................................................. P. trilocularis 44. Ovary 2(3)-locular .................................................................................... 45 45(44). Corolla tube exceeding lobes ................................................................... 46 45. Corolla tube equaling or shorter than lobes .......................................... 49 46(45). Corolla 3.5–7.5 mm long ............................................................ P. trilocularis 46. Corolla 1–2.25 mm long .......................................................................... 47 47(46). Midrib usually slightly sunken on upper surface of leaves; seed scar 1.2– 1.4 cm broad ......................................................................... P. orinocoensis 47. Midrib slightly raised on the upper surface of leaves; seed scar 2–6 mm broad .................................................................................................... 48 48(47). Venation brochidodromous; secondary veins straight ................ P. cladantha 48. Venation eucamptodromous; secondary veins arcuate .............. P. bilocularis 49(45). Leaf apex rounded or truncate; sepals and corolla lobes 4 ... P. atabapoensis 49. Leaf apex acute to acuminate; sepals and corolla lobes 4 or 5 ............. 50 50(49). Seed coat smooth ................................................................. P. amygdalicarpa 50. Seed coat wrinkled or verrucose ............................................................. 51 51(50). Venation conspicuous, eucamptodromous; secondaries strongly convergent; lower surface of leaves usually minutely punctate ...... P. bilocularis 51. Venation inconspicuous, mostly brochidodromous, secondaries parallel or only slightly convergent; lower surface of leaves not punctate ............................................................................................... P. lucumifolia Pouteria ambelaniifolia (Sandwith) T.D. Penn., Fl. Neotrop. Monogr. 52: 362. 1990. —Chrysophyllum ambelaniifolium Sandwith, Kew Bull. 476. 1931. —Chuponcillo. Tree 18–20 m tall; latex white; calyx greenish tan; fruit yellow. Semi-evergreen forests, 200–600 m; Delta Amacuro (Serranía de Imataca), Bolívar (Amaruay-tepui). Guyana, French Guiana, Brazil (Amapá, Amazonas). Pouteria amygdalicarpa (Pittier) T.D. Penn., Fl. Neotrop. Monogr. 52: 255. 1990. —Sideroxylon amygdalicarpum Pittier, Bol. Comerc. Industr. Venez. no. 13: 427. 1921 [reprinted in Arb. Arbust. Venez. 1: 13. 1923]. —Chirin-yo. Tree ca. 6 m tall, 15 cm diameter; outer bark light brown, hard, inner bark brown; latex white. Semideciduous forests on granitic slopes, ca. 400 m; northwestern Bolívar. Parts of northern central Venezuelan

Coastal Cordillera, Mérida; Costa Rica, Panama, Colombia. ◆Fig. 95. The wood of Pouteria amygdalicarpa is heavy, hard, and durable. Pouteria anomala (Pires) T.D. Penn., Fl. Neotrop. Monogr. 52: 344. 1990. —Chrysophyllum anomalum Pires, Bol. Tecn. Inst. Agron. N. 38: 34. 1960. —Sawara kurimi, Yuquito. Tree 14–25 m tall, with dense crown; latex abundant, milky white to yellow-white; fruit green, becoming red, subglobose-globose, edible. Evergreen lowland forests, 100–300 m; Amazonas (Río Cataniapo, San Pedro, Trapichote). Amazonian Brazil (Amazonas, Pará). ◆Fig. 96. Pouteria arcuata T.D. Penn., Fl. Neotrop. Monogr. 52: 371. 1990. —Syzygiopsis sericea Cronquist, Bull. Torrey Bot. Club 73: 471. 1946, non Pouteria sericea (Aiton) Baehni 1942.

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S APOTACEAE

Tree 10–15 m tall; bark gray, scaly, thin; latex present; leaves opposite; fruit globose, yellowish. Evergreen lowland forests, 100– 200 m; Amazonas (Capibara, San Antonio). Endemic. Pouteria atabapoensis (Aubrév.) T.D. Penn., Fl. Neotrop. Monogr. 52: 304. 1990. —Franchetella atabapoensis Aubrév., Mem. New York Bot. Gard. 23: 224. 1972. Tree to 7 m tall; latex milky; flowers yellowish white; fruit green. Sandy, flooded river margins, 100–200 m; Amazonas (confluence of Río Atabapo and Río Orinoco). Brazil (Amazonas: Rio Negro basin). Pouteria bangii (Rusby) T.D. Penn., Fl. Neotrop. Monogr. 52: 288. 1990. —Sideroxylon bangii Rusby, Bull. New York Bot. Gard. 4: 407. 1907. —Paramoyek, Rosado. Tree 12–30 cm tall, 31–40 cm diameter; bark gray-brown, fairly smooth, but with faint shallow longitudinal cracks; latex white; calyx pale cream-yellow or green with purple tip; corolla salmon pink to pink-red; fruit purple, 1-seeded. Evergreen lowland to montane forests, seasonally dry forests, 200–1300 m; Delta Amacuro (Serranía de Imataca), Bolívar (widespread), Amazonas (Cerro Yutajé). Apure; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonia, coastal areas), Bolivia. ◆ Fig. 97. Pouteria bilocularis (Winkler) Baehni, Candollea 9: 229. 1942. —Labatia ?bilocularis Winkler, Repert. Spec. Nov. Regni Veg. 7: 112. 1909. Pseudocladia neblinanensis Aubrév., Mem. New York Bot. Gard. 23: 204. 1972. Tree 10–30 m tall; latex milky white; flowers white or yellow-green; fruit green, edible. Forested slopes, evergreen lowland forests, talus forests, nonflooded and swamp forests, 100–600 m; Bolívar (Cerro Abismo), Amazonas (Río Mawarinuma, Río Yatúa). Colombia, Guyana, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. Pouteria caimito (Ruiz & Pav.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 12: 33. 1882. —Achras caimito Ruiz & Pav., Fl.

Peruv. 3: 18, pl. 240. 1802. —Capure, Melaito, Purgo, Temara, Temare, Temare amarillo Temari, Temure. Lucuma temare Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 241. 1818 [1819]. Tree 1.5–25 m tall, 10–50 cm diameter; latex scant, white; flowers greenish white; fruit green to orange-yellow, 1-seeded, plumsized, edible when ripe. Riparian forests, wet cloud forests, periodically flooded primary forests, secondary forests, intermittent forests on savannas and bluffs, often cultivated near towns or habitations, 100– 1000 m; Delta Amacuro (Caño Arature, near Isla Tigre), Bolívar (La Escalera, upper Río Caura, Serranía de Imataca), Amazonas (widespread). Yaracuy-Falcón border; Costa Rica, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 98. The natural distribution of Pouteria caimito is uncertain due to extensive cultivation for its widely acclaimed edible fruit. Pouteria canaimaensis T.D. Penn., Fl. Neotrop. Monogr. 52: 530. 1990. Shrub 1–1.5 m tall or tree 4–10 m tall, to 10 cm diameter; leaves glaucous, the lower surface silvery; latex scant, milky white; flowers yellow-green to greenish brown; fruit dark brown-ferruginous. Riparian forests, shrub-savanna areas, 300–700 m; Bolívar (Canaima, Cerro Pitón, Río Paragua, Supamo). Endemic. ◆Fig. 93. Pouteria cayennensis (A. DC.) Eyma, Recueil Trav. Bot. Néerl. 33: 174. 1936. —Chrysophyllum cayennensis A. DC., Prodr. 8: 160. 1844. —Amarun-yek. Tree 20–30 m tall, 0.2–1 m diameter; latex milky; buds buff-brown; flowers yellowferruginous; fruit yellow, edible. Evergreen lowland to wet montane forests, 100–1300 m; Bolívar (Macizo del Chimantá, Ptaritepui, Sierra de Lema), Amazonas (Río Mawarinuma, San Carlos de Río Negro). Guyana, French Guiana, Brazil (Amazonas). ◆Fig. 99. Pouteria cladantha Sandwith, Kew Bull. 480. 1931. —Acuchipuru mira, Cacho venado, Pulguillo. Tree to 20 m tall, 20 cm diameter; leaves glossy; latex scant; perianth green-white;

Pouteria 119

fruit round, edible, sweet. Lowland terrafirme forests, 100–300 m; Delta Amacuro (Cerro Muerto), Bolívar (Río Cuyuní basin, San Martin de Turumbán southwest of Tumeremo), Amazonas (San Carlos de Río Negro). Colombia (Vaupés), Guyana, Suriname, French Guiana, Amazonian Peru and Brazil. Pouteria coriacea (Pierre) Pierre in Urb., Symb. Antill. 5: 109. 1904. —Guapeba coriacea Pierre, Not. Bot. Sapot. 42. 1891. Tree to 12 m tall; flowers white or pale green; fruit maturing yellow. Río Negro caatinga, 100–500 m; Amazonas (Piedra Cocuy, upper Río Orinoco, Sierra de la Neblina). Miranda, Mérida, Sucre, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Amazonian Ecuador, northern Amazonian Brazil. Pouteria crassiflora Pires & T.D. Penn., Fl. Neotrop. Monogr. 52: 329. 1990. —Corocuro. Tree to 8 m tall, 10–15 cm diameter; latex white; fruit maturing orange. Seasonally flooded forests along black-water rivers, 100–200 m; Amazonas (San Carlos de Río Negro). Brazil (Amapá, Pará). Pouteria cuspidata (A. DC.) Baehni, Candollea 9: 231. 1942. —Sideroxylon cuspidatum A. DC., Prodr. 8: 183. 1844. Panama, Pacific and Amazonian Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Amazonian Peru and Bolivia; 3 subspecies, all in the flora area. Key to the Subspecies of P. cuspidata 1. Leaves usually 6–15 × 2.3–5 cm, apex acute or narrowly attenuate, lower surface glaucous, secondary veins not impressed on lower surface ........................ ................................. subsp. cuspidata 1. Leaves usually 4.3–7.5 × 1.3–3 cm, apex obtusely cuspidate, acute to shortly narrowly attenuate, lower surface not glaucous, secondary veins impressed on both surfaces ........................... subsp. dura 1. Leaves usually 11–22 × 4.6–8.5 cm, apex obtuse, obtusely cuspidate, rounded, or emarginate, lower surface not glaucous, secondary veins not impressed on lower surface ......................... subsp. robusta

P. cuspidata subsp. cuspidata. —Taymarito. Tree 10–20 m tall; lower surface of leaves glaucous; latex sticky, white; flowers dull pale green; fruit brown, scurfy, ovoid, with soft edible sweet pulp. River edges, forested slopes, gallery forests bordering savannas, 50–500 m; Bolívar (Río Caura, Río Parguaza), Amazonas (Capibara, San Carlos de Río Negro, Santa Barbara del Orinoco). Colombia, Guyana, French Guiana, Amazonian Peru, Brazil, and Bolivia. ◆Fig. 100. P. cuspidata subsp. dura (Eyma) T.D. Penn., Fl. Neotrop. Monogr. 52: 352. 1990. —Pouteria dura Eyma, Recueil Trav. Bot. Néerl. 33: 187. 1936. —Chuponcillo, Pendarito. Tree to 28–30 m tall, 38 cm diameter; latex white; flowers yellow, fragrant. Evergreen lowland to lower montane forests, 50– 500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Río Apacará, Sierra Pakaraima), Amazonas (Río Guainía). Panama, Colombia, Guyana, Suriname, Ecuador, Peru, Amazonian Brazil. P. cuspidata subsp. robusta (Mart. & Eich.) T.D. Penn., Fl. Neotrop. Monogr. 52: 351. 1990. —Sideroxylon robustum Mart. & Eich. in Mart., Fl. Bras. 7: 56. 1863. Oxythece robusta var. brevipetiolasta Monach., Fieldiana, Bot. 28: 486. 1953. Tree to 10–30 m tall, 33 cm diameter; bark brown; latex milky white; flowers whitish cream to yellowish green; fruit brown. Slopes of mixed montane forests, igneous rock beside rivers, 100–400 m; Bolívar (Cerro Ichún, upper Río Paragua), Amazonas (Caño Negro, Cerro Sipapo, Río Coromoto, Río Cunucunuma, San Carlos de Río Negro). Aragua, Sucre; Panama, Colombia, Suriname, French Guiana, Peru, Amazonian Brazil. The fruit of Pouteria cuspidata subsp. robusta is edible, and the hard durable timber is used locally for construction. Pouteria egregia Sandwith, Kew Bull. 479. 1931. —Pulguillo, Pulguillo amarillo. Tree to 20–35 m tall, 27–90 cm diameter; bole cylindrical, buttresses 2.5 m tall, wood hard; leaves gray-green, glaucous, narrow

120

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and revolute; latex sticky, milky white; flowers pale green to yellowish red; fruit yellow, dull orange, or greenish white, 1-seeded, edible, latex present. Evergreen lowland and semi-evergreen forests, 200–700 m; Delta Amacuro (Serranía de Imataca), northeastern Bolívar. Carabobo; Colombia, Guyana, Suriname, French Guiana, Brazil (Maranhão, Pará, Rondonia). ◆Fig. 102. Pouteria elegans (A. DC.) Baehni, Candollea 9: 197. 1942. —Sideroxylon elegans A. DC., Prodr. 8: 183. 1844. —Cachito, Pimúyure (Curripaco), Pico de lombriz, Shibwichi. Small bushy tree 2–20 m tall; trunk twisted; bark dark brown, fissured; lower surface of leaves glaucous or pale; latex copious, sticky, white; flowers fragrant; calyx green; corolla white; fruit pale green to dark ferruginous-brown, with brown pubescence, ovoid, edible. Seasonally flooded riparian forests, nonflooded gallery forests, 100–200 m; Amazonas (widespread). Apure; Amazonian Colombia, Guyana, Amazonian Peru, Brazil, and Bolivia. ◆Fig. 101. Pouteria eugeniifolia (Pierre) Baehni, Candollea 9: 218. 1942. —Micropholis eugeniifolia Pierre, Not. Bot. Sapot. 40. 1891. —Caramacarte, Chicle rosado. Micropholis steyermarkii Monach., Fieldiana, Bot. 28: 481. 1953. Pouteria steyermarkii Monach., Fieldiana, Bot. 28: 482. 1953, nom. altern. Tree 8–36 m tall, 25–56 cm diameter; buttresses to 3 m high; bark gray-brown, shredding; lower surface of leaves glaucous or silvery-green; latex scant or abundant, white; flowers cream; fruit medium dark brown. Lower montane to montane forests, along watercourses, savannas, bluffs, 200–1100 m; Delta Amacuro (Serranía de Imataca), Bolívar (Gran Sabana, Serranía de Imataca, Upata-San Félix), Amazonas (Cerro Sipapo, Cerro Yutajé). Mérida, Miranda, Zulia; Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 103. The timber of Pouteria eugeniifolia is used locally for construction and firewood. Pouteria fimbriata Baehni, Candollea 14: 68. 1952. —Caimito blanco.

Tree to 20 m tall; latex white; flowers greenish white; fruit ripening yellow. Lowland evergreen to montane forests, 100– 1300 m; Bolívar (near El Dorado), Amazonas (Sierra de la Neblina). Guyana, Brazil (central Amazonas). Pouteria gabrielensis (Gilly ex Aubrév.) T.D. Penn., Fl. Neotrop. Monogr. 52: 354. 1990. —Neoxythece gabrielensis Gilly ex Aubrév., Mem. New York Bot. Gard. 23: 221. 1972. Tree to 10 m tall; latex milky; flowers greenish white. Along black-water rivers, Río Negro caatinga, 100–200 m; Amazonas (Río Guainía). Amazonian Colombia and Brazil (Amazonas: Rio Negro basin). Pouteria glomerata (Miq.) Radlk., Sitzungsber. Math.-Phys. Kl. Bayer. Akad. Wiss. München 12: 333. 1882. —Lucuma glomerata Miq. in Mart., Fl. Bras. 7: 81, t. 36, fig. 2. 1863. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, Paraguay; 2 subspecies, both in the flora area. The fruit of Pouteria glomerata is large, sweet, and edible. Key to the Subspecies of P. glomerata 1. Leaf base usually acute, obtuse, or rounded, apex often obtuse, lower surface minutely puberulous (lens) with closely appressed whitish trichomes, usually forming a pellicle, often becoming subglabrous with age ........................ ................................ subsp. glomerata 1. Leaf base and apex narrowly attenuate, lower surface persistently sericeous with yellowish buff indumentum ........... ...................................... subsp. stylosa P. glomerata subsp. glomerata. —Chupón, Temare. Labatia parviflora Pittier, Contrib. Fl. Venez. 41. 1923. Pouteria pittieri Baehni, Candollea 9: 254. 1942. Tree 6–8 m tall; flowers white-cream or yellow; fruit brown, pulp yellow, 1- or 2-

Pouteria 121

seeded, latex present. Seasonally flooded evergreen lowland forests, occasionally nonflooded, 100–200 m; Bolívar (Río Cuyuní basin), Amazonas (Capibara, Tamatama). Widespread elsewhere in Venezuela; throughout the range of the species. P. glomerata subsp. stylosa (Pierre) T.D. Penn., Fl. Neotrop. Monogr. 52: 420. 1990. —Guapeba stylosa Pierre, Not. Bot. Sapot. 42. 1891. Tree 5–40 m tall; bark dark; flowers pale cream to greenish white; fruit maturing yellowish to pale brown. Evergreen lowland to montane forests, 50–200 m; Bolívar (road between Caicara and Puerto Ayacucho, Río Suapure). Falcón, Zulia; Costa Rica, Panama, Colombia, Brazil (Amazonia to Maranhão). Pouteria gomphiifolia (Mart.) Radlk., Sitzungsber. Math.-Phys. Kl. Bayer. Akad. Wiss. München 12: 33. 1882. —Lucuma gomphiefolia Mart. ex Miq. in Mart., Fl. Bras. 7: 78, t. 37, fig. 1. 1863. —Rebalsero, Temarito. Lucuma gomphiaefolia var. blepharantha Mart., Fl. Bras. 7: 78. 1863. Gomphiluma martiana Baill., Hist. Pl. 11: 285. 1891. Tree 15–25 m tall, to 60 cm diameter; bark scaly and slightly pink; latex present; fruit dull yellow-gold, green, or dark brown, pear-like, edible. Evergreen lowland forests of periodically and permanently flooded low ground, 100–200 m; Amazonas (Capibara, Río Siapa, Río Ventuari, San Carlos de Río Negro). Apure; Amazonian Colombia, Peru, and Brazil (Amazonas, Roraima). ◆Fig. 104. Pouteria guianensis Aubl., Pl. Guiana 85, t. 33 (excl. fruct). 1775. —Wa-won-yek, We-won-yek (Arekuna). Labatia tovarensis Engl., Bot. Jahrb. Syst. 12: 515. 1890. —Pouteria tovarensis Klotzsch & Karst. ex Engl. & Prantl, Nat. Pflanzenfam. 4(1) Nachträge: 142. 1897. Pouteria caimito var. strigosa Monach., Fieldiana, Bot. 28: 487. 1953. Tree (5–)12–23 m tall; trunk fluted or with slender buttresses; latex milky white; buds and flowers greenish white to yellow;

fruit ripening orange-yellow. Evergreen lowland riparian to montane forests, 200–1200 m; Bolívar (Gran Sabana), Amazonas (Caño Negro near Cerro Duida). Widespread elsewhere in Venezuela; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil (Amazonian and coastal regions). ◆Fig. 94. The wood of Pouteria guianensis provides useful construction timber. In the past, the latex has been used as a source of balata. The fruit is edible and similar in flavor to that of P. caimito. Pouteria hispida Eyma, Recueil Trav. Bot. Néerl. 33: 177. 1936. —Caimito amarillo, Caimito negro. Tree to 25 m tall, 25 cm diameter; buttresses 1.5 m tall; latex white; flowers greenish white; fruit hispid, ripening yellow or orange with thick, hard flesh, edible. Evergreen lowland forests, lower montane forests on bauxite, 200–500 m range; Delta Amacuro (Serranía de Imataca), Bolívar (Serranía de Imataca, Serranía de los Pijiguaos). Costa Rica, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, and Bolivia. ◆Fig. 108. Pouteria laevigata (Mart.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 14: 457. 1884. —Labatia laevigata Mart., Flora 21(Beibl. 2): 92. 1838 (reprinted as Herb. Fl. Bras. 172). Unbuttressed tree 20 m tall; bark smooth; flowers cream or white; fruit maturing bright yellow, flesh rather tough, sweet, edible. Nonflooded forests, 100–200 m; Amazonas (upper Río Baria, Río Guainía). Amazonian Colombia, French Guiana, Amazonian Peru and Brazil. ◆Fig. 107. Pouteria lucumifolia (Reissek ex Maxim.) T.D. Penn., Fl. Neotrop. Monogr. 52: 282. 1990. —Ilex lucumifolia Reissek ex Maxim., Mém. Acad. Imp. Sci. SaintPétersbourg sér. 7, 29(3): 28. 1881. —Majagua hoja fina, Yuquito. Pseudocladia orinocoensis Aubrév., Mem. New York Bot. Gard. 23: 204. 1972. Pseudocladia cassiquiarensis Aubrév., Mem. New York Bot. Gard. 23: 205. 1972.

122

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Tree 6–20 m tall 10–50 cm diameter or shrub to 2 m tall; outer bark dark brown, fissured, scaly, hard, inner bark fleshy; latex white when present; flowers whitish or green to yellow; fruit usually green, sometimes orange or green-brown, ovoid. Open areas, seasonally flooded forests, high forests on nonflooded ground, forests along river banks, 100–200 m; Amazonas (Río Mawarinuma, Solano, Tamatama). Amazonian Ecuador, Peru, and Brazil. Pouteria maguirei (Aubrév.) T.D. Penn., Fl. Neotrop. Monogr. 52: 448. 1990. —Prieurella maguirei Aubrév., Mem. New York Bot. Gard. 23: 218. 1972. —Cabeza de baba, Cabeza de babocilla, Chudumiro, Palo de babilla. Tree 5–18 m tall with dense crown; latex milky; leaves ± pendent; flowers greenish white; fruit yellow-green to mealy brown, irregulary wrinkled, 7–10 cm long, with edible aril. Seasonally flooded banks of blackwater rivers, 100–200 m; Amazonas (Cerro Yapacana base, Río Pacimoni, Río Temi, San Carlos de Río Negro, Tamatama). Brazil (Amazonas: upper Rio Negro basin). Pouteria orinocoensis (Aubrév.) T.D. Penn., Fl. Neotrop. Monogr. 52: 254. 1990. —Paralabatia orinocoensis Aubrév., Mem. New York Bot. Gard. 23: 205. 1972. —Cacho de venado. Tree 3–15 m tall; latex scant, white; flowers greenish white. Savanna edges, periodically flooded forests, 50–100 m; frequent in northern Bolívar, Amazonas (Isla Ratón, Puerto Ayacucho). Apure. ◆Fig. 109. Pouteria pimichinensis T.D. Penn., Fl. Neotrop. Monogr. 52: 1990. —Neoxythece maguirei Aubrév., Mem. New York Bot. Gard. 23: 223. 1972. Small bushy tree 4–8 m tall or shrub 2–4 m tall; leaves blue- or maroon-green; latex thick, sticky, white; flowers white, fragrant; fruit dark ferruginous-brown, globose. Seasonally flooded black-water riparian forests, savannas, Río Negro caatinga, 100–200 m; southeastern Amazonas. Endemic. ◆Fig. 110. Pouteria plicata T.D. Penn., Fl. Neotrop. Monogr. 52: 456. 1990. —Perguillo.

Tree to 15 m tall, 50 cm diameter; flowers greenish white; fruit maturing yellowish. Periodically flooded forests, 50–200 m; Delta Amacuro (base of Serranía de Imataca), Bolívar (mouth of Río Caura). Apure; Amazonian Peru, Amazonian and northeastern Brazil, Amazonian Bolivia. Pouteria reticulata (Engl.) Eyma, Recueil Trav. Bot. Néerl. 33: 183. 1936. —Chrysophyllum reticulatum Engl., Bot. Jahrb. Syst. 12: 522. 1890. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amapa, Amazonian, Atlantic coastal), Bolivia; 2 subspecies, 1 in Venezuela, 1 in the flora area. P. reticulata subsp. reticulata. —Anamu, Kanke, Mai pihpe. Tree 5–25 m tall, 11–25 cm diameter; latex sticky, white; flowers greenish; fruit maturing orange, red, purple, or black, sweet, edible. Semideciduous forests on granitic slopes, 100–400(–1000) m; Delta Amacuro (Río Grande), Bolívar (between Kavanayén and Río Pacairao, along Río Orinoco), Amazonas (upper Río Casiquiare). Apure, Miranda, Yaracuy, Zulia; Mexico, Central America, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 112. Pouteria rigida (Mart. & Eich.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 12: 333. 1882. —Lucuma rigida Mart. & Eich. in Mart., Fl. Bras. 7: 73. 1863. Venezuela, adjacent Guyana and Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of P. rigida 1. Lower surface of leaves soon glabrous ....................................... subsp. rigida 1. Lower surface of leaves persistently ferruginous-tomentose subsp. tomentosa P. rigida subsp. rigida Neoxythece roraimaensis Aubrév., Adansonia n.s. 4: 230. 1964. Shrub or tree 1–20 m tall, the crown dense; leaves revolute, silvery gray-white;

Pouteria 123

latex scarce or copious, sticky, white; flowers ranging from white, cream, or yellow, to tancolored; calyx tawny, corolla greenish yellow or cream-orange; fruit orange-brown to dark ferruginous, pubescent, ovoid, 1–1.5 × 0.5–1 cm. Rocky, shrubby, dry or wet savannas, shrub islands, open scrub (often on sandstone), elfin forests, gallery forests, mossy forests, 400–2200 m; Bolívar (widespread on Gran Sabana and associated tepuis), Amazonas (upper Río Yatúa, Sierra de la Neblina). Other distribution as in species. P. rigida subsp. tomentosa (Aubrév.) T.D. Penn., Fl. Neotrop. Monogr. 52: 358. 1990. —Neoxythece tomentosa Aubrév., Mem. New York Bot. Gard. 23: 222. 1972. Shrub or small tree 1–6 m tall, the crown dense; leaves revolute, ferruginous; latex white; flowers greenish; calyx ferruginous; fruit ferruginous-brown, lepidote. Forest edges, low scrub, dwarf forests, 1300–1900 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Jaua), Amazonas (Cerro Duida). Endemic. ◆Fig. 106. Pouteria scrobiculata Monach. ex T.D. Penn., Fl. Neotrop. Monogr. 52: 358. 1990. —Pouteria scrobiculata Monach., Fieldiana, Bot. 28: 485. 1953, nom. inval. —Oxythece scrobiculata Monach., Fieldiana, Bot. 28: 484. 1953, nom. inval. —Para-to-re-yek, Tee-u-ben-yek. Tree 4–20 m tall, to 25 cm diameter; lower surface of leaves blue-green to silverygray; latex milky white; flowers buff-white to yellowish green. Forested slopes extending to summits and along ridges, especially riparian forests, humid mossy montane forests, dry, dense, dwarf forests in deep crevices of exposed sandstone, 600–1700 m; Bolívar (Auyán-tepui, Gran Sabana, Ptaritepui, Río Cuyuní, Sierra de Lema), Amazonas (Cerro Parú, Cerro Yutajé, Sierra de la Neblina). Guyana, Brazil (Amazonas: Serra Aracá). ◆Fig. 105. Pouteria sipapoensis T.D. Penn., Fl. Neotrop. Monogr. 52: 367. 1990. Tree 10–15 m tall; latex scant, white; lower leaf surface with ferruginous tomentum; calyx brown, corolla cream. Evergreen

lowland to lower montane forests, 100–500 m; Amazonas (Cerro Sipapo, Río Cuao). Endemic. Pouteria stipitata Cronquist, Lloydia 9: 265. 1946. Tree to 15 m tall; leaves glandular-striate; flowers yellow or greenish white, fragrant; fruit maturing yellow with an unpleasant odor. Gallery forests, ca. 50 m; Bolívar (Caño Orocopiche along road between Ciudad Bolívar and Ciudad Piar). Anzoátegui, Falcón, Lara, Portuguesa, Yaracuy, Zulia; Panama, Colombia, possibly Amazonian Brazil. Pouteria surumuensis Baehni, Candollea 9: 362. 1942. —Makararo. Tree to 10 m tall, 11 cm diameter; latex sticky, free-flowing, white; lower surface of leaves golden brown; flowers pale green to greenish white; fruit golden brown, edible. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma, upper Río Orinoco). Guyana, Amazonian Colombia, Brazil (Amazonas, Roraima). Pouteria torta (Mart.) Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer Akad. Wiss. München 12: 333. 1882. —Labatia torta Mart., Flora 21(Beibl. 2, 4): 94. 1838. Mexico, Central America, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia, Paraguay; 4 subspecies, 1 in Venezeula. P. torta subsp. glabra T.D. Penn., Fl. Neotrop. Monogr. 52: 484. 1990. —Labatia torta Mart., Flora 21(Beibl. 2, 4): 94. 1838 (reprinted as Herb. Fl. Bras. 174). —Bampara amarilla. Tree 12–25 m tall, with dense crown; latex scant, white; fruit yellow to brown with brown irritating trichomes, edible. Semideciduous to evergreen lowland and lower montane forests, 100–800 m; Bolívar (near El Dorado), Amazonas (Río Cataniapo, Río Coro Coro west of Cerro Yutajé, Simarawochi). Mérida-Barinas border; Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 111.

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Pouteria trilocularis Cronquist, Bull. Torrey Bot. Club 73: 469. 1946. Franchetella neblinaensis Aubrév., Mem. New York Bot. Gard. 23: 225. 1972. Tree 18–25 m tall, to 25 cm diameter; latex abundant, fast-flowing, milky white; flowers white or cream-green to yellow; fruit orange-brown, globose. Evergreen lowland forests, along streams flowing over granite outcrops, 50–200 m; Amazonas (near Puerto Ayacucho, Río Coromoto). Widespread elsewhere in Venezuela; Colombia, Amazonian Ecuador, Peru, Brazil, and Bolivia.

corolla and calyx light green; fruit edible, ranging from green with yellow spots, yellow, orange, to brown, with dark brown ferruginous-brown velvety pubescence, 5–8 cm long. Evergreen lowland to montane forests, 100–600(–900) m; Delta Amacuro (Serranía de Imataca), Bolívar (mostly northern portions, also Gran Sabana), Amazonas (Cerro Sipapo, near Puerto Ayacucho, Río Coro Coro west of Cerro Yutajé, Río Manapiare basin). Apure; Guyana, Suriname, French Guiana, Amazonian and eastern coastal Brazil. ◆Fig. 92.

Pouteria ucuqui Pires & R.E. Schult., Bot. Mus. Leafl. 14: 87. 1950. —Hinirji (Curripaco), Yuco, Yuú (Baniwa). Tree 35–40 m tall, to 60 cm diameter; buttresses to 2 m; latex copious, insipid, white; flowers yellowish, strongly scented; fruit fleshy, obovoid, edible, 10–11 cm long, with white latex. Evergreen nonflooded forests, 100–200 m; Amazonas (San Carlos de Río Negro). Colombia (Guainía, Vaupés), Brazil (Amazonas: Rio Negro basin). The flesh of the ripe fruit of Pouteria ucuqui is edible and delicious, forming an important part of the diet of local people at certain times of the year. The seeds have been used to make toy whistles by removing the embryo and drilling several small holes in the seed coat.

Pouteria aff. engleri Eyma, Recueil Trav. Bot. Néerl. 33: 178. 1936. —Punwoyo, Punuwoyo. Tree to 9–20 m tall, 16–30 cm diameter; outer bark fluted, dark brown, hard, inner bark light orange, soft; latex sticky, white; fruit green. Semideciduous forested granitic slopes, 50–400 m; northwestern Bolívar. Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. Two collections from the flora area are tentatively placed in Pouteria engleri: Boom & Grillo 6484 (K) and Boom & Wentzel 6666 (K).

Pouteria venosa (Mart.) Baehni, Candollea 9: 393. 1942. —Achras venosa Mart., Flora 22(Beibl. 1): 4. 1839. Venezuela, Guyana, Suriname, French Guiana, Amazonian and southeastern coastal Brazil; 2 subspecies, 1 in Venezuela and the flora area. P. venosa subsp. amazonica T.D. Penn., Fl. Neotrop. Monogr. 52: 399. 1990. —Achras venosa Mart., Flora 22(Beibl. 1): 4. 1839 (reprinted as Herb. Fl. Bras. 180). —Arbol de vaca, Bampara, Capure, Conserva, Goma de guaca, Juachetuma (Yekwana), Merecurillo, Nispero montañero, Perguillo negro, Purguillo chicle, Purguillo terciopelo, Temare. Tree 10–40 m tall, 18–43 cm diameter; bole fluted; bark dark brown with small scales; heartwood pink; latex copious, sticky, white; flowers frequent, white to pale green;

Pouteria aff. minima T.D. Penn., Fl. Neotrop. Monogr. 52: 285. 1990. Tree; latex milky. Alluvial terraces, 600– 700 m; Bolívar (Río Suapare). Brazil (northwestern Amazonas). Pouteria minima is included for the flora area based on a single, questionable collection, Guánchez et al. 4684 (K). Podoluma pacimoniensis Aubrév., Mem. New York Bot. Gard. 23: 218. 1972. Tree 4–5 m tall; latex milky; fruit red. Savanna margins, 100–200 m; Amazonas (Río Pasimoni). Brazil (Amazonas: Serra Aracá). While the nomenclatural type of Podoluma, P. catocladantha (Eichler) Aubrév., has been placed in the synonymy of Pouteria, P. pacimoniensis is of uncertain affinity and is treated here for lack of better placement. It may belong in Pouteria sect. Franchetella, but if so, then it is isolated and quite unlike any other species. The general appearance of the leaves and their venation suggest Elaeoluma, and it also shares with E. schomburgkiana the tendency to have a small beak above the seed scar.

Pouteria 125

Fig. 92. Pouteria venosa subsp. amazonica

Fig. 93. Pouteria canaimaensis

Fig. 94. Pouteria guianensis

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S APOTACEAE

Fig. 95. Pouteria amygdalicarpa

Fig. 96. Pouteria anomala

Pouteria 127

Fig. 97. Pouteria bangii

Fig. 98. Pouteria caimito

128

S APOTACEAE

Fig. 99. Pouteria cayennensis

Fig. 100. Pouteria cuspidata subsp. cuspidata

Pouteria 129

Fig. 101. Pouteria elegans

Fig. 102. Pouteria egregia

Fig. 103. Pouteria eugeniifolia

130

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Fig. 104. Pouteria gomphiifolia

Fig. 106. Pouteria rigida subsp. tomentosa

Fig. 105. Pouteria scrobiculata

Pouteria 131

Fig. 107. Pouteria laevigata

Fig. 108. Pouteria hispida

132

S APOTACEAE

Fig. 109. Pouteria orinocoensis

Fig. 110. Pouteria pimichinensis

Pouteria 133

Fig. 111. Pouteria torta subsp. glabra

Fig. 112. Pouteria reticulata subsp. reticulata

134

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9. PRADOSIA Liais, Clim. Geol. Geogr. Bot. Bresil 614. 1872. Shrubs or trees. Leaves often opposite or verticillate, less frequently spirally arranged; venation usually eucamptodromous, less frequently brochidodromous, midrib usually sunken on the upper surface, rarely flat or raised, secondaries often impressed on upper surface; intersecondaries usually absent, rarely well developed; tertiaries usually oblique or horizontal; minute paired stipels sometimes present on petiole. Usually cauliflorous or ramiflorous, less frequently with axillary inflorescence. Flowers bisexual. Calyx a single whorl of usually 5 sepals; corolla rotate, tube nearly always shorter than the widely spreading lobes, corolla lobes usually 5. Stamens fixed at top of corolla tube or on base of lobes, exserted; filaments long, geniculate below the apex, and strongly narrowed below insertion of anther; staminodes absent. Disk absent. Ovary (4)5(6)-locular, style short. Fruit a drupe with thinly cartilaginous endocarp, often slightly asymmetric. Seed solitary, with smooth, shining testa and full-length adaxial scar; embryo with thinly plano-convex cotyledons, radicle often exserted or only extending to the surface; endosperm a thin sheath or absent. Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay; 23 species, 6 in Venezuela, 5 of these in the flora area. Key to the Species of Pradosia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5.

Petiole with a pair of minute stipels (1–2 mm long) ................ P. grisebachii Petiole without paired stipels ................................................................... 2 Venation brochidodromous ............................................... P. schomburgkiana Venation eucamptodromous ...................................................................... 3 Intersecondary veins well developed, often extending to the leaf margin ....................................................................................... P. schomburgkiana Intersecondary veins short or absent ....................................................... 4 Lower surface of leaves densely short golden brown-pubescent; fruit 1.5–1.6 cm long ............................................................................ P. beardii Lower surface of leaves not densely golden brown-pubescent; fruit 2– 6.5 cm long ............................................................................................. 5 Petiole 0.8–1.2 cm long; corolla ca. 2.5 mm long .................. P. surinamensis Petiole 1.5 cm long or more; corolla 3–3.5 mm long ................. P. caracasana

Pradosia beardii (Monach.) T.D. Penn., Fl. Neotrop. Monogr. 52: 665. 1990. —Chrysophyllum beardii Monach., Phytologia 3: 159. 1949. —Cakak-oraiyek. Chrysophyllum monachinoanum Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 432. 1966. Glycoxylon rubrum Aubrév., Mem. New York Bot. Gard. 23: 214. 1972. Tree 3–25 m tall or shrub 2–6 m tall; leaves revolute to sulcate, the lower surface ferruginous; latex thick, white; flowers pale to bright green, buff, or yellow-ferruginous, calyx tawny brown; fruit brownish orange to ferruginous, edible. Sandy scrub, shrub is-

lands, savannas, dwarf forests, 700–1800 m; Bolívar (Acopán-tepui, Cerro Guaiquinima, Cerro Guanacoco, Cerro Marutaní, Cerro Sarisariñama, Cerro Venamo and neighboring Cerro Uroi, Los Testigos, Sierra de Lema, Sierra Pakaraima), Amazonas (Cerro Huachamacari slopes). Trinidad, Guyana. ◆Fig. 114. Pradosia caracasana (Pittier) T.D. Penn., Fl. Neotrop. Monogr. 52: 668. 1990. —Oxythece caracasana Pittier, Bol. Comerc. Industr. Venez. no. 13: 425. 1921 [reprinted in Arb. Arbust. Venez. 1: 11. 1923]. —Chupón.

Pradosia 135

Fig. 113. Pradosia schomburgkiana subsp. schomburgkiana

Fig. 114. Pradosia beardii

136

S APOTACEAE

Fig. 115. Pradosia surinamensis

Tree 20–30 m tall; latex thin, milky; flowers pale green; fruit 6–6.5 cm long, densely lenticellate, brown. Evergreen lowland to montane forests, 100–600 m; Bolívar (Altiplanicie de Nuria, Serranía de Imataca, Tumeremo-Anacoco road). Widespread elsewhere in Venezuela; Trinidad. Pradosia grisebachii (Pierre) T.D. Penn., Fl. Neotrop. Monogr. 52: 655. 1990. —Ecclinusa grisebachii Pierre, Not. Bot. Sapot. 57. 1891. —Chupón. Tree to 24 m tall, 80 cm diameter; latex present; fruit maturing dull orange. Evergreen lowland forests, 100–200 m; Delta Amacuro (Serranía de Imataca), Bolívar (El Dorado). Miranda; Trinidad. ◆Fig. 116. Pradosia schomburgkiana (A. DC.) Cronquist, Bull. Torrey Bot. Club 73: 311. 1946. —Chrysophyllum schomburgkianum A. DC., Prodr. 8: 157. 1844. Amazonian Colombia, southern Venezuela, Guyana, Suriname, northern and central Amazonian Brazil; 2 subspecies, both in the flora area.

(Arekuna), Matapalo, Temarito, Temarito banero, Yoquito. Glycoxylon maguirei Aubrév., Mem. New York Bot. Gard. 23: 214. 1972. Dense shrub, treelet, or buttressed tree 3–20 m tall, 7–30 cm diameter; bark lenticular; leaves revolute; latex scant, clear to milky white; flowers fragrant, whitish to yellow or green; fruit yellow or pale green, becoming maroon and black, pubescent, edible, subglobose, 0.8–1 × 1–2 cm. Locally abundant in scrub and savannas on white sand or podsol, low open forests, gallery forests, Mauritia palm swamps, 100–1400 m; widespread in Bolívar and Amazonas. Distribution outside Venezuela as in species. ◆Fig. 113.

Key to the Subspecies of P. schomburgkiana 1. Leaves glabrous; fruit subglabrous ....... ..................... subsp. schomburgkiana 1. Lower surface of leaves finely brown- to grayish-sericeous; fruit appressed puberulous ........................ subsp. sericea P. schomburgkiana subsp. schomburgkiana. —Guayabón, Icaco, Kakka-yek

P. schomburgkiana subsp. sericea T.D. Penn., Fl. Neotrop. Monogr. 52: 660. 1990. —Chrysophyllum schomburgkianum A. DC., Prodr. 8: 157. 1844. Shrub or treelet to 8 m tall, with broad, dense crown; leaves ferruginous; latex sparse or abundant, sticky, transparent, white; flowers greenish yellow; fruit ferruginous-brown, velvety, ovoid, 1.2–2 cm long.

Pradosia 137

Fig. 116. Pradosia grisebachii

Savannas or shrub islands in savannas, periodically flooded savannas, shrubby vegetation on white sand, seasonally dry evergreen forested slopes, 100–500 m; Amazonas (Caño Caname, Caño Cotua, near Maraoa, Río Autana, Río Guayapo. Brazil (Amazonas: Serra Aracá). Pradosia surinamensis (Eyma) T.D. Penn., Fl. Neotrop. Monogr. 52: 652.

1990. —Pouteria surinamensis Eyma, Recueil Trav. Bot. Néerl. 33: 189. 1936. —Chupón, Gateado, Kahka, Kahka yo, Yoron (Arekuna). Tree 8–35 m tall, 20–80 cm diameter; trunk buttressed and fluted at base; outer bark smooth, dark brown-gray, inner bark soft, flesh-colored to orange; latex scant, sticky, white; flowers greenish yellow; fruit pulp yellow, edible. Gallery forests, semideciduous forests on granitic slopes, 50–400 m; northern Bolívar. Guyana, Suriname, French Guiana, Brazil (Pará, Roraima). ◆Fig. 115.

138

S ARRACENIACEAE

SARRACENIACEAE by Paul E. Berry, Ricarda Riina, and Julian A. Steyermark Perennial, insectivorous herbs. Leaves alternate, grouped in basal rosettes or along an erect, scarcely branched stem, with inflated, hollow, pitcher-like traps which are ventrally ridged or winged and usually have a hood-like prolongation or thickened appendages on the dorsal summit; stipules absent or sometimes interpreted as stipular wings on the ventral margin of the pitcher; petiole short, grading upward into the pitcher; pitchers partly filled with water, the lower inner surface smooth, the upper inner surface glandular and with stiff, retrorse trichomes, the epidermis of the outer portion of the trap with nectariferous glands and frequently with stiff, antrorse trichomes. Inflorescence a generally erect raceme with 1–several(–10) nodding or pendent flowers, these showy, bisexual, and actinomorphic. Sepals 3–6, imbricate, persistent, mostly colored and petaloid; petals either 5 or absent, imbricate, deciduous. Stamens (4–)8–20 or more; filaments stout, distinct; anthers 2-locular, basifixed or dorsifixed, poricidal. Ovary superior, with 3 or 5 locules and carpels; placentation axile, sometimes intruded-parietal in upper part of ovary through separation of the dissepiments; style solitary, subentire or shortly 5branched with terminal stigmas, or expanded, peltate, or umbrella-like with a small stigma beneath the tip of each of the 5 lobes; ovules numerous, anatropous. Fruit a loculicidal capsule. Seeds numerous, small, frequently with a wing-like beak; embryo minute; endosperm abundant. Eastern Canada, U.S.A., Venezuela, Guyana, Brazil; 3 genera and 17 species; 1 genus and 10 species in the flora area. 1. HELIAMPHORA Benth., Trans. Linn. Soc. London 18: 429, t. 29. 1840. Herbs with fibrous roots, usually with a rosette habit or with shortened or elongated, monopodial stems to 2(–4) m tall; stems simple or with shortened lateral axes bearing abbreviated rosettes on the side or near the apex to produce a bifurcate appearance, or with elongated basal leafy axes or offshoots. Leaves inflated, hollow, pitcher-like, pale green, maroon, or wine-colored, from ventricose and expanded at the middle to ± tubular, flaring or slightly broadened at the oblique summit, subrevolute at the margin, ventrally ridged or winged; wing with a pore in the seam at its base to control the water level within, dorsally ending at the summit in an expanded, thickened appendage (rarely absent or scarcely developed), which is suborbicular, ovate-oblong, or ligulate-oblong, wine-red or maroon, sweet-smelling, and sessile, narrowed, or stipitate at the base; pitchers glabrous or minutely pubescent on the outer surface, with elongate, parallel veins connecting a transversely reticulate venation, pubescent within with 2 kinds of trichomes, those on the upper part densely crowded, short and retrorse, with the shortest in the uppermost part, the others much more elongated and forming a band at the base of the pubescent zone, below which the tube is completely glabrous, or rarely the pitcher completely glabrous within except for the lowest zone of long trichomes. Inflorescence a scapose, (1)2–10-flowered raceme. Pedicels subtended by a leafy bract with an appendage similar to that of the pitcher or else absent or only cuspidate. Tepals 4–6, petaloid, white, rose, or magenta, often greenish in fruit, oblong to lanceolate, the inner 2 shorter and narrower than the outer ones. Stamens (4–)8– 20; anthers basifixed, erect in bud, then reflexed at anthesis, linear-oblong or oblong, dehiscent through 2 slits at the base of the anthers. Ovary 3-locular; ovules

Heliamphora 139

numerous, pluriseriate; style simple, truncate at the apex; stigma small, obscurely 3-lobed. Seeds obovate, compressed; testa brown, expanded into a membranous brown wing surrounding the seed. Endemic to the Guayana Shield in southern Venezuela, western Guyana, and northern Brazil (Amazonas); 10 species, all in the flora area. Unlike the other two North American genera of Sarraceniaceae, Heliamphora has poricidal anthers and is buzz-pollinated by large bees. There is a high degree of variability and plasticity in the vegetative and even floral characters among members of the genus, and some of the recognized taxa are not sharply differentiated. A more critical review of the genus will be needed to assess if all of these are worthy of recognition. Key to the Species of Heliamphora 1.

1.

2(1).

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

6.

7(6).

7.

Lid of the pitcher emerging directly from the back of the pitcher mouth without any constriction, 4–5 cm long × 3–4 cm wide, triangular and usually upright ................................................................ H. sarracenioides Lid of the pitcher hood-like or at least constricted at the base, smaller than above and bent over the mouth of the pitcher to some degree ................................................................................................................ 2 Back of the lid or hood-like prolongation on the dorsal summit of adult pitchers bearing a hollow structure (follicle) running downward 5– 10 mm ..................................................................................... H. folliculata Lid or hood-like prolongation without the hollow structure described above ...................................................................................................... 3 Lowest floral bract (4.5–)6–21 cm long, as long as or longer than the corresponding pedicel ................................................................................. 4 Lowest floral bract 2–6(–10) cm long, sometimes shorter than the corresponding pedicel .................................................................................... 5 Pitcher lid spoon-shaped, with a sharp contraction at the base; anthers ca. 5 mm long; Bolívar state .............................................. H. chimantensis Pitcher lid rectangular with a very slight contraction at the base; anthers 7–9 mm long; Amazonas state ............................................... H. tatei Inflorescence (2–)8–10-flowered; pitchers (15–)40–50 cm long; Ilú-tepui, Bolívar state ................................................................................. H. ionasii Inflorescences (1)2–7-flowered; pitchers 5–40 cm long; known from several tepuis, including Ilú-tepui ............................................................. 6 Pedicels mostly pubescent; the upper, inner face of the pitcher (excluding the basal ring of trichomes) with some or all trichomes to 2–5 mm long and scattered or ± distantly spaced, or deciduous over a glabrous surface ......................................................................................................... 7 Pedicels mostly glabrous; the upper inner face of the pitcher (excluding the basal ring of trichomes) with a dense, minute, ± uniform velvety indumentum of trichomes 0.5–1 mm long ........................................... 8 Peduncle 20–40 cm long, pedicels 3–4 cm long; anthers ca. 15 per flower, 4 mm long; pitcher lid strongly helmet-shaped, 0.5–1 cm long; known from Auyán-tepui and Macizo del Chimantá .............................. H. minor Peduncle ca. 50 cm long, pedicels ca. 5 cm long; anthers ca. 10 per flower, 7 mm long; pitcher lid cordate, curved, 1–1.5 cm long; known from Sierra de la Neblina ....................................................................... H. hispida

140

8(6).

8.

9(8). 9.

S ARRACENIACEAE

Anthers (4.5–)5–8 mm long; upper pubescent zone of the inside of the pitcher 7–18 cm long, occupying 3/8–1/2 the length of the pitcher, the indument of the upper zone of trichomes mostly 0.5–0.7 mm long, the lower part with longer trichomes 0.7–1 mm long; pitcher appendage longer than broad or as long as broad, 6–35 × 5–30 mm .... H. heterodoxa Anthers 2.5–3.5 mm long; upper pubescent zone of the inside of the pitcher 6–10 cm long, occupying 1/3–3/8 the length of the pitcher, most of the indument of ± uniform trichomes 0.7–1 mm long; pitcher appendage about as broad as long ............................................................ 9 Pitchers 20–32 cm long, 3–3.5 cm wide, narrowly cylindrical in upper half, appendage helmet-shaped, 10–30 × 10–28 mm .............. H. elongata Pitchers 8–18 cm long, 2–6 cm wide, funnel-shaped in upper half, appendage cordate, 5–10 × 5–12 mm ......................................... H. nutans

Heliamphora chimantensis Wistuba, Carow & Harbath, Carnivorous Pl. Newslett. 31: 78. 2002. Plant with branching rhizomes, forming dense colonies; pitchers entirely green with red lid, 20–35 cm tall, upper part glabrous on the inner side; lid spoon-shaped, upright, ending with a sharp tip, the two lobes compressed from the sides near the tip, often touching each other at the front, forming a quasi-helmet; inflorescence 60–65 cm tall, with 3–5 flowers, pedicels 5–13 cm long. Open herbaceous vegetation, often near rivers in the valleys of the tepui summit, 1900– 2100 m; Bolívar (Macizo del Chimantá). Endemic. Heliamphora elongata Nerz, Carnivorous Pl. Newslett. 33: 111. 2004. Plant herbaceous, with branching rhizomes, forming dense clumps; pitcher narrowly ventricose in the lower part, narrowly cylindrical in the upper half, 30–32 cm long, 3–3.5 cm wide; lid helmet-shaped, bent nearly horizontal over the mouth of the pitcher; inflorescence 30–50 cm tall, with 2–5 flowers, pedicels 1.5–3.5 cm long; tepals 4, 3–5 cm long; anthers 11–19, ca. 3 mm long. Exposed areas with herbaceous vegetation on tepui summits, 2500–2700 m; Bolívar (Ilú-tepui, Tramen-tepui). Endemic. Nerz (cited above) mentioned finding putative hybrid individuals intermediate in form between H. elongata and nearby H. ionasii. Heliamphora folliculata Wistuba, Harbath & Carow, Carnivorous Pl. Newslett. 30: 120. 2001.

Plant herbaceous, with branching rhizomes, forming dense clumps; pitcher funnel-shaped in the lower 1/3, cylindrical in the upper part; lid bent sharply to the front, slightly helmet-shaped, not constricted at the base, back of the lid of adult pitchers bearing a hollow structure running downward for 5–10 mm; inflorescence ca. 35 cm tall, with 3 or 4 flowers, pedicels 3–6 cm long. Exposed areas with herbaceous vegetation on tepui summits, 2100–2700 m; Bolívar (Aparamán-tepui, Kamarkawaraitepui, Murisipán-tepui, Tereké-Yuréntepui). Endemic. Heliamphora heterodoxa Steyerm., Fieldiana, Bot. 28: 239. 1951. Plant 30–90 cm tall, often decumbent at base. Forming colonies in sandy or peaty savannas or Bonnetia groves on tepui slopes and summits, (600–)1000–2500 m; Venezuela, Brazil (Roraima); 2 varieties, both in the flora area. Key to the Varieties of H. heterodoxa 1. Pitcher appendage absent or scarcely developed ......... var. exappendiculata 1. Pitcher appendage obviously developed, 3–40 mm long ........... var. heterodoxa H.

heterodoxa var. exappendiculata Maguire & Steyerm., Mem. New York Bot. Gard. 29: 54. 1978. 1000–2500 m; Bolívar (Aprada-tepui, Auyán-tepui, Macizo del Chimantá). Endemic. ◆Fig. 118. A form of this variety, f. glabella Steyerm., was described for plants lacking

Heliamphora 141

trichomes on the upper, inner surface of the pitcher. H. heterodoxa var. heterodoxa Heliamphora heterodoxa var. glabra Maguire, Mem. New York Bot. Gard. 29: 54. 1978. (600–)1200–2500 m; Bolívar (summits of Auyán-tepui, Cerro Venamo, Ilú-tepui, Macizo del Chimantá, Murisipán-tepui, Ptaritepui, Tereké-Yurén-tepui, Uei-tepui, and Yuruaní-tepui). Brazil (Roraima: Serra do Sol). ◆Fig. 119. A form of this variety, f. glabra (Maguire) Steyerm., has been recognized for plants lacking trichomes on the upper, inner surface of the pitcher. Heliamphora hispida Wistuba & Nerz, Carnivorous Pl. Newslett. 29: 37. 2000. Plant with branching rhizomes, forming dense clumps with age; pitchers slightly ventricose, broadly funnel-shaped and expanded in the upper part; lid cordate, very narrow at the base; inflorescence ca. 50 cm tall, pedicels 5 cm long; tepals 4, 5–6 cm long. Open meadows on peaty areas; 1800– 3000 m; Amazonas (Sierra de la Neblina). Probably present on adjacent side of the border in Brazil. Heliamphora ionasii Maguire, Mem. New York Bot. Gard. 29: 54, fig. 53. 1978. Large rosette-forming herb; pitchers ventricose, widely expanded at the mouth; inflorescence to 1 m tall, pedicels 8–12 cm long; tepals 4, 3–6 cm long. Moist sandstone cliff faces and talus slopes, ca. 2600 m; Bolívar (Ilú-tepui). Endemic. ◆Fig. 117. Heliamphora minor Gleason, Brittonia 3: 164. 1939. Plant usually dwarfed, 10–85 cm tall; pitchers ventricose to tubular, 5–22(–30) cm long, often coral red in direct sun; lowest floral bracts 2–6 cm long; tepals 4 or 5. Savannas, bogs, and thickets on tepui summits, 1700–2500 m; Bolívar (Auyán-tepui, Macizo del Chimantá). Endemic. ◆Fig. 121. Heliamphora minor f. laevis Steyerm. was described for plants with the interior of the upper portion of the pitcher glabrous.

Heliamphora nutans Benth., Trans. Linn. Soc. London 18: 432. 1840. Rosette herb 20–50 cm tall; pitchers ventricose to elongated and tubular, 8–20(–30) cm long; peduncle 25–60 cm long, pedicels 2–6 cm long. Wet open savannas and in depressions on exposed sandstone of tepui slopes and summits, 1000–2800 m; Bolívar (Aprada-tepui, Ilú-tepui, Kukenán-tepui, Roraima-tepui). Adjacent Guyana (Mount Roraima). ◆Fig. 120. Heliamphora sarracenioides Wistuba, Harbath & Carow, Carnivorous Pl. Newslett. 34: 4. 2005. Rosette herb with branching rhizomes, forming dense clumps; pitcher funnelshaped in the lower 1/2, ventricose in the middle, and conical in upper 1/3, 20–30 cm tall; lid emerging directly from the back of the pitcher mouth without any constriction, often wider than the pitcher mouth; inflorescence ca. 60 cm tall, with 2 or 3 flowers, pedicels ca. 5 cm long; tepals 4–4.5 cm long; stamens 10 in 1 series, anthers 7–8 mm long × 1.5 mm wide. Exposed areas on tepui summit, elevation not published by the authors; Bolívar (on a tepui of the “northern part of the Gran Sabana” — information purposely withheld by the authors). Endemic. The description of this species sets a new precedent by amateur collectors who collect on tepuis without proper permits and withhold locality information even from government authorities of Venezuela. Similarly, the holotype cited in their publication at VEN was not deposited there as of April, 2005. Because of this, we cannot be certain whether the species described is identical to what Steyermark described as Heliamphora heterodoxa var. exappendiculata, which seems likely. Heliamphora tatei Gleason, Bull. Torrey Bot. Club 58: 368. 1931. Plant to 2(–4) m tall; stems unbranched, with elongated basal offshoots, or with shortened lateral axes bearing abbreviated rosettes; pitchers elongated, tubular, (12–) 15–45 cm long, the appendage conspicuous, 10–40 × 5–42 mm; inflorescence 2–4-flowered; sepals 4–6, 3–7.5 cm long. Venezuela, Brazil (Amazonas); 2 varieties, both in Venezuela.

142

S ARRACENIACEAE

Fig. 117. Heliamphora ionasii

Fig. 118. Heliamphora heterodoxa var. exappendiculata

Fig. 119. Heliamphora heterodoxa var. heterodoxa

Heliamphora 143

Fig. 120. Heliamphora nutans

Fig. 121. Heliamphora minor

Fig. 122. Heliamphora tatei var. tatei

144

S ARRACENIACEAE

Key to the Varieties of H. tatei 1. Upper interior portion of pitcher with trichomes that are not uniform, the indument toward the apical half consisting of trichomes 0.2–1 mm long, the lower portion with more elongate trichomes 1–1.5 mm long; Cerro Aracamuni, Sierra de la Neblina ..................... ....................................... var. neblinae 1. Upper interior portion of pitcher glabrous above the pubescent ring, or if with trichomes, these mainly uniform and 0.8– 2 mm long; Cerro Duida, Cerro Huachamacari, Cerro Marahauka .. var. tatei H. tatei var. neblinae (Maguire) Steyerm., Ann. Missouri Bot. Gard. 71: 311. 1984. —Heliamphora neblinae Maguire, Mem. New York Bot. Gard. 29: 57, fig. 54. 1978. Heliamphora neblinae var. viridis Maguire, Mem. New York Bot. Gard. 29: 57, fig. 54. 1978. Heliamphora tatei var. parva Maguire, Mem. New York Bot. Gard. 29: 59, fig. 49, G–H; fig. 50, A–C. 1978.

Wet, peaty savannas and sandstone escarpments on tepui slopes and summits, 1700–2500 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Adjacent Brazil (Amazonas: Serra da Neblina). Heliamphora tatei var. neblinae varies considerably in habit and in the number and color of the sepals. A form, f. parva (Maguire) Steyerm., has been recognized for plants with pitchers that are externally glabrous. H. tatei var. tatei Heliamphora macdonaldae Gleason, Bull. Torrey Bot. Club 58: 367. 1931. —Heliamphora tatei var. macdonaldae (Gleason) Maguire, Mem. New York Bot. Gard. 29: 57. 1978. Heliamphora tyleri Gleason, Bull. Torrey Bot. Club. 58: 368. 1931. Wet, peaty, treeless or shrubby savannas on tepui slopes and summits, 1200–2400 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 122. A form, f. macdonaldae (Gleason) Steyerm., has been recognized for plants with the upper inside of the pitcher glabrous.

SCROPHULARIACEAE by Noel H. Holmgren and Karl A. Vincent Annual, biennial, or perennial herbs, vines, or shrubs, some hemiparasitic. Leaves entire to pinnately or palmately lobed or parted, opposite, alternate, whorled, or sometimes all basal, without stipules. Inflorescence a spike, raceme, thyrsoid panicle, or the flowers solitary or fascicled in the axils of leaves; bracts foliaceous to reduced; bractlets sometimes present. Flowers bisexual, (4)5-merous, bilabiate to sometimes nearly actinomorphic. Calyx persistent, of 1, 2, 4, or 5 distinct or variously united parts. Corolla sympetalous, 4- or 5-lobed, usually bilabiate, tubular, campanulate, salverform, or rotate. Stamens 2 or 4, rarely 5, didynamous, or with reduced numbers of fertile stamens and vestiges of the others present as staminodes, epipetalous, alternate with the corolla lobes; anthers with (1)2 equal or unequal thecae. Ovary superior, (1)2-locular; ovules anatropous on axile placentas; style slender and entire; stigma solitary and capitate or forked with 2 stigmatic lobes. Fruit a capsule in the flora area, septicidal, loculicidal, or both, some irregularly poricidal or valvate apically, rarely a berry. Seeds usually many and small, rarely few; endosperm usually well developed; embryo small.

S CROPHULARIACEAE 145

Nearly cosmopolitan, mostly in temperate, subtropical, and tropical mountains, less frequently in lowland tropical regions; ca. 200 genera and ca. 3000 species, 18 genera and 42 species in the flora area. Key to the Genera of Scrophulariaceae 1.

Calyx segments united from the base into a well-developed tube or at least connate at anthesis ...................................................................... 2 1. Calyx segments parted to or nearly to the base ...................................... 9 2(1). Pedicels without bractlets ......................................................................... 3 2. Pedicels with bractlets .............................................................................. 5 3(2). Leaves alternate; calyx with 2 lateral lobes, these truncate or rounded .................................................................................................. 9. Castilleja 3. Leaves opposite; calyx 5-lobed, the lobes acute to acuminate ................ 4 4(3). Flowers solitary in the axils of the leaves; calyx 3–7 mm long, not completely enclosing the capsule ................................................. 12. Lindernia 4. Flowers (1)2 or 3 in a cluster in the axils of small subulate bracts; calyx 6–12 mm long, enclosing the capsule ....................................... 17. Torenia 5(2). Corolla campanulate to funnel-shaped; capsule globose, sometimes depressed-globose or subglobose .............................................................. 6 5. Corolla salverform; capsule oblong-ovoid ................................................. 8 6(5). Leaves linear to filiform; pedicels 20–55 mm long; corolla pale pink to lavender or violet; seeds oblong to subspherical ............. 4. Anisantherina 6. Leaves lanceolate to ovate; pedicels 1–65 mm long; corolla yellow to orange; seeds elongate-cylindrical ........................................................... 7 7(6). Pedicels 1–5.5 mm long; calyx broadly campanulate with 5 lobes; stems erect or ascending, self-supporting .............................................. 2. Alectra 7. Pedicels 15–65 mm long; calyx an inflated, ovoid, spathe-like structure with a single, terminal apiculum; stems weak, scandent or climbing ................................................................................................ 18. Vellosiella 8(5). Corolla tube and throat collectively 0.6–1.3 cm long; calyx 4–10 mm long; anthers monothecal ........................................................ 7. Buchnera 8. Corolla tube and throat collectively 6.5–11 cm long; calyx 37–54 mm long; anthers dithecal ............................................................ 11. Escobedia 9(1). Calyx segments distinctly dissimilar ..................................................... 10 9. Calyx segments equal or subequal ......................................................... 12 10(9). Bractlets present at the base of the pedicel; corolla limb yellow ............................................................................................. 13. Mecardonia 10. Bractlets present at the summit of the pedicel (often appearing to be part of the calyx) or absent; corolla limb white, pink, violet, purple, or blue ....................................................................................................... 11 11(10). Calyx of 1 broad, posterior segment and 4 narrower, lateral ones; fertile stamens 2; corolla throat closed by an upraised palate ......... 1. Achetaria 11. Calyx of 3 broad segments (1 posterior and 2 anterio-lateral) and 2 narrower, posterio-lateral ones; fertile stamens (2–)4(5); corolla throat open .............................................................................................. 5. Bacopa 12(9). Calyx segments 4 ..................................................................................... 13 12. Calyx segments 5 ..................................................................................... 15

146

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13(12). Stems firm, erect or ascending; leaves of the lower stem opposite, the upper ones 3-whorled .............................................................. 15. Scoparia 13. Stems flaccid, prostrate, the plants forming mats; leaves all opposite ..... 14 14(13). Pedicels (2–)4.5–10 mm long; stems hirsutulous; stamens 3 or 4; anther thecae 0.3–0.4 mm long; capsules 1.2–3.5 mm long ................... 5. Bacopa 14. Pedicels 0.5–1 mm long; stems glabrous; stamens 2; anther thecae 0.1– 0.2 mm long; capsules 0.8–1.1 mm long ...................... 14. Micranthemum 15(12). Pedicels with bractlets ................................................................. 10. Conobea 15. Pedicels without bractlets ....................................................................... 16 16(15). Leaves alternate ........................................................................... 8. Capraria 16. Leaves opposite or whorled ..................................................................... 17 17(16). Leaves deeply pinnately dissected into filiform divisions....... 6. Benjaminia 17. Leaves entire, crenate, or dentate .......................................................... 18 18(17). Leaves petiolate .......................................................................... 16. Stemodia 18. Leaves sessile ........................................................................................... 19 19(18). Leaves mostly < 3 cm long; stems < 30 cm long; corolla palate upraised with 2 ridges, the tube not bisulcate; capsule ovoid, ellipsoid, or fusiform, distinctly longer than wide .......................................... 12. Lindernia 19. Leaves mostly > 3 cm long; stems > 50 cm long; corolla palate cratershaped with a bifid horn at forward edge, the tube bisulcate in front; capsule globose to depressed-globose ...................................... 3. Angelonia 1. ACHETARIA Cham. & Schltdl., Linnaea 2: 566. 1827. Pubescent herbs, often suffrutescent. Stems quadrangular. Leaves petiolate, opposite, entire, crenate, or toothed, pinnately veined. Inflorescence of solitary, axillary flowers or a spike; pedicels of varying length or absent. Bractlets usually 2 and attached below the calyx, or absent. Calyx of 5 dissimilar segments, parted to the base, the posterior one inserted outside and distinctly larger, the other 4 linear and relatively thin-margined, with the posterio-lateral pair slightly narrower than the anterio-lateral pair; corolla pale blue to violet, distinctly bilabiate, the lower lip 3-lobed, with an upraised palate that nearly closes off the orifice, the upper lip entire to 2-lobed and external in bud. Fertile stamens 2 (the anterior pair); anthers with a knob-like connective separating the thecae; staminodes 2 (the posterior pair), often bearing minute sterile anthers. Style with 2 stigmas. Capsule thinwalled, septicidal and apically loculicidal, broadly ovoid to globose. Seeds numerous, longitudinally ridged with transverse walls forming reticulae. Tropical South America; 5 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Achetaria 1. 1.

Calyx (3–)3.5–5.5 mm long; corolla 5.5–9 mm long with its upper lip erect and lower lip spreading; occurring mostly below 400 m ...... A. guianensis Calyx 1.8–3 mm long; corolla 3–4 mm long with both lips spreading; occurring above 900 m ........................................................ A. scutellarioides

Achetaria guianensis Pennell, Notul. Nat. Acad. Nat. Sci. Philadelphia 46: 5. 1940. Erect or ascending herb, sometimes suffrutescent. Borders of savannas and road-

sides, 100–400 m; northern Bolívar. Anzoátequi, Monagas; Trinidad-Tobago, Guyana, Suriname, French Guiana. ◆Fig. 123. Achetaria guianensis may be conspecific

Alectra 147

with A. erecta (Spreng.) Wettst., which would extend the geographical range to eastern Brazil. Achetaria scutellarioides (Benth.) Wettst. in Engl. & Prantl, Nat. Pflanzenfam. IV. 3b: 74. 1891. —Beyrichia scutellarioides Benth., Scroph. Ind. 9. 1835. Spreading, spindly, much-branched shrub. Wet places in grassy savannas, floodplains, along streambanks, 900–2100 m; Bolívar (Gran Sabana). Brazil (Bahia).

Fig. 123. Achetaria guianensis

2. ALECTRA Thunb., Nov. Gen. Pl. 81. 1784. Erect, hemiparasitic herbs with stiffly scabrid pubescence. Stems usually strict. Leaves opposite or subopposite, entire, crenate, or toothed, parallel-veined. Inflorescence of solitary flowers in the upper leaf axils, sometimes appearing racemose or subspicate; bractlets 2, near the middle or toward the summit of the pedicel. Calyx campanulate, with 5 short lobes, accrescent; corolla yellowish, campanulate, the limb slightly exserted from the calyx, the lobes erect with the lower 3 external in bud, the tube gibbous at the base anteriorly. Stamens 4; filaments glabrous or bearded; anthers U-shaped, the thecae equal or unequal in length. Fruit a globose or depressed-globose capsule. Seeds elongate, narrowly cylindrical-conical, longitudinally elongate-reticulate. Neotropics, Old World; 40 species, 2 in Venezuela, both in the flora area. Key to the Species of Alectra 1. 1.

Leaves 2.4–6.5 cm long, 7–22 mm wide, spreading; pedicels 1–2.5(–3.5) mm long ......................................................................................... A. aspera Leaves 1–2(–2.5) cm long, 1.5–3.7(–5) mm wide, appressed-ascending; pedicels 3–5.5 mm long ................................................................ A. stricta

Alectra aspera (Cham. & Schltdl.) L.O. Williams, Fieldiana, Bot. 34: 118. 1972. —Glossostylis aspera Cham. & Schltdl., Linnaea 3: 23. 1828. Pedicularis melampyroides Rich., Actes Soc. Hist. Nat. Paris 1: 111. 1792. —Melasma melampyroides (Rich.) Pennell in Britton & P. Wilson, Bot. Porto Rico 6: 188. 1925. Scrophularia fluminensis Vell., Fl. Flumin. 263, t. 87. 1825 [1829]. —Alectra fluminensis (Vell.) Stearn, J. Arnold Arbor. 52: 635. 1971.

Perennial herb, ± succulent and brittle. Moist places in pastures and thickets, ca. 100–200 m; western Bolívar (Cerro Gavilán). Central America, West Indies, south to Brazil, Bolivia, and Paraguay. Alectra stricta Benth. in DC., Prodr. 10: 338. 1846. Annual herb with strict, erect stems. Moist places in savannas, 50–400 m; western Bolívar, northeastern Amazonas. Central Venezuela; Brazil (Planalto region) south to Bolivia. ◆Fig. 124.

148

S CROPHULARIACEAE

Fig. 124. Alectra stricta

Fig. 125. Angelonia salicariifolia

Anisantherina 149

3. ANGELONIA Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 92, t. 108. 1812. Annual or perennial herbs and shrubs. Stems terete to quadrangular, sometimes winged. Leaves opposite, rarely subopposite to alternate, sessile or petiolate, serrate or rarely entire. Inflorescence a raceme or of solitary axillary flowers; bracts leaf-like to reduced; bractlets 1, 2, or absent. Calyx subequally 5-parted to near the base, the segments lanceolate to ovate; corolla blue to reddish purple or white, strongly zygomorphic, 5-lobed, the tube short, bisulcate at the base anteriorly, these projections densely glandular-pubescent within, the palate sometimes crater-shaped or sulcate and often with a bifid horn, or else with a laterally compressed ridge, the posterior pair of lobes outermost in bud. Stamens 4, didynamous; upper filaments curved, the lower ones sigmoid; thecae ellipsoid or discoid, borne at the top of the corolla tube. Style gradually tapered to an entire stigma. Fruit a capsule, often globose, dehiscing septicidally and often loculicidally as well. Seeds numerous, conical to pyramidal, deeply reticulate. Neotropics; 26 species, 1 in Venezuela. Angelonia salicariifolia Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 92, t. 108. 1812. —Angelón. Erect, suffrutescent herb. Savannas, open rocky slopes, ca. 50 m; Bolívar (near

Ciudad Bolívar). Elsewhere in northern and western Venezuela; adjacent Colombia, Trinidad-Tobago. ◆Fig. 125. Angelonia salicariifolia is grown as a garden ornamental and sometimes escapes.

4. ANISANTHERINA Pennell, Mem. Torrey Bot. Club 16: 106. 1920. Annual, hemiparasitic herbs. Stems erect, slender, usually branched, hirsute with reflexed-spreading hairs, nonglandular. Leaves opposite, sessile, linear to filiform, occasionally sparingly and weakly toothed, scabridulous on the upper surface and on the revolute margins and midvein beneath. Inflorescence of solitary flowers in the upper leaf axils; pedicels long; bractlets 2, near the middle of the pedicel. Calyx broadly hemispheric or campanulate with 5 lobes, these shorter than the tube; corolla pale pink, pink, lavender, or violet, glandular-pubescent externally with stalked glandular hairs, glabrous within except at the base of the posterior lip, the tube narrow, the throat ventricosely inflated, 5-lobed, the lobes much shorter than the tube and throat, the lower 3 lobes external in bud. Stamens 4, didynamous; filaments distally lanose, especially the anterior pair; thecae unequal, one somewhat reduced. Style slender; stigmas united. Fruit a globose capsule. Seeds oblong to subspherical, reticulate. Southern Mexico, Central America, Cuba, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil; 1 species. Anisantherina hispidula (Mart.) Pennell, Mem. Torrey Bot. Club 16: 106. 1920. —Gerardia hispidula Mart., Nov. Gen. Sp. Pl. 3: 13, t. 207. 1829. —Agalinis hispidula (Mart.) D’Arcy, Ann. Missouri Bot. Gard. 65: 4. 1978.

Herb with few to several divaricate branches. Moist savannas, margins of morichales, often in sandy soils, ca. 50–400 m; northern and western Bolívar, Amazonas (near Puerto Ayacucho). Brazil (Amazonas, Planalto region). ◆Fig. 126.

150

S CROPHULARIACEAE

Fig. 127. Bacopa repens

Fig. 126. Anisantherina hispidula

Fig. 128. Bacopa callitrichoides

Bacopa 151

5. BACOPA Aubl., Hist. Pl. Guiane 128. 1775, nom. cons. Herpestis C.F. Gaertn., Suppl. Carp. 186. 1805 [1807]. Hydranthelium Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 202. 1825. Macuillamia Raf., Neogenyton 2. 1825. Caconapea Cham., Linnaea 8: 28. 1833. Ranaria Cham., Linnaea 8: 30. 1833. Monocardia Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 155. 1920. Prostrate, diffuse, or erect, succulent, aquatic or swamp-inhabiting herbs, mostly annuals. Leaves petiolate or usually sessile, opposite or whorled, entire to toothed or lobed, rarely palmately divided, often glandular-punctate (with glandular epidermal cells that often become sunken upon drying), pinnately or palmately veined. Inflorescence of solitary or fascicled axillary flowers; pedicels of varying length or absent; bractlets (1)2 or absent, at or near apex of the pedicel when present, sometimes appearing to be part of the calyx, situated opposite the basal margins of the posterior calyx segment. Calyx of 5 strongly to slightly dissimilar segments or sometimes 4 weakly differentiated ones parted to the base in 2 whorls; when segments 5, the outer whorl includes the larger posterior one and the slightly smaller or narrower anterio-lateral pair, the inner whorl with the narrowest, posterio-lateral pair; when segments 4, the posterior one slightly larger than the other 3; corolla limb white or violet to blue, the tube or throat sometimes with yellow markings, nearly regular to distinctly bilabiate, the lower lip 3-lobed, the upper lip entire or 2-lobed, external in bud. Fertile stamens mostly 4, sometimes 5, 3, or 2; anthers with 2 thecae and without a well-developed connective. Stigmas 2 or united and capitate. Capsule thin-walled, sometimes membranous, septicidal and loculicidal or mostly loculicidal, ovoid to globose. Seeds numerous, longitudinally ridged with weaker transverse walls forming reticulae. New World tropics and subtropics; ca. 60 species, 15 in Venezuela, 12 of these in the flora area. Key to the Species of Bacopa 1.

1.

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

3.

Leaves filiform to linear, relatively thick and conduplicate-folded; calyx segments barely differentiated; pedicels 1–2.5(–3.5) mm long; flowers often in pairs in each axis, sometimes only one; bractlets present .............................................................................................. B. gratioloides Leaves mostly lanceolate or broader, if narrower neither thick nor conduplicate-folded; calyx segments distinctly differentiated except in B. callitrichoides and some B. repens and B. reptans, flowers in fascicles of 4–8 or solitary; bractlets present or absent ..................................... 2 Pedicels < 1.5 mm long or absent ............................................................. 3 Pedicels > 1.5 mm long .............................................................................. 4 Flowers in fascicles of 4–8 in leaf axils; calyx 1.5–2.1 mm long, the outer 3 segments thick and fleshy, deeply glandular-pitted; corolla 1.6– 2 mm long; stems hispid .................................................. B. monnierioides Flowers solitary in leaf axils; calyx 3.5–6 mm long, the segments neither thick and fleshy nor deeply glandular-pitted; corolla 3–4 mm long; stems glabrous ....................................................................... B. sessiliflora

152

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4(2).

Bractlets present at the summit of the pedicel or attached to the base of the calyx ................................................................................................. 5 4. Bractlets absent (or rudimentary in some B. salzmannii and B. serpyllifolia) ........................................................................................... 8 5(4). Stems prostrate or trailing; calyx 8.5–13.5 mm long; stamens 5 .. B. aquatica 5. Stems erect or ascending; calyx 3–7(–9) mm long; stamens 4 ................ 6 6(5). Leaves petiolate, the blade mostly > 10 mm wide; calyx 5–9 mm long ....................................................................................................... B. stricta 6. Leaves sessile, the blade < 7 mm wide; calyx 3–4.7 mm long ................ 7 7(6). Stems 20–45 cm long; posterior calyx segment ovate; leaves opposite .................................................................................................... B. laxiflora 7. Stems 3–7 cm long; posterior calyx segment (the largest one) lanceolate to broadly lanceolate; leaves opposite to 4-whorled .................. B. reptans 8(4). Calyx 3.5–7.5 mm long, the posterior segment truncate-rounded to cordate at the base, becoming reticulate-venose; capsule firm-walled ... 9 8. Calyx 1.2–4.2 mm long, the posterior segment broadly cuneate at the base, not strongly reticulate; capsule membranous-walled .............. 11 9(8). Stems appressed to ascending-pubescent; outer calyx segments glabrous; pedicels 1.5–6.5 mm long; stamens 2(–4) ................ B. innominata 9. Stems spreading-pubescent; outer calyx segments pubescent; pedicels (4.5–)8–22 mm long; stamens 4 .......................................................... 10 10(9). Pubescence of the stems, pedicels, and calyx-segment margins of fine, yellowish hairs; corolla 3–6(–7) mm long, mostly concealed by the calyx; stems 5–20 cm long ........................................................ B. salzmannii 10. Pubescence of the stems, pedicels, and calyx-segment margins of coarse, whitish, multicellular hairs; corolla 6–7 mm long, the limb exserted from the calyx; stems usually shorter, 3–6 cm long ........... B. serpyllifolia 11(8). Leaves > 12 × 7 mm; calyx 2.7–4.2 mm long, usually consisting of 5 segments, 3 broad ones and 2 narrow ones, sometimes reduced to 4 with one slightly broader than the others; capsule 2.5–3.5 mm long ....................................................................................................... B. repens 11. Leaves < 7 × 5.5 mm; calyx 1.2–2.3 mm long, reduced to 4 segments, one slightly broader than the others; capsule 1.2–1.8 mm long .......................................................................................... B. callitrichoides Bacopa aquatica Aubl., Hist. Pl. Guiane 129, pl. 49. 1775. Prostrate, mat-forming, aquatic herb. Shallow ponds, slow moving streams, marshes, 50–200(–400) m; central Delta Amacuro, northern Bolívar. Central America, Trinidad-Tobago, Guyana, French Guiana, northern Brazil. Bacopa callitrichoides (Kunth) Pennell, Proc. Acad. Nat. Sci. Philadelphia 98: 96. 1946. —Hydranthelium callitrichoides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 203, t. 646. 1825. Small, floating, mat-forming herb. Pools and ponds, 50–200 m; northern Bolívar,

northwestern Amazonas. Adjacent Colombia. ◆Fig. 128. Bacopa gratioloides (Cham.) Chodat & Hassl., Bull. Herb. Boissier, sér. 2, 4: 288. 1904. —Caconapea gratioloides Cham., Linnaea 8: 29. 1833. Stout, erect, perennial, rhizomatous herb. Open wet places, sometimes in shallow water around shores of ponds and lakes, 100–400 m; Bolívar (Río Caroní basin). Guárico; Guyana, Brazil (Planalto region). Bacopa innominata (M. Gómez) Alain, Revista Soc. Cub. Bot. 13: 61. 1956 [1957]. —Conobea innominata M.

Bacopa 153

Gómez, Anales Soc. Esp. Hist. Nat. 23: 178. 1894. Aquatic herb, emergent or prostratefloating. Wet places in mud around drying pools, lake shores, pastures, near sea level to 300 m; Delta Amacuro (near Tucupita), northern Bolívar. Portuguesa, Táchira, Zulia; eastern coastal U.S.A., Antilles, northern Colombia, Brazil, Paraguay.

Prostrate, succulent herb, often floating. Shallow pools or slow-moving streams, stranded on muddy shores, on rocks in pools on granitic outcrops, 50–300 m; Bolívar (near Represa Guri), Amazonas (near Puerto Ayacucho). Barinas, Portuguesa, Zulia; southeastern and western U.S.A., southern Mexico, Central America, West Indies, Colombia, Ecuador, Bolivia. ◆Fig. 127.

Bacopa laxiflora (Benth.) Wettst. ex Edwall, Bol. Commiss. Geogr. Estado São Paulo 13: 180. 1897. —Herpestis laxiflora Benth. in DC., Prodr. 10: 396. 1846. —Mella laxiflora (Benth.) Pennell, Notul. Nat. Acad. Nat. Sci. Philadelphia 46: 1. 1940. Herpestis auriculata B.L. Rob., Proc. Amer. Acad. Arts 26: 172. 1891. —Caconapea auriculata (B.L. Rob.) Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 150. 1920. Erect, divaricately branched herb. Wet open marshes, shores of ponds, 100–400 m; north-central Bolívar, Amazonas (Río Manapiare basin). Carabobo, Portuguesa; Mexico, Central America, Colombia, Guyana, Brazil, Bolivia.

Bacopa reptans (Benth.) Wettst. ex Edwall, Bol. Commiss. Geogr. Estado São Paulo 13: 181. 1897. —Herpestis reptans Benth. in DC., Prodr. 10: 395. 1846. Caconapea debilis Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 151. 1920. Small, mat-forming herb with prostrate stems. Marshy edges of drying ponds, shallow pools, streams, ca. 100 m; Amazonas, (near Puerto Ayacucho and Sanariapo). Guyana south to Brazil (Mato Grosso).

Bacopa monnierioides (Cham.) B.L. Rob., Proc. Amer. Acad. Arts 44: 614. 1909. —Ranaria monnierioides Cham., Linnaea 8: 31. 1833. Herpestis ranaria Benth., Companion Bot. Mag. 2: 57. 1836. Bacopa punctata Pennell, Fieldiana, Bot. 28: 516. 1953, non Engl. 1897. Succulent, erect to trailing herb. Wet places in savannas, on muddy or drying shores of ponds, 50–1000 m; Bolívar (scattered). Central America, Brazil, Bolivia. Bacopa monnierioides is closely related to B. axillaris (Benth.) Standl. of western Venezuela, Colombia, and Ecuador, which has larger calyx segments that lack the fleshiness and the notably sunken glandular cells. Bacopa repens (Sw.) Wettst. in Engl. & Prantl, Nat. Pflanzenfam. IV. 3b: 76. 1891. —Gratiola repens Sw., Prodr. 14. 1788. —Macuillamia repens (Sw.) Pennell, Proc. Acad. Nat. Sci. Philadelphia 75: 9. 1923. Macuillamia limosa Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 158. 1920.

Bacopa salzmannii (Benth.) Wettst. ex Edwall, Bol. Commiss. Geogr. Estado São Paulo 13: 181. 1897. —Herpestis salzmanni Benth., Companion Bot. Mag. 2: 58. 1836. Monocardia humilis Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 157. 1920. Monocardia lilacina Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 156. 1920. Monocardia violacea Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 156. 1920. Small, mat-forming, creeping or sometimes floating herb. Muddy ground around drying pools, marshy places, shallow water in savannas, 50–200 m; along Río Orinoco in Bolívar and northern Amazonas. Coastal Cordillera, Llanos; southern Mexico, Central America, Colombia, Guyana, Peru, Brazil, Bolivia, Paraguay. Bacopa serpyllifolia (Benth.) Pennell, Proc. Acad. Nat. Sci. Philadelphia 98: 98. 1946. —Herpestis serpyllifolia Benth. in DC., Prodr. 10: 398. 1846. Herpestis ciliata Pennell, Notul. Nat. Acad. Nat. Sci. Philadelphia 46: 2. 1940. —Bacopa ciliata (Pennell) Pennell, Proc. Acad. Nat. Sci. Philadelphia 98: 98. 1946. Small, succulent, prostrate herb. Muddy banks of ponds, wet savannas, 50–200 m; northern Bolívar, northern Amazonas. Guyana, Brazil, Paraguay.

154

S CROPHULARIACEAE

Bacopa sessiliflora (Benth.) Pulle, Enum. Vasc. Pl. Surinam 415. 1906. —Herpestis sessiliflora Benth., Companion Bot. Mag. 2: 58. 1836. —Caconapea sessiliflora (Benth.) Pennell, Proc. Acad. Nat. Sci. Philadelphia 75: 11. 1923. Caconapea conferta Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 153. 1920. —Bacopa conferta (Pennell) Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 174. 1936. Erect, glabrous or occasionally glandularpubescent herb. Wet places, often in standing water, 50–400 m; Delta Amacuro, northern Bolívar. Mexico, Central America, Cuba, Hispaniola, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, eastern Brazil.

Bacopa stricta (Schrad.) Wettst. ex Edwall, Bol. Commiss. Geogr. Estado São Paulo 13: 180. 1897. —Herpestis stricta Schrad. in Link, Enum. Hort. Berol. Alt. 2: 142. 1821 [Jan–Jun 1822]. —Caconapea stricta (Schrad.) Britton in Britton & P. Wilson, Bot. Porto Rico 6: 183. 1925. Herpestis domingensis Spreng., Neue Entd. 3: 25. 1822. —Bacopa domingensis (Spreng.) Pennell, Proc. Acad. Nat. Sci. Philadelphia 98: 92. 1946. Erect, succulent herb. Wet places along streambanks, shores of ponds, sometimes emersed in shallow water, ca. 1200 m; Bolívar (Gran Sabana near Luepa). Falcón, Táchira; Hispaniola, Puerto Rico, Colombia, Ecuador, Peru, Brazil.

6. BENJAMINIA Mart. ex Benj. in Mart., Fl. Bras. 10: 255. 1847. Submerged or emergent aquatic herbs, rooting at the nodes. Stems pubescent or glabrate, glandular-punctate. Leaves whorled, finely pinnately dissected, the segments filiform. Inflorescence a solitary, axillary flower; pedicels elongating in fruit; bractlets absent. Calyx of 5 subequal segments, parted to near the base. Corolla pale lilac to blue, bilabiate, the lobes rounded, the upper lip outermost in bud. Stamens 4, didynamous; anthers with 2 thecae, without a well-developed connective. Ovary with a surrounding whorl of bristles at the base; stigma flat and curved. Capsule ovoid, membranous. Seeds numerous, oblong, longitudinally reticulate. Southern Mexico, Central America, Cuba, Venezuela, Trinidad-Tobago, Guyana, Brazil, Bolivia; 1 species. Benjaminia reflexa (Benth.) D’Arcy, Ann. Missouri Bot. Gard. 66: 194. 1979. —Herpestis reflexa Benth. in DC., Prodr. 10: 399. 1846. —Bacopa reflexa (Benth.) Edwall, Bol. Commiss. Geogr. Estado São Paulo 13: 176. 1897, not validly combined. —Naiadothrix reflexa (Benth.) Pennell, Mem. Torrey Bot. Club 16: 106. 1920. Benjaminia utriculariiformis Mart. ex Benj. in Mart., Fl. Bras. 10: 256. 1847, as “urtriculariaeformis.” Herb, often submerged except for the flowers. Pools, shallow margins of lakes, in slow-moving streams, 50–100 m; northern Bolívar, Amazonas (near Puerto Ayacucho and La Esmeralda). Cojedes, Guárico. Distribution as in genus. ◆Fig. 129.

Fig. 129. Benjaminia reflexa

7. BUCHNERA L., Sp. Pl. 630. 1753. Annual or perennial, hemiparasitic herbs, becoming black upon drying. Stems strict or branched, glabrous to pubescent. Leaves opposite, subopposite, or

Buchnera 155

rarely alternate, all cauline, simple, usually sessile, filiform to lanceolate or ovate, entire to dentate, often coriaceous, 1–5-veined from the base. Inflorescence a terminal spike or spicate-raceme; pedicels short and bearing a pair of narrow bractlets. Calyx tubular, 10- or more-veined, glabrous to hispid-scabrid, regularly or irregularly 5-lobed or -toothed, the lobes shorter than the tube, triangular to lanceolate, erect or spreading; corolla white, pink, violet, purple, or blue, salverform, the tube as long as or longer than the calyx and glabrous to softly pubescent externally, pilose or villous within, the lower 3 lobes external in bud. Stamens 4, slightly didynamous, included; anthers with only 1 theca developed. Style entire; stigma clavate. Capsule ovoid to oblong, frequently laterally compressed, loculicidal, included in the calyx. Seeds numerous. Mostly tropical or subtropical in both hemispheres, with 1 species in eastern North America as far north as southern Ontario; ca. 100 species, 8 in Venezuela, 7 of these in the flora area. The species of Buchnera are differentiated by minor characters, which makes identification difficult and sometimes dubious. As many as three species may grow together in a savanna, and mixed collections are not uncommon. Key to the Species of Buchnera 1. 1. 2(1). 2. 3(2).

3.

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

7(5).

7.

Calyx glabrous except for occasional ciliolate lobes ................................ 2 Calyx pubescent ......................................................................................... 4 Calyx 4–5.5 mm long; middle leaves of the stem linear, 1.7–3.4 mm wide ................................................................................................. B. spruceana Calyx 5.8–10 mm long; middle leaves of the stem mostly filiform, 0.5– 1.5(–2) mm wide .................................................................................... 3 Leaves 1-veined, scabrid-verrucose; pedicels 0.6–2.8 mm long; calyx 10veined, often reticulate-venose between the veins in the distal half ................................................................................................... B. palustris Leaves often 3-veined, glabrous or the lower leaves scabrid-hispid; pedicels 0.2–0.7 mm long; calyx 10- or more-veined, not reticulatevenose between the parallel veins ...................................... B. weberbaueri Pubescence of uncinate (hooked at tip) hairs ............................. B. ternifolia Pubescence of straight or curved hairs, but not uncinate ....................... 5 Calyx pubescent between the veins as well as on them, the 10 veins often accompanied by smaller veins between ............................................... 6 Calyx pubescence restricted to the veins, the 10 veins not accompanied by smaller veins between ...................................................................... 7 Calyx (5.5–)6–9 mm long, the tube pubescent all over, the lobes (1–)1.5– 2 mm long; corolla tube pilose on exterior ..................................... B. rosea Calyx 4–5.5 mm long, the tube glabrous below and lightly pubescent above, the lobes 0.5–1.2 mm long; corolla tube glabrous or sometimes lightly pubescent above on exterior ....................................... B. spruceana Calyx veins coarsely pubescent with stiff, spreading, callus-based hairs; plants annual, 15–65 cm tall; corolla white or sometimes with a lavender tinge ....................................................................................... B. pusilla Calyx veins finely pubescent with soft, appressed-ascending, non-callusbased hairs; plants perennial, 55–90 cm tall; corolla pink to violet .................................................................................................. B. longifolia

156

S CROPHULARIACEAE

Buchnera longifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 340. 1817 [1818]. Buchnera elongata Sw., Prodr. 926. 1788. Buchnera macrocarpa Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 342. 1817 [1818]. Erect perennial herb. Open slopes and savannas, 100–300 m; northern Bolívar, Amazonas (near Puerto Ayacucho). Central America, West Indies, Colombia, TrinidadTobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina. Buchnera palustris (Aubl.) Spreng., Syst. Veg. ed. 16, 2: 805. 1825. —Piripea palustris Aubl., Hist. Pl. Guiane 628, pl. 253. 1775. Slender annual herb. Savannas, open places along streams and lake shores, 50– 1400 m; widespread in Bolívar and Amazonas. Carabobo, Guárico, Monagas; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Paraguay. ◆Fig. 130. Buchnera pusilla Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 340. 1817 [1818]. Slender annual herb. Dry open slopes, grassy savannas, open woods, ca. 50–200 m; Bolívar (near Ciudad Piar and El Manteco), Amazonas (near Puerto Ayacucho). Coastal Cordillera; southern Mexico, Central America, Colombia, Guyana, Ecuador, Peru, northern Brazil. Buchnera rosea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 342. 1817 [1818]. Erect perennial herb. Dry, grassy savannas, open rocky hillsides, open forests, 50– 1300 m; widespread in Bolívar and Amazonas. Widespread elsewhere in Venezuela; Panama, Colombia, Trinidad-Tobago, Guyana, Peru, Brazil, Bolivia. Buchnera rosea and B. palustris are the most commonly collected members of Scrophulariaceae in the Venezuelan Guayana. They sometimes grow together in open areas, but B. rosea appears to prefer the better-drained sites. Buchnera spruceana Philcox, Kew Bull. 18: 292. 1965 [1966].

Fig. 130. Buchnera palustris

Erect perennial herb. Dry, sandy savannas, ca. 100 m; Amazonas (near La Esmeralda). Endemic. Buchnera spruceana is very closely related to B. rosea, but the latter appears to be absent from the area around La Esmeralda. Some plants of B. rosea from similar sandy savannas in eastern Bolívar approach B. spruceana morphologically. Buchnera ternifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 341. 1817 [1818]. Buchnera lithospermifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 341. 1817 [1818]. Buchnera lobelioides Cham. & Schltdl., Linnaea 2: 589. 1827. Erect perennial herb with distinctive hooked hairs. Grassy savannas, open areas, ca. 50 m; Bolívar (10 km south of Ciudad Bolívar). Colombia, Brazil, Bolivia, Paraguay, Argentina, Uruguay.

Castilleja 157

Buchnera weberbaueri Diels, Bot. Jahrb. Syst. 37: 430. 1906. Slender, erect, annual herb. Savannas, streambanks, open slopes, 50–1300 m; Delta

Amacuro (Río Amacuro), widespread in Bolívar and northern Amazonas. Central America, Colombia, Trinidad-Tobago, Peru.

8. CAPRARIA L., Sp. Pl. 628. 1753. Erect branched herbs, sometimes suffrutescent. Leaves and stems sparsely hirsute to densely hispid or villous. Stems terete. Leaves alternate, sessile, usually serrate or dentate, prominently pinnately veined. Inflorescence of axillary fascicles of (1)2–4 pedicellate flowers; bractlets absent. Calyx 5-parted to near the base, with linear to narrowly lanceolate segments. Corolla yellowish green or white, campanulate, nearly actinomorphic, 5-lobed, the lobes subequal, the upper 2 external in bud. Stamens (4)5, all fertile; anthers with 2 parallel or divergent thecae. Stigmas united, clavate. Fruit a capsule, septicidal and secondarily loculicidal at the apex, glandular-punctate, the placental mass remaining free standing. Seeds numerous, conical-cylindrical, reticulate in longitudinal rows. New World tropics and subtropics; 5 species, 1 in Venezuela. Capraria biflora L., Sp. Pl. 628. 1753. Capraria biflora f. hirsuta Loes., Bull. Herb. Boissier sér. 2, 3: 284. 1903. Coarse, erect herb, sometimes suffrutescent. Moist disturbed places along roadsides, riverbanks, edges of marshes, near sea level to 300 m; scattered throughout Delta Amacuro (Caño Nabasanuka near Araguao),

Bolívar, and Amazonas. Widespread in northern Venezuela; U.S.A. (southern Florida), Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil. ◆Fig. 131. Capraria biflora is sometimes cultivated for use of the flowers as a medicinal tea for stomach ache and diarrhea.

9. CASTILLEJA Mutis ex L. f., Suppl. Pl. 293. 1781. Perennial or annual hemiparasitic herbs, sometimes shrubs. Stems usually erect, often branched. Leaves alternate, sessile, cauline, entire, or pinnately parted or lobed. Inflorescence a spike or raceme; bracts often leaf-like below and gradually differentiating upward, usually more conspicuously colored than the flowers. Calyx tubular, 2- or 4-lobed, the lobes usually more united laterally into 2 primary lobes, these sometimes entire by complete fusion of the lateral segments. Corolla greenish or the exserted portions colored, tubular-galeate, elongate and narrow, bilabiate, the upper lip entire and galeate, enclosing the anthers, the lower lip external to the upper in bud, somewhat 3-saccate with 3 rudimentary teeth. Stamens 4, didynamous, attached near or above the middle of the corolla tube; thecae unequally placed, the outer longer and attached by the middle and the inner smaller and suspended from its apex, the cells parallel with the galea. Stigmas united and capitate, or sometimes slightly 2-lobed. Fruit a loculicidal capsule, ± asymmetrical, ovate or globose. Seeds numerous, alveolately reticulate. U.S.A. (mainly western), Mexico, Central America, South America, northern Asia; ca. 200 species, 4 in Venezuela, 1 of these in the flora area. Castilleja arvensis Schltdl. & Cham., Linnaea 5: 103. 1830. Annual herb; tips of bracts reddish. Disturbed montane areas, 1500–1600 m; Bolívar (southeast of Kavanayén). Widespread in Venezuelan Andes and Coastal

Cordillera; northeastern Mexico, Central America, nearly throughout South America, introduced in Hispaniola and Hawaii. ◆Fig. 132. Castilleja arvensis is the most widespread of all species of Castilleja.

158

S CROPHULARIACEAE

Fig. 131. Capraria biflora

Fig. 132. Castilleja arvensis

10. CONOBEA Aubl., Hist. Pl. Guiane 639. 1775. Erect to prostrate herbs. Stems terete or quadrangular. Leaves opposite, sessile or petiolate, crenate to serrate, palmately or pinnately veined. Inflorescence of solitary, axillary flowers or racemes; bractlets 1 or 2 at the apex of the pedicel or inserted at the base of the calyx, much shorter than the calyx segments. Calyx segments 5, equal or subequal, parted to near the base; corolla bilabiate, 5-lobed, the upper lip external in bud, the palate pubescent. Stamens 4, didynamous; anthers dithecal. Style branched; stigmas flattened. Capsule globose, septicidal and loculicidal (4-valved), the placental mass free-standing. Seeds numerous, longitudinally ribbed and sometimes with weak cross walls forming reticulae.

Lindernia 159

Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; ca. 4 species, 2 in Venezuela, both in the flora area. Key to the Species of Conobea 1. 1.

Leaves sessile, orbicular or suborbicular, cordate-clasping, 0.5–0.8(–1) cm long and usually slightly wider .......................................... C. aquatica Leaves petiolate, the blade lanceolate, 3.5–10 × 0.7–2(–3.2) cm .............................................................................................. C. scoparioides

Conobea aquatica Aubl., Hist. Pl. Guiane 639, pl. 258. 1775. Prostrate, sprawling, or climbing herb; petals blue. Wet places in savannas, muddy shores of pools, stream banks, 50–500 m; northern and eastern Bolívar, northwestern Amazonas. Elsewhere in eastern Venezuela; Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Amapá, Minas Gerais, Roraima). ◆Fig. 133.

Conobea scoparioides (Cham. & Schltdl.) Benth. in DC., Prodr. 10: 391. 1846. —Sphaerotheca scoparioides Cham. & Schltdl., Linnaea 2: 606. 1827. Erect, strongly aromatic herb. Along rivers, 300–400 m; Amazonas (Salto Yureba in the lower Río Ventuari basin). Mexico, Colombia, Ecuador, Peru, Brazil. Plants of Conobea scoparioides are sometimes used in bathing.

11. ESCOBEDIA Ruiz & Pav., Fl. Peruv. Prodr. 91. 1794. Perennial herbs; roots orange. Stems simple or branched. Leaves opposite, sessile, linear to ovate, coriaceous, palmately to pinnately veined. Inflorescence of racemes with leaf-like bracts; pedicels long and with a pair of bractlets inserted from near the middle to just beneath the calyx. Flowers large and showy, fragrant. Calyx tubular to funnelform, the tube 5–10-veined, equally 5-lobed, the lobes shorter than the tube; corolla white, slightly zygomorphic, salverform, the tube externally glandular-puberulent, rarely glabrous, pilose within, the limb 5-lobed, the lower 3 lobes external in bud. Stamens 4, equal, included; filaments pubescent, attached about midway in the corolla tube; anthers glabrous, with 2 equal thecae, these apiculate to aristate at the base. Style elongate with 2 stigmatic lobes. Capsule oblong-ovoid, loculicidal, included in the calyx. Seeds numerous, linear. Neotropics; 9–11 species, 1 in Venezuela. Escobedia grandiflora (L. f.) Kuntze, Revis. Gen. Pl. 3(3): 231. 1898. —Buchnera grandiflora L. f., Suppl. Pl. 287. 1781. Escobedia parimensis Pennell, Proc. Acad. Nat. Sci. Philadelphia 83: 419. 1931. Erect, coarse herb; flowers large, white.

Edges of Mauritia palm swamps, roadsides, open rocky slopes, 100–2000 m; Bolívar (Cerro Guanay), Amazonas (near Puerto Ayacucho, Sierra Maigualida, Sierra Parima). Venezuelan Andes and Coastal Cordillera; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 134.

12. LINDERNIA All., Mélanges Philos. Math. Soc. Roy. Turin 3: 178. 1766. Vandellia P. Br. ex L., Syst. Nat. ed. 12, 384, 422. 1767. Ilysanthes Raf., Ann. Nat. 13. 1820. Small herbs. Stems quadrangular. Leaves opposite or sometimes 3-whorled, sessile or petioled, denticulate to nearly entire, palmately or pinnately veined. Inflorescence of solitary, axillary flowers; pedicels without bractlets. Calyx 5-parted

160

S CROPHULARIACEAE

Fig. 133. Conobea aquatica

Fig. 134. Escobedia grandiflora

to near the base or united into a tube, the segments or lobes equal; corolla bilabiate, the upper lip short, narrow, erect, 2-lobed, external in bud, the lower lip longer, with 3 spreading lobes, the palate with 2 hairy ridges. Stamens 4 fertile or 2 fertile (posterio-lateral pair) and 2 sterile (anterio-lateral pair), the sterile ones modified into knobby projections from the ridges of the palate; anthers connivent, with 2 divergently spreading thecae. Style branched; stigmas flattened. Capsule oblong-

Lindernia 161

globose, ellipsoid, or fusiform, septicidal, the plate-like septum persisting with the placental mass. Seeds numerous, smooth or transversely lined or shallowly pitted. Cosmopolitan, mostly in Asia and Africa; ca. 70 species, 5 in Venezuela, all in the flora area. Key to the Species of Lindernia 1.

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

4.

Calyx segments united below into a tube, this sometimes splitting into free segments with expansion of the capsule; leaves petiolate, the blade pinnately veined; fertile stamens 4; leaves broadly ovate to suborbicular ................................................................................................. 2 Calyx segments free to near the base; leaves sessile, mostly palmately veined; fertile stamens 2, or if 4, then the leaves lanceolate ............. 3 Calyx 3–4.2 mm long; pedicels 7–25 mm long; capsules 2.8–4.6 mm long, oblong-globose .......................................................................... L. crustacea Calyx 4.5–7 mm long; pedicels 1–3(–4.5) mm long; capsules 6–12 mm long, ellipsoid-fusiform ................................................................ L. diffusa Fertile stamens 4; stem leaves lanceolate, 1–2 mm wide .... L. brachyphylla Fertile stamens 2; stem leaves oblanceolate to suborbicular, 4–11 mm wide ........................................................................................................ 4 Calyx 2.5–5.5 mm long; leaves 1–3 cm long, (ob)lanceolate to (ob)ovate, mostly broadest above the middle and 2 or more times longer than broad ............................................................................................... L. dubia Calyx 1.6–2.3 mm long; leaves 0.5–1 cm long, broadly ovate to suborbicular, broadest below the middle and < 1.5 times longer than broad ................................................................................................ L. microcalyx

Lindernia brachyphylla Pennell, Fieldiana, Bot. 28: 519. 1953. Lindernia barrosorum L.B. Sm., Los Angeles County Mus. Contr. Sci. 23: 5. 1958. Small, glabrous, prostrate aquatic herb. Marshes, muddy shores, streambanks, submerged in ponds and streams, 50–500 m; northern and eastern Bolívar, Amazonas (near Puerto Ayacucho). Portuguesa; Brazil. Lindernia crustacea (L.) F. Muell., Syst. Census Austral. Pl. 1(1): 97. 1882. —Capraria crustacea L., Mant. Pl. 3: 87. 1767. —Torenia crustacea (L.) Cham. & Schltdl., Linnaea 2: 570. 1827. —Vandellia crustacea (L.) Benth., Scroph. Ind. 35. 1835. Small, glabrous to sparsely pubescent, prostrate annual. Muddy or moist places in savannas and along riverbanks, 50–300; scattered in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; southeastern U.S.A., Mexico, Cen-

tral America, West Indies, Peru, Brazil, Bolivia, Africa, native in the Indo-Malayan region. Lindernia diffusa (L.) Wettst. in Engl. & Prantl, Nat. Pflanzenf. IV. 3b: 79. 1891. —Vandellia diffusa L., Mant. Pl. 1: 89. 1767. Small, pubescent, prostrate annual, rooting at the lower nodes. Wet places along disturbed roadsides, ditches, savannas, 50– 1000 m; Bolívar (lowland areas except for near Santa Elena de Uairén), Amazonas (lowlands of Río Orinoco). Portuguesa; southern Mexico, Central America, West Indies, Peru, Brazil, Bolivia, possibly an introduction from tropical Africa. ◆Fig. 136. Lindernia dubia (L.) Pennell, Acad. Nat. Sci. Philadelphia Monogr. 1: 141. 1935. —Gratiola dubia L., Sp. Pl. 17. 1753. —Ilysanthes dubia (L.) Barnhart, Bull. Torrey Bot. Club 26: 376. 1899. Ilysanthes riparia Raf., Ann. Nat. 13. 1820.

162

S CROPHULARIACEAE

Fig. 135. Lindernia microcalyx

Erect, glabrous annual. Muddy or moist places in depressions or along streambanks, ca. 50–300 m; Delta Amacuro (scattered), Amazonas (upper Río Orinoco). Southeastern Canada, U.S.A., Mexico, Central America, West Indies, sporadically distributed throughout much of South America. Lindernia microcalyx Pennell & Stehlé in Stehlé, M. Stehlé & Quentin, Fl. Guadeloupe 2(1): 217. 1938. Lindernia rotundifolia (L.) sensu Standl. & L.O. Williams, Fieldiana, Bot. 34: 122. 1972, non Alston 1931, non Pennell 1938.

Fig. 136. Lindernia diffusa

Prostrate, glabrous herb, rooting at the nodes. Muddy places or in water, near sea level to 50 m; lower Delta Amacuro. Guatemala, West Indies, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 135. The sporadic occurrence throughout most of its range suggests that Lindernia microcalyx is an introduced species. Plants from the flora area have been referred to L. rotundifolia (L.) Alston, which is native to southern India, Ceylon, Mauritius, and Madagascar, but there appears to be sufficient differences to maintain the name L. microcalyx for the Venezuelan plants.

13. MECARDONIA Ruiz & Pav., Fl. Peruv. Prodr. 95. 1794. Much-branched, glabrous herbs. Stems 4-angled. Leaves opposite, narrowed to a sessile or short-petiolate base, sometimes glandular-punctate, pinnately veined. Inflorescence of solitary, axillary flowers; pedicels elongating in fruit;

Micranthemum 163

bractlets 2, at the base of the pedicel. Calyx of 5 dissimilar segments, parted to near the base, in 2 whorls, the outer whorl with 3 distinctly broader segments, consisting of the largest posterior one and 2 slightly smaller anterio-lateral ones, the inner whorl with 2 narrow segments (the posterio-lateral pair); corolla yellow, bilabiate, the lower lip 3-lobed, the upper lip emarginate to distinctly 2-lobed, external in bud, pubescent within at the base of the upper lip. Stamens 4, all fertile, didynamous; anthers with 2 thecae, these held apart by a knob-like connective. Stigmas 2, flattened. Capsule septicidal, the placental mass remaining free-standing. Seeds numerous, reticulate. Southern U.S.A., Central America, West Indies, South America except Chile and Argentina; 10 species, 1 in Venezuela. Mecardonia procumbens (Mill.) Small, Fl. S.E. U.S. 1338. 1903. —Erinus procumbens Mill., Gard. Dict. ed. 8, 6. 1768. —Herpestis procumbens (Mill.) Urb., Symb. Antill. 4: 558. 1911. Herpestis caprarioides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 368. 1817 [1818]. Herpestis chamaedryoides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 369. 1817 [1818]. —Bacopa chamaedryoides (Kunth) Wettst. in Engl. & Prantl, Nat. Pflanzenfam. IV. 3b: 76. 1891. Glabrous, prostrate or decumbent herb, rooting at the lower nodes. Wet places along river banks, in meadows and depressions, near sea level to 300 m; Delta Amacuro (near Capure, north of Pedernales), Bolívar (lower Río Caura), Amazonas (Salto Salas on upper Río Orinoco). Venezuelan Andes and Coastal Cordillera, Zulia; southern U.S.A., Central America, West Indies, South America except Chile and Argentina. ◆Fig. 137. Fig. 137. Mecardonia procumbens

14. MICRANTHEMUM Michx., Fl. Bor.-Amer. 1: 10. 1803. Delicate, prostrate or floating, mat-forming herbs, rooting at the nodes. Stems becoming much-branched. Leaves opposite, sessile or subsessile, entire, palmately veined. Inflorescence of solitary axillary flowers; pedicels often short, in alternating axils of leaves; bractlets absent. Calyx 4-parted to near the base, the segments ± equal. Corolla rotate, bilabiate, 4-lobed by fusion of the posterio-lateral lobes, the resulting posterior lobe (upper lip) short, about 1/2 as long as the anterior 3, the lower lip external in bud. Stamens 2, attached near the top of the corolla tube, exserted; filaments geniculate with a swollen joint at base; anthers with 2 thecae, these separated by a slightly enlarged connective. Ovary 1-locular by lack of a septum distally; style short, branched; stigmas flattened and bent. Capsule

164

S CROPHULARIACEAE

subglobose or globose, membranous and splitting open irregularly or firmer and septicidal. Seeds numerous, distinctly reticulate in longitudinal rows. Eastern U.S.A., Mexico, Central America, West Indies, South America; ca. 4 species, 1 in Venezuela. Micranthemum umbrosum (J.F. Gmel.) S.F. Blake, Rhodora 17: 131. 1915. —Globifera umbrosum J.F. Gmel., Syst. Nat. 2: 32. 1791. Micranthemum orbiculatum Michx., Fl. Bor.-Amer. 1: 10, t. 2. 1803. Micranthemum pilosum Ernst, Flora 57: 215. 1874. Glabrous, creeping, mat-forming annual. Wet places and shallow streams or ponds, often in shade, 0–50 m; Delta Amacuro (near Capure, Tucupita). Widespread elsewhere in Venezuela; southeastern U.S.A., eastern Mexico, Central America, West Indies, Trinidad-Tobago, Guyana, south to Argentina. ◆Fig. 138.

Fig. 138. Micranthemum umbrosum

15. SCOPARIA L., Sp. Pl. 116. 1753. Erect, branched herbs. Leaves in whorls of 3 or 4, sometimes opposite, petiolate, glandular-punctate, serrate-dentate, pinnately veined. Inflorescence of axillary flowers, (1)2 or 3 at a node; bractlets lacking. Calyx 4- (in the flora area) or 5parted to near the base, the segments equal; corolla nearly regular, rotate, 4-lobed by fusion of the 2 posterior lobes, the lobes longer than the tube, the posterior lobe external in bud, the throat densely pilose within on all sides (in the flora area). Stamens 4, subequal, exserted; anthers with 2 thecae. Style exserted; stigmas united, capitate, sometimes slightly bifid. Capsule septicidal and secondarily loculicidal, the placental mass remaining free-standing. Seeds numerous, reticulate. Tropical and subtropical South America, 1 pantropical species; ca. 20 species, 2 in Venezuela, 1 of these in the flora area. Scoparia dulcis L., Sp. Pl. 116. 1753. Erect, perennial herb. Weedy places in open woods, river banks, disturbed fields, and roadsides, near sea level to 1300 m; Delta Amacuro (Caño Macareo), Bolívar (Gran Sabana, Río Orinoco), western Amazonas. Widespread elsewhere in Venezuela; southeastern U.S.A. to southern South America, Old World tropics. ◆Fig. 139. The somewhat less weedy species, Sco-

paria annua Schltdl. & Cham., has been collected elsewhere in Venezuela. It occupies disturbed, moist habitats such as shores of ponds and lakes, stream banks, and cultivated fields and may occur in the Guayana Region. It can be distinguished from S. dulcis by its annual habit, small stature (7– 23 cm tall), 3–7-lobed leaves, calyx with 5 lanceolate, acute segments, and yellow corolla.

16. STEMODIA L., Syst. Nat. ed. 10, 2: 1118, 1374. 1759, nom. cons. Herbs, sometimes suffrutescent. Leaves opposite or whorled, sessile and clasping or petiolate, pinnately veined. Inflorescence of solitary, axillary flowers or in terminal racemes or spikes, glandular-pubescent; bractlets 2, just beneath the calyx, or lacking. Calyx 5-parted to near the base, the segments ± equal, narrowly lanceolate to linear. Corolla bilabiate, the upper lip notched or entire, external in

Stemodia 165

Fig. 139. Scoparia dulcis

Fig. 140. Stemodia foliosa

bud, the lower lip 3-lobed, pilose within on posterior or anterior side or glabrous. Stamens 4, all fertile, didynamous, included; anthers with 2 distinct thecae separated on short arms of the connective. Style branched; stigmas flattened. Capsule ovoid-ellipsoid, septicidal and secondarily loculicidal, the placental mass usually remaining free-standing. Seeds numerous, longitudinally furrowed, pitted, or smooth. Mostly tropical America, also tropical Asia, Africa, and Australia; ca. 30 species, 4 in Venezuela, 1 of these in the flora area.

166

S CROPHULARIACEAE

Stemodia foliosa Benth., J. Bot. (Hooker) 2: 46. 1840. —Melosa. Erect, densely glandular-pubescent herb to 2 m tall. Wet savannas, roadsides, and margins of ponds, lakes, and streams, 50–

400 m; north-central Bolívar. Venezuelan Llanos; Guyana, northeastern Brazil. ◆Fig. 140. The juice from this species is used to get rid of fleas.

17. TORENIA L., Sp. Pl. 619. 1753. Small, branched herbs. Stems quadrangular. Leaves opposite, petiolate, pinnately veined. Inflorescence of axillary fascicles of 2 or more flowers, rarely only 1; bracts small, subulate; pedicels without bractlets. Calyx tubular, enclosing the capsule, the 5 lobes equal; corolla bilabiate, the 2 lobes of the upper lip united nearly to the apex and shorter than the 3 anterior lobes, external in bud. Stamens 4, didynamous; anthers with 2 thecae, connivent in pairs. Stigmas 2, flattened. Capsule fusiform, septicidal, the plate-like septum persisting with the placental mass. Seeds numerous, shallowly pitted. Pantropics; ca. 50 species, 2 in Venezuela, 1 of these in the flora area. Torenia thouarsii (Cham. & Schltdl.) Kuntze, Revis. Gen. Pl. 2: 468. 1891. —Nortenia thouarsii Cham. & Schltdl., Linnaea 3: 18. 1828. Torenia thouarsii var. nivea Pennell, Proc. Acad. Nat. Sci. Philadelphia 72: 164. 1920.

Small, pilose, prostrate or decumbent annual. Wet places in swamps and along river banks, near sea level to 100 m; Delta Amacuro (near Sacupana), Amazonas (south of Puerto Ayacucho). Native to the Old World, introduced and naturalized throughout the Neotropics. ◆Fig. 141.

Fig. 141. Torenia thouarsii

Vellosiella 167

18. VELLOSIELLA Baill., Bull. Mens. Soc. Linn. Paris 90: 715. 1887. Scandent or vining herbs. Stems branched. Leaves opposite, sinuate-toothed to subentire. Inflorescence of solitary flowers in the upper leaf axils; pedicels well developed, often recurved at the tip; bractlets 2, near the summit of the pedicel. Calyx green, entire, spathe-like, inflated, ovoid, with a deep cleft in front and an acuminate to caudate tip in back; corolla orange, funnel-shaped to campanulate, variously exserted from the calyx, the orifice oblique and down-curved, the subequal lobes erect, the lower 3 external in bud. Stamens 4; anthers with 2 parallel thecae. Stigma capitate. Capsule globose, loculicidal. Seeds elongate, cylindrical- to elongate-conical, longitudinally elongate-reticulate. Venezuela, Guyana, central and southern Brazil; 3 species, 1 in Venezuela. Vellosiella spathacea (Oliv.) Melch., Notizbl. Bot. Gart. Berlin-Dahlem 15: 129. 1940. —Melasma spathaceum Oliv., Timehri 5: 200. 1886. Scandent-climbing, ± brittle herb. Slopes of tepuis in cloud forests, often in rocks at base of escarpments, 1800–2600 m; Bolívar (Apacará-tepui, Auyán-tepui, Ilú-tepui, Ptaritepui, Roraima-tepui, Sororopán-tepui), Amazonas (Sierra de la Neblina). Guyana (Mount Roraima). ◆Fig. 142.

Fig. 142. Vellosiella spathacea

168

S IMAROUBACEAE

SIMAROUBACEAE AND PICRAMNIACEAE Using molecular techniques, Fernando et al. (1995) determined that Picramnia and Alvaradoa were not closely related to other genera placed in the Simaroubaceae. As a result, they described a new family, Picramniaceae, to accommodate these genera (Fernando and Quinn 1995). Here we follow this circumscription and treat the families separately. Since previous volumes of the Flora already treated families beginning with “P,” the family treatment for Picramniaceae will immediately follow Simaroubaceae, in which it was originally placed, but outside of the usual alphabetical family order followed in the series. Key to Picramniaceae and Simaroubaceae sensu stricto 1.

1.

Flowers unisexual (plants dioecious); sepals and petals 3–5; stamens 3– 5, equaling and opposite the petals; fruit simple, a berry or samara .............................................................................. Picramniaceae, p. 177 Flowers bisexual or unisexual (plants dioecious in Simarouba); sepals and petals 5; stamens 10, opposite both sepals and petals; fruits aggregate, with 1(2–5) drupes or samaras per flower ........................... ............................................................................. Simaroubaceae, p. 168

References Fernando, E. S., and C. J. Quinn. 1995. Picramniaceae, a new family, and a recircumscription of Simaroubaceae. Taxon 44: 177–181. Fernando, E. S., P. A. Gadek, and C. J. Quinn. 1995. Simaroubaceae, an artificial construct: evidence from rbcL sequence variation. Amer. J. Bot. 82: 92–103.

SIMAROUBACEAE by W. Wayt Thomas and Edivani V. Franceschinelli Trees, shrubs, or subshrubs. Dioecious (Simarouba) or bisexual (Picrolemma, Quassia, and Simaba). Leaves alternate, pinnately compound with 1–29 leaflets or, rarely, simple, the leaflets alternate or opposite, often subcoriaceous with obscure venation; stipules absent. Inflorescences paniculate or racemose, terminal or axillary. Flowers actinomorphic; calyx and corolla (4)5-merous, glabrous to puberulent. Stamens 10; filaments glabrous or puberulent, usually with an adaxial appendage, the appendage free or fused to the filament and represented by a tuft of hairs. Gynoecium borne on a disk or gynophore; carpels 5 and essentially free with a common style and divergent stigmas. Fruit a drupe (technically a druparium) or a samara (samarium), 1(2–5) per flower. Pantropics, with some subtropical or temperate species, especially in Asia; 13 genera and ca. 130 species, 4 genera and 9 species in the flora area.

Simaba 169

Key to the Genera of Simaroubaceae 1. 1. 2(1). 2. 3(2). 3.

Leaf rachis winged .......................................................................... 2. Quassia Leaf rachis not winged .............................................................................. 2 Flowers orange, unisexual; fruits orange; stems hollow ........ 1. Picrolemma Flowers creamy white, unisexual or bisexual; fruit green to maroon or scurfy brown, not orange; stems not hollow ........................................ 3 Flowers unisexual; leaflets alternate to subopposite, usually 9–21 .......... ................................................................................................ 4. Simarouba Flowers bisexual; leaflets opposite, usually 1–7 ........................... 3. Simaba

1. PICROLEMMA Hook. f., Gen. Pl. 1: 312. 1862. Fistulose shrubs or cauliflorous trees with hollow stems. Leaves odd-pinnate or even-pinnate, with 4–9 pairs of leaflets. Inflorescence paniculate, terminal or axillary, or cauliflorous. Flowers 4- or 5-merous, orange, unisexual. Staminodia opposite sepals, stamens opposite petals. Drupes orange and elongate or brownish and egg-shaped. Amazon Basin and lower slopes of the Andes (Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil); 2 species, 1 in Venezuela. Picrolemma sprucei Hook. f., Gen. Pl. 1: 312. 1862. Picrolemma pseudocoffea Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 196. 1925.

Shrub or small tree 1–10 m tall; fruits orange. Evergreen lowland forest understories, 100–300 m; Amazonas (Río Atabapo, Río Negro). Colombia, French Guiana, Peru, Brazil (Amapá, Amazonas). ◆Fig. 143.

2. QUASSIA L., Sp. Pl. ed. 2, 1: 553. 1762. Small tree, shrub, or scandent shrub. Leaves odd-pinnately compound, rachises 1 or occasionally several, terminal or subterminal, winged and jointed at the insertion of the leaflets. Flowers 2–3 cm long, pink to red. Drupes 1–5 per flower. Mexico, Central America, West Indies, northern South America; 1 species. This species is native to northern South America but is planted elsewhere because of its medicinal qualities. Nooteboom (1962) included Simaba, Simarouba, and several Old World genera in a more broadly circumscribed Quassia [Nooteboom, H. P. 1962. Generic delimitation in Simaroubaceae tribus Simaroubeae and a conspectus of the genus Quassia L. Blumea 11: 509–528]. Quassia amara L., Sp. Pl. ed. 2, 1: 553. 1762. —Bejuco barbasco. Small tree, shrub, or scandent shrub. Evergreen lowland forests, 50–300 m; Bolívar (El Dorado, Isla Anacoco, Río Cuyuní). Barinas, Zúlia; Mexico, Central America, West Indies, Colombia, Trinidad-Tobago,

Guyana, Suriname, French Guiana, Brazil (Amazonas). ◆Fig. 144. Quassia amara is native to the Venezuelan Guayana, the Guianas, and northern Brazil; it is used medicinally, especially to combat fevers and digestive disorders.

3. SIMABA Aubl., Hist. Pl. Guiane 400, t. 153. 1775. Shrubs or trees. Leaves odd-pinnate, the leaflets opposite. Inflorescence paniculate, usually erect, puberulent, the pedicels usually subtended by a small pendulous gland. Flowers bisexual. Sepals and petals (4)5. Filaments with an adaxial,

170

S IMAROUBACEAE

Fig. 143. Picrolemma sprucei

Simaba 171

Fig. 144. Quassia amara

slender, usually pubescent, adnate or distally free appendage. Gynophore cylindrical; ovary globose to depressed-globose, of (4)5 separate carpels joined by a common style. Fruit a drupe, usually one per flower. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Africa; 26 species, 6 in Venezuela, all in the flora area.

172

S IMAROUBACEAE

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

2.

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

5(4). 5.

Rachis including petiole 60–115 cm long; leaflets often gland-tipped ....................................................................................................... S. cedron Rachis including petiole 2–25 cm long; leaflets never gland-tipped ...... 2 Inflorescence usually small and few-branched (except S. polyphylla which has a larger inflorescence and usually 5–11 pairs of leaflets); flowers usually not arranged in cymes; pedicels 2–5 mm long; appendage free with free portion 0.6–1.5 mm long or mostly united with 0–0.2 mm free and with 2 apical teeth, equaling or surpassing the ovary ....................................................................................................... 3 Inflorescence usually large, lax, and many-branched, 5–50 cm long (except in S. obovata where they may be as short as 1.5 cm long); flowers usually arranged in cymes with pedicels 4–12 mm long; appendage completely united to the filament and usually with 2 apical teeth, not surpassing ovary (in flowers where ovary is not reduced) .................. 4 Leaves with more than one leaflet (or if only one leaflet, then other leaves of the same plant with more than one leaflet) ..................... S. guianensis Leaves unifoliolate ....................................................................S. monophylla Leaves unifoliolate; leaflet with 2–4 pairs of minute foliar glands, each situated midway between the midvein and the margin ........... S. obovata Leaves (at least most of them) with more than one leaflet; leaflets with 1–4 pairs of minute foliar glands, each one situated at the base of the blade or along margin ........................................................................... 5 Leaflets with glands at base of the blade; flower pedicels 4–12 mm long; fruit flattened ......................................................................... S. orinocensis Leaflets with glands along margin above the base; flower pedicels 4– 6 mm long; fruit ellipsoid or ovoid ........................................ Simaba sp. A

Simaba cedron Planch., London J. Bot. 5: 566. 1846. —Carbón, Casabote, Palo de vela, Uña de pereza, Vela de muerto. Few-branched tree or shrub 1.5–15 m tall; leaflets with an apical gland. Evergreen lowland and riparian forests, forest margins, 50–400 m; Bolívar (Las Trincheras, Represa Guri, Río Caura), Amazonas (Canaripó, Raudal de Atures). Barinas; Costa Rica, Panama, Colombia, Guyana, French Guiana, Suriname, Peru, Brazil, Bolivia. ◆Fig. 145. Simaba guianensis Aubl., Hist. Pl. Guiane 400, t. 153. 1775. Venezuela, Guyana, Suriname, French Guiana, Brazil; probably 4 subspecies (taxonomy not yet resolved), 2 of these in Venezuela, both in the flora area.

Key to the Subspecies of S. guianensis 1. Shrub or small tree, 0.3–7 m tall; leaflets subcoriaceous to coriaceous, seldom revolute; free portion of staminal appendage 0.6–2.1 mm long; fruits yellow to orange ............... subsp. guianensis 1. Shrub 0.2–3 m tall; leaflets usually coriaceous and often strongly revolute; free portion of staminal appendage 0.5– 1.2 mm long; fruits burgundy ................. ...................................... subsp. huberi S. guianensis subsp. guianensis Shrub or small tree 0.3–7 m tall. Guyana, Suriname, French Guiana, Brazil (Amapá). Subspecies guianensis has not yet been verified from Venezuela, though it occurs in nearby Guyana. Elsewhere it is known from lowland forests at 50–500 m elevation.

Simaba 173

Fig. 145. Simaba cedron

174

S IMAROUBACEAE

Fig. 146. Simaba obovata

Fig. 147. Simaba guianensis subsp. huberi

Fig. 148. Simaba orinocensis

Simarouba 175

Simaba guianensis subsp. huberi Francesch. & W. Thomas, Brittonia 52: 311, fig. 1. 2000 [2001]. Shrub or subshrub 0.2–1(–3) m tall. White-sand savannas, 100–200 m; Amazonas (Canaripó, Caño Caname, Caño Corocoro near lower Río Ventuari, near Cerro Autana, near Cerro Yapacana, Pimichín, Río Atabapo above Guarinuma). Endemic. ◆Fig. 147. Simaba monophylla (Oliv.) Cronquist, Lloydia 7: 88. 1944. —Simarouba monophylla Oliv. in Hook., Icon. Pl. 14: 108, t. 1387. 1882. Shrub or subshrub 0.1–3 m tall. Whitesand savannas, 100–1000 m; Bolívar (Ilútepui), Amazonas (Río Sipapo). Guyana. Simaba obovata Spruce ex Engl. in Mart., Fl. Bras. 12(2): 210. 1874. —Limoncillo, Piraquiña. Tree 4–15 m tall. Riverbanks and periodically flooded forests associated with blackwater rivers, 100–200 m; Amazonas (La Esmeralda, Río Atabapo, Río Yatúa, San Carlos de Río Negro, Tama-Tama). Colombia, Brazil (Amazonas, Roraima). ◆Fig. 146.

Simaba orinocensis Kunth in H.B.K., Nov. Gen. Sp. 6 (quarto ed.): 18, t. 514b. 1823. —Almendrón cunavichero, Cerezo, Ciruela, Ciruelillo, Congrillo, Pirequina, Tambak. Simaba multiflora A. Juss., Mém. Mus. Hist. Nat. 12: t. 27. 1825. Shrub or tree 1.5–22 m tall. Seasonally flooded forests, 50–300 m; Delta Amacuro (east-northeast of El Palmar), Bolívar (widespread), Amazonas (Caño Jiboa opposite Minicia, Cerro Yutajé, Isla Borua, La Esmeralda, Raudal de Atures, Río Casiquiare, San Carlos de Río Negro). Apure, Guárico; Guyana, Suriname, French Guiana, Peru, Brazil (Acre, Amazonas, Pará, Roraima), Bolivia. ◆Fig. 148. Simaba sp. A. —Congrillo, Palo azul, Pilón. Tree 10–30 m tall. Evergreen lowland forests, 100–300 m; Delta Amacuro (near El Palmar), Bolívar (El Dorado, Los Patos south of El Manteco). Aragua; Colombia, Brazil (Amazonas, Rondônia). In the flora area this entity is based on Steyermark 89815 (VEN), Steyermark 86685 (VEN), and Pittier 15682 (US, VEN).

4. SIMAROUBA Aubl., Hist. Pl. Guiane 859, pl. 331, 332. 1775, nom. cons., non Simaruba Boehmer 1760. Medium to large trees. Leaflets alternate to subopposite. Inflorescence paniculate, the staminate inforescence larger with more flowers than the pistillate one. Flowers unisexual. Sepals and petals 5. Filaments with an adaxial, pubescent, distally free appendage. Gynophore cylindrical; ovary globose to depressed-globose, with (4)5 separate carpels joined by a common style. Fruit a drupe, 1 or more per flower. U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 6 species, 1 in Venezuela. Simarouba amara Aubl., Hist. Pl Guiane 860. 1775. —Anapora-yek, Anepurayek, Cedro blanco, Cedro dulce, Parrillo, Tú ne yó, Yacu-yek. Tree 3–25(–40) m tall. Evergreen lowland to montane (often secondary) forests, near sea level to 1000 m; Delta Amacuro (Casa Cuyubini, east-northeast of El Palmar, Serranía de Imataca, Tucupita), Bolívar

(Altiplanicie de Nuria, Karaumi, Represa Guri, Río Hacha, Sucre, Uacupe, road from San Félix to Upata), Amazonas (Río Atabapo basin, Cerro Aratitiyope, Sierra de la Neblina). Widespread from Honduras to northern South America, the Amazon Basin and coastal Brazil south to Espírito Santo. ◆Fig. 149.

176

S IMAROUBACEAE

Fig. 149. Simarouba amara

Picramnia 177

PICRAMNIACEAE by José Rubens Pirani Trees or shrubs. Leaves alternate, odd-pinnately compound, the leaflets usually alternate with oblique bases, often with wavy margins; stipules absent. Inflorescences racemes, panicles, or racemose or paniculate simple or compound thyrses, terminal or axillary, rarely cauliflorous, often pendent. Flowers unisexual, actinomorphic, usually < 5 mm long. Calyx and corolla 3–5-merous. Stamens 3–5, opposite petals, exserted. Gynoecium borne on a disk or gynophore; carpels 2–4, united with a very short style and divergent, persistent stigmas. Fruit a berry or a samara. Neotropics; 3 genera and 49 species, 1 genus and 7 species in the flora area. 1. PICRAMNIA Sw., Prodr. 2: 27. 1788, nom. cons. Small trees or shrubs. Leaflets alternate, usually chartaceous, often darkening upon drying, the base of lateral leaflets often oblique. Inflorescence pendent to arching, a simple raceme, either of separate flowers or glomerules of flowers, or a compound thyrse. Flowers unisexual. Stamens or staminodia (if present) 5; filaments unappendaged; intrastaminal disk entire or (4)5-lobed. Staminate flowers with pistillodium usually conical and pubescent. Pistillate flowers with ovary ovoid, glabrous to densely pubescent, carpels 2(–4), the style very short, the stigmas divergent. Fruit a berry, obovoid to ellipsoid, glabrous to glabrate and shiny or short-pubescent, the calyx persistent, the sessile stigmas persistent. Neotropics; 40 species, ca. 10 in Venezuela, 7 in the flora area. Key to the Species of Picramnia 1.

1. 2(1).

2.

3(2).

3. 4(1). 4.

Inflorescence lateral (emerging from a portion of the stem already without leaves) or lateral and terminal inflorescences on the same branch .................................................................................................... 2 Inflorescence terminal ............................................................................... 4 Older stems hollow, housing ants; leaflets with percurrent (arching) secondary veins; staminate flowers long-pedicellate; fruit pilose to glabrescent ............................................................................. P. magnifolia Older stems not hollow, not housing ants; leaflets with secondary veins not percurrent but much-branched and reticulate; staminate flowers sessile or with a pedicel to 1.5 mm long; fruit glabrous ...................... 3 Flowers usually anisomerous, with 5 sepals (rarely 3 or 4), and 3 or 4 petals and 3 or 4 stamens; flowers condensed in dense glomerules of 4–10 flowers; leaflets often abruptly caudate ........................ P. juniniana Flowers isomerous and 5-merous, isolated or in groups of 2 or 3; leaflets never abruptly caudate .............................................................. P. latifolia Plants (leaflets, inflorescence, and fruits) densely puberulent ... P. nuriensis Plants glabrous to sparsely puberulent, or occasionally the fruits puberulent .............................................................................................. 5

Picramnia 177

PICRAMNIACEAE by José Rubens Pirani Trees or shrubs. Leaves alternate, odd-pinnately compound, the leaflets usually alternate with oblique bases, often with wavy margins; stipules absent. Inflorescences racemes, panicles, or racemose or paniculate simple or compound thyrses, terminal or axillary, rarely cauliflorous, often pendent. Flowers unisexual, actinomorphic, usually < 5 mm long. Calyx and corolla 3–5-merous. Stamens 3–5, opposite petals, exserted. Gynoecium borne on a disk or gynophore; carpels 2–4, united with a very short style and divergent, persistent stigmas. Fruit a berry or a samara. Neotropics; 3 genera and 49 species, 1 genus and 7 species in the flora area. 1. PICRAMNIA Sw., Prodr. 2: 27. 1788, nom. cons. Small trees or shrubs. Leaflets alternate, usually chartaceous, often darkening upon drying, the base of lateral leaflets often oblique. Inflorescence pendent to arching, a simple raceme, either of separate flowers or glomerules of flowers, or a compound thyrse. Flowers unisexual. Stamens or staminodia (if present) 5; filaments unappendaged; intrastaminal disk entire or (4)5-lobed. Staminate flowers with pistillodium usually conical and pubescent. Pistillate flowers with ovary ovoid, glabrous to densely pubescent, carpels 2(–4), the style very short, the stigmas divergent. Fruit a berry, obovoid to ellipsoid, glabrous to glabrate and shiny or short-pubescent, the calyx persistent, the sessile stigmas persistent. Neotropics; 40 species, ca. 10 in Venezuela, 7 in the flora area. Key to the Species of Picramnia 1.

1. 2(1).

2.

3(2).

3. 4(1). 4.

Inflorescence lateral (emerging from a portion of the stem already without leaves) or lateral and terminal inflorescences on the same branch .................................................................................................... 2 Inflorescence terminal ............................................................................... 4 Older stems hollow, housing ants; leaflets with percurrent (arching) secondary veins; staminate flowers long-pedicellate; fruit pilose to glabrescent ............................................................................. P. magnifolia Older stems not hollow, not housing ants; leaflets with secondary veins not percurrent but much-branched and reticulate; staminate flowers sessile or with a pedicel to 1.5 mm long; fruit glabrous ...................... 3 Flowers usually anisomerous, with 5 sepals (rarely 3 or 4), and 3 or 4 petals and 3 or 4 stamens; flowers condensed in dense glomerules of 4–10 flowers; leaflets often abruptly caudate ........................ P. juniniana Flowers isomerous and 5-merous, isolated or in groups of 2 or 3; leaflets never abruptly caudate .............................................................. P. latifolia Plants (leaflets, inflorescence, and fruits) densely puberulent ... P. nuriensis Plants glabrous to sparsely puberulent, or occasionally the fruits puberulent .............................................................................................. 5

178

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

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

P ICRAMNIACEAE

Fruits and ovaries puberulent with obtuse (clavate or digitiform) hairs ................................................................................................. P. guianensis Fruits and ovaries glabrous to puberulent, the hairs, if present, subulate or attenuate ........................................................................................... 6 Flowers usually anisomerous, with 5 (rarely 3 or 4) sepals and 3 or 4 petals and 3 or 4 stamens; flowers condensed in dense glomerules of 4–10 flowers; leaflets often abruptly caudate ........................ P. juniniana Flowers isomerous and 5-merous, isolated or in groups of 2 or 3; leaflets never abruptly caudate ......................................................................... 7 Pistillate flowers subsessile; staminate flowers in cymules of 2 or 3; fruit sessile or with a short, thickened pedicel .................................. P. latifolia Pistillate flowers with evident pedicel 1–6 mm long; staminate flowers in cymules of 2–12; fruit clearly pedicellate ........................................ 8 Leaflets subcoriaceous, shiny, drying green to brown, the tertiary venation forming a distinct network ..................................... P. pentandra Leaflets chartaceous, shiny or not on upper surface, dull on lower surface, drying dark brown to black, often leaving reddish marks, the tertiary venation not evident .................................................. P. spruceana

Picramnia guianensis (Aubl.) Jans.-Jac. in Stoffers & Lindeman, Fl. Suriname 5(1): 329. 1979. —Tariri guianensis Aubl., Hist. Pl. Guiane 2(Suppl.): 37, t. 39. 1775. Small tree or shrub 1.2–7 m tall. Expected in Bolívar. Guyana, Suriname, French Guiana, Brazil (Amapá). Picramnia guianensis has not yet been found in Venezuela, but probably occurs there. Elsewhere it is found in forests and scrub forests at elevations of 50–800 m. Picramnia juniniana J.F. Macbr., Candollea 5: 374. 1934. Picramnia tristamina Steyerm., Fieldiana, Bot. 28: 275. 1952. Small tree or shrub 0.8–10 m tall. Evergreen lowland and seasonally flooded riverine forests, 100–300 m; Amazonas (San Fernando de Atabapo to Caño Iguapo). Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 151. Picramnia latifolia Tul., Ann Sci. Nat. sér. 3, 7: 258. 1847. Picramnia macrostachys Klotzsch ex Engl. in Mart., Fl. Bras 12(2): 238. 1874. Small to medium-sized tree 3–10(–20) m tall. Evergreen lowland to lower montane forests, 50–1000 m; Delta Amacuro (northwest of El Palmar), Bolívar (Reserva Forestal Imataca), Amazonas (Raudal de los Guaharibos on upper Río Orinoco). Mexico,

Central America, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Picramnia magnifolia Macbride, Candollea 5: 376. 1934. Picramnia platystachya Killip & Cuatrec., Revista Acad. Colomb. Ci. Exact. 5: 37: 1942. Few-branched shrub or small tree 1–6 m. Evergreen lowland to montane forests, 100– 1600 m; Amazonas (Caño Caname, Caño San Miguel, Cerro Duida, Cerro Huachamacari, Cerro Sipapo, road between Maroa to Yavita, Río Autana, Raudal de Atures, Río Atabapo, Río Atacavi, Río Baría, Río Casiquiare, Río Cataniapo, Río Cunucunuma, Río Negro, Salto Huá on Canal Maturacá, San Fernando de Atabapo, Sierra Imeri). Colombia, Ecuador, Peru, Brazil, Bolivia. Picramnia nuriensis Steyerm., Acta Bot. Venez. 3: 177. 1968. —Jobo. Shrub or small tree 3–5 m tall. Lower montane and montane forests, 300–800 m; Bolívar (Altiplanicie de Nuria, Cerro El Diablo, near Deborah, between El Dorado and Kilómetro 88, Santa Elena de Uairén, east-southeast of Villa Lola). Trujillo; Guyana. Picramnia pentandra Sw., Fl. Ind. Occid. 1: 220. 1797. —Carapilla. Small tree or shrub 1.5–8 m tall. Evergreen lowland semideciduous forests, 200–

Picramnia 179

400 m; Bolívar (east of La Paragua). Distrito Federal, Táchira, Zulia; U.S.A. (Florida), West Indies, Colombia. Picramnia spruceana Engl. in Mart., Fl. Bras. 12(2): 238. 1874. —Picramnia sellowii subsp. spruceana (Engl.) Pirani, Boletim Botânica Univ. São Paulo 12: 132. 1990. Tree or shrub 2–13(–20) m tall; leaves usually drying dark purple or black. Semi-

deciduous to evergreen lowland and seasonally flooded riverine forests, 50–400 m; Bolívar (Distrito Cedeño, Represa Guri), Amazonas (Caño Cumashi tributary of Río Cunucunuma, La Esmeralda, Puerto Ayacucho, Raudal de Atures, Río Atabapo, upper Río Casiquiare, Río Cunucunuma, Río Jenita, Río Mavaca, upper Río Orinoco, San Antonio). Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 150.

Fig. 150. Picramnia spruceana

Fig. 151. Picramnia juniniana

180

S IPARUNACEAE

SIPARUNACEAE by Susanne S. Renner and Gerlinde Hausner Evergreen treelets, trees, or shrubs with spherical oil cells in all parts of the plant. Leaves decussate or in whorls of 3–6, simple, exstipulate, those of a pair occasionally unequal in size, with stellate, lepidote, or simple trichomes, the margin serrulate, denticulate, or entire. Inflorescences axillary or cauliflorous, cymose, sometimes fasciculate. Flowers typically radial, strictly unisexual (plants dioecious or more rarely monoecious). Floral receptacle subglobose or cup-shaped, the gynoecium apocarpic, tepals 4–6, rarely 8, usually minute, in a few species to 10 mm long, sometimes fused to a rim encircling the floral cup, rarely one tepal much longer than the others (in the African Glossocalyx) or tepals forming a calyptra, the flower center ± covered by a membrane (the so-called floral roof or velum) except for a central pore through which stamens or styles emerge. Stamens (1)2–72, dispersed irregularly in the floral cup except in flowers with very few stamens; filaments lacking appendages; anthers with 2 closely adjacent pollen sacs opening by a single apically hinged flap (valve). Carpels 3–35, each immersed in a pocket in the floral cup; styles sometimes postgenitally fused; ovules solitary, basally attached and anatropous, unitegmic, crassinucellate. Mature fruiting receptacles fleshy, globose or pear-shaped, smooth, spiny, or tuberculate, in most species splitting open at maturity to expose the drupelets; drupelets with a thin translucent exo- and mesocarp and a stony endocarp, in dioecious species the drupelets with a red to orange, fleshy appendage at the style base (stylar aril), rarely the drupelets lacking an appendage and the exo- and mesocarp becoming slimy. Tropics; 2 genera and ca. 60 species, 1 genus and 8 species in the flora area. 1. SIPARUNA Aubl., Hist. Pl. Guiane 864. 1775. Trees to 40 m tall and to 120 cm DBH, rarely shrubs. Leaves opposite, rarely in whorls of 3 or 4, entire, dentate, serrate, or crenate, with stellate, simple, or lepidote trichomes, rarely glabrous, papery to coriaceous, strongly aromatic; petioles short or long. Inflorescences mono- or dichasial cymes, axillary or on leafless nodes, inflorescence axes elongate or short, sometimes subsessile flower clusters. Flowers unisexual, usually small; receptacle subglobose, cup-shaped, urceolate, or rarely flask-shaped, with stellate, simple, or lepidote trichomes, rarely glabrous; tepals usually short, triangular, rounded, or spatulate, or forming a rim encircling the floral cup, rarely a calyptra; floral roof in male flowers ± cylindrically raised with a relatively large central pore, in female flowers of monoecious species generally conical with a smaller pore, in female flowers of dioecious species usually differentiated into an outer bulge and an inner tube, tightly sheathing the styles. Stamen filaments flattened, rarely cylindrical, in some species 4 of them laterally adnate forming a tube. Fruiting receptacle smooth, spiny, or with tubercles, when ripe usually, but not always, splitting irregularly and exposing the enclosed seed-like drupelets, in the dioecious species with a fleshy, orange or red stylar aril (see family description). Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 60 species, 13 in Venezuela, 8 of these in the flora area.

Siparuna 181

Key to the Species of Siparuna 1.

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

3.

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

6.

7(5).

7.

Tepals forming a calyptra (dome) that completely encloses the stamens or carpels and after anthesis dehisces circumscissilely; leaves with isolated, silvery, appressed, peltate scales on the upper and lower surfaces; fruiting receptacle almond-shaped or somewhat asymmetric, rarely ovoid, depending on the number of enclosed drupelets, with a distinct tip (where the calyptra fell off), 1.5–3 cm long and 0.7–1.5 cm diameter .................................................................................... S. decipiens Plants not with the above combination of characters ............................. 2 Cymes ca. 1.5 cm long; flowers in subsessile clusters ............................. 3 Cymes ≥ 2 cm long, bifid with regularly spaced flowers or flowers on ± elongate axes ......................................................................................... 4 Shrub or treelet 5–9(–15) m tall and to 20 cm DBH; leaves drying grayish brown; flowers 1.8–3 mm diameter; stamens 10–18, membranaceous, glabrous; fruiting receptacle ± densely covered with stellate trichomes ................................................................................ S. guianensis Subcanopy tree 13–20 m tall and to 10–11 cm DBH; leaves drying light green or yellowish green; flowers 4–5 mm diameter; stamens 27– 30, fleshy, dorsally with a few stellate trichomes; fruiting receptacle persistently velvety pubescent .. ................................................. S. ficoides Plants (populations) dioecious; adult leaves with simple or stellate, ± appressed trichomes on upper surface ......................................... S. bifida Plants monoecious, mature or older leaves glabrous or glabrescent on upper surface ......................................................................................... 5 Fruiting (and flowering) receptacles lacking any outgrowths ................ 6 Fruiting (and flowering) receptacles with fleshy spines, tubercles, or wart-like outgrowths, best seen in mature fruits ................................ 7 Leaves (7–)9–14 cm wide, the secondary veins anastomosing near the leaf margin, the loops inconspicuous; pedicels of staminate flowers to 20 mm long; fruiting receptacles 2–3.5 cm diameter ................ S. cristata Leaves 3.5–10 cm wide, the secondary veins often anastomosing 2–3 mm before reaching the leaf margin and forming conspicuous loops; pedicels of staminate flowers to 13 mm long; fruiting receptacles 0.8– 1 cm diameter .............................................................................. S. obstipa Leaves with 10–14 pairs of secondary veins; leaf apex acuminate, young branchlets densely or loosely reddish brown or yellow pubescent with trichomes to 3 mm long, cymes bifid, 3.5–6.5 cm long, with 30– 80 flowers ..................................................................................... S. reginae Leaves with 6–8(9) pairs of secondary veins; leaf apex cuspidate, young branchlets with silvery peltate scales or stellate-lepidote trichomes, cymes with elongate axes, 2–4 cm long, with 10–15 flowers .................................................................................................. S. cuspidata

Siparuna bifida (Poepp. & Endl.) A. DC., Prodr. 16: 652. 1868. —Citrosma bifida Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 48. 1838, “Citriosma.”

Dioecious treelet, rarely a large tree 2– 11(–20) m tall and to 20 cm DBH; inflorescence bifid, densely covered with minute stellate trichomes, sometimes glabrescent,

182

S IPARUNACEAE

staminate cymes 3.5–5.5 cm long, with 30– 150 regularly spaced subsessile flowers; pistillate cymes 2–3 cm long and with fewer flowers; fruiting receptacle globose, 1–1.5 cm diameter, with short fleshy tubercles and often loosely villous. In primary and secondary forests, usually along banks of streams; 100–1100 m; Bolívar (Distrito Roscio, Santa Elena de Uairén), Amazonas (Río Matacuni, Sierra Parima, Simarawochi, 5–7 km south of Solano on road between San Carlos and Solano). Peru, Brazil, Bolivia. Siparuna cristata (Poepp. & Endl.) A. DC., Prodr. 16: 665. 1868. —Citrosma cristata Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 47, pl. 164. 1838. —Abari, Awadi (Yekwana), “Citriosma.” Monoecious treelet or tree (3–)8–22 m tall and to 45(–60) cm DBH; branchlets lepidote, later glabrous; inflorescence with 3–6 pistillate flowers at base and 15–20 staminate flowers at apex. Evergreen lowland forests to lower montane forests, 100–800 m, rarely to 1200 m; Bolívar (upper Río Caura), Amazonas (near Culebra, Raudal Picure in Río Cunucunuma, Río Padamo, San Carlos de Río Negro, Yavita to Maroa road). Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, expected in Suriname. Siparuna cuspidata (Tul.) A. DC., Prodr. 16: 655. 1868. —Citrosma cuspidata Tul., Ann. Sci. Nat. Bot. sér. 4, 3: 40. 1855. Monoecious treelet or tree 3–20(–35) m tall and to 40 cm DBH; branchlets stellatelepidote; inflorescence 2–4 cm long, with 10– 15 terminally clustered flowers. Evergreen lowland forests, 100–1300 m; Amazonas (San Carlos de Río Negro, Yavita to Maroa road). Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Siparuna decipiens (Tul.) A. DC., Prodr. 16: 643. 1868. —Citrosma decipiens Tul., Monogr. Monim. 368. 1855. —Aguacatillo blanco, Cumanagota, Laurel bobo, “Citriosma.” Conuleum guianense A. Rich., Mém. Soc. Hist. Nat. Paris 4: 391, 406, t. 25. 1823.

Monoecious tree (3–)10–22 m tall and to 25(–30) cm DBH; young branchlets terete or striate; inflorescence 2–6 cm long; fruits ovoid with a pointed tip. Evergreen lowland to lower montane forests, 50–800 m; Delta Amacuro (near Santa Catalina), Bolívar (Altiplanicie de Nuria, between El Manteco and San Pedro de las Bocas, Kilómetro 88, La Escalera, Río Chicanán, near Santa Elena de Uairén), Amazonas (southwestern side of Sierra de la Neblina). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 154. Siparuna ficoides Renner & Hausner, Novon 15: 202. 2005. —Medebadi (Yekwana). Monoecious subcanopy tree 13–20 m tall and to 10–11 cm DBH; young branchlets terete or flattened at the nodes; leaves glabrous, thick-coriaceous, drying light green or yellow-green; inflorescences about 1.5 cm long, with 6–12 flowers, persistently velvety; fruits subglobose, about 2 cm diameter and 1.1 cm long, persistently velvety brownpubescent. Lower montane forests and forest edges, ca. 400 m; Bolívar (Aripao, Botadero Norte). Brazil (Amazonas). Siparuna guianensis Aubl., Hist. Pl. Guiane 865, t. 333. 1775. —Citrosma guianensis (Aubl.) Tul., Monogr. Monim. 361. 1855, “Citriosma.” —Arbol del danto, Culebra, Cumanagoto, Hierba de pasmo, Hoja de bachaco, Palo de bachaco, Paripari, Uatakurán (Arekuna), Yakantayuru. Monoecious shrub or treelet 5–9(–15) m tall. Semideciduous to evergreen lowland, lower montane, and gallery forests, forest edges, open savannas, 50–1200(–1400) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 152. Siparuna guianensis is used by local inhabitants to treat malaria and other fevers. Siparuna obstipa J.F. Macbr., Candollea 5: 354. 1934. Monoecious treelet or tree 3–18 m tall; inflorescences 2–3 cm long, with 20–30 flow-

Siparuna 183

ers, with very fine appressed-stellate trichomes on the cymes, pedicels, and receptacles; fruiting receptacle globose, smooth and glabrous, 0.8–1 cm diameter. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro). Colombia, Peru, Brazil. Siparuna reginae (Tul.) A. DC., Prodr. 16: 654. 1868. —Citriosma reginae Tul., Ann. Sci. Nat. Bot. sér. 4, 3: 39. 1855. —Siparuna micrantha A. DC., J. Bot. 3: 219. 1865. —Siparuna surinamensis Lanj., Recueil Trav. Bot. Néerl. 32: 240. 1935. —Siparuna sancheziana Steyerm., Bol. Soc. Venez. Ci. Nat. 26:

Fig. 152. Siparuna guianensis

419. 1966. —Palo de bachaco, Palo paripari, Paripari. Monoecious treelet or tree 5–22(–40) m tall and to 70 cm DBH; branchlets terete, but flattened below the nodes, densely reddish brown stellate-pubescent; fruiting receptacles globose, with villous-tomentose tubercles or warts. Evergreen lowland to montane forests, 50–1200 m; Bolívar (Aripao, between Divina Pastora and Santa Elena, Río Caura, Río Oris in Río Paragua basin, Sierra de Lema), Amazonas (Caño Cucurital, near Pimichín along road to Yavita, Río Ventuari, around San Carlos de Río Negro, road between Yavita and Maroa). Colombia, Guyana, Suriname, Ecuador, Peru, Brazil. ◆Fig. 153.

Fig. 153. Siparuna reginae

184

S IPARUNACEAE

Fig. 154. Siparuna decipiens

SMILACACEAE by John F. Gaskin and Paul E. Berry Woody or herbaceous vines; stems smooth, scabrous, or armed with spines; roots fibrous or tuberous. Leaves alternate or sometimes opposite, generally petiolate, commonly with a pair of tendrils arising from a short stipular flange or open sheath; blade with 3–9 primary veins arising from or near base of blade. Inflorescence pedunculate, axillary or terminal, in umbels or racemes of umbels. Flowers unisexual (the plants dioecious) or bisexual; perianth segments 6, distinct or shortly connate below, narrow, equal or nearly so; stamens (3–)6(–18), free, the anthers 2-celled; ovary superior, rarely inferior, 1- or 3-locular, with 1 or many ovules in each locule, the placentation axile or parietal; stigmas 3, in female flowers 0–6 staminodes present. Fruit a fleshy berry. Seeds 1–6.

Smilax 185

Widespread in tropical and subtropical regions, also well represented in parts of the northern temperate zones; 13 genera and ca. 330 species, 1 genus and 9 species in the flora area. 1. SMILAX L., Sp. Pl. 1028. 1753. Woody or herbaceous, dioecious vines. Leaves simple, alternate, petiolate, with stipules forming a sheath, the sheath usually terminating in a pair of tendrils; blade broadly cordate to lanceolate with 3–9 subparallel primary veins arising from or near base of blade, the lower order veins reticulate; margin entire, smooth or spiny. Inflorescence axillary, umbellate, or the umbels pseudoracemose on short axillary branchlets. Stamens 6. Ovary 3-locular, with 1 or 2 ovules in each locule, the placentation axile. Fruit a fleshy berry, orange, yellow, or black when mature. Tropical and temperate areas of both hemispheres, with most species concentrated in tropical Asia and the Neotropics; ca. 300 species, 16 in Venezuela, 9 of these in the flora area. Key to the Species of Smilax 1.

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

5.

6(1).

6.

7(6).

7.

Lateral shoots (not individual petioles) with 2 closely placed, overlapping scales at the base on adaxial side (see detail, Fig. 160, page 191) ................................................................................................................ 2 Lateral shoots (not individual petioles) with 1 scale (sometimes bifid) at the base on the adaxial side (see detail, Fig. 155, page 188) .............. 6 Upper branches usually very scabrous, without spines ... S. schomburgkiana Upper branches not usually scabrous, with or without spines ............... 3 Stem and midvein on lower side of leaf pubescent .................. S. triplinervia Stem and lower side of leaf glabrous ........................................................ 4 Mature leaf blade strongly cordate, 5–20 cm long × 8–20 cm wide; flowers usually many (30–70) per receptacle ................................. S. poeppigii Mature leaf blade oblong to ovate, 8–35 cm long × 4–10 cm wide; flowers < 30 per receptacle ................................................................................ 5 Mature leaf blade 15–35 cm long, membranous to subcoriaceous, dull to sublustrous, with higher order venation evident on upper surface; with or without spines on stem .............................................. S. siphilitica Mature leaf blade 8–15 cm long, coriaceous, lustrous and plane (flat, with higher order venation inconspicuous on upper surface); no spines on stem ................................................................................... S. pittieriana Basal internode of branches longer than more distal internodes (2–16 cm long); peduncle shorter than or equal to petiole length; 1200–2300 m ................................................................................................... S. staminea Basal internode of branches shorter than more distal internodes (0.5– 3 cm long); peduncle length longer than petiole or not; 150–1300 (–1650) m ............................................................................................... 7 Pedicels longer than peduncle; leaf without spines, blade ovate to elliptic, often drying green, apex acute; mainly in Delta Amacuro and Bolívar, rarely in northern Amazonas ................................. S. cumanensis Pedicels shorter than peduncle; leaf usually with spines, blade lanceolate to ovate, drying reddish or green, apex mucronate or acute; mainly in Amazonas, rarely in southeastern Bolívar ......................... 8

186

8(7).

8.

S MILACACEAE

Leaf blade lanceolate, apex mucronate or acute, not oblique, dries reddish or green; margin, midvein, and other primary veins on lower surface with spines ........................................................................ S. lappacea Leaf blade ovate to rarely lanceolate, apex acute, often oblique, dries reddish; margin and midvein on lower surface often with spines, but other primary veins without spines ................................... S. maypurensis

Smilax cumanensis Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 783. 1806. —Bejuco de corona, Bejuco corona e cristo, Caijiro, Hala por atrás, Këmëhkë tuku’ya këmën (Panare). Woody vine; stem terete; leaf blade ovate to rarely lanceolate, base acute to rounded, apex acute; mature fruit dark purple or black. Secondary vegetation, low forests, 0– 600 m; Delta Amacuro (Caño Atoiba tributary of Boca Araguao, Caño Simoina, between La Margarita and Puerta Miranda Pedernales, Sacupana), Bolívar (Corozal, 3 km northeast of Hato Las Nieves, La Paragua, island in Lago Guri, 64 km southeast of Pijiguaos, Río Parguaza, San Martín de Turumbán, San Pedro de las Dos Bocas), Amazonas (Isla Ráton, 10 km south of Puerto Ayacucho, San Juan de Manapiare). Widespread elsewhere in Venezuela; Costa Rica, Colombia, Tobago, Guyana, Suriname, French Guiana. ◆Fig. 156. Smilax lappacea Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 777. 1806. Woody vine; stem armed; leaf blade lanceolate, apex mucronate or acute, with spines on margin and primary veins of lower surface; flowers greenish. Low climber in scrub and savanna, 50–300 m; Amazonas (Caño Coro Coro, Puerto Ayacucho, Santa Bárbara del Orinoco). Distrito Federal. ◆Fig. 157. Smilax lappacea seems to intergrade with S. maypurensis with respect to leaf morphology. Smilax maypurensis Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 776. 1806. Creeping or low-climbing woody vine, stem often armed; leaf blade ovate to rarely lanceolate, apex acute and often oblique, base sometimes subcordate, usually with spines on margin and midvein of lower surface; flowers greenish to white; fruit green to

blackish blue. In shrubs and low trees of savannas, also open rocky areas, 50–900 (–1400) m; Bolívar (Río Parguaza), Amazonas (Campamento Asisa, Caño Iguapo, Cerro Duida, between Cerro Morrocoy al Sur and Serranía Colmena, La Esmeralda, Puerto Ayacucho, Río Cunucunuma, Río Pasimoni, Río Sipapo, Río Yureba, Samariapo, San Fernando de Atabapo, Simarawochi, Tobogán de la Selva). Adjacent areas of Colombia. This species seems to intergrade with Smilax lappacea with respect to leaf morphology. Specimens in this group have also been labeled as S. syringoides Griseb. (type from southern Brazil) or S. rufescens Griseb. (type from eastern Brazil), and synonymy has not been resolved. Smilax pittieriana Steyerm., Fieldiana, Bot. 28: 155. 1951. Smilax auraimensis Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 472. 1966. Smilax chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 440. 1967. Woody vine, without spines; leaf blade coriaceous, the upper surface shiny, plane, veins obscure; mature fruit orange. Riparian, scrub, secondary, and montane forests, savannas, thickets, 100–1300(–1700) m; Bolívar (Altiplanicie de Nuria, Amaruaytepui, Apacará-tepui along Río Apacará, Cerro Altamira, Cerro Guaiquinima, Cerro Pitón, El Paují, Gran Sabana, Guayaraca, 7 km northeast of Ikabarú, between Ptaritepui and Sororopán-tepui, Río Oris, Río Tirica above Techiné-merú, between San Ignacio and San Francisco de Yuruaní, Sierra Auraima, Uaipán-tepui), Amazonas (Cerro Aracamuni, Cerro Yutajé, San Carlos de Río Negro, Sierra Parima). Endemic. ◆Fig. 158. Steyermark’s protologue of Smilax pittieriana was an unfortunate confusion of

Smilax 187

two different specimens belonging to two distinct species of Smilax. The holotype, Steyermark 60251 (F), has racemosely arranged male inflorescences, two axillary scales at the base of the branches, and ovate, coriaceous leaf blades with a rounded base and a plane, glossy upper surface. The specimen that Steyermark cited as the “cotype”, Steyermark 60251a (F), actually has no nomenclatural validity as a type. It has remnants of simply umbellate female inflorescences, a single axillary scale at the base of the branches, and narrowly elliptic, membranous leaf blades with an acute base and a reticulately-raised veined, less glossy upper surface; as such, it corresponds to what we are recognizing here as Smilax staminea. Steyermark’s later recognition of Smilax auraimensis and S. chimantensis also stems from his misinterpretation of the type material of S. pittieriana. In his discussions of these species, he was clearly basing his concept of S. pittieriana mainly on the “co-type” specimen, or S. staminea. Some specimens of Smilax pittieriana from the flora area were previously determined as Smilax guianensis Vitman, but this is a species of uncertain affinity that was based on an illustation by Charles Plumier of a plant apparently from Martinique and then attributed to specimens from French Guiana possibly seen by Vitman. Smilax poeppigii Kunth, Enum. Pl. 5: 192. 1850. —Bejuco de corona. Woody, climbing, thick-stemmed vine, lower stems usually spinose; mature leaves strongly cordate, apex acute to acuminate; mature peduncle often flattened, receptacle reniform. Evergreen lowland to lower montane forests, 100–300(–1000) m; Bolívar (northeast of El Manteco, 7 km north of Las Claritas, middle Río Botanamo, Río Paragua between La Paragua and Salto de Auraima), Amazonas (Cerro Aratitiyope, Río Coro Coro, lower Río Guasacavi 1 km upstream from Gallineta, Río Ocamo, 16 km northeast of San Carlos de Río Negro, 12 km south of San Fernando de Atabapo). Anzoátegui, Barinas, Falcón, Trujillo; Peru. ◆Fig. 159. The type specimen of Smilax poeppigii is from Peru, and we consider it to be the clos-

est matching name for specimens from the flora area. The type specimen is similar to the flora area material in peduncle and leaf shape (though specimens we viewed can have strongly cordate mature leaves), but differs in a shorter mature petiole (20 versus 40–50 mm). The collection Berry et al. 6239 (MO) has spines emerging from a tuberous base, and unusual new thick cormose structures with aerial roots about to sprout aerial shoots. The upper stems do not appear to be spiny, which agrees with original description. If specimens from the flora area are not S. poeppigii, then they may be an undescribed species. Liesner 8985 (VEN) has ca. 70 green flowers per receptacle, and the label notes the presence of some spines on the stem. In certain cases, it is difficult to differentiate this species from S. siphilitica, due to similarities in juvenile leaf morphology. Smilax schomburgkiana Kunth, Enum. Pl. 5: 187. 1850. —Tu-pata-yén (Arekuna). Woody vine, stem very scabrous; leaf blade lanceolate, base rounded to acute, apex narrowly acute to acuminate, conspicuously 3-nerved; fruit turning orange at maturity, black when dried. Evergreen lowland to lower montane forests, often secondary vegetation, 50–1200 m; Delta Amacuro (Río Amacuro above San Victor), Bolívar (Amaruay-tepui, 88 and 119 km south of El Dorado, east of Cerro El Picacho, El Pauji, La Escalera, Río Canaracuni, Río Erebato, Río Paragua, Río Torono, Salto Pará), Amazonas (Cerro Aratitiyope, upper Río Baría, Río Cunucunuma, Río Corocoro, Río Mawarimuna, Simarawochi). Guyana, Suriname, French Guiana, Peru, northern Brazil. ◆Fig. 160. Smilax siphilitica Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 780. 1806. —Corona guaica. Smilax duidae Steyerm., Fieldiana, Bot. 28: 154. 1951. High-climbing woody vine or scandent shrub; leaf blade generally lanceolate with rounded to acute base, occasionally ovate with subcordate base; peduncle often flattened when mature; receptacle reniform; inflorescence a raceme of umbels; flowers

188

S MILACACEAE

Fig. 155. Smilax staminea

Fig. 156. Smilax cumanensis

Fig. 157. Smilax lappacea

Smilax 189

Fig. 158. Smilax pittieriana

190

S MILACACEAE

Fig. 159. Smilax poeppigii

Smilax 191

Fig. 160. Smilax schomburgkiana

192

S MILACACEAE

Fig. 161. Smilax siphilitica

Smilax 193

white to pale green; fruit orange to blue gray when mature. Gallery forests, evergreen lowland to lower montane forests, seasonally flooded to nonflooded, 0–1300 m; Delta Amacuro (Caño Capurito, Río Ibaruma, San Victor, Sierra Imataca), Bolívar (Amaruaytepui, north of base of Cerro Guaiquinima, Río Caroní, Río Samay, Río Venamo, 5 km south of San Ignacio de Yuruaní, Serranía de Maigualida), Amazonas (Cerro Duida, Cerro Huachamacari, Maroa, Río Casiquiare, Río Emoni, San Carlos de Río Negro, Serranía Batata, Simarowochi, Victorino). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 161. Specimens from the flora area have also been determined as Smilax aequatorialis (Griseb.) A. DC. (type from Brazil, originally a variety of S. siphilitica), S. febrifuga Kunth (type specimen from Peru), or S. pseudosyphilitica Kunth (type specimen from southern Brazil). Possible synonymy of this complex needs to be better resolved with further study. Smilax staminea Griseb. in Mart., Fl. Bras. 3(1): 11. 1842. Smilax lasseriana Steyerm., Fieldiana, Bot. 28: 154. 1951. Smilax jauaensis Steyerm. & Maguire, Mem. New York Bot. Gard. 23: 854. 1972. Smilax staminea f. obtusata Steyermark, Fieldana, Bot. 28: 156. 1951. Low-climbing woody vine, unarmed; leaf blade 5–7-veined, ovate to lanceolate, apex acute to acuminate, sometimes oblique; flowers white to pale green, fragrant, perianth segment 4–5 mm long. Rocky escarpments, open areas, shrublands, 900–2300 m; Bolívar (Amaruay-tepui, 12 km north of Aparamán-tepui, Auyán-tepui, Cerro Chirikayén, Cerro Guaiquinima, Cerro Guanay, Cerro Jaua, Cerro Marutaní, Cerro Venamo, Kamarkawarai-tepui, northeast of Luepa

between km 134 and km 150, 15 km westsouthwest of Karaurín-tepui, Macizo del Chimantá [Apacará-tepui], Ptari-tepui, Sororopán-tepui, Uaipán-tepui), Amazonas (Cerro Aracamuni, Cerro Asisa, Cerro Duida, Cerro Huachamacari, Cerro Parú, Cerro Yutajé, Sierra de la Neblina, Sierra Parima). Distrito Federal, Sucre; Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 155. Specimens from the flora area have also been determined as Smilax nitida Griseb. (type from southern Brazil). Outside of the flora area, specimens that may be referable to S. staminea have been found in the Andes and in the coastal ranges (Distrito Federal, Sucre). This material has often been determined as S. kunthii Killip & C.V. Morton (= S. floribunda Kunth, type from Ecuador), but we are not certain if these should be treated as a synonym of S. staminea (type specimen also from southern Brazil). Steyermark differentiated both Smilax jauaensis and S. lasseriana from S. staminea in having verruculose-tuberculate versus smooth stems, and S. lasseriana from S. staminea in leaf shape (ovate-oblong versus ovate-lanceolate, acuminate). Verruculose-tuberculate and smooth-stemmed specimens are sympatric on Cerro Jaua, where the type specimen of S. jauaensis was collected, and these morphological differences are judged to be minor variations of one species. Smilax triplinervia Humb. & Bonpl. ex Willd., Sp. Pl. 4(2): 784. 1806. Woody, pubescent, unarmed vine, the lower surface with midvein pubescent. Habitat unknown, ca. 100 m; Amazonas (Río Atabapo). Endemic. This species is known only from the type specimen, and no other pubescent Smilax specimens from the flora area have been found.

194

S OLANACEAE

SOLANACEAE by William G. D’Arcy, Carmen Benítez de Rojas, and Michael H. Nee Herbs, shrubs, vines, or trees, sometimes epiphytic, glabrous or with various kinds of pubescence, sometimes armed. Leaves alternate, sometimes geminate, simple, entire, lobed, or pinnate; stipules absent; petioles mostly present. Inflorescences terminal but mostly appearing axillary, lateral on the stems, or at a branch fork, cymose, variously clustered, sometimes reduced to a single flower; bracts or bracteoles sometimes present. Flowers mostly bisexual, actinomorphic or zygomorphic, mostly 4- or 5-merous. Calyx cupular or tubular, lobed or nearly divided, often accrescent in fruit; corolla sympetalous, rotate, campanulate, tubular, funnelform, or salverform, lobed, in bud valvate, imbricate, or quincuncial. Stamens alternate with the corolla lobes, inserted in the corolla tube, sometimes partly fused; anthers dehiscing longitudinally or by terminal pores, sometimes one or more reduced to staminodes; disk sometimes present. Pistil 1; carpels 2; locules 1–several by false septa, the placentation axile; ovules 1–many in each locule; style solitary. Fruit a berry or capsule. Seeds several to many; embryo straight and central or curved around the periphery of the testa; endosperm abundant. Cosmopolitan (especially abundant in the Neotropics); ca. 95 genera and ca. 2300 species, 16 genera and 79 species in the flora area. Key to the Genera of Solanaceae by William G. D’Arcy 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(3). 5. 6(2). 6. 7(6).

Floral tube long (the flowers longer than wide); fruit a berry or capsule ................................................................................................................ 2 Floral tube short (the flowers shorter than wide); fruit a berry ........... 11 Herbs, erect or clambering; fruit a capsule .............................................. 3 Woody plants, terrestrial or epiphytic; fruit a berry or leathery capsule ................................................................................................................ 6 Diminutive herbs or slender climbers; fruit < 7 mm long; corolla tube < 5 mm wide; calyx < 5 mm long; fertile stamens 2 or 4 ..................... 4 Large-leaved robust herbs; fruit > 7 mm long; corolla tube > 5 mm wide; calyx > 8 mm long; fertile stamens 5 ................................................... 5 Fruit globose, smooth; seeds many ........................................ 13. Schwenckia Fruit falcate-cylindrical, rugulose; seed one ....................... 10. Melananthus Leaves clasping the stem; capsule smooth; seeds < 1 mm across; flowers pinkish .................................................................................... 11. Nicotiana Leaves petiolate; capsule tuberculate or spiny; seeds > 1 mm across; flowers white (violet-blue) ........................................................... 5. Datura Corolla narrowly tubular, < 35 mm long; fruit < 10 mm long ...... 3. Cestrum Corolla apically campanulate, funnelform, or salverform, mostly > 35 mm long; fruit mostly > 10 mm long ........................................................... 7 Terrestrial shrubs; corolla salverform or funnelform .............................. 8

S OLANACEAE 195

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

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

20. 21(19). 21. 22(21). 22.

Woody, high-climbing epiphytes; corolla limb cupular ............................ 9 Corolla salverform, < 5 cm long ................................................. 1. Brunfelsia Corolla funnelform, > 6 cm long ...................................................... 5. Datura Corolla showy, yellow, > 10 cm long; calyx > 7 cm long ............. 14. Solandra Corolla greenish or purplish, < 10 cm long; calyx < 7 cm long ............. 10 Anthers basifixed; filaments elongate; corolla > 3 cm long; berry > 1 cm long; calyx lobed < halfway ......................................................... 9. Markea Anthers dorsifixed; stamens subsessile; corolla < 2 cm long; berry < 1 cm long; calyx lobed > halfway .......................................... 7. Hawkesiophyton Corolla salverform, > 3 × 3 cm; fruit a leathery berry; seeds angular or prismatic; calyx > 14 mm long ............................................... 1. Brunfelsia Corolla campanulate, rotate, or funnelform, < 3 cm long, mostly < 25 mm wide; fruit mostly juicy; seeds compressed, discoid or lenticular; calyx < 14 mm long ....................................................................................... 12 Fruit entirely enclosed in the enlarged calyx; corolla yellow or violet .............................................................................................................. 13 Fruit not enclosed in the calyx, or at least mostly exposed; corolla color various, but not yellow ........................................................................ 14 Corolla violet, lobed 1/3–1/2 way; plants woody ................................. 6. Deprea Corolla yellow, undulate-lobed; plants herbaceous .................... 12. Physalis Climbing plants ....................................................................................... 15 Terrestrial herbs or shrubs ..................................................................... 18 Inflorescence 1- or 2-flowered ................................................................. 16 Inflorescence several- to many-flowered ................................................ 17 Calyx truncate with 10 teeth below the apex ............................ 8. Lycianthes Calyx with 5 apical lobes (teeth) ................................................ 15. Solanum Corolla campanulate; anthers dorsifixed, dehiscing longitudinally ....................................................................................... 7. Hawkesiophyton Corolla rotate; anthers basifixed, dehiscing by terminal pores ........... ................................................................................................. 15. Solanum Plants armed with spines; indumentum of stellate hairs ......... 15. Solanum Plants unarmed; indumentum of simple hairs, stellate hairs, or absent .............................................................................................................. 19 Large plants, > 1.5 m tall ........................................................................ 20 Small plants, < 1.5 m tall ........................................................................ 21 Fruits < 10 mm across; anthers with the dorsal area undifferentiated from the rest of the anther; leaf bases various, but not truncate or cordate; leaves copiously stellate-pubescent .............................. 15. Solanum Fruits > 10 mm across; anthers with an enlarged dorsal connective; leaf bases cordate; leaves with simple hairs or glabrate ....... 4. Cyphomandra Fruits black, yellow, violet, or greenish; anthers dehiscent by terminal pores ........................................................................................ 15. Solanum Fruits red; anthers longitudinally dehiscent ......................................... 22 Fruits juicy; inflorescences of many-flowered subsessile racemes or fascicles; anthers yellow, minutely apiculate ......................... 16. Witheringia Fruits dry or fleshy; inflorescences of solitary or rarely paired flowers; anthers bluish, not apiculate .................................................. 2. Capsicum

196

S OLANACEAE

1. BRUNFELSIA L., Sp. Pl. 191. 1753. by William G. D’Arcy Unarmed shrubs or trees. Indumentum sparse, of simple hairs. Leaves entire, solitary. Inflorescence of solitary or clustered terminal-appearing flowers, sometimes bracteate. Flowers slightly zygomorphyic. Calyx tubular to cupular, 5-lobed part way, somewhat accrescent and often tardily splitting to the base; corolla showy, white, blue, or purplish, often fading to white or yellow, salverform, slightly zygomorphic, the lobes rounded; stamens 5 with one or more reduced to staminodes or absent, inserted on the corolla tube; anthers included, dehiscing longitudinally. Ovary 2-locular basally, apically 1-locular; stigma capitate, sometimes somewhat enlarged. Fruit a leathery berry half-enclosed in the accrescent calyx. Seeds relatively large, prismatic; embryo straight or slightly curved. Neotropics; ca. 50 species, ca. 5 in Venezuela, 2 of these in the flora area. In addition to the two native species keyed below, others that are not native are grown ornamentally in cultivation. Key to the Species of Brunfelsia 1.

1.

Pedicels 8–9 mm long; calyx mostly 15–21 mm long; corolla tube mostly > 30 cm long, the exserted portion mostly longer than the calyx; anthers 1.5–2 mm diameter; seeds 6–10, 6–9 mm long ........... B. imatacana Pedicels 1–6 mm long; calyx 8–20 mm long; corolla tube mostly < 25 cm long, the exserted portion mostly shorter than the calyx; anthers ca. 1 mm diameter; seeds 10–20, 3–5 mm long ............................. B. uniflora

Fig. 162. Brunfelsia imatacana

Capsicum 197

Brunfelsia imatacana Plowman, Fieldiana, Bot. n.s. 8: 4. 1981. —Jazmín del monte, Juan de la calle. Shrub or tree to 5 m tall; flowers blue or white. Evergreen lowland forests, 100–500 m; Bolívar (northwestern Río Cuyuní basin, Serranía de Imataca). Endemic. ◆Fig. 162. Brunfelsia pauciflora (Cham. & Schldl.) Benth. of southeastern Brazil is apparently the closest relative of B. imatacana. Brunfelsia uniflora (Pohl) D. Don, Edinburgh New Philos. J. 85. 1829.

—Franciscea uniflora Pohl, Pl. Bras. Icon. Descr. 1: 2, t. 1. 1826. Shrub or tree to 2 m tall; flowers blue or white. Evergreen lowland and secondary forests, also cultivated in gardens, 50–200 m; Bolívar (Upata). Guárico, Nueva Esparta; Trinidad, Guyana, Brazil, Bolivia, Argentina. The inclusion of Brunfelsia uniflora is based on a specimen [Otto 1003 (W)] that was not seen by T. Plowman but was cited by him (T. Plowman. A revision of the South American species of Brunfelsia (Solanaceae). Fieldiana, Bot., n.s. 39: 1–135. 1998).

2. CAPSICUM L., Sp. Pl. 188. 1753. by Michael H. Nee Herbs or weak shrubs. Indumentum of simple hairs. Leaves solitary or paired, entire. Inflorescences of solitary or paired flowers in the leaf axils or at a branch fork; pedicels ebracteolate. Flowers actinomorphic, 5-merous. Calyx truncate or slightly toothed, hardly accrescent; corolla campanulate to rotate, the lobes often ovate. Stamens 5, alike, inserted low in the corolla; anthers mostly stout, often bluish, dehiscing longitudinally. Ovary 2- or more locular, the placenta often enlarged and fleshy; style slender; stigma small. Fruit a fleshy, bright-colored berry, often spicy-hot with capsacain. Seeds many, compressed; embryo strongly curved. Tropical and subtropical America; ca. 25 species, 3 wild in Venezuela, 2 of these in the flora area. Several species and numerous cultivars are often cultivated in the flora area and worldwide for food, spices, or ornament. Key to the Species of Capsicum 1.

1.

2(1). 2.

Plants cultivated; fruits of various shapes, larger than 15 × 10 mm, held erect or pendulous, usually not readily deciduous when ripe .............................................................................................. Capsicum spp. Plants wild, weedy or sometimes deliberately planted; fruits ovoid or globose, 8–15 × 5–10 mm, held erect, readily deciduous when ripe ................................................................................................................ 2 Flowers and fruits evidently 1 per node; calyx cupular, smoothly curving, 2 mm long in fruit, thin, not ribbed when dry .......................... C. annuum Flowers and fruits often appearing as if 2–several per node; calyx angular in longitudinal section, 2.5–3 mm long in fruit, thickish and somewhat ribbed when dry ............................................................. C. frutescens

Capsicum annuum L., Sp. Pl. 188. 1753. Herb or subshrub 0.5–2 m tall. Native to the southern U.S.A., Mexico, Central America, Colombia, Venezuela, and Ecua-

dor, but domesticated worldwide; 2 varieties, both in Venezuela, 1 of these in the flora area.

198

S OLANACEAE

Fig. 163. Capsicum annuum var. glabriusculum

C. annuum var. glabriusculum (Dunal) Heiser & Pickersgill, Baileya 19: 156. 1975. —Capsicum hispidum var. glabruiusculum Dunal in A. DC., Prodr. 13: 420. 1852. Corollas white; fruit ovoid or globose, 8– 10 × 5–8 mm, red-orange, very hot. Weedy areas, especially around settlements and cultivated areas, sea level to 500 m; widespread thoughout the flora area. Widespread in the northern half of Venezuela; southernmost U.S.A., Mexico, Central America, Colombia, Ecuador. ◆Fig. 163. Variety glabriusculum gave rise to var. annuum, which is the most commonly cultivated taxon in temperate areas and unlikely to grow well in the flora area. Capsicum frutescens L., Sp. Pl. 189. 1753. Herb or subshrub 0.5–2 m tall; corollas

yellow-green; fruit ovoid, 2.5–3 mm long, red-orange, very hot. Weedy areas, especially around settlements, cultivated areas and recent clearings, widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread in northern half of Venezulea; southern Mexico, Central America, West Indies, tropical South America south to Brazil, Bolivia, and Paraguay. Capsicum spp. There are four domesticated species of Capsicum that are characterized by a wide range of fruit sizes, shapes, colors, and degree of “hotness”. The most common one in cultivation in the Venezuelan Guayana is Capsicum chinense Jacq., which was derived from C. frutescens L.; these two intergrade. Other domesticated species are less likely but possible.

3. CESTRUM L., Sp. Pl. 191. 1753; Gen. Pl. ed. 5. 88. 1754. by William G. D’Arcy and Carmen Benítez de Rojas Unarmed trees or shrubs; indumentum of simple or branched hairs. Leaves solitary or paired, simple, entire, petiolate. Inflorescences appearing terminal on the branches or in the leaf axils, mostly several- to many-flowered cymose panicles,

Cestrum 199

sometimes bracteate, mostly bracteolate. Flowers actinomorphic, fragrant. Calyx tubular or campanulate, shortly 4-lobed, persistent and sometimes somewhat accrescent in fruit; corolla tubular with 5 short lobes, sometimes characteristically pubescent inside. Stamens 5, mostly included, inserted on the corolla tube, sometimes geniculate, barbate, or dentate near the point of insertion; anthers alike, dehiscing longitudinally. Ovary sessile or stipitate, 2-locular; ovules few to many; stigma small, sometimes slightly exserted. Fruit a small juicy or fleshy berry, fewto many-seeded. Seeds narrowly ovate, smooth; embryo straight or slightly bent. Neotropics; ca. 175 species, 39 in Venezuela, 11 of these in the flora area. In addition to the species treated here, collections were seen of other species, but the material lacks flowers or is otherwise insufficient either for determination or for description as new species. Key to the Species of Cestrum 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.

Corolla tube pubescent externally. ........................................... C. strigilatum Corolla tube glabrous externally .............................................................. 2 Largest leaves mostly < 3(–4) cm wide, the margins mostly subrevolute; twigs conspicuously tomentulose with ascending-curved hairs ......... 3 Largest leaves broader, mostly > (3–)4 cm long, the margins mostly plane; twigs mostly inconspicuously pubescent .................................. 7 Leaves rounded, mostly < 5(–7) cm long, crowded on the branches, basally truncate or cordate, apically obtuse; filaments free for > 3 mm ................................................................................................ C. tubulosum Leaves ovate to lanceolate or linear, mostly > (5–)7 cm long, spaced on the branches, apically acuminate or acute; filaments free for < 2 mm ................................................................................................................ 4 Leaves linear, < 15 mm wide; flowering calyx < 3 mm long; corolla pubescent at the base within .......................................................... C. neblinense Leaves ovate to lanceolate, > 20 mm wide; flowering calyx > 3 mm long; corolla glabrous ..................................................................................... 5 Mature leaves puberulent beneath, membranous; corolla lobes < 3 mm long; fruits sessile; filaments subsessile ........................... C. alternifolium Mature leaves glabrous, coriaceous; corolla lobes > 3 mm long; fruits on short but evident pedicels; filaments free for > 1 mm ........................ 6 Twigs with a conspicuous yellowish or reddish tomentum; leaf margins somewhat revolute ............................................................ C. subuniflorum Twigs glabrate; leaf margins plane ........................................ C. schulzianum Flowers and fruits in short inflorescences distributed along the stem, mostly not exceeding the petioles; leaves often broadest above the middle ..................................................................................................... 8 Flowers and fruits in inflorescences terminal on branches or elongate, much exceeding the petioles; leaves broadest below the middle ...... 11 Mature leaves membranous, the lower surface puberulent; veins ca. 6 on each side .................................................................................. C. latifolium Mature leaves coriaceous to membranous, the lower surface glabrous; veins 5–16 on each side ......................................................................... 9 Petioles, midribs on lower surface of leaves, and fruits mostly drying black .............................................................................. C. schlechtendahlii Petioles, leaf veins, and fruits drying greenish or brownish ................ 10

200

S OLANACEAE

10(9). Leaves coriaceous; lateral veins 4–8 per side, > 1 cm apart ..... C. glabrescens 10. Leaves membranous; lateral veins numerous per side, < 1 cm apart ............................................................................................... C. racemosum 11(7). Lower surface of mature leaves puberulent; filaments free for < 1 mm ............................................................................................ C. alternifolium 11. Lower surface of mature leaves glabrous; filaments free for > 1 mm .............................................................................................................. 12 12(11). Calyx > 4 mm long; leaves mostly with > 12 lateral veins ..... C. schulzianum 12. Calyx < 4 mm long; leaves mostly with < 12 lateral veins .................... 13 13(12). Inflorescences with several rhomboid or deltoid subfoliaceous bracts, also with minute linear bractlets; inflorescence flexuous and pendulous; upper surface of leaves mostly puberulent, the blade membranous; plants scandent; filaments free for < 2 mm ................... C. reflexum 13. Inflorescences with minute linear bractlets or none; inflorescence erect or stiff; mature leaves glabrous, coriaceous; plants erect; filaments free for > 2.5 mm .................................................................... C. tubulosum Cestrum alternifolium (Jacq.) O.E. Schulz in Urb., Symb. Antill. 6: 270. 1909; 7: 555. 1911. —Ixora alternifolia Jacq., Enum. Syst. Pl. 12. 1760. —Tepe camino. Cestrum alternifolium var. mitanthum O.E. Schulz in Urb., Symb. Antill. 6: 273. 1909. Shrub to 3 m tall; flowers greenish white. Evergreen lowland to lower montane forests, streamsides, 100–600 m; northern Bolívar. Aragua, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Zulia; Central America, Antilles, Colombia. Cestrum glabrescens (C.V. Morton) Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 463. 1967. —Cestrum tenuiflorum var. glabrescens C.V. Morton, Bull. Torrey Bot. Club 58: 465. 1931. Shrub to 2 m tall. Montane forests, rock outcrops on tepui slopes and summits, 1200–1700 m; Bolívar (Auyán-tepui, La Escalera, Macizo del Chimantá [Toronótepui]), Amazonas (Cerro Duida, Cerro Marahuaka). Endemic. ◆Fig. 165. Cestrum latifolium Lam., Tabl. Encycl. 2: 5. 1793 [1794]. —Clavito, Huele de noche, Rabo pelado. Cestrum tenuiflorum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 61. 1818. —Cestrum latifolium var. tenuiflorum

(Kunth) O.E. Schulz in Urb., Symb. Antill. 6: 270. 1909. Cestrum oliganthum Dunal in A. DC., Prodr. 13: 634. 1852. Cestrum oliganthum var. latifolium Dunal in A. DC., Prodr. 13: 634. 1852. Cestrum prieurei Dunal in A. DC., Prodr. 13: 635. 1852. Shrub to 5 m tall with sprawling branches; flowers greenish white. Secondary vegetation, 50–900 m; Delta Amacuro (Caño del Guinipa, Caño Orocoima, Serranía de Imataca upstream from San Victor past Río Matanaima to Salto de Quebradero, road from Tucupita to Las Mulas), northern Bolívar, Amazonas (Cerro Duida, Sierra Parima). Widespread elsewhere in Venezuela; Panama, Jamaica, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 166. Cestrum neblinense D’Arcy & Benítez, Ann. Missouri Bot. Gard. 77: 206. 1990. Shrub 1–2 m tall; flowers small, tubular. Lower to upper montane cloud forests, partly flooded riparian forests, 700–2200 m; Amazonas (uppermost Río Yatúa, Sierra de la Neblina). Endemic. Cestrum racemosum Ruiz & Pav., Fl. Peruv. 2: 29, pl. 154. 1799. —Fruta de paloma, Uvita, Vara blanca. Cestrum grande Pittier, J. Wash. Acad. Sci. 22: 32. 1932. —Cestrum racemosum var. grande (Pittier) Francey, Candollea 6: 275. 1934.

Cestrum 201

Tree or shrub to 15 m tall. Montane forests, ca. 1500 m; Amazonas (Sierra de la Neblina). Anzoátegui, Aragua, Barinas, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Táchira, Yaracuy, Zulia; southern Mexico, Central America, Colombia, Ecuador, Peru, Bolivia. Cestrum reflexum Sendtn. in Mart., Fl. Bras. 10: 218. 1846. Scandent shrub to 2 m tall; flowers whitish. Disturbed areas, 200–300 m; Bolívar (El Palmar). Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 164. Cestrum schlechtendahlii G. Don, Gen. Hist. 4: 482. 1838. Cestrum acuminatissimum Dunal in A. DC., Prodr. 13: 627. 1852. Cestrum megalophyllum Dunal in A. DC., Prodr. 13: 638. 1852.

Cestrum baenitzii Lingelsh., Repert. Spec. Nov. Regni Veg. 7: 248. 1909. Shrub or tree to 4 m tall. Gallery forests, forests on talus slopes, 100–800 m; Bolívar (Amaruay-tepui), Amazonas (between Río Mawarinuma and Río Baría, Río Negro). Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Lara, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; southern Mexico, Central America, Lesser Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia. Cestrum schulzianum Francey, Candollea 6: 272. 1935. Shrub to 3 m tall. Evergreen lowland forests, Mauritia palm swamps, 100–200 m;

Fig. 164. Cestrum reflexum

202

S OLANACEAE

Fig. 165. Cestrum glabrescens

Fig. 166. Cestrum latifolium

Cyphomandra

Amazonas (Río Casiquiare, Río Negro). Colombia, Brazil. Some of the plants under this name were treated by Benítez & D’Arcy (1998) under C. acuminatissimum Dunal, which is a synonym of C. schlechtendahlii. Cestrum strigilatum Ruiz & Pav., Fl. Peruv. 2: 29, pl. 156. 1799. Sprawling or climbing shrub or tree 2–4 m tall. Riparian forests, 200–300 m; Amazonas (Raudal Monserrat on upper Río Orinoco). Aragua, Carabobo, Táchira; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Cestrum subuniflorum Dunal in A. DC. Prodr. 13: 654. 1852. Shrub to 2 m tall. Montane thickets, streamsides, rock outcrops on tepui slopes, 1200–1500 m; Amazonas (Cerro Sipapo, Cerro Yutajé). Brazil.

203

The name Cestrum subuniflorum is used for this taxon with hesitation, as only photographs of the type specimen were available for the determination. This species and Cestrum tubulosum are closely related judging from features of the pubescence and the flowers, but the shape and size of the leaves are strikingly distinct. Cestrum tubulosum Sendtn. in Mart., Fl. Bras. 10: 207. 1849. Shrub to 1 m tall. Stream beds and gallery forests in sandstone or igneous areas, 100–300 m; Bolívar (Río Parguaza), Amazonas (Cerro Marahuaka, Río Coro Coro, Río Manapiare, lower Río Ventuari, San Carlos de Río Negro to Solano road). Brazil, Paraguay. Cestrum tubulosum is very similar to C. schulzianum. The type is from Brazil and may be different from the plants treated here.

4. CYPHOMANDRA Mart. ex Sendtn., Flora 28: 162. 1845. by William G. D’Arcy Unarmed shrubs or trees, often foetid, glabrous to pubescent with simple or dendritic hairs. Leaves solitary or paired, simple or compound, entire, lobed, or pinnate, often basally cordate, sometimes dimorphic; petioles sometimes clasping. Inflorescences terminal, appearing at dichotomies of the stems, simple or branched racemes, sometimes secund or circinnate; pedicels basally articulated. Flowers actinomorphic, mostly 5-merous. Calyx campanulate, the lobes obtuse or obsolete; corollas shallowly or deeply lobed, often with interpetalar membrane. Filaments free or connate; anthers oblong or attenuate, dorsifixed with an enlarged dorsal connective, opening by terminal pores and sometimes longitudinally. Ovary with enlarged placentas and many ovules. Fruit a fleshy or juicy berry, sometimes large. Seeds many, compressed; embryo strongly curved. Southern Mexico, Central America, Colombia, Venezuela, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, cultivated elsewhere; ca. 40 species, 3 in Venezuela, 1 of these in the flora area. This genus is now generally merged into Solanum. Cyphomandra tobagensis Sandwith, Kew Bull. 9: 370. 1938. —Solanum tobagense (Sandwith) Bohs, Taxon 44: 586. 1995. Cyphomandra bolivarensis Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 441. 1966.

Foetid shrub or small tree; flowers pendant, greenish or purplish. Evergreen lowland to lower montane forest edges, 500– 1400 m; Bolívar (southern Gran Sabana, La Danta, Guy-tepui near Perai-tepui). Sucre; Tobago, Guyana. ◆Fig. 167.

204

S OLANACEAE

Fig. 167. Cyphomandra tobagensis

Datura 205

5. DATURA L., Sp. Pl. 179. 1753; Gen. Pl. ed. 5. 83. 1754. Brugmansia Pers., Syn. Pl. 1: 216. 1805. by William G. D’Arcy Unarmed herbs, shrubs, or small trees, sometimes foetid, pubescence mostly of simple hairs. Leaves paired or solitary, often large, simple, entire, repand, lobed or toothed; petiolate. Inflorescences solitary flowers appearing axillary or at dichotomies of the stems, sometimes aggregated, sometimes bracteate; pedicels erect or pendulous. Flowers large, often showy, actinomorphic. Calyx tubular, sometimes inflated or angled, sometimes with circumscissile dehiscence; corolla funnelform to tubular, mostly white, purplish, pinkish, or red, flaccid, the apex sinuate with 5 or 10 lobes. Stamens equal; filaments sometimes flattened and pubescent near insertion; anthers oblong to linear, basifixed, coherent or free, dehiscing longitudinally. Ovary conical, 2- or 4-locular; ovules numerous. Fruit a dry or leathery capsule, ovoid to linear, sometimes apically dehiscent, sometimes covered with tubercules or spines. Seeds large, numerous; embryo curved. Tropical and subtropical America, but probably not native to Venezuela; ca. 20 species, 8 in Venezuela, 1 of these in the flora area. The Datura species that are woody are often separated into the genus Brugmansia; in the flora area they are known only from cultivation, and only

Fig. 168. Datura inoxia

206

S OLANACEAE

Datura suaveolens Willd. has been documented. These species tend to have elongate smooth fruits with only two locules, somewhat distinctive corky seeds, pendulous flowers, and are native to South America, whereas the species in Datura sensu strictu tend to have globose, 4-locular, ornamented fruits, erect flowers, and are native to Mexico and western South America. Some species are cultivated for ornament and others are weeds of disturbed areas. Datura inoxia Mill., Gard. Dict. ed. 8. 1768. Annual herb to 1.5 m tall; flowers large, white, erect. Disturbed areas, near sea level to 300 m; Bolívar (Río Orinoco). Widely dis-

tributed in coastal lowland parts of Venezuela. Probably native to Mexico, naturalized in many tropical countries of both Old and New Worlds. ◆Fig. 168.

6. DEPREA Raf., Sylva Tellur. 57. 1838. by William G. D’Arcy Unarmed herbs or shrubs with simple hairs. Leaves solitary or paired, subentire, petiolate. Inflorescences appearing axillary or fasciculate on the stem, 1–many-flowered, the peduncle reduced. Flowers actinomorphic, small, 5-merous. Calyx campanulate, short-lobed; corolla campanulate, 5-lobed, greenish or purplish. Stamens equal, inserted near the top of the corolla tube; anthers oblong, dehiscing longitudinally. Fruit a juicy berry enveloped by the bladdery accrescent calyx. Seeds many, compressed; embryo strongly curved. Neotropics; ca. 9 species, 2 in Venezuela, 1 of these perhaps in the flora area. Deprea orinocensis (Kunth) Raf., Sylva Tellur. 57. 1838. —Physalis orinocensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 12. 1818. Weak shrub to 1.5 m tall, with simple hairs; flowers small, purplish; fruits bladder-enclosed. Probably riparian vegetation between 50 and 100 m; Bolívar (Río Orinoco). Mérida, Táchira; Costa Rica, Colombia, Ecuador. ◆Fig. 169. No modern collections of Deprea orinocensis are known from the Guayana region, but it is included on the basis of the type collection, Bonpland s.n. (P), supposedly made in the Río Orinoco early in the 19th century, although the habitat is very different from the cloud forests inhabited in the remainder of the range.

Fig. 169. Deprea orinocensis

Hawkesiophyton 207

7. HAWKESIOPHYTON Hunz., Kurtziana 10: 39. 1977. by Carmen Benítez de Rojas Epiphytic shrubs associated with ants. Leaves subopposite, ovate, apically acuminate, basally cuneate, the lower surface minutely punctate. Inflorescences short racemes with thickened axis and scars from fallen pedicels. Flowers actinomorphic, small, yellowish or greenish, 5-merous. Calyx deeply 5-parted; corolla tubular-campanulate, shallowly 5-lobed. Stamens inserted at the top of the corolla tube; filaments shorter than the anthers; anthers dorsifixed, dehiscing longitudinally. Fruit a berry, the persistent calyx deeply lobed. Costa Rica, Panama, Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil; 3 or 4 species, 1 in Venezuela. The genus Hawkesiophyton is often united with Markea. Hawkesiophyton ulei (Dammer) Hunz., Kurtziana 10: 40. 1977. —Ectozoma ulei Dammer, Bot. Jahrb. Syst. 37: 170. 1905. Markea panamensis Standl., J. Arnold Arbor. 11: 127. 1930. —Hawkesiophyton panamense (Standl.) Hunz., Kurtziana 10: 40. 1977.

Woody hemiepiphytic vine; flowers small (7–10 mm), greenish, in short racemes. Evergreen lowland forests, riversides, 100–300 m; Amazonas (Río Cataniapo, Río Mawarinuma). Costa Rica, Panama, Colombia, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 170.

Fig. 170. Hawkesiophyton ulei

208

S OLANACEAE

8. LYCIANTHES (Dunal) Hassler, Annuaire Conserv. Jard. Bot. Genève 20: 173. 1917, nom cons. —Solanum sect. Pachystemonum subsect. Lycianthes Dunal in A. DC., Prodr. 13: 161. 1852. by William G. D’Arcy Mostly shrubs, often climbing, or vines, rarely herbs, unarmed, variously pubescent. Leaves simple, entire or nearly so, often paired. Inflorescences mostly solitary or few-fascicled in the leaf axils. Flowers 5-merous, often opening nocturnally. Calyx campanulate, not splitting at the sinuses in flower or fruit, apically truncate,

Fig. 171. Lycianthes pauciflora

Markea 209

10-ribbed, the ribs usually giving rise to teeth lateral on the calyx wall below the apex, the teeth mostly (0–9)10, often in 2 series; corolla rotate. Stamens often unequal, dehiscing by terminal pores. Ovary 2(–4)-locular; stigma small. Fruit a berry, not enclosed in the calyx. Seeds many, compressed; embryo strongly curved. Mexico, Central America, West Indies, Venezuela, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Asia, Pacific Islands; ca. 200 species, ca. 10 in Venezuela, 1 of these in the flora area. Lycianthes pauciflora (Vahl) Bitter, Abh. Naturwiss. Vereine Bremen 24: 341. 1919. —Solanum pauciflorum Vahl, Eclog. Amer. 1: 20. 1796 [1797]. Lycianthes guianensis (Dunal) Bitter, Abh. Naturwiss. Vereine Bremen 24: 347. 1919. —Solanum guianense Dunal in A. DC., Prodr. 13: 166. 1852.

Unarmed woody vine; flowers white. Evergreen lowland to montane forests, riversides, 100–1200 m; Bolívar (Altiplanicie de Nuria, Cerro Venamo, La Escalera), Amazonas (Río Mawarinuma, Sierra de la Neblina). Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 171.

9. MARKEA Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. by Carmen Benítez de Rojas Epiphytic vines, the branches hanging or creeping, glabrous or pubescent, generally fistulose and associated with ants. Leaves mostly solitary, entire, membranous, chartaceous or coriaceous, petiolate, often grouped in 3s on subterminal short-shoots. Inflorescences solitary or paired flowers, or racemose, often pendulous. Flowers actinomorphic or somewhat zygomorphic, sessile or pedicellate, red, greenish, whitish, or yellowish, often opening nocturnally, 5-merous. Calyx valvate, campanulate or tubular, 5-angled, with 4 or 5 lanceolate or acuminate lobes; corolla imbricate, bilabiate or funnelform, the tube somewhat enlarged, cylindrical and amplified above, with 5 narrow, suborbicular, subequal lobes. Stamens 5, included or slightly exserted; filaments slender, elongate, basally pilose, inserted on the corolla tube; anthers basifixed, ellipsoid or linear with parallel locules, erect, dehiscing longitudinally. Ovary subtended by an evident disk, 2-locular, multi-ovulate; style filiform. Fruit a capsule or berry, hardly juicy, oblong or cylindrical, pointed, 2-locular. Seeds numerous, imbricate, ellipsoidal, hardly compressed; embryo curved or almost straight. Neotropics; ca. 5–8 species, 5 in Venezuela, 4 of these in the flora area. Key to the Species of Markea 1. 1. 2(1). 2. 3(2). 3.

Flowers disposed in racemes or clusters; calyx with main venation conspicuous, dark brown .............................................................. M. longiflora Flowers solitary or paired; calyx venation inconspicuous ...................... 2 Leaves membranous, the margins ciliate; flowers with pedicels > 1.5 cm long ...................................................................................... M. formicarum Leaves coriaceous, eciliate; flowers sessile or subsessile ........................ 3 Leaf rugose, the lower surface bright green with pronounced venation; calyx > 3 cm long; corolla > 9 cm long .................................... M. reticulata Leaves smooth, both surfaces dark green, the venation obscure; calyx < 3 cm long; corolla < 7 cm long ........................................... M. sessiliflora

210

S OLANACEAE

Fig. 172. Markea longiflora

Fig. 173. Markea formicarum

Markea 211

Fig. 174. Markea reticulata

Fig. 175. Markea sessiliflora

212

S OLANACEAE

Markea formicarum Dammer, Bot. Jahrb. Syst. 37: 170. 1906. —Aparo, Batata de lagartilla, Bejuco de arrendajo, Macude. Epiphytic vine; flowers large (6–7 cm long), solitary, yellowish with purple corolla lobes. Evergreen lowland forests, riversides, 100–300 m; widespread in Amazonas. Amazonian Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 173. Markea longiflora Miers, Ann. Mag. Nat. Hist. ser. 2, 4: 186. 1849. Markea camponoti Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 55. 1915. Woody vine 4–10 m high; flowers large (7–8 cm long), greenish, in clusters. Evergreen lowland forests, riversides, 100–500 m; Bolívar (Sierra Pakaraima), Amazonas (upper Río Negro basin). Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 172. Markea reticulata Steyerm. & Maguire,

Mem. New York Bot. Gard. 17(1): 461. 1967. Woody, epiphytic vine; flowers large (9–11 cm long), solitary, greenish. Evergreen lowland to upper montane forests, riversides, 50–2200 m; Bolívar (Cerro Jaua, Macizo del Chimantá, Sororopán-tepui, Uaipán-tepui), Amazonas (Cerro Marahuaka, Río Castanho, Sierra Parima). Brazil (Amazonas). ◆Fig. 174. Markea reticulata is possibly synonymous with M. sessiliflora. Markea sessiliflora Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 56. 1915. Markea porphyrobaphes Sandwith, Bull. Misc. Inform. Kew 1931: 488. 1931. Woody epiphyte; flowers large (4–7 cm long), solitary, greenish. Evergreen lowland to lower montane forests, riversides, 50– 1100 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Guyana, Suriname, French Guiana. ◆Fig. 175.

10. MELANANTHUS Walp., Bot. Zeitung (Berlin) 8: 788. 1850. by Carmen Benítez de Rojas Herbs, subshrubs, or shrubs, pubescent with simple hairs. Leaves simple, entire, narrow, subsessile. Inflorescences racemose or espicate. Flowers diminutive, inconspicuous, zygomorphic. Calyx tubular, 5-lobed; corolla purplish, tubular, 5-lobed but sometimes appearing 15-lobed. Stamens 4, didynamous; anthers longitudinally dehiscent. Ovary 1-locular. Fruit a 1-seeded capsule (achene); embryo straight. Widespread in the Neotropics; ca. 5 species, 2 in Venezuela, 1 of these in the flora area. Melananthus ulei Carvalho, Sellowia 18: 53. 1966. Erect, multistemmed herb or subshrub 50–100 cm tall; flowers tiny. Thickets in gal-

lery forests on sandy and rocky soils, 400– 1300 m; Bolívar (Gran Sabana). Brazil (Piaui). ◆Fig. 176.

11. NICOTIANA L., Sp. Pl. 180. 1753. by William G. D’Arcy Erect herbs or shrubs, often glandular or viscid-pubescent, unarmed. Leaves entire, sessile or petiolate, sometimes large, minor leaves often auriculate or stipule-like. Inflorescences few- to many-flowered panicles; bracts and bracteoles present; pedicels basally articulated. Flowers actinomorphic or zygomorphic, 5merous. Calyx campanulate, mostly 5-lobed; corolla tubular, salverform, or campanulate, prefloration contorted-plicate, rarely imbricate. Stamens 5, equal or not; filaments often pubescent, sometimes geniculate; anthers longitudinally dehiscent; hypogenous nectariferous disk present. Ovary 2-locular; ovules axillary, numerous.

Melananthus 213

Fig. 176. Melananthus ulei

214

S OLANACEAE

Fig. 177. Nicotiana tabacum

Physalis 215

Fruit an apically dehiscent capsule subtended by the slightly accrescent but not enveloping calyx. Seeds many, minute; embryo straight or bent. Tropics, subtropics, more rarely in temperate regions; 60 species, 2 in Venezuela, 1 of these in the flora area. Nicotiana tabacum L., Sp. Pl. 180. 1753. —Tabaco. Robust viscid herb with large leaves; flowers pinkish or whitish. Cultivated, sometimes escaping near villages, near sea level to 1800 m; scattered in Delta Amacuro,

Bolívar, and Amazonas. Widespread in northern Venezuela, native to western South America, cultivated elsewhere. ◆Fig. 177. The leaf of Nicotiana tabacum is smoked, and alkaloids extracted from the plants are sometimes used as an insecticide.

12. PHYSALIS L., Sp. Pl. 182. 1753. by William G. D’Arcy Unarmed herbs or shrubs; indumentum mostly of simple hairs. Leaves simple, mostly somewhat toothed. Inflorescences mostly solitary, often nodding, pedicellate flowers in a leaf axil or branch fork. Flowers 5-merous. Calyx deeply lobed; corolla mostly yellow or white, sometimes with a dark eye, funnelform to rotate, the margin nearly entire. Stamens 5, alike, inserted in the corolla throat; filaments sometimes pubescent; anthers dehiscing longitudinally. Ovary 2-locular; style slender; stigma small. Fruit a juicy or fleshy berry enclosed in the inflated, bladdery accrescent calyx. Seeds many, compressed; embryo strongly curved. Cosmopolitan [Canada, U.S.A., Mexico (center of diversity), widespread in the Neotropics and the Old World]; ca. 100 species, 5 in Venezuela, 4 of these in the flora area. Key to the Species of Physalis 1. 1. 2(1). 2. 3(1). 3.

Fruiting calyx strongly 5-angled; corolla yellow with a strongly contrasting dark eye; lobes of young fruiting calyx subulate ........................... 2 Fruiting calyx rounded or indistinctly 10-angled; corolla yellow, often without a contrasting eye; lobes of young fruiting calyx deltoid ........ 3 Fruiting calyx glabrous; stems basally glabrate; flowering pedicels glabrate ........................................................................................ P. cordata Fruiting calyx pubescent; stems with some long hairs near the base; flowering pedicels mostly conspicuously pubescent ............... P. pubescens Anthers > 2 mm long; angles of fruiting calyx smooth, glabrous; corolla > 6 mm wide ............................................................................... P. angulata Anthers < 2 mm long; angles of fruiting calyx often bumpy and with stout hairs; corolla < 6 mm across ............................................ P. lagascae

Physalis angulata L., Sp. Pl. 183. 1753. —Huevo de sapo, Topotopo. Herb to 1 m tall; flowers yellow. Ruderal weed, near sea level to 200 m; Delta Amacuro (Capure, Pedernales), Bolívar, Amazonas (Río Orinoco at Nericagua). Widespread throughout northern Venezuela; cosmopolitan. ◆Fig. 178.

Physalis cordata Mill., Gard. Dict. ed. 8. 1768. Herb to 1 m tall; flowers yellow with a dark eye. Ruderal weed, 200–400 m; Delta Amacuro, Bolívar (El Manteco, Upata). Mérida, Miranda, Nueva Esparta, Zulia; Canada, U.S.A., Mexico, Central America, West Indies, Trinidad, scattered in Colombia, Ecuador, Peru, and Brazil.

216

S OLANACEAE

Physalis lagascae Roem. & Schult., Syst. Veg. 4: 679. 1819. Annual herb to 60 cm tall; flowers yellow, sometimes with a dark eye. Ruderal weed, 50–300 m; Bolívar (La Camilera, Los Aceites), Amazonas (Departmento Atures). Anzoátegui, Aragua; Mexico, Central America, West Indies, scattered in Colombia, Peru, Brazil, and Bolivia.

Fig. 178. Physalis angulata

Physalis pubescens L., Sp. Pl. 183. 1753. Physalis surinamensis auct. non Miq.: sensu Steyerm., Acta Bot. Venez. 3: 72, 157. 1968. Herb to 1 m tall; flowers yellow with a dark eye. Ruderal weed, 100–900 m; Delta Amacuro (El Palmar), Bolívar (Altiplanicie de Nuria, Cerro Pichacho, Kavanayén), Amazonas (scattered). Throughout Venezuela; nearly cosmopolitan.

13. SCHWENCKIA L., Gen. Pl. ed. 4: 577 (567). 1764, “Schwenkia.” Chaetochilus Vahl, Enum. Pl. 1: 101. 1804. by Carmen Benítez de Rojas Erect, creeping or climbing herbs, sometimes suffruticose or shrubs with flexuous stems, glabrous or pubescent with simple hairs. Leaves alternate, simple, entire, petiolate or subsessile, solitary or fasciculate. Inflorescences solitary flowers, groups of 2 or 3, racemose or paniculate, open or dense; bracts present or not; pedicels filiform, sometimes enlarged with an evident articulation. Flowers slightly zygomorphic. Calyx valvate, tubular or campanulate, 5-dentate or 5-fid; corolla plicate, tubular, the tube erect, somewhat expanded upward, straight or curved, the throat not expanded, the limb reduced, 5-lobed, the median lobe (lacinia) much-reduced or enlarged, denticulate or acicular, the adjacent lateral lobes often united, the united structure sometimes emarginate or prolonged into a petaloid extension extending beyond the median lobes. Stamens 4 and didynamous or 2 or 3 with staminodes; filaments filiform or laminar, inserted in the corolla tube; anthers oblong, broadly oblong, ovoid, or narrowly ellipsoid, 2-locular, longitudinally dehiscent, inserted or exserted. Ovary situated on a cupular or annular disk, 2-locular; ovules numerous; style subexserted, slightly compressed apically; stigmatic portion enlarged. Fruit a capsule dehiscent by 2 entire valves. Calyx persistent and the pedicel somewhat enlarged. Seeds minute, rounded or angled, rugose or muricate; embryo straight. Tropical and subtropical America, introduced and naturalized in Africa; ca. 25–30 species, 9 in Venezuela, 7 of these in the flora area. Key to the Species of Schwenckia 1. 1.

Fertile stamens 4, didynamous; erect herbs or subshrubs ..................... 2 Fertile stamens 2, staminodes 3; vines or herbs ..................................... 3

Schwenckia 217

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

6.

Corollas < 3 mm long, straight distally; leaf bases narrowly cordate, the lobes embracing the stem; capsules < 3 mm across .............. S. micrantha Corollas > 10 mm long, somewhat curved distally; leaf bases attenuate to broadly attenuate; capsules > 3 mm across ..................... S. heterantha Stamens exserted; corollas 2.5–3 cm long, the median corolla lobes 5, minute, denticulate; vines ................................................ S. grandiflora Stamens included; corollas 9–15 mm long; the median corolla lobes 2– 5, claviform; erect or reclining herb or shrub ...................................... 4 Leaves fasciculate in branch axils, the margins slightly wavy ..... S. elegans Leaves scattered along the stems, the margins entire ............................ 5 Corolla somewhat curved distally; plants glabrate ...................... S. glabrata Corolla straight; plants pubescent ........................................................... 6 Plants creeping; pubescence generally silvery green, sericeous, subtomentose; fruiting pedicel inserted directly into the rachis of the inflorescence; leaves mostly firmly papyraceous ........................ S. huberi Plants erect or reclining; pubescence yellowish or whitish (not silvery), plants often glabrate; fruiting pedicels articulated on a ± conspicuous peduncle; leaves mostly membranous or firmly membranous ................................................................................................ S. americana

Schwenckia americana L., Gen. Pl. ed. 6: 567. 1764. —Sardina, Sauce sabanero. Schwenckia guineensis Schumach. & Thonn. in Schumach., Beskr. Guin. Pl. 8. 1827. Schwenckia hirta Klotzsch ex Benth. in A. DC., Prodr. 10: 194. 1846. —Schwenckia americana var. hirta (Klotzsch ex Benth.) Carvalho, Rodriguésia 29: 429. 1978. Schwenckia guianensis Benth. in A. DC., Prodr. 10: 195. 1846. Schwenckia americana var. angustifolia J.A. Schmidt in Mart., Fl. Bras 8(1): 251. 1862. Erect weedy herb 40–100 cm tall; flowers small, greenish, purplish, or yellowish. Wooded savannas, 50–900 m; Bolívar (between Río Caroní and Ciudad Bolívar, Santa Elena), Amazonas (road from Puerto Ayacucho to El Burro, road from Puerto Ayacucho to Gavilán, Raudales de Atures, 8 km southeast of San Carlos de Río Negro). Aragua, Distrito Federal, Guárico, Sucre, Táchira, Trujillo; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina, introduced into tropical Africa. ◆Fig. 179. Schwenckia elegans Carvalho, Loefgrenia 33: 1. 1969.

Branched subshrub, mostly growing in colonies; flowers small greenish or purplish. Savannas, gallery forests, 300–1300 m; widespread in Bolívar, Amazonas (road from Maroa to Yavita). Guyana, French Guiana, Brazil (Roraima). Schwenckia glabrata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 374, t. 178. 1817 [1818]. —Sardina. Schwenckia patens Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 374, t. 179. 1817 [1818]. Branched herb or shrub to 2.5 m tall; flowers small, greenish yellow or orange. Disturbed places in low forests, river banks, 100–600 m; widespread in Bolívar (Río Paragua, Río Paramichi, Río Uroi, Sierra de Lema, Sierra Maigualida), Amazonas (Río Yatua, San Carlos de Río Negro, San Carlos to Solano road, Yavita). Venezuelan Coastal Cordillera; Colombia, Brazil (Amazonas). Schwenckia grandiflora Benth. in A. DC., Prodr. 10: 193. 1846. Schwenckia ulei Dammer, Notizbl. Königl. Bot. Gart. Berlin 6: 187. 1914. Twining climber; flowers greenish yellow. Semidisturbed areas, river banks, 50–500 m; Bolívar (between Cerro Guaiquinima and Río Toronó, Cerro Sipapo, Río Caura), Ama-

218

S OLANACEAE

Fig. 179. Schwenckia americana

Schwenckia 219

Fig. 180. Schwenckia grandiflora

220

S OLANACEAE

Fig. 181. Schwenckia huberi

Solandra 221

zonas (Río Mavaca, Río Ocamo, Río Siapa, Río Tama-Tama, Río Yapacana). Monagas; Guyana, Suriname, Peru, Brazil, Argentina. ◆Fig. 180. Schwenckia heterantha Carvalho, Loefgrenia 33: 2. 1969. Erect herb 40–60 cm tall; flowers pale green or purplish. Granitic outcrops (lajas), 100–200 m; Amazonas (Caño Cupaven, upper Río Autana and Río Guayapo). Brazil (Goiás, Mato Grosso).

Schwenckia huberi Benítez, Ann. Missouri Bot. Gard. 77: 412. 1990. Creeping herb; flowers dark brownish or purplish. Sandy savannas, 50–100 m; Amazonas (just north of Puerto Ayacucho). Endemic. ◆Fig. 181. Schwenckia micrantha Benth. in A. DC., Prodr. 10: 195. 1846. Herb or subshrub; flowers small, yellow. River beaches, near sea level to 50 m; Delta Amacuro (lower Río Orinoco), Bolívar (Ciudad Bolívar). Brazil.

14. SOLANDRA Sw., Kongl. Vetensk. Acad. Nya Handl. 8: 300. 1787, nom. cons. by William G. D’Arcy Hemiepiphytic shrubs. Leaves entire, mostly coriaceous. Inflorescences solitary flowers among the leaves, ebracteate; pedicels short and stout. Flowers large, slightly zygomorphic, 5-merous. Calyx tubular, lobed, sometimes angled, not accrescent, splitting deeply; corolla fleshy, campanulate to goblet-shaped, white, yellow, or bluish, darker in age; the lobes entire to laciniate or fimbriate. Stamens subequal, inserted high on the corolla tube, sometimes pubescent; anthers basifixed, longitudinally dehiscent. Ovary 4-locular, partly inferior; style elongate,

Fig. 182. Solandra grandiflora

222

S OLANACEAE

slender, sometimes exserted. Fruit a leathery conical berry. Seeds many, compressed or angular; embryo curved. Neotropics; ca. 10 species, 2 in Venezuela, both in the flora area. One or two other species are perhaps cultivated in gardens. Key to the Species of Solandra 1. 1.

Corolla mostly < 23 cm long; filaments inserted < 13 cm from the corolla base; corolla tube shorter than the cup ............................... S. grandiflora Corolla mostly > 23 cm long; filaments inserted > 13 cm from the corolla base; corolla tube longer than the cup .................................... S. longiflora

Solandra grandiflora Sw., Kongl. Vetensk. Acad. Nya Handl. 8: 300, t. 11. 1787. —Trompeta de angel. High-climbing hemiepiphytic shrub; flowers large, yellow. Riparian forests, cultivated in gardens, 100–400 m; expected in Amazonas (recorded from Río Castanho, a tributary of the Río Padauiri in the upper Río Negro basin in adjacent Brazil). Aragua, Distrito Federal, Falcón; Antilles, Colombia, Brazil. ◆Fig. 182.

Solandra longiflora Tussac, Fl. Antill. 2: 49. 1818. High-climbing hemiepiphytic shrub. Mature montane forests, 1200–1600 m; Amazonas (Cerro Huachamacari, Sierra de la Neblina). Panama, West Indies, Suriname, Ecuador. The record of Solandra longiflora from Estado Falcón, by Bernardello and Hunziker, Nord. J. Bot. 7: 647. 1987, was based on an erroneous determination of the collection Steyermark 99277 (MO).

15. SOLANUM L., Sp. Pl. 184. 1753; Gen Pl. ed. 5. 1754. by Michael H. Nee and William G. D’Arcy Trees, shrubs, vines, or herbs, glabrous or pubescent with simple or variously branched or stellate hairs, these with lateral rays and a vertical midpoint, sometimes armed. Leaves alternate or geminate, simple and entire to lobed or compound; petioles present or not. Inflorescences terminal, appearing axillary, extraaxillary, or pseudoterminal, basically cymose, racemose, paniculate, or of solitary flowers; pedicels present or not; mostly ebracteate. Flowers mostly actinomorphic, mostly 5-merous, bisexual or unisexual (plants monoecious). Calyx usually campanulate or cupular; corolla rotate or sometimes campanulate, shallowly or deeply lobed, often with abundant plicate interpetalar tissue. Stamens inserted at the base of the corolla tube; filaments basally united at point of insertion, mostly equal; anthers oblong or attenuate, opening by terminal pores and sometimes later by longitudinal slits; nectary absent. Ovary 2-carpellate, sometimes 4-locular by formation of false septa; ovules mostly numerous; style small, exserted or not. Fruit a berry, juicy or dry. Seeds mostly numerous, compressed; embryo strongly curved. Cosmopolitan (best represented in the Neotropics); ca. 1500 species, ca. 65 in Venezuela, 39 of these in the flora area. Key to the Species of Solanum 1. 1. 2(1).

Plants with all simple hairs or glabrous .................................................. 2 Plants with branched or stellate hairs or peltate scales on the lower surfaces of leaves ...................................................................................... 11 High-climbing vines; inflorescence terminal; corolla often showy, blue or purple (rarely white) ............................................................................. 3

Solanum 223

2. 3(2). 3. 4(3). 4. 5(2). 5. 6(5). 6. 7(6). 7. 8(7). 8. 9(8). 9. 10(9).

10.

11(1). 11. 12(11). 12. 13(12). 13. 14(13).

14.

Herbs, shrubs, trees; inflorescences along the stem or opposite leaves or branches; corolla white or yellowish .................................................... 5 Leaves all simple; stamens unequal in length ................................ S. pensile At least some leaves deeply lobed or compound; stamens equal in length ................................................................................................................ 4 Fruits globose, red; flowers numerous, forming a showy blue or purplish (white) panicle; leaves glabrous ...................................... S. seaforthianum Fruits ellipsoid, whitish or greenish; few flowers opening at one time, white; leaves pubescent ............................................................ S. caripense Herbs; inflorescences lateral on the stems between the nodes ............................................................................................. S. americanum Herbs, shrubs, or trees; inflorescences opposite the leaves or branches ................................................................................................................ 6 Herbs; fruits ovoid and pointed; inflorescences at a di- or trichotomy of the stem; at least some leaves lobed ................................... S. allophyllum Shrubs or trees; fruits globose; inflorescences opposite the leaves; leaves all entire ................................................................................................. 7 Leaves glabrous, subcoriaceous; fruiting pedicels stout, woody; peduncle mostly < 3 cm long ................................................................... S. arboreum Lower surfaces of young leaves with hairs, at least in the main vein axils; fruiting pedicels slender; peduncles various ................................. 8 Anthers > 3.5 mm long; calyx truncate with apiculate lobes; inflorescences < 5 cm long ............................................................... S. leucocarpon Anthers < 3 mm long; calyx with deltoid lobes; inflorescences sometimes elongate, to 10 cm long .......................................................................... 9 Leaf pubescence even, denser near the base and middle; corolla < 1 cm long ................................................................................... S. oppositifolium Leaf pubescence only along the major veins or in the vein axils; corolla > 1 cm long ........................................................................................... 10 Stems unwinged, the bark dark reddish brown, peeling on older stems; leaves dark green and brittle in live plants, drying black; inflorescence axes 5–16 cm long, not fleshy ................................ S. campaniforme Stems winged, the bark light green; leaves repandly rugose, somewhat fleshy, drying olive green; inflorescence axes 1–1.6 cm long, somewhat fleshy ......................................................................................... S. tepuiense Anthers oblong with relatively large terminal pores; plants not prickly; leaves entire ......................................................................................... 12 Anthers slender, attenuate, with small terminal pores; plants mostly prickly; leaves mostly sinuate or dentate .......................................... 20 High-climbing vines, mostly glabrate .............................................. S. pensile Erect shrubs, mostly densely pubescent overall .................................... 13 Leaves with peltate-scutellate hairs or scales with rays at least partly fused ..................................................................................................... 14 Leaves with outwardly pointing or multi-angulate hairs, the rays distinct ...................................................................................................... 15 Leaves copious, crowded, mostly > 6 cm long, apically acute or slightly cuspidate; veins mostly > 6 on each side, mostly solitary; plants muchbranched ............................................................................. S. swartzianum Leaves sparse, separated, mostly < 6 cm long, apically long-cuspidate;

224

S OLANACEAE

15(13). 15. 16(15). 16. 17(16). 17. 18(15). 18. 19(18). 19. 20(11). 20. 21(20). 21.

22(20).

22.

23(22). 23.

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

veins mostly < 6 on each side, mostly unequal-paired; plants fewbranched ..................................................................................... S. davidsei Inflorescence simple or 1-branched, small and lax at the leaf axils or subterminal .......................................................................................... 16 Inflorescence several- to many-branched, large and held above the foliage ........................................................................................................ 18 Calyx with well-defined, deltoid or oblong lobes; leaves concolorous, the upper surface abundantly pubescent; fruit glabrous ............. S. umbratile Calyx subtruncate, without well-defined lobes; leaves discolorous, glabrescent; fruit sparingly pubescent .................................................... 17 Peduncles short, mostly unbranched; fruits > 7.5 mm across; inflorescence held beneath the associated leaf .................. S. schlechtendalianum Peduncles elongate, mostly branched; fruits < 7.5 mm across; inflorescence beside or above the associated leaf .................................. S. badilloi Hairs of the lower surface of leaves mostly sessile or nearly so .....S. rugosum Hairs of the lower surface of leaves mostly short-stalked .................... 19 Leaves mostly broadly elliptic, the lower surface soft-tomentose; hairs whitish; ovary and style glabrous ................................................ S. bicolor Leaves mostly lanceolate, the lower surface scabridulous; hairs yellowish or ferruginous; ovary and base of style tomentose ............. S. asperum Upper surface of leaves with simple hairs, spines mostly straight and slender; leaves mostly lobed 1/3–2/3 way to the midvein .................... 21 Leaves with only stellate hairs, spines mostly stout-based and recurved or wanting, mostly lobed < 1/3 way to the midvein ............................ 22 Corolla yellow-green; fruit 1.5 cm diameter, globose; seeds strongly flattened, winged; leaves with the lobes mostly entire ............ S. acerifolium Corolla blue-purple; fruit 3.5–5.5 cm diameter, globose or lobed; seeds slightly flattened, unwinged; leaves with main lobes secondarily lobed or toothed ............................................................................. S. mammosum Fruit glabrous, ovary glabrate or soon glabrescent; leaves pubescent but mostly not densely tomentose, often subentire; shrubs, often sprawling, or woody vines .............................................................................. 23 Fruit and ovary tomentose, sometimes tardily glabrescent; leaves mostly densely tomentose on the lower surface, often conspicuously sinuate or lobed; erect shrubs or small trees .................................................. 31 Leaves coriaceous, the upper surface glabrate and lustrous, the lower surface with hairs mostly stalked; corolla purple ............................. 24 Leaves membranous, the upper surface tomentose, not lustrous, the lower surface with hairs mostly sessile; corolla white, violet, or purple .............................................................................................................. 25 Young parts mostly unarmed; small tree ................................. S. altissimum Young parts mostly armed with recurved spines; woody vines ... S. coriaceum Corolla purple; flower buds mostly slender and curved ............ S. subinerme Corolla whitish; flower buds mostly short, mostly straight .................. 26 Spines few, mostly straight and restricted to stems; leaves narrowly lanceolate; plants of banks of swift-flowing streams ............S. schomburgkii Spines mostly copious, recurved; leaves ovate-lanceolate or wider; upland plants, not riparian ..................................................................... 27

Solanum 225

27(26). Anthers < 4 mm long; fruit 7–9 mm across, soft; seeds 1.5 mm long; leaves sessile with cuneate bases ......................................... S. jamaicense 27. Anthers > 5 mm long; fruit 7–30 mm across with firm rind; seeds 2–6 mm long; leaves mostly petiolate with curved, not cuneate bases ... 28 28(27). Upper surface of at least some leaves with acicular prickles on the midvein, the lower surface often without recurved spines; corolla or calyx often with shaggy pubescence outside, the stellate hairs with long midpoints ............................................................ S. monachophyllum 28. Upper surface of leaves unarmed, the lower surface always with recurved spines; corolla lacking shaggy pubescence, the midpoints of the stellate hairs mostly shorter than the rays ....................................... 29 29(28). Leaves mostly entire, narrowly lanceolate; inflorescence unarmed; flower buds ± symmetrical; stems with numerous, uniform, small (1–2 mm) recurved spines; pedicels slender and erect in fruit ......... S. lanceifolium 29. Leaves lobed or entire, narrowly to broadly ovate; inflorescence parts often with spinulose bristles; flower buds curved; stems with sparse, unequal spines 2–8 mm long; pedicels thickened and curved in fruit .............................................................................................................. 30 30(29). Leaves firm to subcoriaceous, subentire, the margin revolute; fruits 2– 3 cm across; seeds 3.5–6 mm long ............................................. S. aturense 30. Leaves membranous, mostly angulate or lobed, the margin plane; fruits 1–1.5 cm across; seeds 2–3 mm long ...................................... S. adhaerens 31(22). Vines or scrambling shrubs; spines mostly recurved ............................ 32 31. Erect shrubs or small trees; spines mostly straight .............................. 35 32(31). Plants whitish-tomentose; fruits > 2.5 cm across ....................... S. oocarpum 32. Plants yellowish- or ferruginous-tomentose, especially on young parts; fruits < 2 cm across ............................................................................. 33 33(32). Plants fine-pubescent, the stellate hairs with short midpoints or none; upper surface of leaves scabrous ....................................... S. rufistellatum 33. Plants pilose, the stellate hairs with long midpoints; leaves scabrous or not ......................................................................................................... 34 34(33). Leaves broadly ovate, basally acute to subcordate, the upper surface softly tomentose ...................................................................... S. velutinum 34. Leaves obovate to elliptic, basally acute to rounded, the upper surface scabrous ...................................................................................... S. nuricum 35(31). Calyx truncate; leaves often with acicular spines; stellate hairs of ovary with short midpoints about equaling the rays; fruits evenly finetomentose, < 2 cm across ............................................... S. stramonifolium 35. Calyx deltoid; leaves with recurved or broad-based spines or none; stellate hairs of ovary with long midpoints mostly exceeding the rays; fruits coarsely or unevenly pubescent, > 2 cm across ....................... 36 36(35). Plants bristly with long-stalked hairs bearing rays at their tips; anthers pubescent ................................................................................... S. crinitum 36. Plants without bristles, stellate hairs mostly with the rays at the base; anthers glabrous .................................................................................. 37 37(36). Leaves strongly discolorous, the upper surface glabrate ...................... 38 37. Leaves concolorous or slightly discolorous, the upper surface pubescent overall .................................................................................................. 39

226

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38(37). Minor venation of the lower surface of leaf evident, the indumentum yellowish to brownish; fruits mostly < 16 mm across, sparingly pubescent; corolla narrowly lobed to near the base; anthers glabrous ................................................................................................ S. paludosum 38. Minor venation of the lower surface of leaf obscured by the whitish to yellowish indumentum; fruits mostly > 16 mm across, fine-tomentose overall; corolla lobed < 1/2 way down, the lobes deltoid; anthers dorsally pubescent......................................................................... S. oocarpum 39(37). Fruits small, < 2.5 cm across; leaves mostly < 25 cm long ............. S. hirtum 39. Fruits large, > 2.5 cm across; leaves often > 25 cm long ....................... 40 40(39). Leaves mostly subentire, basally rounded, armature of stems of short (2–4 mm), stout, recurved spines ........................................ S. vestissimum 40. Leaves mostly lobed, basally truncate; armature of stems when present of slightly curved, acicular spines 2–8 mm long ............... S. sessiliflorum Solanum acerifolium Dunal, Sol. Syn. 41. 1816. Solanum quinquangulare Roem. & Schult., Syst. Veg. 4: 669. 1819. Solanum arcuatum Sendtn. in Mart., Fl. Bras. 10: 60. 1846. Solanum hastatum Sendtn. in Mart., Fl. Bras. 10: 61. 1846. Shrub 0.5–3 m tall; flowers white; fruits yellow. Forest clearings, ravines, 500–1800 m; Bolívar (Gran Sabana), Amazonas (Sierra de la Neblina). Southern Mexico, Central America, Colombia, Trinidad, Ecuador, northern Peru, eastern Brazil. Solanum adhaerens Roem. & Schult., Syst. Veg. 4: 669. 1819. Solanum enoplocalyx Dunal in A. DC., Prodr. 13: 222. 1852. Solanum humboldtii Dunal in A. DC., Prodr. 13: 216. 1852., non Willd. 1803. Solanum donnellsmithii J. Coulter, Bot. Gaz. (Crawfordsville) 16: 144. 1891. Sprawling vine with recurved spines; flowers white. Disturbed areas, 50–1200 m; widespread in Delta Amacuro and northern Bolívar. Common in northern Venezuela; Mexico, Central America, Colombia, Trinidad, Guyana. Solanum allophyllum (Miers) Standl., J. Wash. Acad. Sci. 17: 16. 1927. —Pionandra allophylla Miers in Seem., Bot. Voy. Herald 174: 1854. —Cyphomandra allophylla (Miers) Hemsl., Biol. Cent.-Amer., Bot. 2: 417. 1882.

Herb or weak shrub to 1 m tall; flowers whitish. Understory in lowland seasonal forests, tall savannas, 200–300 m; northern Bolívar. Zulia; Honduras to Panama, Colombia. ◆Fig. 184. Solanum altissimum Benítez, Ernstia 19: 13. 1893. —Solanum altissimum Pittier et al., Cat. Fl. Venez. 2: 372. 1947, nom. nud. —Tupiro de danto. Tree to 6 m tall or more; flowers large, blue-purple. Evergreen lowland forests, 100–200 m; Amazonas (upper Río Negro basin). Eastern Colombia, Peru, western Amazonian Brazil. ◆Fig. 185. Solanum americanum Mill., Gard. Dict. ed. 8. 1768. —Yerba mora. Solanum nodiflorum Jacq., Collecteana 2: 288. 1788 [1789]. Solanum pterocaulon Dunal, Hist. Nat. Solanum 153. 1813. Solanum oleraceum Lam., Encycl. suppl. 3: 750. 1813 [1814]. Unarmed herb to 1 m tall; flowers small, white; fruits green or black. Disturbed areas, 100–400(–1700) m; widespread in Bolívar. Common elsewhere in lowland Venezuela; throughout lowland tropical and warm temperate regions. ◆Fig. 183. Solanum americanum is distinguished from other widespread species (not present in the flora area) by anthers < 2 mm long. Solanum macrotonum Bitter (S. gollmeri Bitter) is one of these species with larger anthers that occurs in Venezuela, usually above 1000 m. Sometimes the fruit or foliage

Solanum 227

of Solanum americanum is eaten. Plants usually contain enough alkaloids to render them slightly or severely toxic. The leaves are also used in poultices for ringworm. Solanum arboreum Dunal, Sol. Syn. 20. 1816. Solanum dolichostylum O.E. Schulz in Urb., Symb. Antill. 6: 158. 1909. Solanum ripivagum Pittier et al., Cat. Fl. Venez. 2: 382. 1947, nom. nud. Rhizomatous shrub to 3 m tall; flowers small, white. Evergreen lowland forests, 200–400 m; Delta Amacuro (east of El Palmar), Bolívar (El Palmar, Los Patos, Río Toro), Amazonas (Departamento Atures). Anzoátegui, Aragua, Distrito Federal, Lara, Mérida, Monagas, Nueva Esparta, Táchira, Trujillo, Yaracuy; Nicaragua to Panama, northern Colombia, Trinidad. ◆Fig. 186. Solanum asperum Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. —Tabaquero peluo, Tabaquillo. Solanum salviifolium auct., doubtfully Lam., Tabl. Encycl. 2: 14. 1794. Solanum radula Vahl, Eclog. Amer. 2: 16. 1798. Tree to 6 m or more tall; leaves asperous to the touch; flowers white. Secondary growth thickets, near sea level to 1200 m; widespread in Delta Amacuro, Bolívar (Altiplanicie de Nuria, Cerro Bolívar, Distrito Cedeño, Gran Sabana). Aragua, Anzoátegui, Distrito Federal, Falcón, Miranda, Nueva Esparta, Sucre, Yaracuy; Belize, Panama, northern Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 187. Solanum aturense Dunal, Sol. Syn. 32. 1816. —Uña de gato. Solanum asperrimum Bitter & O. Moritz, Repert. Spec. Nov. Regni Veg. 16: 393. 1920. Solanum secundum Bitter & O. Moritz, Repert. Spec. Nov. Regni Veg. 16: 391. 1920. Solanum siparunoides Ewan, Nat. Hist. Misc. 94: 1. 1952. Robust armed vine; flowers violet; fruits red to orange. Evergreen lowland forests, 100–300 m; Bolívar (El Dorado, Río Bo-

tanamo), Amazonas (Departamento Atures). Apure, Aragua, Distrito Federal, Mérida, Miranda, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Colombia, Peru. ◆Fig. 189. Solanum badilloi Benítez, Ernstia 44: 1. 1987. —Pasita. Shrub to 3 m tall; leaves discolorous; flowers white. Disturbed areas in moist forests, 100–200 m; Amazonas (near confluence of Río Casiquiare and Río Negro). Endemic. Solanum badilloi may be conspecific with S. distichophyllum Sendtn. of western Brazilian Amazonia (Amazonas, Acre, Rondônia) and northern Bolivia. Solanum bicolor Roem. & Schult., Syst. Veg. 4: 661. 1819. —Tabacón borrachera, Tabaquillo. Solanum callicarpifolium Kunth & Bouché, Index Sem. (Berlin) app. 1845: 10. 1845. Shrub to 3 m tall; corolla white, anthers yellow. Early secondary growth in deciduous forest areas, 50–400(–1200) m; northern Bolívar and Amazonas. Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Miranda, Portuguesa, Sucre, Yaracuy; northern Colombia, Lesser Antilles (St. Vincent), Trinidad, Guyana, Ecuador. ◆Fig. 190. Solanum hazenii Britton has been reported from the flora area, but it is most likely a misidentification of S. bicolor. Solanum campaniforme Roem. & Schult., Syst. Veg. 4: 662. 1819. —Pica pica. Shrub to 5 m tall; flowers white. Thickets or forest patches, often on sandy soil in savannas, 200–1200 m; Bolívar (Altiplanicie de Nuria, Ciudad Piar, Miamo). Anzoátegui; Guyana, French Guiana, eastern coastal Brazil, Bolivia. ◆Fig. 193. Solanum caripense Dunal, Sol. Syn. 8. 1816. Climbing herb to 2 m tall; flowers white. Naturally disturbed sites, 1500–1700 m; Amazonas (Ceror Yutajé). Mérida, Monagas; Nicaragua to Panama, Colombia, Ecuador, Peru.

228

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Solanum coriaceum Dunal, Hist. Nat. Solanum 197, t. 14. 1813. Robust vine to 20 m high or sprawling shrub 2–4 m tall; flowers blue or purple. Evergreen montane forests, 1200–1400 m; Bolívar (La Escalera to Luepa). Guyana, Suriname, French Guiana, northeastern Brazil (Amapa, Pará, Roraima). ◆Fig. 188. Solanum crinitum Lam., Tabl. Encycl. 2: 20. 1794. Solanum macranthum Dunal in Lam., Encycl. suppl. 3: 775. 1813 [1814]. Solanum formosum Dunal, Sol. Syn. 35. 1816. Solanum jubatum Roem. & Schult., Syst. Veg. 4: 669. 1819. —Solanum cyananthum var. jubatum (Roem. & Schult.) Dunal in A. DC., Prodr. 13: 316. 1852. Solanum cyananthum Dunal in A. DC., Prodr. 13: 316. 1819. Shrub or tree to 6 m tall with bristly twigs and inflorescences; flowers large, showy, blue-purple; fruits large, pubescent. Disturbed wet forests, savanna edges, 100– 1000 m; widespread in Bolívar and Amazonas. Apure; eastern Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 195. A similar species, Solanum wrightii Benth., from the Bolivian Andes, is cultivated for ornament in Venezuela and may be expected; it is much less tomentose and becomes a taller tree. Solanum davidsei Carvalho, Ann. Missouri Bot. Garden 78: 229, figs. 4, 14D. 1991. Shrub to 1.5 m tall; flowers white. Upland savannas, 1300–1400 m; Bolívar (north of Luepa and 50 km west of Luepa on road to Kavanayén). Endemic. Solanum hirtum Vahl, Symb. Bot. 2: 40. 1791. —Huevo de gato. Solanum flavescens Dunal in Lam., Encycl. suppl. 3: 778. 1813 [1814]. Solanum orenocense Dunal, Sol. Syn. 42. 1816. Solanum echinocarpum Bredem. ex Roem. & Schult., Syst. Veg. 4: 659. 1819. Solanum flavescens var. albescens Dunal in A. DC., Prodr. 13: 254. 1852.

Solanum obtusifrons Pittier et al., Cat. Fl. Venez. 2: 371. 1947, nom. nud. Solanum timotense Pittier et al., Cat. Fl. Venez. 2: 371. 1947, nom. nud. Shrub to 2 m tall; flowers white; fruits tomentose. Disturbed deciduous forests, 100– 300 m; Bolívar (km 190 on road from Caicara del Orinoco to San Juan de Manapiare). Occurring generally elsewhere in Venezuela; Mexico, Central America, northern Colombia, Trinidad. ◆Fig. 197. The furry, unpalatable fruits of Solanum hirtum are sometimes eaten. Solanum jamaicense Mill., Gard. Dict. ed. 8. 1768. Shrub to 2 m tall; flowers small, white; fruits red. Disturbed deciduous lowland forests, near sea level to 200 m; Delta Amacuro (Boca de Cuyubini), Bolívar (Distrito Heres). Apure, Barinas, Carabobo, Distrito Federal, Guárico, Miranda, Monagas, Sucre, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Solanum lanceifolium Jacq., Collectanea 2: 286. 1788 [1789]. Solanum obscurum herb. Willd., doubtfully Vahl, Symb. Bot. 2: 41. 1791. Solanum brachyacanthum Dunal, Sol. Syn. 32. 1816. —Solanum lanceifolium var. brachyacanthum (Dunal) O.E. Schulz in Urb., Symb. Antill. 6: 249. 1909. Solanum umbrosum Dunal, Sol. Syn. 33. 1816. Solanum topirito Pittier et al., Cat. Fl. Venez. 2: 370. 1947, nom. nud. Sprawling shrub or vine to many m long; flowers purple or blue and white; fruits orange or red. Openings in deciduous lowland forests, often on limestone, 50–500 m; Delta Amacuro (between La Margarita and Puerto Miranda), Bolívar (Altiplanice de Nuria, Distrito Piar). Distrito Federal, Falcón, Lara, Miranda, Nueva Esparta, Táchira, Trujillo; Mexico, Central America, Antilles, Colombia, Trinidad. ◆Fig. 194. Solanum leucocarpon Dunal in Lam., Encycl. suppl. 3: 756. 1813 [1814]. Solanum surinamense Steud., Flora 26: 764. 1843.

Solanum 229

Solanum triste var. crassipes Dunal in A. DC., Prodr. 13: 148. 1852. Shrub or tree to 6 m tall; flowers white. Disturbed evergreen forests, 50–1500 m; Bolívar (Santa María de Erebato, Sierra de Maigualida, Túriba), Amazonas (Oromaná, Puerto Ayacucho, San Juan de Manapiare). Barinas, Táchira, Yaracuy; Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 196. Solanum leucocarpon is perhaps the only species of sect. Geminata centered in lowland evergreen forests of Amazonia. Solanum mammosum L., Sp. Pl. 187. 1753. —Manzana del diablo. Solanum platanifolium Hook., Bot. Mag. 53: t. 2618. 1826. Solanum globiferum Dunal in A. DC., Prodr. 13: 251. 1852. Shrub or herb to 2 m tall; flowers bluepurple; fruits large, yellow, with round “udders.” Cultivated around dwellings, and escaped or perhaps native, 200–400 m; Bolívar (Santa María de Erebato). Scattered in northern half of Venezuela; southern Mexico, Central America, Antilles, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 199. Solanum mammosum is probably sparingly cultivated throughout the area as an ornamental, and for medicinal and insecticide uses by rural peoples, as well as being at the edge of the native range. Solanum monachophyllum Dunal, Sol. Syn. 33. 1816. Solanum molle Dunal, Sol. Syn. 28. 1816, non Burman 1768. Solanum javitense Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 42. 1818, nom. superfl. Solanum sacupanense Rusby, Descr. S. Amer. Pl. 115. 1920. Solanum bolivarense Pittier et al., Cat. Fl. Venez. 2: 369. 1947, nom. nud. Slender armed shrublet to 2 m tall; flowers white. Edges of savannas, gallery forests, especially along watercourses and seasonally flooded areas, near sea level to 500 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Ranges through Llanos of Venezuela; Colombia, Guyana, Suriname, French Guiana, northern Brazil.

At times it is difficult to distinguish Solanum monachophyllum from S. subinerme (with violet flowers) and S. adhaerens (more viny and with more strictly curved spines). Solanum nuricum M. Nee, Novon 4: 285. 1994. Shrub or woody vine with pilose, accrescent calyx. Forests, forest edges, 300– 700 m; Bolívar (Altiplanicie de Nuria). Endemic. Solanum oocarpum Sendtn. in Mart., Fl. Bras. 10: 106, t. 7, fig. 28–30. 1846. Solanum orientale Benítez, Ernstia 23: 8. 1984. Scrambling shrub or vine to 6 m tall; flowers purple; fruits large, fine-pubescent. Disturbed moist forests, 100–500 m; northern Bolívar. Anzoátegui, Sucre; Brazil (Amazonas: Rio Branco, Goiás). ◆Fig. 200. Solanum oocarpum is closely related to S. crinitum, which is bristly-setose on young parts. Solanum oppositifolium Ruiz & Pav., Fl. Peruv. 2: 35. 1799. Solanum puberuloba Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 444. 1966. Shrub to 5 m tall; flowers whitish. Understory in moist forests, 100–800 m; Delta Amacuro (Serranía de Imataca), Bolívar (El Palmar), Amazonas (Cerro Sipapo, Río Cataniapo, San Fernando de Atabapo). Eastern Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 203. Solanum paludosum Moric., Pl. Nouv. Amér. 29, t. 20. 1837, non Dunal 1952. Shrub to 3.5 m tall; flowers white to violet. Evergreen lowland to montane forests, 200–1700 m; widespread in Bolívar. Guyana, Suriname, French Guiana, eastern Brazil. ◆Fig. 201. Solanum pensile Sendtn. in Mart., Fl. Bras. 10: 50. 1846. Solanum scandens L., Pl. Surin. 5. 1775, non Mill. 1768. Solanum scandens L. f., Suppl. Pl. 147. 1781 [1782], non Mill. 1768. Solanum ipomoea Sendtn. in Mart., Fl. Bras. 10: 50. 1846.

230

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Solanum laetum Miq., Stirp. Surinam. Select. 135. 1850 [1851], non Kunze 1842. Solanum sempervirens Dunal in A. DC., Prodr. 13: 88. 1852, non Mill. 1768. Solanum styracoides Rusby, Mem. Torrey Bot. Club 4: 230. 1897. Solanum miquelii C.V. Morton, Contr. U.S. Natl. Herb. 29(1): 43. 1944. Solanum granelianum D’Arcy, Ann. Missouri Bot. Gard. 60: 758. 1972 [1973]. Solanum prehensile Pittier et al., Cat. Fl. Venez. 2: 380. 1947, nom. nud. High-climbing vine; flowers showy violet or purple. Wet forests, near sea level to 300 m; Delta Amacuro (Río Cuyubini), Bolívar (El Manteco to San Pedro de las Dos Bocas), Amazonas (Caño Yapacana, Puerto Ayacucho). Aragua, Carabobo, Mérida; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 198. Solanum pensile shows a greater variation in pubescence than most other species in the family. Pubescence varies from glabrous to pubescent, and most specimens have simple hairs, but some specimens bear scattered, minute, branched hairs. Solanum rufistellatum Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 445. 1966. Sparingly armed shrub or vine with rusty indumentum; flowers white. Semideciduous forests, disturbed areas of montane forests, 200–1200 m; Bolívar (Cerro Venamo to Luepa, between Tumeremo and Bochinche). Endemic. ◆Fig. 191.

—Solanum geminifolium Schlect., Linnaea 5: 112. 1830, non Thonn. 1827. Unarmed shrub to 3 m tall; flowers white. Disturbed moist forests, 50–800 m; Bolívar (Altiplanicie de Nuria, Río Canaracuni, Santa María de Erebato), Amazonas (base of Sierra de la Neblina, Sierra Parima, Tobogán de la Selva near Coromoto). Barinas, Distrito Federal, Miranda, Monagas, Portuguesa, Yaracuy, Zulia; southern Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northwestern Argentina. ◆Fig. 192. The positioning of the inflorescence along the underside of the associated leaf in Solanum schlechtendalianum is shared only with S. swartzianum in the flora area. Solanum schomburgkii Sendtn. in Mart., Fl. Bras. 10: 55. 1846. Solanum hostmannii Dunal in A. DC., Prodr. 13: 184. 1852. Solanum boissieri Dunal in A. DC., Prodr. 13: 188. 1852. Erect herb with narrow leaves. Banks of swift-moving streams, 50–100 m; Bolívar (Río Caroní). Apure; Guyana, Suriname, French Guiana, Brazil (Pará, Roraima).

Solanum rugosum Dunal in A. DC., Prodr. 13: 108. 1852. Shrub to 5 m tall, with unpleasant odor; flowers white. Disturbed openings in wet forests, 100–1200 m; Bolívar (Altiplanicie de Nuria, Cerro Marutaní, El Palmar, Río Grande), Amazonas (Mavaca, Río Ventuari, Santa Rosa de Ucata, Sierra Parima, Tamatama). Aragua, Carabobo, Distrito Federal, Yaracuy; Guatemala to Panama, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 204.

Solanum seaforthianum Andrews, Bot. Repos. 8, t. 504. 1808. —Viudita. Solanum cyrrhosum Dunal, Sol. Syn. 9, t. 91. 1816. Solanum salignum Roem. & Schult., Syst. Veg. 4: 663. 1819. Solanum prunifolium Roem. & Schult., Syst. Veg. 4: 662. 1819. High-climbing unarmed vine; flowers showy, blue, purple, or white; fruits red. Seaside and forest openings, sea level to 500 m; Delta Amacuro, Bolívar (Altipanicie de Nuria), Amazonas (scattered). Carabobo, Distrito Federal, Falcón, Miranda, Táchira, Zulia; Mexico, Central America, Antilles, apparently native in northern South America, Colombia, Trinidad, French Guiana, Brazil, Paraguay. ◆Fig. 207. Solanum seaforthianum is commonly cultivated for ornament in most parts of Venezuela and worldwide in the tropics.

Solanum schlechtendalianum Walp., Repert. Bot. Syst. 3: 61. 1844.

Solanum sessiliflorum Dunal in Lam., Encycl. suppl. 3: 775. 1813 [1814]. —Túpiru.

Solanum 231

Solanum topiro Dunal, Sol. Syn. 10, t. 92, fig. 1. 1816. Solanum arecunarum Pittier et al., Cat. Fl. Venez. 2: 372. 1947, nom. nud. Shrub to 2 m tall; flowers white; fruits tomentose, to 9.5 cm across. Disturbed and open areas, often cultivated in “conucos,” 50–900 m; southern Bolívar, western Amazonas. Colombia, Ecuador, Peru, northwestern Brazil. ◆Fig. 208. The fruits of Solanum sessiliflorum are often eaten fresh, as juice, or cooked. Solanum stramonifolium Jacq., Misc. Austriac. 2: 298. 1781. Solanum toxicarium Rich., Actes Soc. Hist. Nat. Paris. 1: 107. 1792. Solanum platyphyllum Dunal, Sol. Syn. 38. 1816. Solanum orinocense Dunal, Sol. Syn. 42. 1816. Solanum undecimangulare Roem. & Schult., Syst. Veg. 4: 669. 1819. Solanum trichocarpum Miq., Stirp. Surinam. Select. 134. 1850 [1851]. Solanum demerarense Dunal in DC, Prodr. 13: 257. 1852. Solanum demerarense var. inerme Dunal in A. DC., Prodr. 13: 257. 1852. —Solanum stramonifolium var. inerme (Dunal) Whalen, Gentes Herb. 12: 76. 1981. Sparingly armed shrub to 2 m tall; flowers white. Disturbed areas of forests and savannas, near sea level to 1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Monagas; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, northern Peru, Brazil. ◆Fig. 206. Solanum subinerme Jacq., Enum. Syst. Pl. 15. 1760. —Cocuna, Cucuna, Tupirito. Solanum juripeba Rich., Actes Soc. Hist. Nat. Paris. 1: 107. 1792. Solanum heterophyllum Lam., Tabl. Encycl. 3: 22. 1799. Solanum styracifolium Roem. & Schult., Syst. Veg. 4: 663. 1819. Solanum stamineum Steud., Flora 26: 764. 1848. Shrub to 3.5 m tall, with recurved prickles; flowers violet; fruits red or orange. Ag-

gressive in disturbed areas of humid forests, near sea level to 1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Throughout Venezuela; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil, Bolivia. ◆Fig. 205. Solanum swartzianum Roem. & Schult., Syst. Veg. 4: 602. 1819. Shrub to 6 m tall; flowers white; fruits partly enclosed in the calyx. Streamsides and disturbed areas of deciduous lowland forests, 900–1800 m; Bolívar (Gran Sabana, Santa Elena to Icabarú), Amazonas (Sierra de la Neblina). Eastern non-Amazonian Brazil (and Roraima). Solanum tepuiense S. Knapp, Novon 1: 125. 1991. Shrub ca. 2.5 m tall; flowers white. Tepui slopes, ca. 1800 m; Bolívar (Sororopántepui). Endemic. Solanum umbratile I.M. Johnst., Proc. Amer. Acad. Arts 40: 695. 1905. Shrub or small vine; flowers white. Open woodlands or secondary vegetation, 200–600 m; northeastern Bolívar. Miranda, Nueva Esparta, Portuguesa, Yaracuy. Solanum velutinum Dunal, Sol. Syn. 33. 1816. —Huevo de gato amarillo. Solanum rhamphidacanthum Bitter, Repert. Spec. Nov. Regni Veg. 12: 436. 1913. Solanum confusum Pittier et al., Cat. Fl. Venez. 2: 368. 1947, nom. nud. Solanum fraternorum Pittier et al., Cat. Fl. Venez. 2: 369. 1947, nom. nud. Solanum lanugiflorum Pittier et al., Cat. Fl. Venez. 2: 369. 1947, nom. nud. Solanum mesopliarthrum Pittier et al., Cat. Fl. Venez. 2: 368. 1947, nom. nud. —Solanum mesopliarthrum Benítez & Steyerm., Pittieria 7: 20. 1978. Scrambling shrub to 5 m tall; flowers lilac. Lower montane forests, 300–1700 m; Bolívar (Altiplanicie de Nuria, Cerro Bolívar, Gran Sabana, Macizo del Chimantá, southeast of Upata). Venezuelan coastal ranges; Guyana, Suriname, French Guiana, circum-Amazonian eastern Brazil. ◆Fig. 202.

232

S OLANACEAE

Fig. 183. Solanum americanum

Fig. 184. Solanum allophyllum

Solanum 233

Fig. 185. Solanum altissimum

Fig. 186. Solanum arboreum

234

S OLANACEAE

Fig. 187. Solanum asperum

Fig. 188. Solanum coriaceum

Solanum 235

Fig. 189. Solanum aturense

Fig. 190. Solanum bicolor

236

S OLANACEAE

Fig. 191. Solanum rufistellatum

Fig. 192. Solanum schlechtendalianum

Solanum 237

Fig. 193. Solanum campaniforme

Fig. 194. Solanum lanceifolium

238

S OLANACEAE

Fig. 195. Solanum crinitum

Solanum 239

Fig. 196. Solanum leucocarpon

Fig. 197. Solanum hirtum

240

S OLANACEAE

Fig. 198. Solanum pensile

Fig. 199. Solanum mammosum

Solanum 241

Fig. 200. Solanum oocarpum

Fig. 201. Solanum paludosum

242

S OLANACEAE

Fig. 203. Solanum oppositifolium Fig. 202. Solanum velutinum

Solanum 243

Fig. 204. Solanum rugosum

Fig. 205. Solanum subinerme

244

S OLANACEAE

Fig. 206. Solanum stramonifolium

Fig. 207. Solanum seaforthianum

Solanum 245

Fig. 208. Solanum sessiliflorum

Solanum vestissimum Dunal in A. DC., Prodr. 13: 322. 1852. Solanum scabridum Dunal in A. DC., Prodr. 13: 253. 1852. Solanum hyporrhodium Pittier et al., Cat. Fl. Venez. 2: 370. 1947, nom. nud., non A. Braun & Bouché 1853.

Shrub or tree to 5 m tall; flowers violet to white. Montane forests, 300–1500 m; Amazonas (Sierra de la Neblina, Sierra Parima). Aragua, Barinas, Falcón, Mérida, Miranda, Táchira, Trujillo, Yaracuy, Zulia; Andean Colombia.

246

S OLANACEAE

16. WITHERINGIA L’Hér., Sert. Angl. 33, t. 1. 1789. by William G. D’Arcy Small unarmed shrubs or herbs; indument of simple or branched hairs. Leaves entire or sinuate-dentate. Inflorescences several-flowered fascicles in the leaf axils, ebracteate, pedicels curving down in flower, upward in fruit. Flowers 4or 5-merous, calyx apically truncate, sometimes accrescent, corolla campanulate to rotate, the lobes valvate. Stamens alike, inserted at or above the middle of the corolla tube; filaments mostly pubescent; anthers mostly stout, sometimes with an apiculate connective; dehiscing longitudinally. Ovary 2-locular; style slender; stigma small. Fruit a juicy berry. Seeds many, compressed; embryo strongly curved. Mexico, Central America, Jamaica, Colombia, French Guiana, Ecuador, Peru, Bolivia; ca. 10 species, 1 or 2 in Venezuela, 1 of these in the flora area. Witheringia solanacea L’Hér., Sert. Angl. 33, t. 1. 1789. —Saracha solanacea (L’Hér.) Miers, Ann. Mag. Nat. Hist. ser. 2, 3: 451. 1849. Witheringia macrophylla Dunal, Sol. Syn. 1. 1816. —Brachistus macrophyllus (Dunal) Miers, Ann. Mag. Nat. Hist. ser. 2, 3: 263. 1849. —Bassovia macrophylla (Dunal) J. Coulter, Bot. Gaz. (Crawfordsville) 16: 145. 1891. —Capsicum solanaceum var. macrophylla (Dunal) Kuntze, Revis. Gen. Pl. 2: 450. 1891. —Capsicum macrophyllum (Dunal) Standl., J. Wash. Acad. Sci. 17: 16. 1927.

Sicklera tetrandra A. Braun & Bouché, Index Sem. (Berlin) app. 1853: 10. 1853. Herb, sometimes hollow-stemmed, rarely arborescent and to 1.5 m tall. Lower montane forests, ca. 1200 m; Amazonas (Sierra de la Neblina). Aragua, Carabobo, Distrito Federal, Lara, Falcón, Mérida, Miranda, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Jamaica, Colombia, French Guiana, Ecuador, Peru, Bolivia. ◆Fig. 209.

Fig. 209. Witheringia solanacea

Sphenoclea 247

SPHENOCLEACEAE by Bruce A. Stein Annual herbs. Stems spongy, fistulous. Leaves alternate; short-petiolate; blades simple and entire. Inflorescence terminal, an erect, densely congested spike. Flowers small, 2–3 mm across, bisexual, sessile, subtended by a bract and a pair of bracteoles; hypanthium laterally compressed. Sepals 5, united at the base, imbricate, persistent. Corolla actinomorphic, campanulate, the 5 equal lobes imbricate. Stamens free, adnate to the corolla near the middle; filaments short; anthers rounded, appearing peltate. Ovary inferior, bilocular; stigma capitate, sessile or on very short style. Fruit a circumscissilely dehiscent capsule. Seeds numerous, minute, oblong, longitudinally ridged with indistinct pits. Native to Africa (perhaps Asia also), introduced and naturalized into the tropics and subtropics of the New World; 1 genus and 2 species; 1 species in the flora area.

Fig. 210. Sphenoclea zeylanica

248

S PHENOCLEACEAE

1. SPHENOCLEA Gaertn., Fruct. Sem. Pl. 1: 113, pl. 24, fig. 5. 1788, nom. cons. Characters as in the family. Pantropics; 2 species, 1 in Venezuela. Sphenoclea zeylanica Gaertn., Fruct. Sem. Pl. 1: 113, pl. 24, fig. 5. 1788. Weedy herb; flowers white. Moist disturbed sites, seasonally submerged river beaches, near sea level to 100 m; Delta Amacuro (Caño Mánamo south of Tucupita, Sacupana), Bolívar (Puerto Paéz), Amazonas (south of Puerto Ayacucho on

road to Samariapo, island in front of Isla Ratón). Llanos, Apure, Miranda, Portuguesa, Zulia; southeastern U.S.A., Mexico, Honduras, Costa Rica, Panama, Puerto Rico, Trinidad, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 210.

STERCULIACEAE by Carmen L. Cristóbal, Janice G. Saunders, and Paul E. Berry Shrubs, subshrubs, or trees, rarely vines or herbs, monoecious, dioecious, or polygamous; stems unarmed or occasionally aculeate, with stellate, simple, or glandular hairs. Leaves alternate, simple, or sometimes palmately compound, stipulate, petiolate, rarely sessile; margins serrate, 3–5-lobed, or entire. Inflorescences 1–many-flowered, axillary or terminal, sometimes cauliflorous. Flowers homostylous or heterostylous, actinomorphic, zygomorphic, or asymmetric but appearing zygomorphic. Calyx valvate, the sepals (2–)5(6), basally connate or free; corolla contorted or imbricate, the petals caducous or marcescent, (4)5 or none, similar or dissimilar, flat with the claw free, adherant or adnate to the staminal tube base, or cucullate with the claw appressed to the staminal tube margin, in this case the blade present or absent, sometimes smaller than the clawed base, simple or bifid. Androgynophore present or absent, sometimes longer than the corolla. Stamens opposite the petals, 5–many, free or more often with connate filaments forming a tube, sometimes with 5 groups bearing 2 or 3 anthers, anthers with 2(3) thecae, free or connate as a tiny head; staminodes 5 or none, alternating with the stamens or in an internal whorl. Gynophore sometimes developed. Ovary superior, (1–)5(–10 or 12)-carpellate, carpels 1–many-ovulate, placentation axile; styles (1–) 5(–10 or 12). Fruit usually a capsule, sometimes berry-like, drupaceous, pluri-folliculate or samaroid, with or without stinging hairs; locules 1–many-seeded. Seeds smooth or verrucose, tuberculate, rarely winged; endosperm present or absent; cotyledons foliaceous, often bilobed and plicate. Pantropics; ca. 70 genera and 1100 species, 10 genera and 48 species in the flora area. Pterygota colombiana Cuatrec. occurs natively in Venezuela in Zulia state, and species of Cola and Abroma are occasionally found cultivated.

Ayenia 249

Key to the Genera of Sterculiaceae 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(3). 7. 8(7). 8. 9(8).

9.

Flowers without petals; fruit woody and 5-folliculate................. 7. Sterculia Flowers with petals; fruit capsular, berry-like, or drupaceous ............... 2 Leaves palmately compound; fruit indehiscent, many-seeded, 10-ribbed ................................................................................................... 5. Herrania Leaves simple; fruits various .................................................................... 3 Petals flattened, spathulate, not appressed to the staminal tube margin ................................................................................................................ 4 Petals cucullate, appressed to the apical staminal tube margin ............ 7 Seeds winged; anthers 1-thecate, reniform; trees .................. 9. Uladendron Seeds not winged; anthers 2-thecate, oblong; trees, shrubs, or vines .... 5 Androgynophore longer than the perianth; capsule cylindric, often spirallike ............................................................................................ 4. Helicteres Androgynophore absent; capsule obovoid, globose, or pyramidal .......... 6 Carpels and styles 5; petals lilac, violet, or rarely white ............ 6. Melochia Carpels and styles 1; petals generally yellow ........................... 10. Waltheria Blade of petals bifid, yellow; fruit woody, indehiscent or with 5 spiral fissures; stamens 15 in 5 groups with 3 anthers each ............... 3. Guazuma Blade of petals entire and red, or absent; fruits various ........................ 8 Trees; fruit drupaceous, ovoid, smooth or ribbed, many-seeded; stamens 10–15 in 5 groups bearing 2 or 3 anthers each .................... 8. Theobroma Shrubs or vines; fruit capsular, globose or subglobose, 5-seeded, forming 5 cocci; stamens 5 .................................................................................. 9 Anthers with 3 thecae; stems smooth, cylindrical; cocci dehiscent; claw of petal with filiform base, longer laminar portion appressed to the staminal tube; blade of petal absent or shorter than the clawed region, claviform, red; shrubs or subshrubs, rarely trees ....................... 1. Ayenia Anthers with 2 thecae; stems aculeate or smooth, cylindrical or angled; cocci dehiscent or indehiscent; claw of petal shorter than blade, purplish, green-yellow, or white; shrubs, subshrubs, or vines ..... 2. Byttneria

1. AYENIA L., Kongl. Svenska Vetensk. Akad. Handl. 17: 23. 1756. Shrubs or erect to prostrate subshrubs, rarely trees. Stems smooth, cylindrical. Leaves alternate, simple, petiolate, membranaceous or rarely coriaceous, with stellate or simple hairs; stipules subulate, caducous; margins serrate, dentate, or entire; venation noticeable abaxially, rarely with a nectary at the base of the main vein. Inflorescences short, axillary. Flowers pedicellate, bisexual, actinomorphic, small. Sepals 5, ovate, basally connate; petals 5, red or pink, mostly clawed, lower portion of the claw filiform and longer than the upper portion, this bilobed, triangular, or rhomboidal and appressed to the staminal tube margin; blade of the petals absent or shorter than the claw, claviform, rarely spathulate, red, glandular. Androgynophore present or absent. Staminal tube urceolate, membranaceous or somewhat fleshy; stamens 5; anthers with 3 thecae, sessile or at the end of the free portion of the filament; staminodes 5, alternate with the petals, glabrous, lobes of the staminodes cucullate and recurved. Ovary 5-carpellate, included in the staminal tube; style simple; stigma capitate, exserted, stigmatic lobes rounded. Capsule 5-coccate, globose or somewhat flattened, small; cocci dehiscent, subwoody,

250

S TERCULIACEAE

pubescent, lacking stinging hooks. Seeds without endosperm, 1 per locule, ovoid, maroon, corrugated, and tuberculate, rarely smooth; cotyledons foliaceous, bilobed, rolled at the hypocotylar axis. Neotropics; 81 species, 3 in Venezuela, 1 of these in the flora area. Ayenia magna L., Syst. Nat. ed. 10, 2: 1247. 1759. Subshrub; leaves ovate, mostly petiolate. Deciduous forests, ca. 50 m; Bolívar (Ciudad

Fig. 211. Ayenia magna

Bolívar). Aragua, Trujillo; Antilles, south in South America to Brazil (Mato Grosso), Bolivia, Paraguay (Chaco). ◆Fig. 211.

Byttneria 251

2. BYTTNERIA Loefl., Iter. Hispan. 313. 1758, nom. cons. Shrubs, erect or climbing subshrubs, or vines. Stems cylindrical or ribbed, smooth or with recurved spines. Leaves alternate, simple, membranaceous or coriaceous, with stellate and/or simple trichomes, often petiolate; stipules subulate, fugacious; margins entire, serrate, or crenate; nectaries 1–5 on the basal veins of the abaxial surface, smooth or with a median longitudinal aperture. Inflorescences axillary, rarely terminal, cymes abbreviated or with well-developed axes. Flowers bisexual, actinomorphic, small. Sepals 5, ovate, connate at the base; petals 5, purple, green-yellow, or white, clawed, the claw cucullate, with a narrow foot and fleshy hood or membranaceous with a wing on each side, appressed to the margin of the staminal tube or on the outer surface; blade of the petals longer than the claw, erect, fleshy, claviform, cylindrical, rarely flat, glabrous or pubescent (the trichomes then glandular). Androgynophore absent. Staminal tube urceolate, fleshy or membranaceous; stamens 5, anthers 2-thecate, sessile or with a short filament, sometimes divergent; staminodia 5, alternate with the petals, lobes surpassing the stamens, recurved, fleshy or membranaceous, glabrous or pubescent. Ovary 5-carpellate, included in the staminal tube; style simple, somewhat longer than the staminal tube, capitate, stigmatic lobes not very evident. Capsule 5-coccate; cocci dehiscent or indehiscent, woody or subwoody, globose, rarely ellipsoid, covered by scattered or congested, persistent or caducous stinging hooks, sometimes with mammiliform projections. Seeds without endosperm, 1 per locule, rugose, verrucose, or smooth; cotyledons foliaceous, bilobed, folded and rolled at the hypocotylar axis. Pantropics; ca. 133 species, 13 in Venezuela, 10 of these in the flora area. Key to the Species of Byttneria 1.

1.

2(1).

2.

3(1). 3.

Erect shrubs or subshrubs 0.4–2 m tall; leaves lanceolate or linear, ca. 7 times as long as wide; venation prominent on both surfaces of the leaf; petiole flat; foliar nectary with a median longitudinal hole, filiform; stems 4- or 5-angled, with prominent ribs ........................................... 2 Vines or prostrate shrubs; leaves wider than above; venation prominent only on the abaxial surface; petiole terete or semiterete, well-defined; foliar nectary apparently smooth or porose, differing from the rest of the midvein by the absence of trichomes, different color, or by being slightly concave; stems usually not angled or ribbed ......................... 3 Leaves rigid, lanceolate, entire or denticulate near the apex, the petiole distinct; stems solid, with prominent yellowish, recurved hooks present on the petiole and the midvein of the blade ................... B. scabra Leaves membranaceous, linear or narrowly lanceolate, the boundary between the blade and the petiole indistinguishable; young stems hollow, unarmed, rarely with very tiny hooks ............................. B. genistella Unarmed vines; mericarps dehiscent, with fine, acute, small hooks; leaf margins entire; leaf blades glabrous or lightly pubescent .................. 4 Armed shrubs; mericarps indehiscent, generally without hooks; leaf margins entire or dentate; leaves uniformly pubescent or with trichomes on the midvein .......................................................................... 6

252

4(3). 4.

5(4).

5.

6(3). 6. 7(6).

7.

8(6).

8.

9(8).

9.

S TERCULIACEAE

Leaves broadly ovate, the base rounded or subcordate; nectary ca. 1 mm long; petals glabrous, the blade flat in cross section ........... B. catalpifolia Leaves ovate, oval-lanceolate, oblong, or oblong-lanceolate, the base rounded to acute; nectary ca. 2 mm long; petals pubescent, the blade rounded in cross section ........................................................................ 5 Leaves matte, attenuate at both apex and base, those on flowering branches commonly shorter and more narrowly lanceolate than those on sterile branches; nectary at the base of midvein .................... B. piresii Leaves glossy, the apex acuminate, the base rounded; leaves on fertile branches similar to ones on sterile branches; nectary between the second or third pairs of lateral veins .......................................... B. uaupensis Leaves coriaceous, glabrous or with a tuft of hairs at the union of the midvein and the secondary veins, margins entire .............................. 7 Leaves membranaceous, evenly pubescent, margins dentate ................ 8 Leaves oval-lanceolate, attenuate at the apex, adaxial surface greener than the abaxial surface; nectary 1, yellow, glossy, porose, close to the secondary veins on abaxial surface; blade of petals claviform, pubescent .............................................................................................. B. obliqua Leaves oval or oblong, subobtuse or obtuse at the apex, concolorous; nectary at the base of the midvein or partially on the petiole; blade of petals long, lanceolate with a row of trichomes on each side .............................................................................................. B. rhamnifolia Nectaries 3–5, on the basal foliar veins; stems and leaves with small stellate trichomes; blade of petals noticeably dilated on the upper portion, acute; fruit globose, woody, covered by mammiliform processes, pointed at the apex, with very tiny hairs .............................. B. divaricata Nectary 1, on the base of the midvein; stems and leaves with small stellate or simple trichomes; blade of petals gradually dilated at the apex or cylindrical, lightly pubescent or pubescent only at the base ......... 9 Nectary with clear irregular pores; blade of petals gradually dilated toward the apex, lightly pubescent; stems and leaves with small stellate trichomes; fruits not known ................................................... B. caripensis Nectary with conspicuous pores; blade of the petals cylindrical, acute, lightly pubescent at the base; stems hollow and ribbed; stems and leaves with simple trichomes; fruits covered with aristate processes ............................................................................................. B. aristeguietae

Byttneria aristeguietae Cristóbal, Bonplandia 4: 39, fig. 20. 1976. Climber; fruits globose, pale green, covered by aristate processes. Open places on granitic formations, lowland forests, 50–200 m; Bolívar (Moitaco), Amazonas (north and south of Puerto Ayacucho). Guárico; Colombia. Byttneria caripensis Cristóbal, Bonplandia 6: 126, figs. 1, 2. 1989. —Espina. Subshrub, flowering branches delicate; leaves ca. 7 cm long, with tiny hooks. Riparian habitats, 50–100 m; Bolívar, (Laguna de

Caripo near mouth of Río Suapure). Endemic. Byttneria catalpifolia Jacq., Pl. Hort. Schoenbr. 1: 21, t. 46. 1797, “Buetneria catalpaefolia.” Mexico, Central America, West Indies, Trinidad, Colombia, Venezuela, Ecuador, Peru, northeastern Brazil, Bolivia; 2 subspecies, 1 in Venezuela. B. catalpifolia subsp. catalpifolia Vine. Leaves well developed, wide, usually petiolate; inflorescences terminal; flow-

Byttneria 253

ers fragrant, white. Evergreen lowland forests, 200–300 m; Amazonas (Río Ugueto). Widespread in northern Venezuela; Mexico, Central America, West Indies, Colombia, Ecuador, Peru, northeastern Brazil, Bolivia. Byttneria divaricata Benth., J. Bot. (Hooker) 4: 124. 1842. Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazon basin; 2 varieties, 1 in Venezuela. B. divaricata var. divaricata. —Quejido. Byttneria venezuelensis Steyerm., Acta Bot. Venez. 3(1–4): 183. 1968. Climber. Flowers small, axillary, condensed, purple. Evergreen lowland and riparian forests, sometimes in secondary vegetation, near sea level to 200 m; Delta Amacuro (Río Acure between La Margarita and Puerto Miranda). Zulia; other distribution as in species. ◆Fig. 212.

Byttneria genistella Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 333. 1862. Annual subshrub; branches simple, fewleaved, flowering branches with very reduced leaves, sessile. Lowland savannas, 50–300 m; Bolívar (Altamira, Caicara, Río Paragua), Amazonas (Cerro Ualipano, Puerto Ayacucho, Raudal de Maipures, upper Río Orinoco, Río Samariapo). Guárico, Monagas; Colombia, French Guiana, northern and central Brazil. ◆Fig. 213. Byttneria obliqua Benth., J. Bot. (Hooker) 4: 124. 1842. Shrub 1–3 m tall; flowers purple; fruits woody, covered with clustered, truncate, cylindrical processes. Seasonally flooded forests along black-water rivers, 50–100 m; Amazonas (close to Amazonas in Colombian Río Guaviare near Río Atabapo). Colombia, Amazonian Brazil. ◆Fig. 214.

Fig. 212. Byttneria divaricata var. divaricata

254

S TERCULIACEAE

Fig. 213. Byttneria genistella

Fig. 214. Byttneria obliqua

Fig. 215. Byttneria piresii

Guazuma 255

Byttneria piresii Cristóbal, Bonplandia 4: 380, fig. 96. 1976. —Bejuco de lora. Vine; flowering branches generally leafy, shorter and narrower than those in sterile branches. Evergreen lowland forests, 100– 300 m; Amazonas (La Esmeralda). Amazonian Brazil. ◆Fig. 215. Byttneria rhamnifolia Benth., Hooker’s J. Bot. 3: 164. 1851. Shrub; flowers pendent, purple. Riparian habitats, 50–200 m; Bolívar (Los Icacos ca. 10 km from Ciudad Bolívar, middle Río Orinoco), Amazonas (Isla Ratón, Puerto Ayacucho, banks of Río Orinoco). Widespread in northern South America to northern Argentina.

Byttneria scabra L., Syst. Nat. ed. 10, 2: 939. 1759. Rhizomatous subshrub, few-leaved; inflorescences broad, terminal, paniculiform. Moist savannas, along rivers, 50–900 m; Bolívar (Río Ocoine at southeastern base of Auyán-tepui, San Ignacio, Santa Elena de Uairén), Amazonas (Guachapana above Santa Bárbara del Orinoco, Puerto Ayacucho). Anzoátegui, Barinas; Colombia, Trinidad, northern South America to Uruguay. Byttneria uaupensis Spruce ex K. Schum. in Mart., Fl. Bras. 12(3): 93. 1886. Unarmed liana. Disturbed lowland forests, 100–300 m; Bolívar (below Cerro Abismo, Río Samay in Río Icabarú basin). Colombia (Río Vaupés), Guyana, Brazil.

3. GUAZUMA Mill., Gard. Dict. Abr. ed. 4, 596. 1754. Unarmed trees or shrubs. Leaves petiolate, simple, alternate, distichous, ovate or oval-lanceolate, usually the base asymmetric, membranaceous or subcoriaceous, subglabrous to tomentose, with stellate or rarely simple hairs; venation prominent on the abaxial surface; stipules small, subulate, deciduous. Cymes wide or contracted, axillary or terminal. Flowers pedicellate, bisexual, actinomorphic, small. Calyx 2- or 3-lobed, the sepals ovate, concave, generally reflexed, green-yellow; petals 5, yellow or pink, clawed, the claw cucullate with 5 prominent veins in the central third, with a slit on the margin that joins at the free portion of the staminal process; blade of petals membranaceous, bifid, with linear branches, longer than claw, undulate. Staminal tube campanulate, stamens 15 in 5 groups with 3 anthers each, with a free small portion at the end, opposite the petals; staminodia 5, triangular, recurved, alternate with petals. Pistil included on the stami-

Fig. 216. Guazuma ulmifolia var. ulmifolia

256

S TERCULIACEAE

nal tube; ovary 5-carpellate, pluriovulate, sessile or on a short gynophore, style simple, stigmatic branches 5, filiform. Capsule many-seeded, small, woody, verrucose, indehiscent or with 5 slits, or dehiscent and covered by long plumose structures. Seeds small, pale maroon, irregularly compressed; endosperm scarce; cotyledons foliaceous, suborbicular, rolled at the hypocotylar axis. Neotropics; 3 species, 1 in Venezuela. Guazuma ulmifolia Lam., Encycl. 3: 52. 1789. Tree. Mexico, Central America, Antilles, northern South America south to northern Argentina; 2 varieties, both in the flora area. Key to the Varieties of G. ulmifolia 1. Fruits flattened to subglobose, partially loculicidally dehiscent by 5 slits, aromatic, the adaxial surface glossy, the abaxial surface glabrous to tomentellous ............................. var. tomentella 1. Fruits ovoid to subglobose, indehiscent, nonaromatic, the adaxial surface matte to subglossy, the abaxial surface with scattered to congested stellate hairs ...................................... var. ulmifolia G. ulmifolia var. tomentella K. Schum. in Mart., Fl. Bras. 12(3): 81. 1886. —Guácimo, Guácimo montañero, Guácimo negro, Luma. Semideciduous forests, sometimes secondary forests, possibly cultivated, 50–500

m; Delta Amacuro (east of Río Grande northeast of El Palmar, 13 km east-southeast of Sierra Imataca), Bolívar (Altiplanicie de Nuria, 25 km east of El Dorado, 20–35 km southwest of El Manteco, La Paloma in Río Cuyubini basin, Río Asa above Raudal Cotúa, Río Cuyubini, Temblador), Amazonas (Gavilán, San Juan de Manapiare). Central America, Antilles, northern South America to northeastern Argentina. G. ulmifolia var. ulmifolia. —Guácimo, Guácimo blanco. Deciduous to evergreen lowland and riparian forests, 50–300 m, Bolívar (Caicara, between Hato El Manguito and Río Oronata 7 km west of El Manteco, Moitaco, banks of Río Orinoco), Amazonas (upper Río Orinoco). Widespread in northern Venezuela; Mexico, Central America, northern South America to northeastern Argentina. ◆Fig. 216. An infusion of the bark of Guazuma ulmifolia var. ulmifolia is used as a medicine for dysentery.

4. HELICTERES L., Sp. Pl. 963. 1753. Shrubs erect, unarmed, rarely trees. Leaves simple, sometimes lobed, serrate, with simple, glandular, stellate, and dendritic hairs, petiolate; stipules subulate, rarely laciniate, caducous or persistent. Cincinni simple, rarely double, 2–manyflowered, opposite the leaves, axillary or terminal. Flowers bisexual, often showy, erect and subactinomorphic or oblique to the stem and apparently zygomorphic with the corolla asymmetric; nectaries at the base of the pedicels, 1–many per flower, rounded, black, glossy, or absent; bracts 2 per flower. Calyx gamosepalous, tubular, inflated or campanulate, ferruginous, cinereous, green or red, 5-dentate or bilabiate with 3 upper teeth and 2 lower teeth, rarely the teeth all fused or irregularly opening during anthesis; nectary basal on the inner face, margin 2-dentate or irregular; petals 5, free, red, purplish blue, white, or green, longer than the calyx or included, clawed, the claw somewhat fleshy and complex at the apex often with wing-like processes; petals subequal or the upper 2 more complex and the 3 lower subequal or different; blade spathulate, membranaceous, rarely absent. Androgynophore erect or incurved, often surpassing the perianth or included.

Helicteres 257

Androecium and gynoecium parallel to the androgynophore or inclined inward; stamens (6–)10(–100); filaments free or partially fused; anthers 2-thecate, the thecae divergent, aligned, erect, or sigmoid, separate or congested on a small head; staminodia 5, spathulate or lanceolate, free or fused at the base of staminal tube in its inner side. Ovary 5-carpellate, the carpels multiovulate; style simple, exserted. Capsule cylindric, sometimes woody, dehiscent or indehiscent, spiraled or erect, glabrous to shortly woolly. Seeds small, irregularly compressed, pale brown, lightly tuberculate, sometimes with a wing along the raphe; cotyledons foliaceous, bilobed, reflexed and plicate, surrounding the hypocotylar axis; endosperm scarce. Pantropics; ca. 50 species (38 American), 6 in Venezuela, 5 of these in the flora area. Key to the Species of Helicteres 1.

1.

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

4.

Androgynophore incurved, ca. 5 times longer than the calyx; nectaries 1–3, glossy, black, located between the pedicels; flowers oblique, apparently zygomorphic (corolla asymmetric); calyx tubular-campanulate, cinereous, somewhat fleshy .......................................................... 2 Androgynophore erect, ca. 2 times longer than calyx; inflorescences without nectaries; flowers subactinomorphic; calyx tubular, red or red-orange, inflated, membranaceous .................................................. 3 Petals linear, green; calyx with 5 strong veins, teeth grouped in 2 lips ........................................................................................... H. baruensis Petals widely spathulate, red, showy; calyx without veins, with 5 teeth separated ................................................................................. H. brevispira Petals white, included in the tubular calyx, bracts linear; stamens 6 .............................................................................................. H. heptandra Petals red, longer than calyx; bracts ovate or lanceolate; stamens 6, 7, or 10 ............................................................................................................ 4 Stems, leaves, and calyx hispid, with large dendritic hairs; leaves distichous, densely grouped, base cordate, asymmetric, venation depressed on the adaxial surface and less noticeable on the abaxial side; inflorescence hidden by leaves and red bracts; stamens 6 or 7 ..... H. pentandra Stems, leaves, and calyx with stellate hairs, short, not dendritic; leaves more loosely spaced, with base rounded and symmetric, venation not depressed on the adaxial surface and not noticeable at the abaxial surface, not obscuring the inflorescence; stamens 10 ........... H. guazumifolia

Helicteres baruensis Jacq., Enum. Syst. Pl. 30. 1760. Shrub 2–3 m tall. Savannas and scrub, 50–400 m; Bolívar (Altiplanicie de Nuria, El Manganeso south of El Palmar, Laguna Larga 8 km southeast of Upata, San Martín de Turumbán southwest of Tumeremo, Upata). Carabobo, Distrito Federal, Falcón, Lara, Nueva Esparta, Sucre, Zulia; Mexico, Central America, Trinidad, Colombia, Guyana, Suriname, northeastern Brazil. ◆Fig. 217.

Helicteres brevispira A. St.-Hil., Fl. Bras. Merid. 1(7): 274, pl. 54. 1828. Shrub 2–4 m tall. Forests on granitic outcrops, 50–200 m; Amazonas (around Puerto Ayacucho). Colombia, Brazil, Bolivia, Paraguay. Helicteres guazumifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 304. 1821 [1822]. —Pepino, Tornillo. Shrub 1–4 m tall, showy at flowering. River banks, savannas, often weedy, 50–200

258

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Fig. 217. Helicteres baruensis

Fig. 218. Helicteres guazumifolia

Fig. 219. Helicteres pentandra

Herrania 259

m; Bolívar (road between Caicara and El Burro, Ciudad Bolívar, El Tigre, northwest of El Manteco along shores of Lago Guri, between La Guavina and Maniapure, Paso de Caruachi, Puerto Ordaz, Río Cuchivero, Río Parguaza, Río Yuruari, between San Félix and Los Castillos de Guayana, San Pedro de las Dos Bocas), Amazonas (Puerto Ayacucho, Puerto Venado, Raudal de Atures, Río Cataniapo, Río Samariapo). Anzoátegui, Apure, Aragua, Barinas, Cojedes, Miranda, Monagas, Guárico, Portuguesa, Sucre, Yaracuy, Zulia; widespread in Neotropics. ◆Fig. 218. Helicteres heptandra L.B. Sm., Contr. Gray Herb. 117: 36, pl. 2, figs. 48, 49. 1937. Helicteres brachypetala Leane, Bradea 4:

56, fig. 1. 1984. Shrub to 2.5 m tall; flowers showy, orange. Deciduous forests, 200–300 m; Bolívar (islands in Represa Guri). Northeastern Brazil. Helicteres pentandra L., Mant. Pl. 2: 294. 1772. —Tornillo. Helicteres pronifolia Rich., Actes Soc. Hist. Nat. Paris 1: 11. 1792. Helicteres bracteosa Mart., Flora 22(Biebl. 1): 15. 1839. Shrub. Evergreen to semideciduous lowland forests and savannas, 50–400 m; Bolívar (base of Auyán-tepui, Guacaima, Hato la Vergareña, La Paragua, 10 km from Los Castillos de Guayana). Monagas; southern Colombia, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 219.

5. HERRANIA Goudot, Ann. Sci. Nat. Bot. sér. 3, 2: 230. 1844. Lightia R.H. Schomb., Linnaea 20: 757. 1847. Brotobroma H. Karst. & Triana, Nuev. Esp. Fl. Neogranad. 1. 1854. Small unarmed, erect trees 2–9 m tall, 5–30 cm diameter, branched or singlestemmed. Leaves apical on the stems, alternate, large, palmatisect, often ferruginous-tomentose; leaflets 4–9, pinnatinerved, obovate-lanceolate or oval-lanceolate, acute, serrate, lobate to pinnatisect, rarely entire, with stellate hairs, scabrous or smooth, often yellowish, rarely glabrous, mostly thick-petiolate; stipules caducous or persistent, 1.5–5 cm long, tomentose. Flowers bisexual, actinomorphic, in fewor many-flowered fascicles, cauliflorous, generally arranged at the bottom or at apex of trunk. Calyx valvate, 3–5-dentate, patelliform or subcymbiform; petals (4)5(6), purple, rarely white, clawed, the claw cucullate, striate, often papillose or granulate, appressed to the staminal tube margin hiding the stamens, 5–12 mm × 4–9 mm, blade (ligule) reflexed, pendulous, glabrous, membranaceous, linear or filiform, 15–180 mm × 1–3 mm. Staminal tube short; stamens in 5 groups with 2 or 3 stamens each; anthers 2-thecate; staminodia 5, alternate with the petals, showy, lanceolate or elliptic, apex acute or tridentate, petaloid, reflexed, hiding the stamens and the claw of petals, granulose or verrucose on both faces, purplish, red or yellowish, rarely white, 6–25 mm × 5–10 mm. Fruit yellowish or green-yellow, capsular or berry-like, ellipsoid, ovoid, or globose, 8–14 cm long, with 5 noticeable primary ribs and 5 secondary ones that are generally finer, intercostal veins very noticeable. Seeds 25–100 or more, covered by mucilaginous pulp. Costa Rica, Nicaragua, Colombia, Venezuela, Suriname, French Guiana, Peru, Brazil; 20 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Herrania 1.

Leaf segments dentate, teeth acute with concave sides, at the end of a pilose, prolonged appendix of the vein 1–2 mm long; hairs setiform, > 1 mm long or hirsute on both sides, but more abundant at the abaxial surface ................................................................................. H. camargoana

260

1.

S TERCULIACEAE

Leaf segments pinnately lobed, basal lobes acute, the sinuses as deep as 1/3–1/2 the distance to the midvein, adaxial surface with scarce hairs but denser along the main veins, abaxial surface with evenly dispersed hairs ........................................................................... H. lemniscata

Fig. 220. Herrania lemniscata

Melochia 261

Herrania camargoana R.E. Schult., Bot. Mus. Leafl. 14: 120, pl. 29, 32. 1950. Small tree. Evergreen lowland forests, 100–200 m; Amazonas (upper Río Negro basin). Colombia, Brazil. Herrania lemniscata (M.R. Schomb.) R.E. Schult., Caldasia 2: 13. 1943. —Lightia lemniscata M.R. Schomb., Rep. Brit.

Assoc. Advancem. Sci. 2: 71. 1844. —Cacao montañero. Small tree; fruits edible. Evergreen lowland to lower montane forests, near sea level to 1100 m; Delta Amacuro (near mouth of Río Orinoco), Bolívar (Cerro Pakaraima, Río Pacairao near Santa Teresita de Kavanayén, Salto de Pará in middle Río Caura). Colombia, Suriname, French Guiana. ◆Fig. 220.

6. MELOCHIA L., Sp. Pl. 674[“774”], 1753, nom. cons. Visenia Houtt., Nat. Hist. 2(8): 308. 1777. Riedlea Vent., Mem. Cl. Sci. Math. Inst. Natl. France 8: 2. 1807, nom. illeg. Mougeotia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 316, 1821.[1823]. Anamorpha H. Karst. & Triana, Nuev. Esp. Fl. Neogranad. 9. 1854. Physocodon Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(1): 212. 1858. Unarmed shrubs or subshrubs, erect, climbing, or prostrate, rarely trees. Leaves petiolate, simple, alternate, oval, ovate, suborbicular, lanceolate, or linear, membranaceous, serrate, dentate, or subentire, glabrous to tomentose or sericeous, hairs stellate, simple, bifurcate and glandular or often mixed; venation on the lower surface prominent, secondary veins sometimes parallel to the margin; stipules triangular, subulate, oval, or suborbicular, caducous or persistent. Inflorescences few- to many-flowered, axillary, opposite the leaves, or terminal, broad or contracted in sessile or pedunculate glomeruli. Flowers pedicellate or sessile, bisexual, actinomorphic, small, generally heterostylous. Calyx persistent, sometimes accrescent, 5-dentate, teeth triangular or largely acuminate, sinus acute or obtuse then teeth small and distant; corolla contorted, the petals 5, obovate or spathulate, free or fused at the base of the staminal tube, lilac, pink, purplish, yellow, or white. Androgynophore absent. Stamens 5; filaments fused at the base or along them; anthers dithecate, extrorse; staminodia absent, rarely 5, dentiform, alternating with the stamens. Ovary generally 5-carpellate, carpels fused, biovulate; styles 5, filiform, free or fused at the base, stigmatic papillae toward the apex. Capsule small, pubescent, subglobose, ovoid, or pyramidal, with wing-like, acute to rounded processes; dehiscence loculicidal, septicidal, or both, often disintegrating at maturity. Seeds 1 or 2 per locule, pale brown to black, smooth, lateral sides flattened, dorsal side convex, with endosperm; cotyledons suborbicular, flattened. Pantropics; ca. 56 species, mostly American, 16 in Venezuela, 9 of these in the flora area. Key to the Species of Melochia 1. 1. 2(1). 2. 3(1).

Fruits pyramidal; teeth of calyx largely triangular, sinus acute ............ 2 Fruits globose or subglobose; teeth of calyx triangular or not ................ 3 Leaves ovate, tomentose; petals violaceous .............................. M. tomentosa Leaves obovate, obovate-lanceolate, or oblong, obtuse, glabrous or subglabrous; petals white, sometimes pink ................................ M. parvifolia Fruit completely septicidal, fruit falling apart, cocci with long, simple hairs; leaves ovate or oval-lanceolate, acute, lightly pubescent, hairs generally simple, scattered; flowers in dense axillary glomeruli; teeth of calyx triangular, sinus acute ............................................... M. nodiflora

262

3.

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

5.

6(4). 6. 7(6). 7. 8(7). 8.

S TERCULIACEAE

Fruit at first loculicidal, all along the dorsal suture, usually later septicidal; flowers in dense leafy nodes or laxer clusters; teeth of calyx triangular to distantly spaced, the sinus acute or obtuse ....................... 4 Teeth of calyx largely triangular, sinus acute .......................................... 5 Teeth of calyx distant, sinus wide, obtuse ................................................ 6 Inflorescences dense in leafy nodes at the naked end of branches; plants villous, stellate hairs short and setiform hairs; leaves to 8.5 × 4 cm; subshrubs from prostrate to erect 0.25–1 m; petals purplish or pink ...................................................................................................... M. villosa Inflorescences axillary, wide; leaves to 15 × 8 cm, stellate hairs dispersed, more abundant at the abaxial surface; climber to 4 m tall, petals white or yellowish white .................................................... M. ulmifolia Leaves linear, glabrous or subglabrous with stellate, minute hairs, denticulate, teeth remote; inflorescence wide, terminal ........ M. graminifolia Leaves ovate or oval, rarely lanceolate, serrate; inflorescence axillary ..... 7 Leaves velutinous-cinereous, covered by very short, stellate hairs, oval; peduncle to 2 cm long, pedicel 3–6 mm long ............................ M. arenosa Leaves subglabrous, hairs simple, scattered, ovate to lanceolate; flowers congested along branches ..................................................................... 8 Fruits with many capitate, glandular hairs ........................... M. manducata Fruits with simple hairs, setiform and stellate, very small ..... M. melissifolia

Melochia arenosa Benth., J. Bot. (Hooker) 4: 127. 1842. —Orégano de playa. Melochia cinerea A. St.-Hil. & Naudin, Ann. Sci. Nat. Bot. sér. 2, 18: 35. 1842. Riedlea littoralis Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 73. 1845; Riedlea littoralis K. Schum. in Mart., Fl. Bras. 12(3): 44. 1886, nom. superfl., “Riedleia.” Melochia llanorum Cuatrec., Revista Acad. Colomb. Ci. Exact. 8: 487. 1952. Shrub to 2.5 m tall, grayish; flowers pink. Seasonally flooded savannas or in open places along river banks, sometimes weedy, near sea level to 500 m; Delta Amacuro (Orocoima in front of Isla El Toro), Bolívar (La Paragua, near Maripa, mouth of Río Paragua, mouth of Río Parguaza, Río Taratá at km 147–148 south of El Dorado, 6 km northeast of Tumeremo), Amazonas (Isla Ratón, around Puerto Ayacucho, Puerto Samariapo, Raudal de Atures savannas near San Juan de Manapiare). Guárico, Monagas; Cuba, Colombia, Guyana, Brazil, Bolivia, Paraguay. ◆Fig. 221. Melochia graminifolia A. St.-Hil., Fl. Bras. Merid. 1: 160. 1825. —Ojo de Grillo.

Quamoclit rodriguesii J.I. Falcão, Publ. Inst. Nac. Pesq. Amazonica Bot. 25: 9, fig. 2. 1968. Subshrub to 1 m tall. Seasonally flooded savannas, 50–200 m; Bolívar (savanna of La Paragua, Potrerito on Río Paragua, Río Paragua near Chorrera de Aguacanta, mouth of Río Paragua), Amazonas (Raudal de Atures, Samariapo, San Juan de Manapiare and Río Manapiare basin, savannas of Santa Bárbara del Orinoco). Colombia, Guyana, Brazil, Bolivia, northern Paraguay. Melochia manducata C. Wright, Anales Acad. Ci. Med. Habana 5: 241. 1868. Melochia glandulifera Standl., Contr. U.S. Natl. Herb. 23(3): 803. 1923. Subshrub. Seasonally flooded areas, sandy soils, along roads, near sea level to 100 m; Delta Amacuro (between Tucupita and La Horqueta). Barinas, Portuguesa, Zulia; Mexico, Central America, Greater Antilles, Colombia, Ecuador, Peru, Amazonian Brazil. Melochia melissifolia Benth., J. Bot. (Hooker) 4: 129. 1842, “melissaefolia.” Melochia concinna Miq., Linnaea 22: 467. 1849.

Melochia 263

Shrub; stem red. Open well-drained places, sometimes weedy, 50–200 m; Amazons (El Venado, Samariapo). Costa Rica, Panama, Suriname, French Guiana, Brazil, Africa. Melochia nodiflora Sw., Prodr. 97. 1788. Melochia carpinifolia J.C. Wendl., Bot. Beob. 52. 1798. Riedlea urticifolia Turcz., Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 31(1): 209. 1858, “Riedleja urticaefolia.” Melochia conglobata Sessé & Moc., Pl. Nov. Hisp. 1: 106. 1889. Subshrub; flowers pink, yellow, or white. Open places, 50–200 m; Bolívar (El Cuchivero, Moitaco). Miranda; Mexico, Central America, West Indies, Brazil (Ceará, Río de Janeiro). ◆Fig. 222. Melochia parvifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 325. 1821 [1823]. Cuba, Colombia, Venezuela, Guyana, Brazil, Paraguay, Argentina; 2 varieties, 1 of these in Venezuela and the flora area. M. parvifolia var. parvifolia. —Bretónica. Melochia scordiifolia Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(1): 207. 1858. Melochia obovata Urb., Symb. Antill. 9: 233. 1924. Shrub 1.5–2 m tall. Savannas, open places, granitic outcrops, weedy areas, 30– 300 m; Delta Amacuro (Boca de Macareo, Caño de Uricoa, Castillos de Guayana, Sacupana, between Tucupita and Las Mulas), Bolívar (Ciudad Bolívar, near Guasipati and El Manteco, Hato Guanaraparo, Hato La Vergareña, Isla de Anacoco on Río Cuyuní, La Paragua, La Peluca, Santa Isabel), Amazonas (port of Puerto Ayacucho). Distribution as in species. Melochia tomentosa L., Syst. Nat. ed. 10, 2: 1140. 1759. Shrub; leaves cinereous; veins straight and parallel to the margin. Southern U.S.A., Mexico, Central America, West Indies, Colombia, south in South America to northern Brazil; 4 varieties, 3 in Venezuela, 2 of these in the flora area.

Key to the Varieties of M. tomentosa 1. Leaf blade typically 1–3 × 2 cm, petiole < 1 cm long; stems much more branched and canescent than other variety .......... ..................................... var. frutescens 1. Leaf blade typically 2–6 × 0.7–4 cm, petiole generally > 1 cm long ........................ ..................................... var. tomentosa M. tomentosa var. frutescens DC., Prodr. 1: 490. 1824. —Bretónica. Deciduous forests, 20–50 m; Bolívar (between San Félix and Puerto Ordaz). Mérida, Lara, Nueva Esparta, Sucre, Trujillo; Mexico, Central America, West Indies, Colombia. M. tomentosa var. tomentosa Savannas, scrub, sometimes cultivated, 20–50 m; Bolívar (Ciudad Bolívar). Carabobo, Distrito Federal, Falcón, Mérida. Southern U.S.A., Mexico, Central America, West Indies, northern South America. We follow A. Goldberg (Contr. U.S. Natl. Herb. 34(5): 329. 1987), in recognizing two varieties of Melochia tomentosa from the flora area. There are too few specimens, however, to provide a clear differentiation between them. Melochia ulmifolia Benth., J. Bot. (Hooker) 4: 129. 1841. Riedlea ramuliflora Miq., Linnaea 19: 132. 1847, “Riedleia.” Riedlea dichotoma Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(1): 208. 1858, “Riedleja.” Melochia benthamii K. Schum. in Mart., Fl. Bras. 12(3): 40, pl. 27. 1886. Melochia carthaginensis Willd. ex K. Schum. in Mart., Fl. Bras. 12(3): 40. 1886. Subshrub. 50–100 m; Bolívar (La Unión). Colombia, Suriname, French Guiana, Brazil. ◆Fig. 223. Melochia ulmifolia has only been found once in Venezuela. Melochia villosa (Mill.) Fawc. & Rendle, Fl. Jamaica 5: 165. 1926. —Sida villosa Mill., Gard. Dict. ed. 8. 1768. Melochia hirsuta Cav., Diss. 6: 323, pl. 175, fig. 1. 1788.

264

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Fig. 221. Melochia arenosa

Fig. 222. Melochia nodiflora

Fig. 223. Melochia ulmifolia

Sterculia 265

Riedlea heterotricha Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(1): 211. 1858, “Riedleja.” 3 varieties, 2 in Venezuela, both in the flora area.

It is difficult to distinguish var. tomentosa from the typical variety because of the gradual variation in the amount of hair type. M.

Key to the Varieties of M. villosa 1. Leaves and stems grayish, tomentose or villous-tomentose, with stellate hairs and long simple hairs, more abundant on the main veins ....... var. tomentosa 1. Leaves and stems hirsute to subglabrous, mainly with simple hairs .......................................... var. villosa M. villosa var. tomentosa (K. Schum.) Goldberg, Contr. U.S. Natl. Herb. 34(5): 287. 1967. —Melochia hirsuta var. tomentosa K. Schum. in Mart., Fl. Bras. 12(3): 47. 1886. Flooded and rocky savannas, 50–200 m; Bolívar (expected). Apure; Brazil, Paraguay, northeastern Argentina.

villosa var. villosa. —Bretónica morada, Hierba San Juan. Savannas, river banks, scrub, savannas surrounding Mauritia palm swamps, pastures, weedy areas, 50–900 m; Delta Amacuro (between Castillos de Guayana and Piacoa), Bolívar (northwest of El Manteco, Encrucijada de El Pao, Hato de Nuria east of Miamo, Hato Terecay Isla del Cazabe on Río Paragua, La Prisión, Lago Guri, Santa Elena de Uairén, Tunaca south of Caicara, Túriba, near Upata), Amazonas (La Esmeralda, near Puerto Ayacucho, upper Río Ventuari at Sabana del Oso, road to Samariapo, San Carlos de río Negro basin of Río Manapiare, Santa Bárbara del Orinoco). U.S.A., Mexico, Central America, West Indies, south in South America to northern Paraguay.

7. STERCULIA L., Sp. Pl. 1007. 1753. Trees to 40 m tall, often with tabular projections at base of trunk; branches robust to the apices where the leaves and inflorescences are grouped, basally with prominent leaf scars and axillary branches with 1 or more caducuous prophylls. Leaves large, petiolate, simple, elliptic or ovate, 3–5-lobed or palmately compound, surface smooth or bullate, stipules fugaceous, generally only accompanied by foliar primordia, the upper leaf surface pubescent to glabrous, the lower leaf surface homotrichous or heterotrichous (with trichomes of one size or two different sizes), trichomes stellate; venation generally prominent. Inflorescences axillary, paniculate, broad, many-flowered or rarely few-flowered, axes with thick glandular trichomes at the base, often reddish. Flowers small, pedicellate, diclinous, unisexual (plants monoecious or dioecious), actinomorphic, with a single perianth whorl. Calyx to 20 mm long, campanulate or urceolate, 5-dentate, longer than the sexual whorls, sepals shortly connate at the base, the inner face generally with a scaly appendage located more or less in the middle and with trichomes of different kinds, stellate, others glandular (vermiform), whose length and distribution characterizes some of the species, outer face glandular; pistillate flowers with a short, curved androgynophore; ovary spherical, pilose, 5-carpellar with the carpel united, but separating in fruit, the carpels 2–many-seeded; style simple, stigma capitate, lobed, fleshy, the rudimentary androecium surrounding the base of the ovary; staminate flowers with a straight or curved androgynophore, glabrous or pubescent, the staminal tube short, stamens 10–15, sessile or subsessile, anthers 2thecate, tightly grouped and forming a headlike structure that encloses the rudimentary ovary. Fruit generally woody, large, pendulous, 5 follicles or fewer by abortion, 1–many-seeded, dehiscent along the suture, inner surface with rigid simple or

266

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stellate trichomes, often urticating; seeds pendulous, smooth, episperm hard; embryo straight, inverted; cotyledons fleshy, accompanied by a variable amount of endosperm. Pantropics; ca. 200 species, ca. 8 in Venezuela, 6 of these in the flora area. Key to the Species of Sterculia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5.

Leaves palmately lobed ................................................................... S. apetala Leaves not palmately lobed, margins entire or crenate .......................... 2 Inflorescence with < 10 flowers ................................................. S. abbreviata Inflorescence with > 10 flowers ................................................................ 3 Indument of the lower leaf surface with trichomes of 2 sizes; leaf margins irregularly crenate ......................................................... S. amazonica Indument of the lower leaf surface with trichomes of 1 size; leaf margins entire ...................................................................................................... 4 Indument of the lower leaf surface dense, difficult to see the individual trichomes .................................................................................... S. pruriens Indument of the lower leaf surface moderately dense, easy to see the individual trichomes ................................................................................. 5 Interior base of the calyx glabrous .................................................... S. kayae Vermiform (glandular) trichomes present at the interior base of the calyx .................................................................................................. S. rugosa

Sterculia abbreviata E.L. Taylor ex Mondragón, Acta Bot. Venez. 28(1): 136. 2005. —Cojón de Berraco, Pica-pica. Tree 10–30 m tall; perianth red with pale yellow tips and margins; inflorescence with < 10 flowers. Nonflooded riparian forests, near sea level to 200 m; Delta Amacuro (Curiapo), Amazonas (San Pedro del Cataniapo). Aragua, Mérida, Miranda. Sterculia amazonica E.L. Taylor ex Mondragón, Acta Bot. Venez. 28(1): 138. 2005. Tree 20–40 m tall; short vermiform trichomes in the fruit, hypophyll with trichomes of two different sizes. Evergreen lowland forests, 100–300 m; Bolívar (Río Adaua in upper Río Caura basin, Río Tabaro in Río Nichare basin), Amazonas (Río Ocamo). Endemic. Sterculia apetala (Jacq.) H. Karst, Fl. Columb. 2: 35, t. 118. 1862. —Helicteres apetala Jacq., Enum. Syst. Pl. 30. 1760. —Camaruco. Tree 15–35 m tall; leaves palmately lobed. Semideciduous lowland forests, edge of Mauritia palm swamps, also planted in and around towns, near sea level to 100 m;

Delta Amacuro (10 km northeast of San Félix), Bolívar (Ciudad Bolívar, Morichal San Isidro). Widespread in northern Venezuela; Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 224. Sterculia kayae P.E. Berry, nom. nov. pro Sterculia parviflora (Ducke) E.L. Taylor, Mongr. Syst. Bot. Missouri Bot. Gard. 45: 1259. 1993, nom. illeg., non Sterculia parviflora Roxb. & G. Don 1831. —Sterculia pruriens var. parviflora Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 130. 1925. —Guayare, Zapatero. Tree to 25 m tall; flowers reddish in the center, petal tips tan. Evergreen lowland to lower montane forests, nonflooded riparian forests, 50–600 m; Delta Amacuro (Caño Araguabisi, 22 km southwest of Wausa near Caño Cariabo), Bolívar (Río Urbani near Las Trincheras, Serranía de los Pijiguaos), Amazonas (upper Río Baría, Río Coromoto at Tobagán de la Selva, Río Sipapo, 4 km east of San Carlos de Río Negro, San Pedro del Cataniapo, base of Sierra de la Neblina). Colombia, Brazil. This species is renamed after Kay Yatskievych, key editor for this flora series.

Sterculia 267

Fig. 224. Sterculia apetala

Fig. 225. Sterculia rugosa

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Fig. 226. Sterculia pruriens

Sterculia pruriens (Aubl.) K. Schum. in Mart., Fl. Bras. 12(3): 8. 1886. —Ivira pruriens Aubl., Hist. Pl. Guiane 695, t. 279. 1775. Tree to 30 m tall; flowers yellowish. Evergreen lowland forests, gallery forests, 100–

300 m; Delta Amacuro (Río Toro), Bolívar (20–35 km southwest of El Manteco on road to San Pedro de las Dos Bocas, east of El Palmar on road to La Tigra, Represa Guri east of Ciudad Piar, Río Asa, Río Urbani near Las Trincheras, near Salto Pará),

Theobroma 269

Amazonas (Río Mawarinuma at base of Sierra de la Neblina). Mérida, Miranda; French Guiana, Ecuador, Peru, Brazil. ◆Fig. 226. Sterculia rugosa R. Br. in Benn., Pl. Jav. Rar. 229. 1844. —Guaraunera, Kerepo-

yek (Arekuna), Macagua. Tree to 35 m tall; leaves coriaceous; fruits reddish. Tall evergreen lowland to lower montane forests, 100–600 m; Bolívar (Cerro Paují, km 30 south of el Dorado, Urimán), Amazonas (headwaters of Caño Yagua). Colombia, Guyana, Ecuador, Brazil. ◆Fig. 225.

8. THEOBROMA L., Sp. Pl. 782. 1753. Cacao Mill., Gard. Dict. Abr. ed. 4, 1754. Tribroma O.F. Cook, J. Wash. Acad. Sci. 5: 288. 1915. by Carmen L. Cristóbal and Paul E. Berry Unarmed trees or shrubs, branches whorled. Leaves alternate, simple, petiolate, entire or sinuate or dentate near the apex, pinnately or palmately veined, oval, oblong, or ovate, base rounded, subcordate, or cuneate, sometimes asymmetric, apex acute or apiculate, chartaceous or coriaceous, often the abaxial surface covered by long, or short, hyaline, and red-stellate hairs intermixed; stipules caducous or persistent. Cymes at the leaf axil or usually cauliflorous on trunk prominences or thick branches. Flowers pedicellate, bisexual, actinomorphic, purplish, reddish, or yellowish. Calyx valvate, sepals 5, connate at the base; petals 5, clawed, the claw cucullate, ± fleshy with prominent veins and the end appressed to the border of the staminal tube, hiding the stamens; blade spathulate or sessile and rounded, entire, flat, fleshy or membranous, rarely absent. Staminal tube cylindric; stamens in 5 groups, each having 2 or 3 anthers; staminodia 5, showy, as long or longer than petals, subulate, linear-oblong, or obovate, erect or reflexed, fleshy or membranaceous, pubescent or glabrous. Fruit 6–35 × 5–12 cm, bacciform, yellow, light brown, or green when mature, fleshy or woody. Seeds covered by mucilaginous pulpy layer, irregularly compressed, 15–40 × 10–12 mm. Cotyledons erect, thick, corrugate; endosperm scarce. Neotropics; 22 species, 5 species in Venezuela, all of these in the flora area. Some species of Theobroma are cultivated for the seeds and pulp, which are used to make chocolate, fruit drinks, and ice cream. Key to the Species of Theobroma 1. 1. 2(1).

2.

3(1).

3.

Leaves mostly glabrous on both sides ...................................................... 2 Lower leaf surface with an indumentum of 1 or 2 kinds of trichomes ... 3 Leaves firmly coriaceous, regularly penniveined, with 10–14 secondary veins on each side, usually 20–30 × 7–10 cm; fruits ovate-oblong, ± pentagonal or decagonal, glabrous, with fleshy relatively thick pericarp; flowers on trunk and branchlets ........................................... T. cacao Leaves chartaceous with 3–6 pairs of secondary veins on each side; fruits ovoid or subglobose, with hard-costate, reticulate, tomentose pericarp; flowers on branchlets ............................................... T. microcarpum Indument of the abaxial surface with 2 kinds of trichomes: very small, intricate, fine white ones covering the areole, and long, thick, reddish trichomes on the minor and major veins; fruits 7.5–11.5 × 5–6.5 cm, pericarp coriaceous to woody ............................................... T. subincanum Indument of the abaxial surface very uniform, with one type of trichome; fruits large, woody ................................................................................. 4

270

4(3).

4.

S TERCULIACEAE

Leaves chartaceous, abaxial surface cinereous, evenly covered by very small, intricate, sericeous, stellate trichomes; leaf base cordate, wide, with 3 pairs of secondary veins; young growth not ferruginous-lanate; fruits strongly costate-reticulate ................................................. T. bicolor Leaves coriaceous, abaxial surface with the first to fourth vein orders glabrous, areole covered by very small, sericeous, stellate trichomes; leaf base obtuse to subcordate, narrow; young vegetative growth with ferruginous, floccose-lanate, deciduous indument; fruits large, brownish-tomentose, smooth, 16–25 × 10–12 cm ....................... T. grandiflorum

Theobroma bicolor Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 104, pl. 30. 1806. Tree 4–5 m tall, the wood soft. Riparian forests, 200–300 m; Amazonas (Río Ugueto). Cultivated from Mexico to Brazil and Bolivia, probably originated in Central America.

Culebra, Río Cataniapo, Río Sipapo, San Carlos de Río Negro, Santa Rosa de Amanadona, Yavita, Yutajé). Colombia, Brazil, cultivated in many countries. ◆Fig. 227. The fruits of Theobroma grandiflorum are used to make ice cream and a delicious drink similar to that made from Annona species.

Theobroma cacao L., Sp. Pl. 782. 1753. Mexico, Central America, tropical South America; 2 subspecies, 1 of these in the flora area. Subspecies cacao is limited to Central America.

Theobroma microcarpum Mart. in Buchn., Repert. 35: 24. 1830. —Cacao de monte, Kayani (Yanomami). Tree 4–20 m tall. Riparian forests, 100– 300 m; Amazonas (Río Mavaca). Colombia, Brazil.

T. cacao subsp. sphaerocarpum (A. Chev.) Cuatrec., Contr. U. S. Natl. Herb. 35(6): 515. 1964. —Theobroma sphaerocarpa A. Chev., Veg. Util. Afr. Trop. Fr. 4: 12. 1908. Small tree; fruit ellipsoid to almost globose or oblong, 5- or 10-furrowed. Riparian forests, 50–200 m; Bolívar (Temblador), Amazonas (upper Río Orinoco, Tamatama). Widely cultivated in the tropics, especially in South America and Africa. ◆Fig. 229. The seeds of Theobroma cacao are used to make chocolate.

Theobroma subincanum Mart. in Buchner, Repert. Pharm. 35: 23. 1830. —Cacao montero, Cacao-rana, Jaguajeshi (Yanmomai), Mauirruda (Bare), Mawanani (Yekwana), Padamá (Arekuna) Theobroma guianense (Aubl.) Gmel., Syst. Veg. ed. 13, 2: 1151. 1796. —Cacao guianensis Aubl., Hist. Pl. Guiane 684. 1775, pro parte (leaves). Theobroma sylvestris (Aubl.) G. Don, Gen. Syst. 1: 622. 1831, non Mart. 1830. —Cacao sylvestris Aubl., Hist. Pl. Guiane 687. 1775. Tree to 12 m tall. Evergreen lowland forests, along rivers, 100–600 m; Bolívar (Río Icabarú), Amazonas (base of Cerro Duida, La Esmeralda, Maroa and Maroa-Yavita road, Río Casiquiare, Río Guainía, base camp on Río Mawarinuma at base of Sierra de la Neblina, Río Metacuni, upper Río Orinoco, Río Padamo, Salto Yureba on Caño Yureba, Tamatama). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 228.

Theobroma grandiflorum (Willd. ex Spreng.) K. Schum. in Mart., Fl. Bras. 12(3): 76, pl. 8. 1886. —Bubroma grandiflorum Willd. ex Spreng., Syst. Veg. 3: 332. 1826. —Cupuasú, Orinaré Bareua (Piaroa). Tree 4–15 m tall; fruits oblongoid, brownvelutinous, subwoody. Moist evergreen lowland forests, often cultivated in backyards, 50–400 m; Amazonas (Capibara, near

Theobroma 271

Fig. 227. Theobroma grandiflorum

Fig. 228. Theobroma subincanum

Fig. 229. Theobroma cacao subsp. sphaerocarpum

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9. ULADENDRON Marc.-Berti, Pittiera 3: 9. 1971. Trees to 30 m tall. Trunks unarmed, usually without buttresses. Leaves simple, often shallowly lobed and remotely dentate, palmately 5–9-veined from their cordate bases, sparsely stellate-tomentose on upper surface, more densely so on lower surface; stipules linear-triangular. Flowers moderately large, apparently actinomorphic, axillary, solitary or paired; pedicels 3-bracteolate. Calyx tubular, expanded toward apex, 5-lobed; petals spathulate, adnate to base of staminal tube, creamy white, glabrous or nearly so within, densely stellate-tomentose externally. Stamens numerous, fused for most of their length into a tube; anthers borne on distal half of tube; free portions of staminal filaments short, simple or bifid, each ultimate segment bearing a single, reniform, monothecate (1-locular) anther. Ovary superior, 5-locular; locules biovulate; style simple; stigma capitate, 5-sulcate. Fruits capsular, ovoid, loculicidally dehiscent by 5 valves. Seeds alate, 2 per locule. Venezuela; 1 species endemic to the flora area. Uladendron codesuri Marc.-Berti, Pittiera 3: 10. 1971. —Algodoncillo, Oreja del tigre. Large tree to 30 m tall and 1 m diameter. Evergreen lowland and lower montane forests, 50–300 m; Bolívar (near la Prisión on the Río Caura, near Túriba). Endemic. ◆Fig. 230. The wood of Uladendron codesuri reportedly is used for house construction along the Río Caura.

Fig. 230. Uladendron codesuri

Waltheria 273

10. WALTHERIA L., Sp. Pl. 673. 1753. Lophanthus J.R. Forster & G. Forst., Char. Gen. Pl. 27, t. 14. 1775. Astropus Spreng., Neue Entd. 3: 64. 1822. Sitella L.B. Bailey, Gentes Herb. 4(9): 349. figs. 222. 1940.

by Janice G. Saunders Unarmed shrubs, subshrubs, suffrutices, or herbs, rarely trees; branches usually smooth, furrowed, or slightly tuberculate, usually covered by stellate hairs, or simple, or capitate-glandular, sessile-glandular, or intermixed. Leaves alternate, spiral or distichous, simple, petiolate, serrate, dentate, or crenate; stipules linear, subulate, half-ovate, subcordiform, deciduous, rarely persistent. Inflorescences usually many-flowered, axillary or terminal, sessile to pedunculate, cymose; bracteoles 2 or 3, free to variously fused; primary cyme with bracts usually (3)4(5), free, or middle 2 variously fused, rarely 3 mostly fused or all fused into an involucral cup around the flower pair. Flowers usually sessile or pedicels to 1 mm long, usually distylous, or homostylous (derived monomorphic), actinomorphic, 2–15 mm diam. Calyx campanulate, turbinate, or tubular, sepals 5, basally fused, 5-toothed, 5-lobed, or 5-laciniate, the teeth, lobes or lacinae triangular, subulate, or ovate, acute to acuminate. Corolla patent or spreading, yellow, a few species outside the flora area also red basally or purple distally, or entirely white, persistent; petals 5, flat, or lower portion slightly folded inward laterally, or margin incurved apically but not hooded, spathulate, obdeltate, obtriangular, oblanceolate, elliptic, or obovate, membranaceous or opaque, interior glabrous or pilose, pubescent, outer surface stellate or glabrous, rarely apical margin with stellate to simple trichomes, with the claw basally to entirely adherent, rarely adnate, to the staminal tube base. Androgynophore absent. Stamens 5; filaments basally to entirely fused forming a membranaceous or opaque thick tube; anthers dithecal, oblong, extrorse, parallel, dehiscence longitudinal; staminodia usually absent. Ovary sessile or subsessile, 1-carpellate, 2-ovulate (one aborts); style 1, lateral or eccentric, shorter or surpassing the calyx; stigma penicillate, plumose, clavate or filiform-papillose, with papillae or branches 0.1–1 mm long arrayed on stigmatic column 0.2–2.5 mm long. Capsule spineless, protruding from or rarely enclosed within persistent, rarely accrescent calyx, 1-locular, loculicidal or operculate, obovoid, obovoid-oblong, turbinate, sometimes laterally apiculate. Seed 1, obovoid or obpyramidal, at times apically ridged and bisulcate or circumferentially sulcate, smooth or verrucose, with endosperm; embryo straight; cotyledons flat. Thrum pollen: exine spinulose, grains medium to large size, 40–58(–62) µm equatorial diameter; pin or homostylous pollen: exine reticulate, grains medium to large size, 29–54 µm diameter. Intermorph differences in grain size and shape, average aperture number, aperture size, and aperture type. United States (Arizona, New Mexico, Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay, western Africa, Madagascar, Oceania (Hawaii [Maui], Marquesas Islands, Society Islands), China (Hainan), Malaysia (Kuala Kelantin), Australia; ca. 60 species, 8 in Venezuela, all of these in the flora area. The greatest diversity of Waltheria is in Brazil and Mexico. There are 6 Old World species and W. indica sensu lato is a pantropical weed.

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Distylous collections from Bolívar that could not be assigned to species include Fernandez 2783, CTES, between Mina Potosi and Hato Candelario; and Maguire and Wurdack 34419A, NY, from Cerro Bolívar, slopes above MorrisonKnudsen camp and Pilot Plant. The latter is similar to Stoffers 174, NY, from Rupununi, Guyana. Key to the Species of Waltheria 1.

1.

2(1).

2.

3(2).

3. 4(3).

4.

5(4).

5.

Herbs; stems with long, simple, appressed trichomes, but gland-tipped hairs, stellate trichomes lacking; stipules (1)1.5–4 mm wide, demidiate-cordiform (half heart-shaped); capsule operculate, (rarely tardily loculicidal), thick hardened bony endocarp at apex; seed verrucose (wart-like protuberances) .............................................. W. operculata Perennials; stems with stellate or glandular capitate trichomes predominant, simple ones present or absent; stipules < 1 mm wide, demidiateovate, narrowly-triangular, linear-triangular, linear; capsule loculicidal, apical hardened endocarp thin or absent; seed smooth or not verrucose ................................................................................................ 2 Leaves spirally arranged; capsule with hardened endocarp at apex inside; petals lacking brown stripes when fresh or deposits along veins when dried, not venose; all flower morphs with stamen filaments connate either basally, midway or mostly, but free at apex and incised or arcuate between free filaments ............................................................ 3 Leaves distichous; capsule interior without hardened endocarp; fresh petals brown-striped (dried showing collateral brown deposits along veins), venose; pin flower stamen filaments forming an entirely fused tube straight across the apex between anthers (thrum flower stamens connate about 1/3–1/2 length) .................................................................. 7 Stems usually sticky from long red (yellow) gland-tipped hairs; seed subapically circumferentially depressed (mucronate tip broadly truncate in lateral view) .................................................................... W. viscosissima Stems not sticky, gland-tipped hairs short, scarce, obscure, or absent; seed bisulcate, rounded, truncate or obtuse at apex ........................... 4 Four bracts around flower pair very unequal, widest one ovate (length to width ratio 2–2.5:1), usually 1–2 mm wide; petal claw adherent to basal stamen tube for 0.7–1.7 mm ....................................................... 5 Four bracts around flower pair equal or subequal, widest one lanceolate or linear, rarely narrowly ovate (length to width ratio 7.7–3:1), < 1 mm wide; petal claw adherent to basal stamen tube for 0.3–0.4 mm ................................................................................................................ 6 Prostrate shrubs; leaves downy puberulent, lateral margins with mostly dark red, multitiered-capitate glandular hairs 0.2–0.4 mm long, with red cross traces in stalk, generally extended beyond stellate trichomes that are usually equal-rayed, short and flat to slightly spreading; flowers homostylous .......................................................... W. carmensarae Suffruticose, declining or erect subshrubs; leaves sericeous, lateral margins with long stellate trichomes, unequally rayed, generally obscuring usually yellow, peach, or rose long and/or short gland-tipped hairs, their tiers in tip and traces in stalk less obvious; flowers usually distylous .................................................................................... W. albicans

Waltheria 275

6(4).

6.

7(2).

7. 8(7).

8.

Shrub 1–2 m tall; leaves tomentose; bracts, bracteoles distinct, entire, without minute red capitate gland-tipped hairs; flowers homostylous, sessile .......................................................................... W. indica sensu lato Shrub to 0.5 m tall; leaves stellate-pubescent, glabrescent; bracts with middle 2 fused 0.6–0.9 mm, or distinct; bracteoles 3, variably fused, together either 2-cleft and 1-divided, divergent, or 2-lobed with one tip forked, falcate; bracts and bracteoles with minute (0.2 mm) red capitate gland-tipped hairs; flowers distylous (pin stamens 2.2– 2.5 mm, stretched pistil 3–3.4 mm long), pedicels 0.3–0.4 mm long ...................................................................................................... W. collina Bracts surrounding flower pair together entirely connate forming an urn-like or cupuliform involucre 5.5–11 mm wide, irregularly serratedentate at apex, accrescent, papery; fertile branch racemiform, mostly simple, large-leaved, reduced leaves few, elliptic; involucres in loose clusters 2–3 cm long, branchlets simple, disperse; calyx lobes 1.1– 1.5(–1.6) mm wide; anthers (1–)1.1–1.5 mm long; calyx tube (3–)4– 6 mm long; (Río Chaviripa to lower Río Orinoco to Ciudad Bolívar) ........................................................................ W. involucrata sensu stricto Four bracts around flower pair free and entire, or variously fused but only to 3/4 length, margins variable ...................................................... 8 Four bracts surrounding flower pair free, largest 2–4 mm wide, widest above middle, spathulate or oblanceolate, margins entire, even, not accrescent; fertile branch becoming deltate, upper portion leafless or few reduced narrowly ovate leaves, its basal lateral shoots to 6–15 × 4–5 cm, gradually shorter to apex, at nodes 2 or 3 branchlets, first clustered together, subsympodial, later freer, more spaced, some becoming long-pedunculate, floral clusters closely aggregate, oblong, curling laterally, becoming loose underneath with free paniculiform branches; calyx lobes (1.2–)1.6–1.9 mm wide; anthers 0.8–1(–1.2) mm long; calyx tube 3.5–4.2 mm (Colombia) .............. W. berteroi sensu stricto Characters erratic, with different combinations or intermediate states or both partial states of those in couplet leads 7(2) and 8(7), variable, each collection different, not keyable, respectively placed by principal components; (lower Río Orinoco from Moitaco to Los Castillos, or Isla Anacoco—see W. involucrata s.l.; middle Río Orinoco from Samariapo to Los Pijiguaos—see both); for further distinctions see discussion below ....................... Intermediates between W. berteroi and W. involucrata

Intermediates between W. berteroi and W. involucrata Intermediates between W. berteroi and W. involucrata have the largest bract 4–6 mm wide, semiaccrescent or not, mostly with 4 bracts surrounding flower pair with 2 free and 2 middle bracts fused 1/2–3/4 of length, or with 3 bracts connate 1/3–3/4 length and 1 bract free, or all 4 bracts connate for 3/4 length, or very rarely all 4 distinct; bracts usually widest midlength, variously shaped, elliptical, orbicular, obovate, ovate, oval, asymmetric, margins usually irregularly serrate entirely or partially, very rarely entire throughout; flowering branch spike-like, simple or with numerous foliose fertile lateral branches, these progressively shorter from base to apex but their leaf size larger toward apex, or occurring only at midlength,

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at leaf nodes flower clusters few, interspersed unevenly among large leaves, or numerous subsessile rounded aggregates interspersed evenly among leaves progressively smaller or progressively larger, or oblong, very bracteose, with bracteoles also semiaccrescent, short pedunculate clusters somewhat erratically disposed but numerous. Waltheria albicans Turcz., Bull. Soc. Imp. Naturalistes Moscou 31(1): 214. 1858. —Cadillo. Suffruticose to 0.6 m tall or erect subshrub to 1 m tall; single to few-culmed, branches long from base, gradually shorter to tip; flowering in the morning; calyx lobe bases with 1(2) pair(s) of prominent, red or yellow arched secondary veins; petals to 5 × 3 mm, obtriangular, apical margin with a few simple, forked hairs, midvein raised, anthers 0.5–0.7 mm long; stigma oblong, distended-plumose (branches arising from column for > 1.5 mm); capsule base with sessile glandular hairs. Granitic outcrops and surrounding rocky savanna slopes, sandy savannas, (50–)100–600 m; Bolívar (Cerro Arimagua, Cerro El Médano south of Caicara, Cerro Picacho on Altiplanicie de Nuria north of Tumeremo, Ciudad Bolívar, Fundo La Trinidad, Hato Morichito), Amazonas (Isla Ratón). Anzoátegui, Aragua, Zulia; Mexico, Colombia, Guyana, Brazil, Bolivia, Paraguay, Argentina. Distylous plants occur in Mexico, Colombia, Venezuela (Cerro Pichaco, Cerro El Médano), Guyana, Brazil, Bolivia, and Paraguay. Homostylous plants occur in Venezuela, Paraguay, and Argentina. Waltheria albicans sensu stricto is distylous, with well-separated stigma and anthers, only common in Brazil within the grid formed by 5°–18°S and 39–48°W. Waltheria albicans in the flora area occurs in the Caroní lowlands, the peneplain of Serranía de Imataca, and Isla Ratón. The flora area plants have shorter, sparser gland-tipped trichomes, smaller petals, less separation between stigma and anthers (some homostylous), shorter anthers, at times with red anther connective (Steyermark 88903), and capsules with the lower membranous portion being brown translucent, not clearcolored and transparent. Waltheria albicans is variable, and homostylous flowered and/or erect plants may be hybrids with W. indica sensu lato or con-

fused with the related but poorly known Waltheria paniculata Benth., which is homostylous, small-flowered and occurs in moist savannas in Guyana. Waltheria albicans is depicted in the F type photo of Schomburgk 500 (distributed as W. paniculata). Waltheria berteroi (Spreng.) J.G. Saunders, Novon 15: 365. 2005. —Malachra berterii Spreng., Syst. Veg. 3: 94. 1826. Waltheria subcordata Standl., Contr. U.S. Natl. Herb. 18: 117. 1916. Distylous shrub or small tree to 3 m tall; leaves 3–15 × 3–10 cm; petal claw adherent to stamen tube for 0.4–0.7 mm. Granitic outcrops, savannas, semideciduous forests, sometimes weedy areas, 80–600 m; Bolívar (between Puerto Ayacucho and Caicara, mouth of Río Parguaza, near mouth of Río Parhueña, lower Río Suapure, Serranía de los Pijiguaos), Amazonas (Gavilán, Puerto Ayacucho to Samariapo, Raudal do Garza, Río Cataniapo). Falcón, Zulia; Colombia (Magdalena, Cesar, Santander, Atlántico, La Guajira). ◆Fig. 231. Waltheria berteroi sensu lato occurs discontinuously from 5°–12°N, 66°30´–75°W, at 10–1600 m elevation. Waltheria berteroi sensu stricto is from the Sierra Nevada de Santa Marta and vicinities in northern Colombia. Waltheria berteroi sensu lato includes the Magdalena River mouth, the La Guajira peninsula in Colombia, and Lake Maracaibo Basin rim in Venezuela, 10–100 m elevation, where intermediate with W. glomerata, and the middle Río Orinoco vicinity in Venezuela and Colombia, where intermediates occur with W. involucrata. The flora area collections of Waltheria berteroi are all intermediate with W. involucrata. They differ from Colombian non-intermediate W. berteroi by wider bracts, with the largest bract 4–6 mm versus 2–4 mm wide, variously shaped, the widest one oval, oval-rhombate, rarely obovate, but not spathulate, oblanceolate, variously fused versus distinct, margins irregularly

Waltheria 277

serrulate, undulate, or partly entire versus entire and even, the fertile branch portion less deltate-shaped, less ramose, often more foliose. Intermediates most similar to W. berteroi (W. berteroi >> W. involucrata) occur along the Samariapo–El Burro road from 10 km north of Río Parhueña (western Bolívar) south to Samariapo (northwestern Amazonas), specifically at 27 km south of Puerto Ayacucho, and 47 km north of there (e.g., Gröger 541, MO; Stergios 3170, CTES; Gröger & Barcroft 295, MO). However, this same transect fosters many more intermediates designated under W. involucrata (see below). Intermediates midway between the two species, but a little closer to Waltheria berteroi than to W. involucrata, occur northward from the vicinity of Río Parhueña (Bolívar), specifically at 39 km north of Puerto Ayacucho, to Los Pijiguaos at 100– 150(–600) m elevation along river margins, the road or crystalline laja (e.g., Aymard & Stergios 3340, MO; Gröger 216, MO; Gröger 827, MO; Guánchez & Huber 4822, CTES; Trujillo 11785, MY; Wurdack & Monachino 39763, TEX). Trujillo 11785 (MY) has 3 semiaccrescent bracts fused halfway, oblong, obovate, the largest 3–5 mm wide, and one distinct but linear-rhombate, narrow, 1.3 mm wide, margins of 2 uneven, 1 entire, 1 once-toothed laterally; a second node has bracts similarly fused but more accrescent, margins uneven, serrulate. Amyard & Stergios 3340 (MO) has the middle 2 bracts fused 3/4 way, the outer 2 distinct, all ovalrhombate, 6 mm wide, semiaccrescent, margins uneven, parts minutely serrate. That collection, the northernmost extension of W. berteroi intermediates in the flora area, is separated by 50 km from the western limit of W. involucrata sensu stricto at Caño Chiviripa. Huber 588 illustrates the general aspect of Waltheria berteroi sensu stricto, differing by leaves narrower, long-acuminate-tipped, a fertile branch less deltate, lacking the elongated basal shoots to 9–15 × 4–5 cm superceding subtending leaves typical of later stages, floral clusters later curling, and bracts broader, and ovate. Waltheria carmensarae J.G. Saunders, Darwiniana 43: (in press). 2005. Prostrate homostylous shrub to 1 m long; xylopodium to 2 cm wide; leaf apices ob-

tusely angled–rounded; inflorescences in age disarticulate, leaving persistent bracteoles, 2, widely divergent, narrowly triangular, and branchlet nodes enlarged, triangular; flowering in the morning; petals narrowly obtriangular, margin glabrous, claw adherent to stamen tube 1.3–1.7 mm. Sandy shrub savannas mixed with savanna scrub, 50–100 m; Bolívar (between Caicara and Ciudad Bolívar, 46 km northeast of Río Aro near Agua Blanca). Southern Paraguay, northern Argentina. Waltheria carmensarae is known from a few localities in Argentina and adjacent Paraguay from sand dunes, sandy banks, or loose sandy soil near the Paraná and Paraguay rivers. A single disjunct collection (Davidse 4495, MO) occurs in the flora area, from “very sandy soil, in dry TrachypogonCuratella-Byrsonima savanna.” It differs by stems being straight, subterete and woody to the tips rather than slightly flexuose and decurrent for the ultimate 6–10 nodes. It shares the distinctive leaves, bracteal shape, xylopodium, and prostrate habit, sand-loving habitat of the Argentinean populations. Waltheria collina K. Schum. in Mart., Fl. Bras. 12(3): 63. 1886. Shrub to 50 cm tall, stems 2 mm diameter; leaves heteromorphic, oval, rhomboid, 2–3 × 0.7–1 cm, serrate, tooth side and length subequal, costal vein thick; bracts fused, unequal, the largest ovate-elliptic, 0.7 mm wide, length to width ratio 5.3–5.5:1, or distinct, equal, linear-lanceolate, 0.4 mm wide; pedicels 0.3–0.4 mm long; calyx lobed halfway, multi-areolate; petals papillose. Savannas, dry scrub, 50–200 m; Bolívar (south of Paso de Caruachi in Río Caroní basin). Colombia, Brazil. Distribution represented by single variable specimens, but bracteole shape and fusion (including variability for both), serrulate margins, reddish color, and minute red capitate glandular hairs are characteristic. The flora area collection (Holst & al. 1975, MO) differs from the type (Riedel 64, LE) by its 3 bracteoles at times with one segment distinct, lobes deeper, bracts that are equalshaped or if unequal-shaped like the type, then fused versus distinct, lack of filiform non-capitate sessile glandular hairs, inflo-

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rescences short-pedunculate versus subsessile, calyx 3.1–3.3 mm versus 3.8–4.3 mm long, anthers 0.8 mm versus 0.4–0.5 mm long, their base less sagittate, less connective, style 1.1–1.4 mm long, glabrous versus 1.9 mm long, once-stellate. The species is closely related to W. indica. Waltheria indica L., Sp. Pl. 673. 1753. Waltheria americana L., Sp. Pl. 673. 1753. Homostylous, erect, tomentose, suffrutescent herb or subshrub to 2 m tall; calyx lobe bases multiple-areolate; petals obovate or oblanceolate, apices rounded-obtuse; stamen free; filaments 0–0.1 mm; stigma plumose, lanceolate or penicillate, conical. Curatella savannas, disturbed areas, very weedy, 50–400 m; Delta Amacuro (Los Castillos), Bolívar (between Guasipati and El Cintillo, Guri and Caño La Becerra, Hato Morochito), Amazonas (Puerto Ayacucho). Widespread in northern Venezuela; tropics and subtropics worldwide. ◆Fig. 232. Waltheria indica sensu lato is in need of a critical revision. It is not monophyletic based upon preliminary morphological studies (Saunders unpubl. data). Most plants in the flora area are more similar to W. albicans (e.g., Zuloaga et al. 4479, SI), W. paniculata and W. detonsa (e.g., Amyard & Cuello 3845, CTES), with a distended-plumose stigma and the corneous (horny) endocarp limited to the capsule apex. Other plants now included in W. indica sensu lato are more similar to W. petiolata or W. rotundifolia, characterized by a penicillate stigma and usually complete extent of capsule corneous (horny) endocarp. Kohler (1971, 1976) has found pollen aperture number and type differences within W. indica sensu lato. Waltheria involucrata Benth., J. Bot. (Hooker) 4: 125. 1842. —Sitella involucrata (Benth.) L.H. Bailey, Gentes Herb. 4(9): 349, figs. 222, 223. 1940. Distylous, broad, multicaulous shrub to 3 m tall; leaves 4–17 × 2–11 cm; inflorescences showy, with cupuliform expanded involucres; flowers diurnal; petal claw basally adherent for 0.2–0.7 mm to stamen tube. Semideciduous forests, dry savannas, riparian woods, river borders on granitic out-

crops, near sea level to 400 m; Delta Amacuro (Los Castillos de Guayana) Bolívar (Camarica, Caño Chaviripa near San Pablo, Ciudad Bolívar, Guaniamo, Isla Anacoco, Moitaco, Paso de Caruachi, Puerto Ordaz, Río Caroní toward Upata, Río Caura, Río Cuyuní, Río Perro de Agua, Río Venamo, San Felix to Upata at Los Negros, San Martin de Turumbán southwest of Tumeremo), Amazonas (Gavilán, Las Pavas, Puerto Ayacucho to Samariapo). Monagas, Zulia; Guyana, Brazil (Roraima). ◆Fig. 233. Waltheria involucrata sensu lato, including intermediates with W. berteroi, discontinuously occurs between 2°–10°N, 58°–71°W, mostly near the perimeter of the Venezuelan Guayana. Waltheria involucrata sensu stricto occurs in the Guyanan Rupununi and Brazilian Roraima region between 2°40´–4°10´N, 59°–62°W, 100–300 m elevation, within the triangular area formed by Ilha de Maraca in Brazil and Surama and Karaudanawa in Guyana, in Guyana along the Ireng River below Mount Roraima, the lower Río Orinoco, and at Mene Grande, Zulia in Venezuela. In the flora area W. involucrata sensu stricto occurs in northern Bolívar between Ciudad Bolívar west to near Caicara either near the lower Río Orinoco (Ciudad Bolívar, Moitaco), or its connecting rivers (Río Caura near Maripa, Caño Cocuiza, Guaniamo near Perro de Agua, Río Chaviripa near San Pablo), at low elevations of near sea level to 70 m or rarely (López-Palacios A520, NY) at 100–280 m near the base of the Guyanan Shield Drainage near Guaniamo and Perro de Agua (e.g., Bailey & Bailey 1427, US; Holt & Gehringer 163, MO; Trujillo 5894, F; Fernández 4254, MO). Otherwise, most flora area collections are intermediate with Waltheria berteroi. Among intermediates only slightly different (W. involucrata >> W. berteroi) are those with an undivided shallowly 4-lobed involucral cup (Davidse & Gonzalez 16427, MO) or one that is 3-parted, fused for 1/3 its length but with one bract free, one 5.5 mm wide, parts rounded, with margins entire rather than irregularly serrate, a second node has bracts with 3 fused variously 1/3 or halfway, uneven margins (Gentry 10738, MO). Gentry 10738 also has axillary inflorescences dispersed among large leaves, even larger toward branch apices, with clusters subses-

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sile, glomerulate, globose, leaves coarsely toothed, petioles long, thin. These intermediates occur in northeastern Bolívar and Delta Amacuro either between the area enclosed by connecting Puerto Ordaz-UpataLos Castillos, along streams or dry stream beds in nonflooded evergreen, deciduous forests or semideciduous riparian forests (e.g., Davidse & Gonzalez 16427, MO; Gentry 10738, MO; Holst 1995, MO; Lasser 38086, VEN) or in eastern Bolívar near Isla Anacoco in nonflooded evergreen forest (e.g., Delascio 8585, VEN; Fernández 2617, NY; Stergios 5009, MO) at 50–200(–400) m. Intermediates about midway between the two species but slightly closer to W. involucrata (W. involucrata ≥ W. berteroi), in some plants characterized by the middle 2 of 4 bracts fused 1/2–3/4 length, laterals free, or axillary glomerulate leafy inflorescences with expanded bracts, occur in northwestern Amazonas between Río Parhueña and Samariapo near the Río Orinoco, particularly to 30 km around Puerto Ayacucho, often at the border of sandy savanna-low forested hills, often on, among, or along granitic laja, at 75–100 m elevation (e.g., Castillo 750, CTES; Davidse 2847, MO; Davidse & Huber 14964, MO; Davidse & Huber 15337, MO; Guánchez 1525, MO Huber 588, US; Huber 1274, NY; Kral & Boom 71723, MO; Liesner 17239, CTES; Maguire & al. 36073, VEN; Romero & Melgueiro 2020, TEX). Some examples are here described. Davidse 2847 (MO), 20 km south of Puerto Ayacucho, has irregular dentation on leaves, the fertile branch with crowded, dense axillary flower clusters along the main stem and on a few short lateral branches that are unorderly in length and placement, the upper flowering branch foliose with leaves small rather than greatly reduced, floral clusters remaining subsessile or short-pedunculate, glomerulate, rounded or oblong, bracts with 2 lateral ones distinct and the middle 2 fused 3/4 their length, all expanded, 5 mm wide, rounded or roundedspathulate, irregularly serrate, or entire, papery. Davidse & Huber 14964 (MO), 8 km south of Puerto Ayacucho, has leaves with variable irregular margins, a ramose fertile portion but with leaves increasing in size toward tips of lateral branches, flower clusters amorphous, shaggy, bracteose, 2- or 3-

parted bracteoles also expanded, papery, overlying bracts, bracts with middle 2 fused 3/4 length, very accrescent, each 5 mm wide, imbricate, undulate, slightly serrate or entire, the lateral ones free, a little accrescent, 3 mm wide, oval, oblong-spathulate, subentire. Waltheria involucrata flowers are nectiferous and diurnal, open until after 5 pm. Probable pollinators observed in Roraima include Lepidoptera (Lycaenidae, Nymphalidae, Junonia), vespid wasps (Polybiini), and sphecid wasps (Sphex) (unpubl. data, determined by J. Neff (Hymenoptera) or P. de Vries (Lepidoptera). Bailey (1940), naming his Río Orinoco collection 1427!, segregated Waltheria involucrata Benth. as Sitella involucrata based on Bentham’s same type (Schomburgk 722, K!) by its accrescent bracteal involucral cup and arborescent habit. The latter trait is common to related W. glomerata and W. berteroi as well. Both characters are found together in some species of Melochia (sect. Physodium), segregated generically by some, but not together in Hermannia, Hermannia inflata, a shrub with inflated bracteal cup, and not segregated generically. However, in those genera, the involucral cup surrounds a single flower rather than 2 flowers as Waltheria involucrata. Based upon the sporadic appearance of cupuliform involucres and arborescent habit in its tribe, and the coherence of other characters Bailey gave (stalked bracteal cups, sessile glomerules, short petals, hirsute capsule, flagelliform branches), character combinations variously found in the rest of Waltheria, in my opinion, Sitella involucrata belongs in Waltheria; therefore the genus Sitella is not recognized. Waltheria operculata Rose, Contr. U.S. National Herb. 5: 183. 1899. Turnera valleana Standl. & L.O. Williams, Ceiba 1(12): 88. 1950. Homostylous herbaceous annual to 60 cm; branches erect, basally whorled, spreading, dichotomous above, strigose; leaves lanceolate to oblong, scattered simple-setose above, on veins below, basal palmate veins obscure; inflorescences capitate; flowering in the morning; petal oblong, 3–8 × 0.7–2 mm, claw adherent 1–2 mm to stamen tube;

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Fig. 231. Waltheria berteroi

Fig. 232. Waltheria indica

Fig. 233. Waltheria involucrata

S TRELITZIACEAE 281

stigma distended-plumose, 1.2–2.4 mm long, papillae disperse, unilateral; capsule endocarp osseous (bony) for apical 0.5–1 mm (operculum), membranous below except base; seed obpyramidal, apex ridged, bisulcate. 50 m, Bolívar (Ciudad Bolívar). Mexico (Oaxaca), Honduras (Choluteca), Colombia (Santa Marta), central and eastern Brazil (Bahía, Ceará, Goias, Mato Grosso, Minas Gerais, Paraiba, Pernambuco, Piauí, Rio Grande de Norte, Rio de Janeiro), northern Bolivia (Santa Cruz), northern Paraguay (Alto Paraguay). Scarcely known (1 or 2 localities per country) outside Brazil, where common in caatinga, open ground, thin-soiled, moist sandy places. Singly known in Venezuela (Sprague s.n., K, 1898). Brazilian collections were formerly confused with Waltheria bracteosa A. St.-Hil. or W. macropoda Turcz., Brazilian endemics. It is distinguished from both by habit, flowers homostylous, anthers < 0.8 mm long, stigma 0.4–0.5 mm wide, lanceolate. Review of the Honduran (Choluteca) isotype (Standley 24564, NY!) of Turnera valleana Standl. & L.O. Williams (Turneraceae) reveals it to be the same species as the Mexican (Oaxaca-Chiapas) holotype (Nelson 2876, US!) of Waltheria operculata Rose, based upon characters in the above description. Waltheria viscosissima A. St.-Hil., Fl. Bras. Merid. 1: 150. 1825.

Waltheria hirsuta C. Presl, Reliq. Haenk. 2. 152. 1835. Waltheria tubiflora Klotzsch, Linnaea 14: 300. 1840. Waltheria machrisiantha L.B. Sm., Los Angeles County Mus. Contr. Sci. 23: 10, figs. 5, 6. 1958. Usually distylous, or homostylous, scandent or erect shrub to 2 m; stems reddish; upper surface of leaves stellate, lower surface simple-hirsute, stellate-velutinous; flowers pedicellate, sessile, closing midafternoon; petals 5.5–10 mm × 1.4–4.6 mm, obdeltate, glabrous, midvein raised; claw completely adherent for 2–3 mm to stamen tube apex; stigma distended-plumose, 0.8–1.5 × 0.8–1.2 mm, branches 0.2–0.7 mm long; capsule loculicidal, tearing, base persistent; seed 2.1–2.7 × 1.4–2.0 mm. Semideciduous forests, tree clumps in sandy savannas, sandy Curatella savannas, 50–200 m; Bolívar (Moitaco), Amazonas (Puerto Ayacucho). Anzoátegui, Apure, Guárico, Portuguesa, Sucre, Zulia; Mexico, Colombia, Guyana, northern and central Brazil, Bolivia (Santa Cruz), Paraguay (Amambay, Concepción). Waltheria viscosissima exhibits a sporadic largely discontinuous distribution, most often in western Mexico, or Brazil between 10°–18°S, 36°–60°W. It is disjunctly distributed between Mexico, where homostylous, and South America, where it is distylous. However, near Santa Marta, Colombia, apparently at the base of Sierra Nevada de Santa Marta, both morphs occur.

STRELITZIACEAE by Lennart L. Andersson Rhizomatic plants, when mature often with a woody trunk that has a cluster of leaves on top, sometimes acaulescent with dichotomously branched stems (Strelitzia). Leaves alternate, distichous, held in fan-like clusters, differentiated into sheath, petiole (sometimes indistinct), and blade; sheath not forming distinct pseudostems, without a distinct ligule; blade entire, with a coarse midrib and closely set, parallel, slightly S-shaped (sigmoid) lateral veins which are interconnected by tertiary cross veins. Inflorescences terminal or lateral, structurally welldefined, of much-condensed circinnate flower clusters in axils of spathaceous bracts; flowers bracteolate; bracteoles keeled. Flowers zygomorphic, basically 3-merous,

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Fig. 234. Phenakospermum guyannense

5-cyclic, heterochlamydeous. Sepals free, equal or subequal; petals shortly fused at base, 2 paired, these larger and sometimes much different in shape from the third one (Strelitzia). Androeceum of 5 or 6 stamens; staminodes never present. Ovary inferior, 3-locular; placentation axile; ovules numerous in each locule, anatropous. Fruit a loculicidal, woody capsule; seeds operculate, with a conspicuously bright-colored aril.

S TYRACACEAE 283

Southern coast of Africa, Madagascar, and adjacent islands, tropical South America; 3 genera and 6 or 7 species, 1 species in the flora area. 1. PHENAKOSPERMUM Endl., Gen. Pl. 3: 229. 1837. Tall plants, the old ones with leaves in a fan-like cluster on top of an unbranched woody trunk. Leaf buds opening in a pseudo-calyptrate fashion, the mature leaf appearing torn off at tip. Inflorescence erect, terminal, on a peduncle covered by scale-like leaves, with several circinnate flower clusters in the axils of spathaceous bracts. Flowers bisexual. Sepals free and equal; petals cream-colored, fused near the base, the 2 anterior similar to the sepals, the posterior one similar in shape but smaller. Stamens 5, staminode lacking. Fruit a woody, dehiscent capsule. Seeds many in each locule, arillate; aril red, dissected into thread-like lobes. Neotropics; 1 species. Phenakospermum guyannense (Rich.) Endl., Bot. Zeitung (Berlin) 3: 345. 1845. —Urania guyannensis Rich., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 15: 21, t. 6. 1831. Gigantic caulescent herb to 12 m tall. Forest edges, clearings, swamp forests, 50–700

m; Bolívar (Caño Pablo near Salto Pará, Cerro Abismo), Amazonas (8 km south of Puerto Ayacucho, Río Mawarinuma, Río Yureba, east of San Carlos de Río Negro). Apure; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 234.

STYRACACEAE by Peter W. Fritsch and Julian A. Steyermark Trees or shrubs with stellate trichomes or scales. Leaves alternate, simple, pinnately veined, without stipules. Inflorescences paniculate, racemose, or cymose, or flowers rarely solitary, lateral or terminal; bracts small or absent; bracteoles 1 or 2 or absent. Flowers actinomorphic, bisexual or rarely unisexual, with a hypanthium adnate to the ovary wall. Calyx gamosepalous, (0–)2–9-toothed or -lobed; corolla gamopetalous, the tube usually much shorter than the lobes, or rarely the petals nearly distinct; lobes (4)5(–8), valvate or imbricate in bud. Stamens usually twice as many as, or rarely to 4 times as many or equal to the number of corolla lobes, attached to the base of the corolla or higher, 1-seriate, ± equal or of 2 lengths; free portion of filaments completely distinct or proximally connate; anthers basifixed, introrse, 2-locular, longitudinally dehiscent, linear; ovary 2–4(5)-carpellate, 2–4(5)-septate at the base but usually 1-locular through the distal attenuation of the septa, nearly superior to completely inferior, placentation axile or rarely basal; style 1, simple, filiform, hollow; ovules erect to pendulous, 1–30 in each locule. Fruit capsular, nut-like (dry and indehiscent), or drupaceous, calyx persistent. Seeds 1–many, brown, endosperm abundant; embryo straight or rarely curved. U.S.A., Mexico, Central America, South America, the Mediterranean region, eastern Asia, southeastern Asia and adjacent islands; 11 genera and ca. 160 species, 1 genus and 11 species in the flora area.

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1. STYRAX L., Sp. Pl. 444. 1753. Trees or shrubs with stellate trichomes or scales, evergreen (in the flora area) or deciduous. Leaf blades entire (in the flora area) or toothed. Inflorescences lateral or terminal, simple or sparsely branched, racemose or cymose (in the flora area), rarely 1-flowered. Flowers bisexual (in the flora area) or rarely unisexual, fragrant, with a short hypanthium adnate to ovary wall only at the base. Calyx cupuliform, (0–)5(–9)-toothed. Corolla gamopetalous, white, pink, or rarely yellow, the lobes usually 5, valvate (in the flora area) or imbricate in bud. Stamens usually 10, adnate to the corolla at the base of the lobes (in the flora area) or rarely in the tube; free portion of filaments distinct or proximally connate, the ventral side glabrous to (in the flora area) stellate-pubescent or lepidote; ovary 3-carpellate, septate at the base but 1-locular through the distal attenuation of the septa, essentially superior, placental obturators usually present; ovules 1 to ca. 8 in each locule. Fruits drupaceous (in the flora area), loculicidal capsules dehiscent via 3 valves, or nut-like, depressed-globose to ellipsoid. Seeds 1(–3), completely filling fruit cavity, globose to ellipsoid; testa indurate, usually smooth with 3(–6) longitudinal grooves and conspicuous hilum. Distribution as in family; ca. 130 species, 17 species in Venezuela, 11 of these in the flora area. Key to the Species of Styrax 1. 1. 2(1). 2. 3(2).

3.

4(3).

4.

5(2).

5.

Lateral leaf veins glabrous or at most with moderately dense vestiture on the lower surface, most of the vein surface evident ....................... 2 Lateral leaf veins stellate-tomentose on the lower surface, the surface of the veins obscured ................................................................................. 7 Calyx, pedicels, and inflorescence branches lepidote; lower surface of leaf blades lepidote or glabrous ............................................................ 3 Calyx, pedicels, inflorescence branches, and lower surface of leaf blades stellate-pubescent .................................................................................. 5 Lower surface of leaf blades glabrous or nearly so; anther sacs tapered at apex; longest filament trichomes 1.2–2 mm long; drupe 12–18 mm long ........................................................................................... S. glabratus Lower surface of leaf blades densely lepidote; anther sacs nontapered at apex; longest filament trichomes 0.6–1.1 mm long; drupe 8–13 mm long ......................................................................................................... 4 Larger leaf blades 6.6–10.5 × 2.9–5.8 cm; drupe 11–13 × 7–8 mm; domatia present at junction of lateral veins and midrib on at least some leaves; longer petioles 6–11 mm long; lower pedicels 5–9 mm long ................................................................................................. S. glaber Larger leaf blades 2.7–4.6 × 1.4–2.6 cm; drupe 8–9 × 5–7 mm; domatia absent; longer petioles 3–6 mm long; lower pedicels 3–5 mm long .......................................................................................... S. wurdackiorum Larger leaf blades 21–31 × 7.4–11.2 cm; lateral leaf veins 10–15 per side; anther sacs tapered at apex; longest filament trichomes 1.3–2 mm long ......................................................................................... S. guyanensis Larger leaf blades 4.8–8.2 × 1.9–4.6 cm; lateral leaf veins 4–7 per side; anther sacs nontapered at apex; longest filament trichomes 0.9– 1.1 mm long ........................................................................................... 6

Styrax 285

6(5).

Larger leaf blades 1.9–3.3 cm wide, coriaceous; lateral leaf veins impressed to sulcate on upper surface, those nearest the leaf base subhorizontal to an angle of 35°; lower surface of leaves with thick tomentum .......................................................................... S. guaiquinimae 6. Larger leaf blades 3.2–4.6 cm wide, chartaceous to subcoriaceous; lateral leaf veins plane with upper surface, those nearest the leaf base arched-ascending at angle of 35–60°; lower surface of leaves with thin tomentum ..................................................................... S. longipedicellatus 7(1). Lower surface of leaves with 8–15-armed dark orange-brown stellate trichomes scattered among the easily visible white tomentum; drupe truncate at apex ......................................................................... S. neblinae 7. Lower surface of leaves with 18–38-armed light-orange to orange-brown stellate trichomes partially to completely obscuring the white tomentum beneath; drupe convex at apex or fruit unknown ........................ 8 8(7). Longer petioles 16–20 mm long; leaf blades thick-chartaceous; calyx uniformly grayish brown-tomentose .................................... S. sipapoanus 8. Longer petioles 4–12 mm long; leaf blades subcoriaceous to coriaceous; calyx tawny brown-tomentose with scattered orange-brown trichomes or uniformly orange-brown-tomentose ................................................. 9 9(8). Lower pedicels 2–3 mm long; inflorescences 1.5–3 cm long; drupe 5– 6 mm wide; calyx tawny brown-tomentose with scattered orangebrown trichomes ........................................................................... S. duidae 9. Lower pedicels 6–10 mm long; inflorescences 3–6 cm long; drupe 7–8 mm wide (fruit unknown in S. yutajensis); calyx uniformly orange-browntomentose ............................................................................................. 10 10(9). Larger leaf blades 8.1–10 × 4.8–6.1 cm; anthers 3.5–4 mm long; corolla lobes spreading; lower pedicels 6–8 mm long; inflorescence branches 1.1–1.5 mm wide; filaments with delicate, interwoven trichomes .............................................................................................. S. guanayanus 10. Larger leaf blades 5.9–7.1 × 2.8–4.1 cm; anthers 6–7 mm long; corolla lobes reflexed to revolute; lower pedicels 8–10 mm long; inflorescence branches 0.7–0.9 mm wide; filaments with stiff, ± straight trichomes ................................................................................................. S. yutajensis Styrax duidae Steyerm., Fieldiana, Bot. 28: 491, fig. 110. 1953. Shrub to 3 m tall; leaf blades elliptic to ovate, the larger 4.9–14.8 × 2.8–4.6 cm, margin revolute, domatia absent; calyx 3.5–5 × 3.5–6 mm, margin curved between the teeth at anthesis; corolla lobes spreading; longest filament trichomes 0.9–1.3 mm long; anther sacs nontapered at apex. Venezuela; 2 subspecies, both in the flora area. Key to the Subspecies of S. duidae 1. Longer petioles 6–9 mm long; larger leaf blades 8.8–14.8 × 3.5–4.6 cm, 2.5–3.2 times as long as wide; lateral leaf veins 7–9 per side .................. subsp. duidae

1. Longer petioles 4–5 mm long; larger leaf blades 4.9–7.2 × 2.8–3.8 cm, 1.6–2.3 times as long as wide; lateral leaf veins 5–7 per side .................. subsp. paruae S. duidae subsp. duidae Dwarf open thickets on sandstone substrate, 1000–1400 m; Amazonas (Cerro Duida). Endemic. S. duidae subsp. paruae Maguire, Mem. New York Bot. Gard. 29: 213, fig. 98A– G. 1978. Along margin of sandstone table mountain in shrubby thickets, ca. 2000 m; Amazonas (Cerro Parú). Endemic. ◆Fig. 237.

286

S TYRACACEAE

Styrax glaber Sw., Prodr. 74. 1788. Styrax occidentalis Sw. ex Thunb., Styrac. 8. 1813. Styrax subleprosum Klotzsch in M.R. Schomb., Reis. Br. Guiana 3: 1087. 1849, nom. nud. Styrax roraimae Perkins, Bot. Jahrb. Syst. 31: 478. 1902. Styrax micrasterus Perkins, Repert. Spec. Nov. Regni Veg. 2: 24. 1906. Styrax glaber var. micranthus Perkins, Pflanzenr. IV(Heft 241): 63. 1907. Styrax fanshawei Sandwith, Kew Bull. 1948: 321. 1948. Styrax costanus Steyerm., Fieldiana, Bot. 28: 490. 1953. Styrax tepuiensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17: 456, fig. 7. 1967, “tepuiense.” Shrub or tree to 15 m tall; leaf blades chartaceous to thick-chartaceous, elliptic to slightly oblanceolate, margin usually not revolute; inflorescences 2–7 cm long; calyx 4–5 × 4–6 mm, grayish green, margin straight between the teeth at anthesis; corolla lobes reflexed to revolute; longest filament trichomes 0.9–1.1 mm long. Dwarf riparian forests, shrub islands near sandstone rocks, open dwarf forests overlying sandstone substrate, 500–2200 m; Bolívar (Auyán-tepui, Kukenán-tepui, Macizo del Chimantá, Río Majagua). Anzoátegui, Monagas, Nueva Esparta, Sucre, Trujillo; Lesser Antilles, Trinidad, Tobago, Guyana. ◆ Fig. 240. Styrax glabratus Schott in Spreng., Syst. Veg. 4(2): 406. 1827. —Strigilia glabrata (Schott) Miers, Ann. Mag. Nat. Hist. ser. 3, 3: 283. 1859. Styrax psilophyllus A. DC., Prodr. 8: 266. 1844, “psilophyllum.” —Strigilia psilophylla (A. DC.) Miers, Ann. Mag. Nat. Hist. ser. 3, 3: 283. 1859. Styrax lauraceus Perkins, Bot. Jahrb. Syst. 31: 478. 1902. Styrax squamulosus M.F. Silva, Acta Amazon. 1: 23. 1971, “squamulosa.” Tree to 20 m tall; leaf blades chartaceous, 9–19 × 2.5–6 cm, elliptic to oblanceolate, margin not revolute, domatia often present on lower surface at junction of lateral veins and midrib; inflorescences 2–7 cm long;

lower pedicels 2–13 mm long; calyx 4–7 × 4– 6 mm, grayish green-lepidote, margin straight between the teeth at anthesis; corolla lobes reflexed to revolute. Rich montane forests on tepui slopes, 1000–1800 m; Amazonas (Cerro Cuao, Sierra de la Neblina). Distrito Federal; Costa Rica, Colombia, Suriname, French Guiana, Brazil. Styrax guaiquinimae (Maguire & Steyerm.) P.W. Fritsch, Novon 14: 48. 2004. —Styrax tepuiensis subsp. guaiquinimae Maguire & Steyerm., Mem. New York Bot. Gard. 29: 218, fig. 103A– F. 1978. Shrub to 3 m tall; larger leaf blades 5.1– 5.6 cm long, elliptic to occasionally ovate, margin revolute, tomentum on lower surface grayish green with scattered conspicuous golden yellow trichomes, domatia absent; inflorescences 2.5–3 cm long; lower pedicels 4– 6 mm long; calyx 4–5 × 5–6 mm, tawny green- to tawny-tomentose, margin straight between the teeth at anthesis; corolla lobes spreading. Shrub islands and rocky savannas with sandstone, 1400–2000 m; Bolívar (Cerro Guaiquinima). Endemic. Styrax guanayanus Maguire & K.D. Phelps, Bol. Soc. Venez. Ci. Nat. 14: 15. 1952, “guanayana.” Styrax jauaensis Steyerm., Bol. Soc. Venez. Ci. Nat. 32: 390, fig. 24A–E. 1976. Shrub or tree to 4 m tall; leaf blades broadly elliptic to ovate-elliptic, margin revolute, domatia absent; calyx 4–6 × 6–7 mm, margin curved between the teeth at anthesis; longest filament trichomes 1.1–1.3 mm long; anther sacs nontapered at apex. Dwarf forests on sandstone ridges and rocky slopes, and bordering streams, 1700–2300 m; Bolívar (Cerro Guanay, Cerro Jaua), Amazonas (Cerro Guanay). Endemic. ◆Fig. 235. A collection from Cerro Jaua (Steyermark et al. 109574) has been described as Styrax jauaensis Steyerm. The reported difference between S. jauaensis and S. guanayanus seems to be of minor significance, and in the absence of flowering material the taxonomic status of S. jauaensis is uncertain.

Styrax 287

Styrax guyanensis A. DC. in DC., Prodr. 8: 261. 1844, “guyanense.” —Strigilia guianensis (A. DC.) Miers, Ann. Mag. Nat. Hist. ser. 3, 3: 282. 1859, “guayanensis.” Styrax guyanensis var. japurensis Seub. in Mart., Fl. Bras. 7: 188. 1868, “guianense var. japurense.” Tree to 15 m tall; leaf blades chartaceous, oblong-elliptic, margin plane, tomentum on lower surface grayish green with scattered orange-brown trichomes, domatia absent; inflorescences 2–4 cm long; lower pedicels 2–6 mm long; calyx 5–6 × 3–5 mm, grayish brown-tomentose, margin straight between the teeth at anthesis; corolla lobes reflexed to revolute. Riparian forests, 100–300 m; Bolívar (Salto Pará), Amazonas (Caño Minicia in middle Río Orinoco basin, Río Casiquiare, Río Negro near Santa Lucia, Río Orinoco near Río Atabapo, Río Pasimoni, Río Vasiva). Colombia, Guyana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 236. Styrax longipedicellatus Steyerm., Acta Bot. Venez. 10: 242. 1975. Styrax tepuiensis subsp. huachamacarii Maguire & Steyerm., Mem. New York Bot. Gard. 29: 218, fig. 104A–G. 1978. Styrax filii Maguire, Mem. New York Bot. Gard. 29: 220, fig. 105A–H. 1978. Shrub or tree to 10 m tall; larger leaf blades 4.8–8.2 cm long, ovate to obovate, margin revolute, tomentum on lower surface white with very sparsely scattered and inconspicuous golden yellow trichomes, domatia absent; inflorescences 1.5–5 cm long; lower pedicels 5–9 mm long; calyx 4–4.5 × 4– 5 mm, grayish green- to tawny brown-tomentose, margin straight between the teeth at anthesis; corolla lobes spreading. Dwarf riparian forests, 800–1800 m; Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari). Northern Brazil (Amazonas: Serra da Neblina). ◆Fig. 238. Styrax neblinae (Maguire) P.W. Fritsch, Novon 14: 49. 2004. —Styrax duidae subsp. neblinae Maguire, Mem. New York Bot. Gard. 29: 210, fig. 96A–P, fig. 97A–F. 1978. Shrub or tree to 6 m tall; leaf blades subcoriaceous to coriaceous, the larger 6.3–14.4 × 3–7.8 cm, elliptic, domatia absent; lower

pedicels 5–7 mm long; calyx 5–6 × 5–6 mm, tawny brown-tomentose with scattered orange-brown trichomes, margin curved between the teeth at anthesis; corolla lobes spreading; longest filament trichomes 1.5– 1.75 mm long; anther sacs nontapered at apex. Dwarf open thickets and dry forests on sandstone substrate, 400–2100 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Endemic. Styrax sipapoanus Maguire, Mem. New York Bot. Gard. 29: 215, fig. 99A–L, fig. 100A–F, fig. 101A–G. 1978, “sipapoana.” Small tree; larger leaf blades 8.5–11.7 × 3.9–5.7 cm, ovate-elliptic, margin slightly revolute, domatia absent; inflorescences 3–6 cm long; lower pedicels 5–7 mm long; calyx 5–6 × 6 mm, grayish brown-tomentose, margin straight between the teeth at anthesis; corolla lobes spreading; anther sacs nontapered at apex. Dwarf woodlands, riparian forests, 1500–2100 m; Amazonas (Cerro Sipapo). Endemic. Styrax wurdackiorum Steyerm., Acta Bot. Venez. 10: 240. 1975. Shrub to 1.5 m tall; leaf blades coriaceous, elliptic to ovate-elliptic, margin revolute; inflorescences 1–3 cm long; calyx 2.5–4 × 4–4.5 mm, grayish green-lepidote, margin straight between the teeth at anthesis; corolla lobes recurved; longest filament trichomes 0.6–0.9 mm long. Shrubby thickets, dwarf shrub formations, rocky sandstone promontories in savanna formations, 1200– 1400 m; Bolívar (Río Aponguao basin). Endemic. ◆Fig. 239. Styrax yutajensis (Maguire) P.W. Fritsch, Novon 14: 56. 2004. —Styrax guanayanus var. yutajensis Maguire, Mem. New York Bot. Gard. 29: 207, fig. 95A– G. 1978, “guanayana.” Shrub or tree to 3 m tall; leaf blades elliptic to ovate-elliptic, margin revolute, domatia absent; calyx 5 × 5.5–6 mm, margin straight or slightly curved between the teeth at anthesis; longest filament trichomes 1.1– 1.3 mm long; anther sacs nontapered at apex. Dwarf forests on sandstone ridges and rocky areas bordering streams, 1300–2100 m; Amazonas (Cerro Coro Coro, Cerro Yutajé). Endemic.

288

S TYRACACEAE

Fig. 235. Styrax guanayanus

Fig. 237. Styrax duidae subsp. paruae

Fig. 236. Styrax guyanensis

Styrax 289

Fig. 238. Styrax longipedicellatus

Fig. 239. Styrax wurdackiorum

Fig. 240. Styrax glaber

290

S YMPLOCACEAE

SYMPLOCACEAE by Julian A. Steyermark and Paul E. Berry Shrubs or trees, glabrous or pubescent with multicellular trichomes or stalked glands. Leaves alternate, simple, entire or serrulate, exstipulate. Inflorescence axillary or terminal, racemose, cymose, paniculate, spicate, or fasciculate, few- to many-flowered, or reduced to 1 flower. Flowers fragrant, subtended by a bract and 2 bracteoles, perfect or rarely unisexual, regular, epigynous or infrequently semi-epigynous. Calyx (3)5-lobed, the lobes valvate or imbricate, basally connate, persistent in fruit. Corolla sympetalous, usually white or cream-colored, connate below into a short tube, 5–11-lobed, the lobes imbricate, in 1 or 2 series. Stamens 4–many; filaments attached from the middle to the base of the corolla, or free, usually in > 1 series or in fascicles alternate with the corolla lobes; anthers longitudinally dehiscent; pollen short-colpate. Pistil 1; ovary inferior or semi-inferior, usually 2–5-locular, placentation axile with 2–4 pendulous ovules in each locule; style slender, often surrounded by a disk; stigma simple, capitate or 2–5-lobed. Fruit ± fleshy, a berry or drupaceous, crowned by the persistent calyx, usually 1-seeded; endosperm abundant; embryo straight or curved. Tropical and subtropical America, southern and eastern Asia, Australia, East Indies; 1 genus and ca. 300 species, 11 species in the flora area. 1. SYMPLOCOS Jacq., Enum. Syst. Pl. 5. 1760. Characters and distribution as in family; ca. 300 species, 17 in Venezuela, 11 of these in the flora area. Key to the Species of Symplocos 1.

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

3.

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

Pubescence of lax or spreading trichomes on upper part of the branchlets, petioles, and lower surface of leaves, and/or on the lower veins; ovary 4-locular ............................................................................................. S. ulei Pubescence, if present, of appressed or sericeous trichomes; ovary 3- or 5-locular ................................................................................................. 2 Calyx lobes acute to acuminate at apex; style pubescent ........................ 3 Calyx lobes obtuse to rounded at apex; style glabrous to pubescent ....... 5 Inflorescence sessile or nearly so, shorter than the petiole, 2- or 3-flowered; calyx lobes 1.8–2 mm long, corolla 7 mm long; leaves minutely and distinctly crenulate ....................................................... S. acananensis Inflorescence equal to or longer than the petiole, shortly pedunculate, 5– 8-flowered; calyx lobes 2.5–3 mm long; corolla 8–9 mm long; leaves entire to subentire ..................................................................................... 4 Leaf margins entire; leaves pubescent on lower surface ......... S. guianensis Leaf margins subentire; leaves (sub)glabrous on lower surface .................... ............................................................................................S. schomburgkii Ovary 5-locular .......................................................................................... 6 Ovary 3-locular .......................................................................................... 7 Leaves acuminate at apex, acute at base, glabrous on lower surface; petiole glabrous throughout .................................................... S. martinicensis

Symplocos 291

6.

Leaves obtuse at apex, rounded or obtuse at the base, sparsely pubescent on the veins of the lower surface; petiole strigillose above ............................................................................................ S. chimantensis 7(5). Calyx glabrous without, except for the ciliate lobes; filaments not connate, free or nearly so to and at the base; leaves completely glabrous on lower surface; petioles glabrous .......................................... S. pycnophylla 7. Calyx appressed-pubescent, strigillose, or sericeous on exterior, at least on the lobes; filaments free above, connate below; leaves (at least the lower midrib) appressed-pubescent; petioles appressed-pubescent, sericeous, or glabrous ............................................................................ 8 8(7). Calyx lobes longer than the calyx tube; leaves rounded to obtuse at apex, rounded or obtuse at base, 3.5–6 cm long ............................... S. jauaensis 8. Calyx lobes shorter than or equal to the calyx tube; leaves acute or acuminate at apex, acute at base, 7–10 cm long ............................................ 9 9(8). Petioles glabrous; corolla 7–8 mm long; style pubescent in lower part; corolla 3–4 times the length of the calyx ...................................... S. nitens 9. Petioles pubescent, corolla 3–4.5 mm long; style glabrous; corolla 11/3 times the length of the calyx ......................................................................... 10 10(9). Fruit densely strigillose, ca. 5 mm long; stamens 25–40; lower surface of leaf sparsely appressed-pubescent to glabrescent; bracts suborbicular, rounded or obtuse; corolla 3 mm long ............................... S. yapacanensis 10. Fruit glabrate, 6–8 mm long; stamens 20–25; lower surface of leaf glabrous; bracts acute; corolla 4–4.5 mm long .............................. S. neblinae Symplocos acananensis Steyerm., Ann. Missouri Bot. Gard. 74: 656. 1987. Shrub ca. 1.5 m tall; leaf blades ellipticoblong, 4.5–8 × 2.5–3.5 cm; flowers fasciculate, calyx green, corolla white. Gallery forests bordering savannas, ca. 500 m; Bolívar (Río Acanán). Endemic. ◆Fig. 242.

land forests near streams and roadsides, 50– 100 m; Amazonas (30 km south of Samariapo near Río Sipapo). Guyana, Suriname, French Guiana, Brazil.

Symplocos chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 459. 1967. Tree ca. 5 m tall; leaves coriaceous, conspicuously rugose, oval-obovate or broadly obovate, 9.5–15.5 × 5–8 cm, strongly revolute; corolla 10–12.5 mm long; stamens 4–5seriate. Dwarf forests on tepui slopes and summits, 1900–2100 m; Bolívar (Macizo del Chimantá [Torono-tepui]). Endemic.

Symplocos jauaensis Steyerm. & Maguire, Mem. New York. Bot. Gard. 23: 876, fig. 12. 1972. Shrub 2–4 m tall; leaves coriaceous, 3.5–6 × 1.5–3 cm, strongly revolute, the lower surface silvery or pale green; flowers 3 or 4 in a congested raceme; calyx rose green; fruit oblong, ca. 10 mm long. Dwarf forests, sandstone ledges along small streams, rocky places among shrubby-herbaceous vegetation, 1300–2100 m; Bolívar (summit of Cerro Jaua), Amazonas (summit of Cerro Yaví). Endemic. ◆Fig. 246.

Symplocos guianensis (Aubl.) Gurcke in Engl. & Prantl, Nat. Pflanzenfam. I. 4: 172. 1891. —Ciponima guianensis Aubl., Hist. Pl. Guiane 566, t. 567. 1775. Shrub ca. 2 m tall; leaves chartaceous, 8– 11 × 2–4 cm, rusty brown-pubescent on stems and lower surface of leaves; flowers numerous in a congested raceme. Evergreen low-

Symplocos martinicensis Jacq., Enum. Syst. Pl. 24. 1760. Tree to 8 m tall; leaves coriaceous, ellipticoblong, 5–14 × 3–6 cm, entire to crenulate; corolla tube 10–12 mm long; fruit 9–10 mm long. Montane forests at base of sandstone escarpments, ca. 700 m; Bolívar (La Escalera). Venezuelan Coastal Cordillera;

292

S YMPLOCACEAE

Fig. 241. Symplocos yapacanensis

Fig. 242. Symplocos acananensis

Fig. 243. Symplocos neblinae

Symplocos 293

Fig. 244. Symplocos ulei

Fig. 245. Symplocos pycnophylla

Fig. 246. Symplocos jauaensis

294

S YMPLOCACEAE

Fig. 247. Symplocos schomburgkii

common in West Indies, Trinidad-Tobago, Guyana. Symplocos neblinae Maguire & Steyerm., Mem. New York Bot. Gard. 29: 229, figs. 109D, E. 1978. Shrub or tree to 8 m tall; leaves chartaceous, elliptic, elliptic-oblanceolate, or elliptic-lanceolate, 11–16 × 3–6.5 cm, entire; inflorescence mostly 3–6-flowered. Dwarf forests and wet upland shrubby savannas, 1800–2200 m; Amazonas (summit of Sierra de la Neblina). Endemic. ◆Fig. 243. Symplocos nitens (Pohl) Benth., Trans. Linn. Soc. London 18: 232. 1841.

—Stemmatosiphon nitens Pohl, Pl. Bras. Icon. Descr. 2: 88, t. 158. 1830. Venezuela, Brazil; 2 varieties, 1 in Venezuela. S. nitens var. claussenii Brand in Engl., Pflanzenr. IV. 242(6): 86. 1901. Tree to ca. 9 m tall; leaves obovate-oblong, 6–8 × 3.5–4.5 cm, subentire or scarcely undulate, glabrous on both sides. Evergreen lowland riparian forests, 100–200 m; Amazonas (Río Casiquiare, Río Pasiba, Río Pasimoni). Brazil. Variety claussenii differs from typical Symplocos nitens of northern and central Brazil in its smaller leaf blades.

T ACCACEAE 295

Symplocos pycnophylla Sleumer, Repert. Spec. Nov. Regni Veg. 42: 266. 1937. Symplocos roraimensis Steyerm., Fieldiana, Bot. 28: 494. 1953. Shrub or tree 0.5–7 m tall; leaves obovate or elliptic-obovate, rounded to obtusely acute at apex, cuneately acute at base; inflorescence closely racemosely flowered, ca. 2.5 cm long. Dwarf forests along streams, gallery forests, edges of sandstone escarpment, upland rocky savannas, (1300–)1700–2500 m; Bolívar (Ilú-tepui, Kamarkaibaray-tepui, near Kavanayén, Macizo del Chimantá, Roraimatepui), Amazonas (Cerro Parú, Sierra de la Neblina). Venezuelan Andes; Andean Colombia. ◆Fig. 245. Symplocos schomburgkii Klotsch ex Brand in Engl., Pflanzenr. IV. 242(6): 83. 1901. Shrub 3–4 m tall, usually completely glabrous; leaves coriaceous, 6–11 × 2.5–4 cm; fruit 5–6 mm long. Gallery forests and thickets bordering savannas, 500–900 m; Bolívar (Gran Sabana, drainages of Río Acanán, Río Yuruaní, and upper Río Aponguao). Probably in adjacent Guyana (the Roraima specimens collected by Schomburgk should be recorded for Venezuela and not Guyana). ◆Fig. 247.

Pending future study, Symplocos schomburgkii may eventually be considered synonymous with the older name S. guianensis (Aubl.) Gurke, which differs from S. schomburgkii in having more pubescent leaves and entire leaf margins. Symplocos ulei Brand, Notizbl. Königl. Bot. Gart. Berlin 6: 172. 1914. Shrub 1–4 m tall; leaves ovate or ovate- or oblong-elliptic, abruptly short-acuminate at apex, rounded to subacute at base, 5–13 × 1.5–4(–5) cm; inflorescence densely pubescent. Gallery forests bordering savannas, 300–1300 m; Bolívar (Gran Sabana in basins of Río Acanán, Río Mapaurá, Río Yuruaní, and lower Río Caroní). Guyana, adjacent Brazil. ◆Fig. 244. Symplocos ulei is variable in shape, base, and apex of leaf blade, as well as in the pubescence and margin of the leaf blade. Symplocos yapacanensis Steyerm., Acta Bot. Venez. 10(1): 242. 1975. Small tree ca. 3 m tall; leaves coriaceous, broadly elliptic, obtusely acuminate at apex, cuneately acute at base, 5–9 × 2.3–4.5 cm, entire. Dwarf forests on rocky tepui summit, 1000–1200 m; Amazonas (summit of Cerro Yapacana). Endemic. ◆Fig. 241.

TACCACEAE by James S. Miller Perennial herbs from a tuberous to elongate, starchy rhizome. Leaves basal, entire or pinnately lobed, large, long-petiolate. Inflorescences umbellate, borne on an elongate peduncle, subtended by an involucrate whorl of bracts, the inner often thin and thread-like. Flowers actinomorphic, epigynous, 3-merous, bisexual, the perianth not clearly differentiated into sepals and petals. Tepals 6, alternating in 2 whorls, petaloid, usually dark colored, often basally connate in a short tube. Stamens 6, in 2 whorls; filaments short, basally adnate to the tepals, flattened and somewhat petaloid, the connective broad; anthers bithecal, dehiscing longitudinally. Ovary inferior, 3-carpellate, unilocular, the placentation parietal; ovules numerous, anatropous to campylotropous; style short; stigmas 3, somewhat petaloid. Fruit baccate or rarely a 3-valved, loculicidal capsule. Seeds numerous; endosperm abundant. Pantropics, centered in southeastern Asia, with a single species known in the New World; 1 genus and 10 species, 1 species in the flora area.

296

T ACCACEAE

1. TACCA J.R. Forst. & G. Forster, Char. Gen. Pl. 69. 1775. Characters and distribution as in the family. Tacca parkeri Seem., Fl. Vit. 102. 1865. Tacca sprucei Benth. in Benth. & Hook. f., Gen. Pl. 3: 741. 1883. Tacca parkeri var. lanceolata H. Limpr. in Engl., Pflanzenr. IV. 42: 21. 1928. Tacca ulei H. Limpr. in Engl., Pflanzenr. IV. 42: 22. 1928.

Rhizomatous herb; flowers purple. Moist white-sand savannas, small streams, edges of lowland forests, 100–400 m; Amazonas (Caño San Miguel, Río Atacavi, Río Pasimoni, Río Sipapo, lower Río Ventuari, Solano). Guyana, adjacent Brazil. ◆Fig. 248.

Fig. 248. Tacca parkeri

Tepuianthus 297

TEPUIANTHACEAE by Paul E. Berry and Zachary S. Rogers Trees or shrubs, with bitter bark that can be peeled off in strips. Leaves alternate or opposite, simple, entire, pinnately veined, exstipulate. Inflorescences cymose, terminal or subterminal. Flowers actinomorphic, staminate or bisexual (plants androdioecious). Sepals 5, free, imbricate; petals 5, free, imbricate, yellow, clawed at the base. Nectary disk extrastaminal, with 5–10 discrete, contiguous glands; glands glabrous externally, usually pubescent internally. Stamens 5–16(–22), in 1–3 cycles, alternate with the petals when unicyclic; anthers sagittate, versatile and dorsifixed, bithecate, introrse, longitudinally dehiscing; connective sometimes modified; pollen 3-colporate, exine reticulate. Receptacle densely hirsute; ovary superior, compound, carpels and locules 3; styles 3, bifid; ovules 1 per locule, anatropous, pendulous, raphe ventral. Fruit a loculicidal capsule, densely sericeous. Seeds glabrous; endosperm abundant; embryo small. Southeastern Colombia (Vaupés), southern Venezuela, northern Amazonian Brazil (restricted to the Guayana Shield); 1 genus and 6 species, 4 species in Venezuela, all in the flora area. Wurdack & Horn (BSA abstract, 2001) performed a parsimony analysis of sequence data from 18S rDNA, atpB, and rbcL, which placed Tepuianthus within the Malvales, sister to, or at the base of, Thymelaeaceae sensu lato (i.e., including Octolepidoideae). Tepuianthus is, in fact, extremely similar to the New Caledonian Solmsia (Thymelaeaceae, Octolepidoideae; 1 or 2 species), down to the emarginate, tough leaves with a venose and glossy upper leaf surface and a sericeous indumentum on the lower surface. However, the flowers of Tepuianthus, unlike those of Thymelaeaceae, possess a true corolla, which is composed of 5 clawed, yellow petals (J.W. Horn. Bot. J. Linn. Soc. 144: 1–40. 2004). Close phylogenetic relationships of other Guayana Shield-centered genera with Malesian and Australasian groups are shown by Archytaea (with Ploiarium), Pentamerista (with Tetramerista), and Pakaraimaea and Pseudomonotes (with Old World Dipterocarpaceae). 1. TEPUIANTHUS Maguire & Steyerm., Mem. New York Bot. Gard. 32: 9. 1981. Characters and distribution as in family; 6 species, 4 species in Venezuela, all in the flora area. Tepuianthus colombianus Maguire & Steyerm., from Cerro Isibukuri in Departamento Vaupés, Colombia, may be conspecific with T. savannensis. The sixth species, T. aracensis Steyerm. & Maguire, is endemic to Serra Aracá, in northern Estado Amazonas, Brazil. Key to the Species of Tepuianthus 1.

1.

Inflorescence erect, congested; peduncles to 4 mm long; pedicels 1– 2 mm long; midrib on lower surface of leaf with spreading to loosely ascending trichomes; secondary and tertiary venation on lower surface of leaf generally evident ........................................................................ 2 Inflorescence pendent, laxly flowered; peduncles 7–15 mm long; pedicels 2–8 mm long; midrib on lower surface of leaf with finely appressed, minute trichomes; secondary and tertiary venation on lower surface of leaf mostly concealed by a fine tomentum ............................................ 3

298

2(1).

2.

3(1). 3.

T EPUIANTHACEAE

Stamens 5; sepals acute to acuminate at apex; secondary leaf veins 12– 15 per side, 3–4 mm apart in lower half; leaf blade mucronate; shrubs 0.5–3 m tall .................................................................... T. auyantepuiensis Stamens 12–16; sepals obtuse to rounded at apex; secondary leaf veins 16–23 per side, 1–2 mm apart in lower half; leaf blade scarcely or not mucronate; trees 3–10 m tall ............................................ T. yapacanensis Anther connective entire at apex; petals 3.5–5 mm long; pedicels 2– 4 mm long ..................................................................... T. sarisariñamensis Anther connective multifimbrillate or erose at apex; petals 5.5–6 mm long; pedicels 4–7 mm long ................................................. T. savannensis

Tepuianthus auyantepuiensis Maguire & Steyerm., Mem. New York Bot. Gard. 32: 10, figs. 1, 2. 1981. Shrub 0.5–2.5(–3) m tall; leaves obovate, 1.6–5 × 1–2.1 cm, petioles 3–7 mm long; flowers 2.5–4.5 × 2.5–3 mm, sepals 2–4.5 × 1.2–2 mm; fruits 6–10 × 5–8 mm. Locally abundant in dense colonies in tepui scrub savannas, 1600–2200 m; Bolívar (Auyántepui). Endemic.

Tepuianthus sarisariñamensis Maguire & Steyerm., Mem. New York Bot. Gard. 32: 16. 1981. Branched tree or shrub 3–7 m tall; leaves obovate or oblanceolate, 1.2–5.8 × 0.7–1.6 cm, petioles (4–)6–10 mm long; flowers ca. 3.5 × 3.5 mm, sepals ca. 3.2–4 × 2–2.8 mm; fruits ca. 7 × 6 mm. Dwarf forests on tepui summits, (700–)1100–1500 m. Venezuela; 2 subspecies, both in the flora area.

Fig. 249. Tepuianthus sarisariñamensis subsp. duidensis

Tepuianthus 299

Fig. 250. Tepuianthus savannensis

Key to the Subspecies of T. sarisariñamensis 1. Pubescence of lower surface of leaf, petioles, and stem apex minutely and closely appressed, not loosely ascending at tip; reticulation of upper surface of leaf with small, regular areoles ................................. subsp. duidensis 1. Pubescence of lower surface of leaf, petioles, and stem apex densely silky with slightly ascending tips; reticulation of upper surface of leaf with larger, irregular areoles ... subsp. sarisariñamensis T. sarisariñamensis subsp. duidensis Maguire & Steyerm., Mem. New York Bot. Gard. 32: 18. 1981. Open savannas and forest patches, (700–) 1100–1500 m; Amazonas (Cerro Duida, Cerro Parú). Endemic. ◆Fig. 249. T. sarisariñamensis subsp. sarisariñamensis Dwarf forests on tepui summits, 1300– 1400 m; Bolívar (Cerro Sarisariñama). Endemic.

Tepuianthus savannensis Maguire & Steyerm., Mem. New York Bot. Gard. 32: 14. 1981. Small tree 2–8 m tall; leaves obovate, oblanceolate, or rarely elliptic, 1.5–5.3 × 1– 2.1 cm, petioles 2–5 mm long; flowers 3–4.5 × 4–5 mm, sepals 3–4.5 × 2.5–3.5 mm; fruits 7–10 × 7–10 mm. In and along edges of white-sand savannas, 100–600(–800) m; Amazonas (Caño Yagua, Cerro Vinilla, Río Autana, Río Guayapo, lower Río Ventuari). Colombia (Vaupés). ◆Fig. 250. Tepuianthus yapacanensis Maguire & Steyerm., Mem. New York Bot. Gard. 32: 12, fig. 3. 1981. Branched tree 8–10 m tall; leaves obovate or oblanceolate, 3–4.5 × 1–1.8 cm, petioles 3–5 mm long; flowers ca. 4.5 × 5 mm, sepals 2.1–4 × 1.5–2.5 mm; fruits ca. 6 × 5 mm. Dominant in tepui summit dwarf forests, 1000–1200 m; Amazonas (Cerro Yapacana). Endemic.

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TERNSTROEMIACEAE by Paul E. Berry and Anna L. Weitzman Evergreen trees or shrubs, accumulating aluminum; indumentum of unicellular hairs. Leaves distichous or spiral, conduplicate-involute or supervolute; usually evergreen and coriaceous; margins entire or toothed, a small, deciduous seta near the apex of each tooth; stipules absent. Inflorescences axillary, rarely pseudo-terminal. Flowers solitary or fasciculate (in short racemes of up to 15 flowers), bisexual or unisexual (plants then dioecious), pedicellate, regular, 5-merous, usually < 2 cm diameter, pedicel with ± apical prophylls. Sepals quincuncial, connate or distinct, persistent in fruit, usually thick, concave, subequal (unequal); petals distinct or connate basally, quincuncial, thick or membranous. Stamens 5–many, free, connate or adnate to corolla basally; filaments equal or unequal, wide and flat or thickened, often narrowing and incurved apically, shorter than to 4 times as long as the anthers; anthers basifixed, introrse, oblong, opening by longitudinal slits (flaps or pore-like slits), connective often elongated. Carpels (2)3–5(6); ovary superior (inferior), placentation axile (apical; parietal); ovules anatropous to campylotropous, many per carpel, rarely 1 or 2; style simple or with apical style branches, or divided into separate stylodia; stigmas usually separate. Fruit a berry (irregularly dehiscent or a loculicidal capsule). Seeds 1–numerous, reniform, small, reticulate or areolate, or larger, ellipsoid, ± smooth; endosperm slight to copious; embryo horseshoe-shaped to ± straight. Tropics and subtropics, mostly in Asia and the Neotropics, a few in Africa; 12 genera and ca. 340 species, 2 genera and 21 species in the flora area. This family was included by Cronquist in the Theaceae, but has since been shown to be quite distinct from that family (see discussion under Theaceae). It has alternatively been treated by Stevens (Angiosperm Phylogeny Website; http:// www.mobot.org/MOBOT/research/APweb/) in the Pentaphylaceae. Key to the Genera of Ternstroemiaceae 1.

1.

Leaves distichous, spread along twigs, usually at least slightly sinuous; flowers in axillary racemes, rarely solitary; corolla urceolate; fruits indehiscent; seeds minute (< 1.5 mm long), brown or black, without sarcotesta .................................................................................... 1. Freziera Leaves spiraled, generally entire, usually clustered near branch apices (pseudoverticillate); flowers usually solitary and axillary; corolla campanulate or spreading; fruits irregularly dehiscent; seeds larger than above, often with reddish sarcotesta ................................. 2. Ternstroemia

1. FREZIERA Willd., Sp. Pl. 2: 1179, nom. cons. Eroteum Sw., Prodr. 85. 1788. Lettsomia Ruiz & Pav., Fl. Peruv. Prodr. 4: 77. 1794. —Eurya sect. Freziera Szyszyl. in Engl. & Prantl, Nat. Pflanzenfam. III. 6: 190. 1893. Eurya subg. Freziera Melchior in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 148. 1925. Patascoya Urb., Ber. Deutsch. Bot. Ges. 14: 283. 1896. Killipiodendron Kobuski, J. Arnold Arbor. 23: 231. 1942. Trees or shrubs, frequently pubescent, dioecious, stems usually zigzag, often with paired wings or ridges decurrent from petiole base extending through two in-

Freziera 301

ternodes. Leaves distichous; blades often pubescent above and/or below; margin usually sinuous; petiole, when present, winged. Inflorescence axillary, short-racemose or rarely single-flowered with 1–15 flowers; pedicel usually under 1 cm long. Flowers to 15 mm long, actinomorphic, functionally unisexual, though staminate flowers appear bisexual and carpellate flowers contain staminodia; bracteoles 2, persistent, closely subtending calyx. Sepals distinct, broadly ovate, nearly always convex and sclerotic (at least centrally), the outer surface often pubescent; corolla urceolate, petals distinct or lightly connate basally, imbricate, membranaceous in basal 1/4–1/2, sclerotic and appearing waxy above. Stamens (in staminate flowers) (8–)15–35(–48), uniseriate; filaments free or adnate basally, equal or unequal, often bent; anthers basifixed, usually equal, connective usually extending into a short apiculus; staminodia (in carpellate flowers) (6–)15–35(–38), uniseriate, free or adnate basally, flattened, usually equal. Gynoecium (in staminate and carpellate flowers) broadly pyriform to conical, (2)3–5(6)-carpellate, with locules containing (0 or 2–6–)16–100(–150) ovules; style tapering; stigmatic lobes flaring (or erect), with abaxial and adaxial (or only adaxial) surfaces papillate (staminate flowers usually have narrower gynoecia with fewer ovules and erect stigmatic lobes). Fruit usually a fleshy berry (or a pyrene), when ripe, reportedly black to purple to dark blue, but usually collected immature, green, tapering abruptly into persistent style. Seeds irregularly reniform; embryo slightly curved or rarely horseshoe-shaped (hippocrepiform), (0–3–)25–60(–128) per locule, dark red (or brown or black); testa reticulate (or areolate). Southern Mexico, Central America, Colombia, Venezuela, Guayana, Ecuador, Peru, Bolivia, the West Indies; 56 species, 11 in Venezuela, 3 of these in the flora area. Key to the Species of Freziera 1.

1.

2(1).

2.

Branchlets terete; leaf blades with ca. 11 lateral veins and 34–44 marginal serrations per side; indumentum of mixed curled and straight hairs ...................................................................................... F. roraimensis Branchlets square or flattened, often with 4 wings; leaf blades with 15– 30 lateral veins and (46–)60–130 marginal serrations per side; indumentum, if present, entirely of straight hairs ............................. 2 Leaf base usually unequal, at most slightly revolute; midrib and petiole only rounded below; leaf blade usually large (10–)12–20(–30) × (3–)5– 10(–19) cm ............................................................................... F. calophylla Leaf base equal, strongly revolute; midrib and petiole usually keeled below; leaf blade smaller (4.1–)9.2–14.8 × (2.1–)2.9–4.9(–6) cm .... F. carinata

Freziera calophylla Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 261. 1862. —Eroteum calophyllum (Triana & Planch.) O. Kuntze, Revis. Gen. Pl. 1: 62. 1891, “Erotium.” —Eurya calophylla (Triana & Planch.) Szyszyl. in Engl. & Prantl, Nat. Pflanzenfam. III. 6: 190. 1893. Freziera hieronymi Kobuski, Ann. Missouri Bot. Gard. 25: 355. 1938, “hieronyma,” for Eroteum nitida Hieron.

Freziera cuatrecasasii Kobuski, J. Arnold Arbor. 34: 136. 1953. Freziera esmeraldana Little, Phytologia, 18: 466. 1969. Tree 8–25 m tall. Montane forests, ca. 1300 m; Amazonas (headwaters of Río Orinoco in Sierra Parima). Panama, Colombia, Ecuador, Peru. Freziera calophylla is morphologically heterogeneous. All specimens share a conspicuously canaliculate, wide midrib. In

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addition, most specimens have erect (or only slightly involute) petiole wings, small flowers, and large coriaceous leaves with regular, nearly straight lateral veins. ◆Fig. 251. Freziera carinata Weitzman, J. Arnold Arbor. 68: 326. 1987. Small to medium sized tree characterized by its winged stems and prominently keeled petioles and midribs; leaf blades auriculate at the base, quickly becoming revolute and the blades then appearing attenuate. MonFig. 251 Freziera calophylla

tane forests on steep slopes, (700–)1700– 2300(–2600) m; Bolívar (Aparamán-tepui, Auyán-tepui, Cerro Guaiquinima, Cerro Guanay, Cerro Jaua, Cerro Sarisariñama, Macizo del Chimantá, Ptarí-tepui, upper Río Venamo, Roraima-tepui, Sierra de Lema), Amazonas (Cerro Aracamuni, Cerro Huachamacari, Cerro Marahuaka at Salto los Monos, Sierra de la Neblina, Yutajé). Endemic. ◆Fig. 252. Freziera roraimensis Tul., Ann. Sci. Nat. Bot. sér. 3, 8: 339. 1847. —Eurya roraimensis (Tul.) Szyszyl. in Engl. & Prantl, Nat. Pflanzenfam. III. 6: 190. 1893. Lettsomia guianensis Klotzsch ex M.R. Schomb., Reis. Br.-Guiana 3: 1092. 1848, nom. nud. —Freziera guianensis Klotzsch ex Wawra in Mart., Fl. Bras. 12(1): 284. 1886. —Eroteum guianense (Klotzsch ex Wawra) Kuntze, Rev. Gen. Pl. 1: 62. 1891, as “Erotium.” —Eurya guianensis (Klotzsch ex Wawra) Szyszyl. in Engl. & Prantl, Nat. Pflanzenfam. III. 6: 190. 1893.

Fig. 252. Freziera carinata

Ternstroemia 303

Probably a small tree, with a dense pubescence of partly curly and partly straight hairs. Probably tepui slope forest, elevation

not recorded; Bolívar (slopes of Roraimatepui but Venezuelan or Guyanan side not specified). Guyana.

2. TERNSTROEMIA Mutis ex L. f., Suppl. 39, 264. 1782, nom. cons. Shrubs or trees. Leaves pseudoverticillate, often with black dots on the lower surface; leaf margin entire or less often crenulate. Flowers single, paired, or fasciculate, in the axils of nonexpanded leaves, rather large (usually > 1.5 cm across), bracteolate, the bracteoles 2(4), opposite or subopposite; sepals 5(6), imbricate, persistent; petals 5(6), free or basally connate. Stamens many, 3-seriate; filaments connate, shorter than the anthers; anthers retuse to apiculate, base hardly raised. Ovary 1–3-locular, or 4–6-locular with false septae, 1–20 pendulous ovules per locule; style 1, entire or rarely deeply 2- or 3-parted. Fruit indehiscent, irregularly or less often valvately or circumscissilely dehiscent. Seeds usually > 4 mm long, reddish brown; testa fleshy. Tropical and subtropical America, Sri Lanka to southeastern and eastern Asia, (most species), Africa (2 species); ca. 100 species, ca. 22 in Venezuela, 18 of these in the flora area. Ternstroemia is a genus that has not been examined in any depth for South America since the study of Kobuski (C. E. Kobuski. J. Arnold Arbor. 23: 298–343. 1942). Since then a great many additional collections have been made, such that the genus is seriously in need of a taxonomic revision for the entire region. Because of the lack of a revised treatment and the poor understanding of species limits in the genus, it was decided not to provide a diagnostic key to the species at this time. Instead we are providing a list of the taxa described from the Venezuelan Guayana from the literature, as well as species described from neighboring areas that are likely to be found in the flora area. We know that there are a number of as yet undescribed species from the flora area as well, but we will mostly leave these for the attention of future workers in Ternstroemia. Ternstroemia brevistyla Kobuski, J. Arnold Arbor. 23: 310. 1942. Shrub with grayish, terete branchlets; leaves coriaceous, obovate to elliptic, 3–5.5. × 2–3.2 cm, apex rounded, base widely cuneate, margin entire except for crenulate apex; petiole 4–8 mm long; fruits subgloboid, ca. 6 mm diameter, 3(4)-locular, each locule 1-seeded; style 1 mm long, stigma subcapitate, 3-crenate; pedicels 6–8 mm long. Tepui slope scrub and montane shrubland, 1000–1300 m; Bolívar (slopes of Auyántepui, El Jardín between Parupa and Kavanayén, Río Karuai on southwest slopes of Ptari-tepui, Río Yuruaní). Endemic. Ternstroemia campinicola B.M. Boom, Brittonia 41: 138. 1989. Shrub or small tree 1–6 m tall; branchlets opposite or fasciculate; leaves oblanceolate to obovate, 3–6 × 1–3 cm, dull-surfaced,

margins black-glandular on lower surface; petiole 2–9 mm long; flowers 1–6-fasciculate; peduncles 4–14 mm long; fruits 2-locular, 4-seeded or fewer; persistent style ca. 5 mm long. White-sand savannas, 100–300 m; Amazonas (Caño Yagua, base of Cerro Yapacana, lower Río Siapa, Sabana Hechimoni in Río Siapa basin, San Antonio del Orinoco, Yavita). Colombia (Caquetá, Vaupés), Brazil (Amazonas, Roraima). Ternstroemia candolleana Wawra in Mart., Fl. Bras. 12(1): 273. 1886. Ternstroemia candolleana var. rotundata Wawra in Mart., Fl. Bras. 12(1): 274. 1886. Shrub; leaves chartaceous to subcoriaceous, oblong-obovate, 7–10 × 1.5–3 cm, apex obtuse to acuminate, base attenuate, shiny on upper surface, ferruginous on lower surface; petiole ca. 1 cm long; flowers soli-

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tary, pedicels 2–4 cm long, filiform; ovary 3locular, each locule with 2 ovules; style 7–12 mm long; stigma punctiform; fruit 1–1.5 cm long, sulcate, usually 6-seeded. Evergreen lowland and riparian forests, 100–200 m; Amazonas (Río Guainía above mouth of Río Casiquiare). Colombia (Guainía), Guyana, Brazil (Amazonas). Ternstroemia crassifolia Benth., London J. Bot. 2: 363. 1843. Shrub; leaves coriaceous, obovate, 4–6 × 1.5–3 cm, apex obtuse to rounded, base cuneate, margin subrevolute, veins obscure, petiole 8–10 mm long; flowers solitary or few-fasciculate, at end of branchlets; pedicels ca. 6 mm long; ovary 2-locular, each locule with 1 or 2 ovules; style shorter than the ovary; stigma discoid, 2-lobed. Montane scrub, savannas, 900–1400 m; Bolívar (Gran Sabana, between Parupa and Kavanayén at El Jardín, near Roraima-tepui). Guyana. ◆Fig. 255. Ternstroemia crassifolia is purportedly close to T. discoidea. Ternstroemia discoidea Gleason, Bull. Torrey Bot. Club 58: 398. 1931. Ternstroemia monosperma Gleason, Bull. Torrey Bot. Club 58: 399. 1931. Shrub ca. 3 m tall, young branches verticillate; leaves coriaceous, elliptic-oblong or cuneate-oblong, 3–5.5 × 1–2.5 cm, apex rounded-retuse, base cuneate, margin finely crenulate, somewhat revolute, dark punctate on lower surface; petiole 5–8 mm long; flowers solitary, white, pedicels to 28 mm long; ovary apparently 1-locular, stigma peltate; fruit ca 13 mm long, 1- or 2-seeded. Tepui summit scrub vegetation, 1400–2200 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 256. Ternstroemia distyla Kobuski, J. Arnold Arbor. 23: 317. 1942. Shrub with terete, subverticillate branchlets; leaves coriaceous, elliptic-lanceolate, 4.5–6.5 × 1–1.5 cm, apex and base acute, margin denticulate-crenulate, lower surface punctate; petioles 5–7 mm long; fruit solitary, conical, 2-locular, 4-seeded; pedicels slender, often recurved, 13–17 mm long; style ca. 1 mm long, 2-parted; stigmas 2, subcapitate-reniform. Upper tepui slopes and

summit scrub, 1400–2000 m; Bolívar (Auyántepui), Amazonas (Cerro Aracamuni). Guyana. Ternstroemia duidae Gleason, Bull. Torrey Bot. Club 58: 400. 1931. Ternstroemia paucifolia Gleason, Bull. Torrey Bot. Club 58: 401. 1931. Ternstroemia duidae f. latifolia Kobuski, J. Arnold Arbor. 23: 324. 1942. Shrub ca. 2 m tall, with slender, verticillate branchlets; leaves coriaceous, linearoblanceolate, 3–5 × 0.4–0.6(–1.5) cm, apex obtuse, base long-cuneate, margin subentire; petiole to 4 mm long; flowers few, solitary, pedicels slender, recurved, to 3 cm long; ovary conical, ca. 2 mm long, 2-locular, fewovulate, tapering into a stout style 5–6 mm long; stigma punctiform; calyx and corolla pink-white; fruit ca. 13 mm diameter, 6–8seeded. Tepui summit savannas and scrub, 1300–2000 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 257. Ternstroemia dura Gleason, Bull. Torrey Bot. Club 58: 401. 1931. Shrub with subverticillate, fissured branchlets; leaves thick-coriaceous, ellipticoblong, to 3 × 1.2 cm, apex obtuse and slightly retuse, base attenuate, margin revolute; petiole 1–2 mm long; fruit solitary, 2locular, usually 4-seeded; pedicels stout, to 3 cm long; style entire, ca. 9 mm long, stigma punctiform. Tepui summit savanna and scrub, 1300–2200 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 253. Ternstroemia guanchezii B.M. Boom, Ann. Missouri Bot. Gard. 76: 970 .1989. Shrub 2–3 m tall with graceful, subopposite to opposite branchlets; leaves clustered in the apical 3–5 cm of the branchlets, oblanceolate to narrowly elliptic, 5–8.5 × 1– 3 cm, apex acute to acuminate, base acuminate to attenuate; petiole 0.2–1.0 mm long; flowers to 7 per axillary fascicle; peduncles to 10 mm long; ovary 2-locular with 2 ovules per locule. Forests along streams along granitic outcrops, 100–600 m; Amazonas (Raudal Manteco in Caño Cabeza de Manteco in Río Autana basin, slope of Cerro Aracamuni). Endemic. Ternstroemia laevigata Wawra in Mart., Fl. Bras. 12(1): 281, pl. 55. 1886.

Ternstroemia 305

Shrub with terete, smooth branchlets; leaves coriaceous, oblong-obovate, 5–8 × 2–3 cm, pale green on upper surface, reddish on lower surface, both surfaces smooth and epunctate, margin subrevolute and denticulate; petiole ca. 1 cm long; flowers few, peduncles 1.5–3 cm long; ovary globose, striate, 2-locular, each locule 2-seeded; style filiform; stigma minute and punctiform. Presumably montane forests, elevation not recorded; Bolívar (area surrounding Roraima-tepui). Guyana. Ternstroemia maguirei B.M. Boom, Ann. Missouri Bot. Gard. 76: 973. 1989. Shrub or small tree 1–6 m tall; branchlets fasciculate, angular in cross section; leaves distributed along branchlets, elliptic to ovate or obovate, 7.5–14 × 4–7 cm, apex acute to rounded; petioles 5–15 mm long; flowers subterminal in pairs; peduncles 11–50 mm long; ovary 2-locular with 8 ovules per locule; fruits smooth, depressed-ovoid, ca. 15 × 16 mm, fruit wall 2.5–3 mm thick, apparently indehiscent. Exposed montane rock outcrops, tepui slopes and ridges, 800–2400 m; Amazonas (Cerro Aracamuni, Cerro Duida, Sierra de la Neblina). Probably also on the Brazilian side of Serra da Neblina (Amazonas). Ternstroemia punctata (Aubl.) Sw., Prod. 81. 1788. —Taonabo punctata Aubl., Hist. Pl. Guiane 571. 1775. Ternstroemia revoluta Splitg. in Hoeven & de Vriese, Tijdschr. Nat. Gescheid. 9: 99. 1842. Small tree 3–8 m tall, with twisted, nonverticillate branchlets; leaves stiffly coriaceous, obovate, 1–6(–8) × 2–3(–4) cm, apex obtuse to rounded and emarginate, base cuneate, margin entire and revolute, veins clearly evident on upper surface, lower surface dark punctate; petiole 3–5 mm long; flowers solitary, pedicels 2–3 cm long; ovary 3-locular, each locule with 3 or 4 ovules; style 3-parted; stigmas 3, punctiform. Tepui slope scrub, 1000–1400 m; Bolívar (Auyán-tepui, Roraima-tepui). Guyana, Suriname, French Guiana, Brazil. Ternstroemia pungens Gleason, Bull. Torrey Bot. Club 58: 400. 1931. Shrub with terete, striate branchlets; leaves coriaceous, obovate to oblong-obovate,

to 5.5 × 3 cm, apex rounded to retuse, base cuneate, margin subentire or finely crenulate, often revolute, lower surface reddish and black-punctate; petiole 3–4 mm long; calyx lobes sharp-pointed, 13–15 mm long; fruit on long, slender, recurved pedicels to 3 cm long, subglobose, ca. 1 cm diameter, 2-locular, 4–6-seeded; style persistent, 6–7 mm long, 2parted. Montane to lower montane forests and outcrops, 300–1900 m; Bolívar (west of Canaima, Cerro Guaquinima, Gran Sabana, west of Kavanayén, north of Las Bonitas, slopes of Ptari-tepui, Río Cucurital, middle Río Paragua, Río Trueno north of Cerro Guaiquinima, south of Roraima-tepui, top of Salto Aicha at base of Uaipán-tepui), Amazonas (Cerro Sipapo, Cerro Duida, Cerro Parú, Sierra de la Neblina, lower slopes of Sierra Unturán). Suriname(?), Brazil. ◆Fig. 258. Ternstroemia retusifolia Kobuski, J. Arnold Arbor. 23: 328. 1942. Shrub with terete, verticillate branchlets; leaves grouped at branch tips, thick-coriaceous, 2.5–3 × 1–1.3 cm, apex rounded and retuse, base attenuate, margin crenulate and plane, punctate on lower surface; petiole 3–5 mm long; fruits axillary, or congested near branch tips, globose, 2-locular, each locule 3seeded; pedicels 1.5–2 cm long; style persistent, 5–6 mm long. Tepui slope scrub, 1600– 2000 m; Bolívar (Auyán-tepui). Endemic. Ternstroemia tristyla Gleason, Bull. Torrey Bot. Club 58: 398. 1931. Shrub with terete branchlets; leaves coriaceous, sessile, 2.5–3 × 1–1.3 cm, apex rounded and minutely retuse, base cuneate, margin subrevolute with minute glandular teeth, veins hardly visible; flowers solitary, pedicels compressed, ca. 1.5 cm long; ovary 2locular, each locule with 5 or 6 ovules; style ca. 7 mm long, 2-parted for ca. 3 mm, the 2 stigmas punctiform. Tepui slope and summit scrub, 1300–1500 m; Amazonas (Cerro Duida). Endemic. Ternstroemia tristyla is supposedly closely related to T. punctata, with sessile, smaller, cuneate leaves that are less punctate on the lower surface. Ternstroemia verticillata Klotsch ex Wawra in Mart., Fl. Bras. 12(1): 272. 1886.

306

T ERNSTROEMIACEAE

Small tree with 5- or 6-verticillate branchlets; leaves coriacrous, obovate to cuneate-oblong, 2–4 × 1–2 cm, often 3–5-verticillate, apex truncate to retuse, pale green and fuscous on upper surface, granularpunctate on lower surface, veins inconspicuous, margin glandular-crenulate near the apex, subsessile or petiole to 5 mm long; flowers solitary, petals connate for 5 mm basally; pedicels ca. 1 cm long, graceful; ovary 2-locular, each locule with 2 ovules, style ca. 4 mm long; stigma punctiform, 2-lobed. Dense tepui scrub, 1800–2400 m; Bolívar (Auyántepui, Macizo del Chimantá [Abacapá-tepui, Acopán-tepui, Toronó-tepui]). Guyana, Suriname. Ternstroemia sp. A Shrub or small tree 2–4 m tall; leaves coriaceous, glaucous on upper side when dried, broadly elliptic to orbicular, 8–12 × 6–8 cm; flowers on pedicels 1–4 cm long; petals cream, anthers yellow. Edges of forest patches near savannas, ridges, (400–)1200– 2000 m; Amazonas (Cerro Parú, Cerro Vinilla, Cerro Yutajé, Sierra de la Neblina). Brazil (Amazonas: side of Serra Neblina). ◆Fig. 254. Ternstroemia sp. B Shrub or small tree 2–3 m tall; leaves tightly grouped at branch tips, narrowly oblanceolate, 3–5 × 1–2.5 cm. Seasonally flooded black-water shrub islands in whitesand savannas, 100–200 m; Amazonas (Cerro Yapacana, west of Pimichín, Río Guasacavi). Colombia (Guainía). Species of Ternstroemia described from the Guianas and likely to occur in the Venezuelan Guayana Ternstroemia browniana Kobuski, J. Arnold Arbor. 23: 309. 1942. Shrub to 6–20 m tall with terete, subverticillate branchlets; leaves coriaceous, obovate, 4–6 × 1.5–2.5 cm, apex rounded or obtuse, base attenuate, margin crenulate, subrevolute, veins inconspicuous, petioles 3– 5(–7) mm long; flowers solitary, pedicels 2– 2.5 cm long, slender; ovary conical, 3-locular, locules 1-ovulate, style > 25 mm long, stigma

punctiform. Possibly in eastern Bolívar (known from lowland areas in Guyana, Suriname, and possibly French Guiana). Ternstroemia browniana is said by Kobuski to be most closely related to T. delicatula. Ternstroemia delicatula Choisy, Mém. Soc. Phys. Genève 14: 106. 1855. Shrub; leaves membranous, obovate-elliptic, 5–9 × 2–4 cm, apex obtuse or shortly acuminate, base attenuate, crenulate in upper half, lateral veins prominulous on both surfaces; petioles 6–15 mm long; flowers numerous, crowded; peduncle slender, 1–2 cm long, recurved; ovary 3-locular, each locule with one ovule, contracted into a style ca. 4 mm long; stigma subcapitate, 3-crenate. Possibly in eastern Bolívar (known from low to mid-elevations in Trinidad, Guyana, Suriname, and French Guiana). Ternstroemia dentata (Aubl.) Sw., Prodr. 81. 1788. —Taonabo dentata Aubl., Hist. Pl. Guiane 569. 1775. Tree or shrub; leaves coriaceous, obovate to elliptic-obovate, 6–13 × 2–7 cm, apex obtusely acuminate, base cuneate, margin serrate, 8–12 pairs of secondary veins, lower surface sparsely dark-punctate; petiole 7– 10(–20) mm long; flowers solitary or 2-fasciculate; pedicel 13–25 mm long; ovary 2- or 4-locular, 4-ovulate, tapering through the 2.5–3 mm long style into a punctiform stigma; fruit subglobose, ca. 1 cm diameter, 4-seeded. Possibly in eastern Bolívar (known from lowland areas in Guyana, Suriname, French Guiana, and Brazil). Ternstroemia gleasoniana Kobuski, J. Arnold Arbor. 23: 305. 1942. Shrub 3–5 m tall with thick, terete branchlets; leaves thick-coriaceous, oblongelliptic or oblong-obovate, 18–21 × 7–8 cm, apex obtuse or subrounded, abruptly acuminate, base obtusely cuneate, margin subentire, subrevolute, 18–20 pairs of secondary veins; petioles thick, 2–2.5 cm long; flowers congested toward branchlet tips; pedicels thick, 6–8 mm long; ovary 1-locular, each locule with few ovules; style entire, 4 mm long, stigma punctiform. Possibly in eastern

Ternstroemia 307

Bolívar (described from Potaro Landing along Potaro River in Guyana). Ternstroemia grandiosa Kobuski, J. Arnold Arbor. 23: 318. 1942. Tree ca. 12 m tall, branchlets thick and terete; leaves thickly coriaceous, oblong-obovate, 13–19 × 5–8 cm, apex obtuse to abruptly acuminate, base broadly cuneate, punctate on lower surface, margin subentire, veins in 18–20 pairs; petioles 14–25 mm long; flowers solitary, pedicels 1–2 cm long; ovary subconical, ca. 2 mm long, 2-locular, each locule with 2 ovules; style 2-parted, ca. 7 mm long; stigmas 2, punctiform. Possibly in eastern Bolívar (known from upper Mazaruni District in Guyana, along trail from Kuruoung River to Kamarang). Ternstroemia longipes Klotsch ex Wawra in Mart., Fl. Bras. 12(1): 277. 1886. Shrub; leaves submembranous, elliptic, 4–6 × 1–2 cm, acute at both ends, margin un-

Fig. 253. Ternstroemia dura

dulate, glandular-crenulate, veins obscure, petiole ca. 1 cm long; flowers few, solitary, internodal; pedicels slender, ca. 4 cm long; ovary 2-locular, each locule with 2 ovules; style terete, thick, shorter than the ovary; stigma deeply bisulcate, or divided into 2 substipatate, orbicular stigmas. Possibly in eastern Bolívar (the type specimen is a Schomburgk collection from adjacent Guyana). Ternstroemia schomburgkiana Benth., London J. Bot. 2: 362. 1842. Shrub with terete, grayish branchlets; leaves coriaceous, obovate, to 7 × 3 cm, apex obtuse to rounded, base obtusely cuneate; petiole 6–7 mm long; flowers 4- or 5-aggregate at the branchlet tips; pedicels 7–8 mm long; ovary 3-locular, each locule with 3 or 4 ovules; style longer than the ovary. Probably in Bolívar near base of Roraima-tepui (the type specimen is from close to Roraima on the Guyana side, also found in Suriname).

Fig. 254. Ternstroemia sp. A

308

T ERNSTROEMIACEAE

Fig. 255. Ternstroemia crassifolia

Fig. 256. Ternstroemia discoidea

Fig. 258. Ternstroemia pungens

Fig. 257. Ternstroemia duidae

Pentamerista 309

TETRAMERISTACEAE by James S. Miller and Paul E. Berry Trees or shrubs. Leaves alternate, simple, entire to toothed, coriaceous, (sub) sessile, estipulate. Inflorescences axillary, umbellate to corymbose racemes with a whorl of subfoliaceous bracts, or flowers solitary, each flower subtended by a pair of persistent or caducous, sometimes petaloid bracteoles. Flowers bisexual, actinomorphic, hypogynous, 4-merous (Tetramerista) or 5-merous (Pentamerista and Pelliciera). Sepals 4 and imbricate or 5 and quincuncial, glandular-pitted on the inner surface. Petals 4 and imbricate or 5 and quincuncial, free. Stamens 4 or 5, alternate with the petals; filaments basally connate, flattened; anthers bithecal, dehiscing longitudinally; pollen tricolporate. Ovary superior, 2-, 4-, or 5-carpellate, 1-, 4-, or 5-locular, the placentation axile-basal or subapical and pendulous; style terminal, simple; stigma minutely 4- or 5-lobed or bifid and punctate; ovules one per locule, anatropous to campylotropous. Fruits baccate (or dry and indehiscent), 1-, 4-, or 5-seeded; seeds large, sharply pointed in Pellicieria, with abundant endosperm. Coastal Central America, Pacific coastal Colombia, Venezuela, southern Malaya, Sumatra, Borneo; 3 genera and 5 species, 1 species in the flora area. Tetramerista occurs in southern Malaya, Sumatra, and Borneo, and Pentamerista is known only from Amazonas state in Venezuela and adjacent Colombia. The presence of closely related genera restricted to the central part of the Guayana Shield and to white-sand habitats in Malesia is paralleled by Ploiarium and Archytaea in the Bonnetiaceae, and is of great interest phytogeographically. Recently, the monotypic and New World mangrove genus Pelliceria has been placed in the Tetrameristaceae, based on molecular and floral evidence. 1. PENTAMERISTA Maguire, Mem. New York Bot. Gard. 23: 187. 1972. Dwarf shrub or tree to 15 m tall, the twigs angular. Leaves alternate, simple, entire, glabrous, lustrous, coriaceous, sessile or subsessile. Inflorescence axillary, a long pedunculate raceme, the bracts caducous, the flowers densely clustered at the end of the rachis, each subtended by 1 or 2 bracteoles. Flowers bisexual, actinomorphic, hypogynous. Sepals 5, quincuncial, glandular on the inner surface; petals 5, quincuncial, oblong. Stamens 5, alternate with the petals; filaments basally connate, flattened; anthers oblong-sagittate, dehiscing longitudinally. Ovary superior, 5-carpellate, 5-locular; glabrous; style terminal, terete, with a minutely 5-lobed stigma; ovules one per locule, anatropous. Fruits baccate, obloid, 5-seeded; seeds with abundant endosperm. Pentamerista is a monotypic genus known only from southwestern Venezuela and adjacent Colombia. Pentamerista neotropica Maguire in Maguire et al., Mem. New York Bot. Gard. 23: 187. 1972. Dwarf shrub or tree 1–15 m tall; flowers yellow-green; fruits purple-black. Savannas,

savanna margins, forest margins on white sand, 100–200 m; Amazonas (Caño Caname, base of Cerro Yapacana, Guarinuma, Río Pasimoni basin). Adjacent Colombia. ◆Fig. 259.

310

T ETRAMERISTACEAE

Fig. 259. Pentamerista neotropica

THEACEAE [with segregates now in BONNETIACEAE and TERNSTROEMIACEAE] Cronquist (1981) treated the Theaceae as a broad alliance containing four subfamilies, namely Asteropeiodieae, Bonnetioideae, Ternstroemioideae, and Theoideae. Subsequent studies have shown these groups to be quite distinct and not part of a monophyletic clade (Stevens 2001 onward). Some genera included by Cronquist in the Bonnetioideae, such as Caraipa and Neotatea, are now placed in Clusiaceae (Weitzman and Stevens 1997). Here we follow the circumscription outlined in Weitzman and Stevens (1997) and Weitzman et al. (2004), which recognizes Bonnetiaceae with 3 genera, Ternstroemiaceae with 12 genera, and Theaceae with 7 genera. Since previous volumes of the flora already treated families beginning

T HEACEAE 311

with “B,” the family treatment for Bonnetiaceae immediately follows the Theaceae, where it was formerly included, but outside of the usual alphabetical family order followed in this series. The Ternstroemiaceae treatment, on the other hand, is in its correct alphabetical order in this volume.

References Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. New York: Columbia University Press. Stevens, P. F. (2001 onward). Angiosperm Phylogeny Website. Version 5, May 2004. http://www.mobot.org/MOBOT/research/APweb/ Weitzman, A. L., and P. F. Stevens. 1997. Notes on the circumscription of Bonnetiaceae and Clusiaceae, with taxa and new combinations. BioLlania, Ed. Espec. 6: 551–564. Weitzman, A. L., S. Dressler, and P. F. Stevens. 2004. Ternstroemiaceae. Pp. 450– 460 in Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. VI. Flowering Plants. Dicotyledons. Celastrales, Oxalidales, Rosales, Cornales, Ericales. Berlin: Springer. Key to the Families Bonnetiaceae, Ternstroemiaceae, and Theaceae 1. 1. 2(1).

2.

Anthers basifixed; fruits indehiscent or irregularly dehiscent ........................................................................ Ternstroemiaceae, p. 300 Anthers versatile; fruits regularly dehiscent capsules ........................... 2 Fruits loculicidally dehiscent; seeds with large wing on upper end of seed; branches rarely thick; leaves usually spread along branches ........................................................................................ Theaceae, p. 311 Fruits septicidally dehiscent; seeds wingless or with very small wing; stems and/or branches often thick (pachycaulous); leaves often clustered near branch apices ..................................Bonnetiaceae, p. 313

THEACEAE by Anna L. Weitzman Trees or shrubs, glabrous or pubescent, the indumentum of unicellular hairs. Leaves spiral or distichous, simple, usually evergreen, usually coriaceous, margins toothed or more rarely entire, with a small, deciduous gland associated with each tooth; stipules absent. Flowers hermaphroditic, solitary, axillary, large and showy, prophylls 2, or several bracteoles intergrading with calyx and corolla. Calyx of 5 or more sepals, imbricate, connate or distinct basally, often persistent in fruit, usually thick, concave, equal to unequal; corolla of 5 (to numerous) petals, distinct or connate basally, imbricate. Stamens 20+, free (connate), often adnate to base of corolla, anthers versatile, opening by longitudinal slits; pollen tricolporate, pseudopollen present. Gynoecium syncarpous, (3–)5(–10)-carpellate; ovary

312

T HEACEAE

superior, placentation axile (apically axile, or axile-apical); ovules/carpel 2–few, ovules bitegmic, tenuinucellate; styles distinct or connate; stigmas usually lobed. Fruit a loculicidal capsule (irregularly dehiscent or a drupe), columella persistent or sometimes lacking; seeds few, sometimes winged; testa vascularized, ± lignified; endosperm nuclear, usually slight; embryo straight. Southeastern U.S.A., Mexico, Central America, the Caribbean, tropical South America, Southeast Asia (most speciose), Indo-Malesia; ca. 7 genera and 200–460 species, 1 species in the flora area. Generic limits in Theaceae are difficult, and relationships are still poorly understood. Gordonia is treated here in a broad sense based on morphology, although some recent molecular studies suggest that Laplacea should be separated from it. 1. GORDONIA Ellis, Philos. Trans. 60: 518, t. 11. 1770, nom. cons. Haemocharis Salisb., Parad. Lond. sub n. 56, 1806; Salisb. ex Mart., Nov. Gen. Sp. Pl. 1: 106. 1824 [1826]. Laplacea Kunth in H.B.K., Nov. Gen. Sp. 5: 160. 1821 [1822]. Lindleya Nees, Flora, 4: 299. 1821. Wikstroemia Schrad., Götting. Gel. Anzeig 1: 710. 1821. Trees or shrubs, often sericeous, dioecious or hermaphroditic. Leaves coriaceous (to membranaceous), symmetrical or asymmetrical, usually rounded or obtuse apically, tapering to the base to a short petiole or sessile, margins entire to sinuous. Flowers solitary, in axils of the upper leaves; bracteoles 2 or more, thick, unequal, somewhat sepaloid, quickly caducous. Sepals thick, usually sericeous, unequal, graduating in size and shape from bracteoles to petals, caducous (or rarely persistent). Corolla white or whitish, campanulate, petals unequal, usually membranaceous, usually emarginate apically. Stamens many; filaments adnate to the base of the petals (rarely connate their entire length forming a staminal tube and free from the petals); anthers versatile. Gynoecium usually (4)5(6–10)-carpellate, usually sericeous; ovules 4 or more per locule. Fruits ellipsoid to oblong, ± woody, loculicidal capsule with persistent columella, usually with 5 grooves, splitting from the apex downward. Seeds ellipsoid, flat or compressed, with a large, apical, oblong, membranaceous wing; endosperm in a thin layer; embryo fairly large, slightly curved. Eastern North America, Mexico, Central America, West Indies, tropical South America, Australasia; ca. 7 species, 1 in Venezuela and the flora area. Gordonia fruticosa (Schrad.) H. Keng, Gard. Bull. Straits Settlem. 33: 310. 1980. —Wikstroemia fruticosa Schrad., Götting. Gel. Anzeig 1: 711. 1821. —?Lindleya fruticosa Nees, Flora 4: 299. 1821. —Laplacea fruticosa (Schrad.) Kobuski, J. Arnold Arbor. 28: 437. 1947. Lindleya semiserrata Nees, Flora 4: 328. 1821, nom. nud., as syn. —Gordonia semiserrata (Nees) Spreng., Syst. Veg. 4(cur. post.): 260, 408. 1827. —Laplacea semiserrata (Nees) Cambess. in A. St.Hil., Fl. Bras. Merid. 1: 300. 1827.

Laplacea pubescens Planch. & Linden ex Triana & Planch., Ann. Sci. Nat. sér. 4, 18: 269. 1862. —Haemocharis pubescens (Planch. & Linden) Linden & Planch., Trois. Voy. Linden. [Bot. Pl. Columb.] 1: 59. 1863. Laplacea camelliaefolia Triana & Planch., Ann. Sci. Nat. sér. 4, 18: 270. 1862. —Wikstroemia camelliaefolia (Triana & Planch.) S.F. Blake, Contr. Gray Herb. n.s. 53: 39. 1918. —Laplacea pubescens var. camelliaefolia (Triana & Planch.) Kobuski, J. Arnold Arbor. 31: 427. 1950.

B ONNETIACEAE 313

Fig. 260. Gordonia fruticosa

Haemocharis caracasana Linden & Planch., Trois. Voy. Linden [Bot., Pl. Columb.] 1: 59. 1863. —Laplacea caracasana Klotsch & H. Karst. ex Wawra in Mart., Fl. Bras. 12(1): 289. 1866, nom. illeg., as syn. Laplacea fruticosa var. chimantae Steyerm., Ann. Missouri Bot. Gard. 74: 650. 1987. Laplacea pubescens var. minor Steyerm., Ann. Missouri Bot. Gard. 74: 652. 1987. Shrub to tree 3–15 m tall; flowers white. Tepui slope forests, 1000–2000 m; Bolívar (Cerro Sarisariñama, Serranía de los Pijiguaos, Macizo del Chimantá [Agparamántepui, Chimantá-tepui]), Amazonas (Cerro Duida, Sierra de la Neblina). Widely distributed in the Venezuelan Andes and Coastal

Cordilleras; Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 260. Gordonia fruticosa is a very variable species, which is treated here in a broad sense. The characters that have been used to define species and subspecies in the past have been shown (Weitzman, unpublished) to vary both without any correlation between characters and, in most cases, without gaps that can be used to define character states.

BONNETIACEAE by Anna L. Weitzman Trees or shrubs, small to medium-sized, ± subpachycaulous; glabrous. Leaves alternate, spiral, simple, estipulate, convolute, usually crowded toward apex of branches; petiole short or absent; blades with close, ascending lateral veins, margins initially setulose. Flowers showy, solitary or in ± cymose inflorescences; pedicels with 2 prophylls or several bracts; flowers bisexual, cyclic; sepals 5, unequal, free, quincuncial; petals 5, contorted, free. Stamens numerous; filaments slender, free or basally connate into 5 antepetalous bundles; anthers short, versatile; fasciclode present or absent. Ovary 3(–5)-locular, with numerous orderly arranged ovules on biseriate axile placentae; styles free or united into a branched or simple style; stigmas papillate. Fruits septicidal capsules with numerous seeds and a persistent central column; seeds with scanty endosperm; embryo straight.

314

B ONNETIACEAE

Cuba, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, southeast Asia, western Malesia, Moluccas, New Guinea; 3 genera and 34 species, 2 genera and 28 species in the flora area. The Old World genus Ploiarium contains 3 species. Cronquist (1981) treated Archytaea, Bonnetia, and Ploiarium as members of the Theaceae (subfamily Bonnetioideae), together with the latex-producing Neotatea and other genera such as Caraipa and Kielmeyera. The possession of xanthones, floral morphology, seed testa anatomy, and wood anatomy sets the first three genera apart from the Theaceae and aligns them with Clusiaceae and Hypericaceae in the Malpighiales. Molecular DNA sequence studies also place Bonnetiaceae far from the Theaceae and close to Clusiaceae, while Neotatea, Caraipa, and Kielmeyera are all placed in the Clusiaceae. Key to the Genera of Bonnetiaceae 1.

1.

Ovary 5-locular; stamens pentadelphous, often persistent in fruit; capsule dehiscent from base; marginal disciform glands usually visible on leaves and/or bracts ................................................................. 1. Archytaea Ovary usually 3-locular; stamens free, deciduous; capsule dehiscent from top; disciform bracts absent ..................................................... 2. Bonnetia

1. ARCHYTAEA Mart., Nov. Gen. Sp. Pl. 1: 116. 1826. Evergreen trees or shrubs. Leaves alternate, clustered near branch apices, coriaceous, sessile, often asymmetrical, often with marginal, disciform glands. Inflorescence axillary, near apex, fasciculate or racemose with 3–10 flowers; peduncles flattened, becoming wider apically; pedicels subtended by a single bract often with at least one pair of marginal disciform glands. Calyx persistent; corolla free, white or pink. Stamens numerous, persistent, in 5 antesepalous fascicles; anthers minute, versatile; glands 5, alternating with the stamen fascicles. Ovary glabrous, 5-locular, with numerous ovules/locule; style simple, persistent; stigma punctate. Capsule septicidal, splitting from the base, with persistent columella. Seeds numerous, linear, imbricate, exalbuminous. Southeastern Colombia, southern Venezuela, western Guyana, northern Brazil (Amazonas, Roraima); 2 species, both in the flora area. On morphological grounds, this genus is more closely related to the Malaysian genus Ploiarium than to the sympatric Bonnetia. Key to the Species of Archytaea 1.

1.

Leaves linear-lanceolate, acute, 6–10 × 0.5–1.9 cm; lateral veins often inconspicuous; peduncles 2–6 cm long, commonly < 1/2 the length of the subtending leaves; bracts inconspicuous; petals white or slightly pinkish .................................................................................. A. angustifolia Leaves broadly oblanceolate, usually acuminate (or obtuse), 8–16 × 2.5– 5.5 cm; lateral veins conspicuous; peduncles 4–12 cm long, often equaling or exceeding the leaves; bracts prominent; petals pink ....... A. triflora

Archytaea 315

Fig. 261. Archytaea angustifolia

Archytaea angustifolia Maguire, Mem. New York Bot. Gard. 23: 137. 1972. Prostrate to erect shrub or tree 0.2–10 m tall. Seasonally flooded black-water river banks, white-sand savannas, 50–200 m; Amazonas (basins of Río Atabapo, Río Atacavi, Río Caname, Río Casiquiare, Río Guainía, Río Guasacavi, Río Negro, Río Pacimoni, Río San Miguel, and Río Temi). Colombia (Meta and Guainía). ◆Fig. 261. Archytaea triflora Mart., Nov. Gen. Sp. Pl. 1: 117. 1826. —Conók-pichoray, Yaradek (Pemón). Archytaea multiflora Benth., London J. Bot. 2: 363. 1843. Shrub or tree 2–10 m tall. Lower montane to montane moist forests, gallery forests, upland savannas, rarely in seasonally flooded savannas, 300–2000 m, usually above 800 m; Bolívar (Cerro Guaiquinima, Cerro Jaua, Gran Sabana, Kavanayén, Macizo del Chimantá [Acopán-tepui], Salto Eutobarima on Río Caroní, Sierra de Lema), Amazonas (Caño Culebra on slopes of Cerro Duida, slopes of Cerro Aracamuni, lower slopes of Cerro Marahuaca, Cerro Vinilla). Colombia (Caquetá, Vaupés), Guyana (Ayanganna, Kaieteur Plateau, Mount Roraima, Pakaraima Mountains), Brazil (Serra Aracá, Serra Tepequem, Serra Tunui). ◆Fig. 262.

Fig. 262. Archytaea triflora

Both Kobuski (C. Kobuski. J. Arnold Arbor. 31: 196–207. 1950) and Maguire (B. Maguire. Mem. New York Bot. Gard. 23: 1– 832. 1972) thought that Archytaea multiflora and A. triflora are very closely related, but chose not to combine them without more study. Kobuski did not see any of the Martius material on which the name A. triflora is based, but he maintained both species because he thought that the bracts of A. triflora were not foliaceous and lacked “glands” (= domatia, see above), since they were not described nor illustrated by Martius (Martius & Zuccarini, 1826). Although the bracts of all three specimens in the Munich herbarium are at the smaller end (0.5–1 cm) of the range (to 2 cm) of bract length in A. multiflora, they are similar in shape to the leaves and have 2 marginal domatia. Maguire (1972) saw the Martius material now held at Munich and New York. In his key to species of Archytaea he used only “peduncles 3-flowered” versus “peduncles multiflowered” as opposing characters for A. triflora and A. multiflora respectively. Other characters in the key were listed only for the

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latter species. However, in the discussion he noted the “ample bracteoles” of Archytaea triflora and, in the key he used “strongly bracted” for A. multiflora. Although both Kobuski and Maguire thought that A. triflora had only 3 flowers per inflorescence, while A. multiflora had 5 or more, all three type specimens of A. triflora at Munich have

some inflorescences with > 3 pedicel scars, and one inflorescence on the specimen which Maguire designated as the holotype retains 5 pedicels. While inflorescences of A. multiflora more commonly have > 3 flowers in an inflorescence, inflorescences with only 3 flowers are frequent. For these reasons only one species is recognized here.

2. BONNETIA Mart. ex Nees & Mart., Nov. Acta Phys.-Med. Acad. Leop.-Carol 12(1): 36. 1824, nom. cons. Kieseria Nees, Flora 4: 298. 1821. Neblinaria Maguire, Mem. New York Bot. Gard. 23: 155. 1972. Neogleasonia Maguire, Mem. New York Bot. Gard. 23: 158. 1972, pro parte. Acopanea Steyerm., Ann. Missouri Bot. Gard. 71: 323. 1984. Trees or shrubs, or underground-stemmed rosette plants, without latex. Stems and/or branches often subpachycaulous. Leaves evergreen, coriaceous, often asymmetrical, entire, often clustered near branch apices, sessile or petiolate; margins often with minute, deciduous setae; veins pinnate or parallel. Flowers solitary or inflorescence axillary, cymose, racemose, or paniculate; peduncles ancipitous or terete; pedicels with persistent or deciduous, single or whorled, foliar or nonfoliar bracteoles. Sepals persistent; petals convolute, flabelliform or obovate. Stamens numerous; filaments adnate to the base of the ovary and otherwise free; anthers dehiscing longitudinally or by 2 pores at the base. Ovary 3(4)-locular; stylodia 3, or style simple and then sometimes apically branched; stigmas papillate. Capsule septicidal, columnella persistent. Seeds linear, elongated above and below into a small membranous wing. Neotropics (mainly in the Guayana Shield area, also in Cuba, Amazonian and eastern-coastal Brazil, Andean Venezuela, Ecuador, and Peru); 29 species, 26 in Venezuela, all of these in the flora area. According to Weitzman and Stevens (A. L. Weitzman and P. F. Stevens. BioLlania Edición Especiál 6: 551–564. 1997), Bonnetia and its former segregate genera listed above in synonymy all share trilacunar leaf nodes with endodermis around the leaf traces, which is lacking in Archytaea and Ploiarium. They also have leaf margins that are minutely serrulate, though often appearing entire, and all Bonnetia species lack the domatia on the leaf margins that characterize the other two genera. Bonnetia has a multiseriate androecium that is adnate to the base of the ovary, while stamens of Archytaea and Ploiarium are pentadelphous, with the 5 bundles opposite the petals. Finally, Bonnetia has 3-locular ovaries, while the ovaries of the other two genera are 5-locular. There are only 3 species in the genus that do not occur in Venezuela: Bonnetia cubensis (Britton) R.A. Howard from eastern Cuba, B. stricta Mart. from Bahia to Rio de Janeiro states, Brazil, and B. holostyla Huber from Vaupés, Colombia. Key to the Species of Bonnetia 1. 1. 2(1). 2.

Flowers in compound inflorescences ......................................................... 2 Flowers solitary or few-flowered inflorescences ....................................... 7 Styles free to the base ................................................................ B. kathleenae Styles branched in the upper 1/3–1/2 ........................................................... 3

Bonnetia 317

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

7(1). 7. 8(7). 8. 9(8). 9. 10(9). 10. 11(10). 11. 12(8). 12. 13(12). 13. 14(7). 14. 15(14). 15. 16(14). 16. 17(16). 17. 18(17). 18. 19(17). 19. 20(19). 20. 21(19).

Inflorescence congested, pedicels < 10 mm long ....................................... 4 Inflorescences open, pedicels > 10 mm long ............................................. 5 Anthers broader than or as long as broad, subreniform; ovary incompletely trilocular .................................................................. B. euryanthera Anthers longer than broad, ovoid, ellipsoid, or suborbicular; ovary completely trilocular ...................................................................... B. jauaensis Leaf base rounded to slightly cordate, somewhat clasping ........ B. cordifolia Leaf base cuneate to acute, not clasping .................................................. 6 Leaves asymmetrical, > 1.5 times longer than broad; sepals 6–8(–10) mm long; petals oblong-obovate, 12–20 × 12–15 mm .................. B. paniculata Leaves usually symmetrical, sometimes slightly asymmetrical, < 1.5 times longer than broad; sepals 12–16 mm long; petals broadly obovate, 22– 25 × 28–34 mm ................................................................................ B. celiae Style divided to the base, or at least upper 2/3 .......................................... 8 Style undivided or divided only apically ................................................. 14 Corolla white or pink ................................................................................. 9 Corolla yellow ........................................................................................... 12 Leaves < 2 cm long ........................................................................ B. roraimae Leaves > 2 cm long ................................................................................... 10 Leaves with veins pinnate, recticulate ................................... B. bolivarensis Leaves with veins closely parallel ........................................................... 11 Pedicels 25–40 × 1–3 mm, terete or slightly flattened; petals 1–1.8 cm long ......................................................................................... B. fasciculata Pedicels 4–5 × 4–5 mm, flattened; petals 4–5 cm long ........... B. multinervia Leaves < 3 cm long ...................................................................... B. huberiana Leaves > 3 cm long ................................................................................... 13 Leaves obovate; pedicels 30–60 mm long ....................................... B. tristyla Leaves lanceolate; pedicels 4–6 mm long .................................. B. ptariensis Style entire ............................................................................................... 15 Style divided above .................................................................................. 16 Flowers sessile; leaves 1.3–3.5 × 1.5–2 cm ................................ B. tepuiensis Flowers on pedicels ca. 1 cm long; leaves 2–2.5 × ca. 1 cm ............ B. liesneri Mainly acaulescent subshrubs, with long stolons and peduncles ................................................................................................... B. ahogadoi Trees or shrubs, without long stolons ..................................................... 17 Leaves closely parallel veined; few-branched, pachycaulous trees or shrubs ................................................................................................... 18 Leaves pinnately veined; well-branched, nonpachycaulous trees or shrubs ................................................................................................... 19 Leaves navicular, venation not emergent; peduncles 10–15 cm long; pedicels 0.5–1 cm long ...................................................... B. maguireorum Leaves plane, venation emergent; peduncles 6–8 cm long; pedicels ca. 2 cm long ................................................................................. B. rubicunda Flowers yellow .......................................................................................... 20 Flowers pink or white .............................................................................. 21 Leaves 1–1.5 × 0.5–1 cm, base attenuate; pedicels subterete ...... B. wurdackii Leaves larger, ca. 8 × 4 cm, base rounded; pedicel flattened-winged ............................................................................................. B. steyermarkii Leaves < 2 cm wide .................................................................................. 22

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21. 22(21). 22. 23(22). 23. 24(21).

Leaves > 3 cm wide .................................................................................. 24 Leaves < twice as long as wide ...................................................... B. neblinae Leaves > twice as long as wide ................................................................ 23 Leaves ca. 12–27 × 5–8 mm ................................................... B. chimantensis Leaves ca. 4.5–6.5 × 0.7–1.6 cm ................................................ B. lanceifolia Pedicels strongly ancipitous; leaves punctate; 4 sepaloid bracteoles closely subtending calyx .......................................................... B. roseiflora 24. Pedicels terete or only slightly flattened; leaves not punctate; bracteoles absent, caducous, or lower on pedicel ................................................. 25 25(24). Leaves generally spread along the branches, not usually tightly clustered at the branch tips; sepals rounded (to obtuse) at apex .............. B. sessilis 25. Leaves clustered at the branch tips; sepals acute at apex ..................... 26 26(25). Pedicels 2–3.5 mm thick, < 3 cm long; sepals ca. 26 × 11 mm ........ B. crassa 26. Pedicels 1–1.5 mm thick, 3–4 cm long; sepals ca. 15–20 × 3–5 mm .................................................................................................... B. neblinae Bonnetia ahogadoi (Steyerm.) A.L. Weitzman & P.F. Stevens, BioLlania Edición Especiál 6: 559. 1997. —Acopanea ahogadoi Steyerm., Ann. Missouri Bot. Gard. 71: 323, fig. 10. 1984. Stoloniferous subshrub with basal rosettes; peduncle reddish, emerging below the rosette or between the lower leaves; calyx red-purple; corolla pink; stamens orange. Forming dense colonies in wet, rocky tepui bogs and meadows, 1900–2000 m; Bolívar (southeastern summit of Acopán-tepui in Macizo del Chimantá). Endemic. ◆Fig. 272. Bonnetia bolivarensis Steyerm., Ann. Missouri Bot. Gard. 74: 647. 1987. Shrub 1–2 m tall. Rocky tepui summit scrub, 2300–2400 m; Bolívar (Ptari-tepui). Endemic. Bonnetia bolivarensis has white or pink flowers and seemingly only differs from B. ptariensis in that the latter has yellow flowers. They are both known only from Ptaritepui. Two specimens from nearby Aparamán-tepui and Murisipán-tepui are sterile and may belong to one of these two species.

Bonnetia toronoensis Steyerm., Ann. Missouri Bot. Gard. 71: 327. 1984. Shrub 1–2.5 m tall; petals white-pink. Tepui meadows and rocky areas, 2000–2500 m; Bolívar (Aprada-tepui, Macizo del Chimantá, Uaipán-tepui). Endemic. Bonnetia cordifolia Maguire, Bol. Soc. Venez. Ci. Nat. 22: 357. 1976. Single-stemmed, woody shrub ca. 2 m tall; petals white with broad, dull yellow stripe down center on outside. Slightly raised hummocks in tepui meadows, 1400–1500 m; Bolívar (Cerro Guanacoco). Endemic.

Bonnetia celiae Maguire, Mem. New York Bot. Gard. 23: 143, fig. 22. 1972. Shrub or tree 1–5 m tall; petals white to deep pink. Rocky tepui slopes and summit, 1500–2200 m; Amazonas (Cerro Yutajé). Endemic.

Bonnetia crassa Gleason, Bull. Torrey Bot. Club 58: 395. 1931. Shrubby (rarely spreading or sprawling), 0.5–5 (rarely to 15) m tall; petals pink. Tepui slopes and summit on rock or in meadows and forests, 1300–2100 m; Amazonas (Cerro Coro-Coro, Cerro Duida, Cerro Guanay, Cerro Parú, Cerro Sipapo, Cerro Yutajé). Endemic. Some specimens attributable to Bonnetia crassa appear to intergrade with B. sessilis. The sepals in B. crassa are longer (> 2 cm) and acute at the apex while in B. sessilis they are < 2 cm and rounded to obtuse. Nearly all specimens of B. crassa have been collected in Amazonas, however, while B. sessilis is more common in Bolívar and Guyana; it is widespread in Amazonas, along with adjacent areas of Colombia and Brazil.

Bonnetia chimantensis Steyerm., Ann. Missouri Bot. Gard. 71: 326. 1984.

Bonnetia euryanthera Steyerm., Ann. Missouri Bot. Gard. 74: 647, fig. 7. 1987.

Bonnetia 319

Shrub ca. 1 m tall; petals white-pink. On exposed tepui summits in shrubland and meadows, 2000–2100 m; Bolívar (Cerro Jaua). Endemic. Bonnetia fasciculata A.L. Weitzman & P.F. Stevens, BioLlania Edición Especiál 6: 559. 1997. —Neogleasonia wurdackii Maguire, Mem. New York Bot. Gard. 23: 160. 1972, non Bonnetia wurdackii Maguire, Mem. New York Bot. Gard. 23: 147. 1972. Shrub 1–2 m tall, somewhat pachycaulous, leaves in tight rosettes at branch tips; peduncles purple-red, petals white to pink. Dense tepui scrub on rocky tepui summit, 2000–2500 m; Bolívar (Macizo del Chimantá [Apacará-tepui]). Endemic. Bonnetia huberiana Steyerm., Ann. Missouri Bot. Gard. 71: 326. 1984. Shrub 0.5–3 m tall, densely branched, globose, forming dense cushions; leaves ca. 2 × 0.5 cm, clustered at branch tips; calyx green, petals yellow. Rocky outcrops near tepui meadows, along river edges, 1800–2500 m; Bolívar (Macizo del Chimantá [Acopán-tepui, Angasima-tepui, Chimantá-tepui]). Endemic. Bonnetia jauaensis Maguire, Bol. Soc. Venez. Ci. Nat. 22: 358. 1976. Shrub or tree 1–10 m tall; petals whitepink, 2–2.5 cm long. Upper slopes and summit of tepui in montane scrub forests, 1300– 1800 m; Bolívar (Cerro Sarisariñama). Endemic. Bonnetia kathleenae Lasser, Bol. Soc. Venez. Ci. Nat. 10: 327. 1946. Small weak tree or scandent shrub to 5 m tall; petals white-pink. Montane forests and savannas on tepui slopes, (400–)1400–1500 m; Amazonas (Cerro Sipapo, hills at foot of Cerro Sipapo 10 km north of Cerro Autana and 1 km east of Lago Leopoldo). Endemic. ◆Fig. 267. Bonnetia lanceifolia Kobuski, J. Arnold Arbor. 29: 412. 1948. Bonnetia guaiquinimae Steyerm., Ann. Missouri Bot. Gard. 74: 649. 1987. Branched shrub 0.2–1.5 m tall; petals white. Open tepui meadows and shrublands, 700–1800 m; Bolívar (Cerro Guaiquinima). Endemic. ◆Fig. 266.

Bonnetia liesneri Steyerm., Ann. Missouri Bot. Gard. 76: 969. 1989. Shrub ca. 0.5 m tall with spreading branches. Tepui meadows, 1500–1600 m; Amazonas (Cerro Aracamuni). Endemic. Bonnetia liesneri is known only from a collection with old fruit. Bonnetia maguireorum Steyerm., Ann. Missouri Bot. Gard. 71: 326. 1984. Neblinaria celiae Maguire, Mem. New York Bot. Gard. 23: 157. 1972, non Bonnetia celiae Maguire 1972. Shrub or tree 0.3–3 m tall, sparselybranched, thick, corky-stemmed, with large, prominent leaf scars; petals pink. Montane scrub and boggy open meadows on tepui slopes, 1700–2000 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 275. Bonnetia multinervia (Maguire) Steyerm., Ann. Missouri Bot. Gard. 71: 326. 1984. —Neogleasonia multinervia Maguire, Mem. New York Bot. Gard. 23: 158. 1972. Shrub 0.3–2 m tall, monopodial or slightly branched, often with pachycaulous stems; petals white-pink. Tepui meadows, rock outcrops, 1600–2300 m; Bolívar (Ilú-tepui, Sierra de Lema, Yuruaní-tepui). Endemic. ◆Fig. 274. Bonnetia neblinae Maguire, Mem. New York Bot. Gard. 23: 148, fig. 23. 1972. Sprawling shrub to tree 0.2–8(–15) m tall; petals pink. Edges of boggy tepui meadows and dwarf forests, (1100–)1700–2300 m; Amazonas (Cerro Parú, Sierra de la Neblina). Brazil (Amazonas: adjacent side of Serra da Neblina). ◆Fig. 265. Bonnetia paniculata Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5: 63. 1861. Bonnetia parviflora Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5: 63. 1861. Bonnetia dinizii Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 7: 302. 1913. Shrub or small tree 2–6(–15) m tall, sparingly branched; petals white-pink. Lowland white-sand savannas, montane savannas, lowland to montane forest margins, 100– 2200 m; Bolívar (Amaruay-tepui, Auyántepui, Cerro Guaiquinima, Gran Sabana, Kavanayén near Oparumá, Macizo del

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Chimantá, Ptari-tepui, Roraima-tepui, Upuigma-tepui), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Cerro Yapacana, Cerro Yutajé, Raudal Gallineta on Río Siapa, Río Cuao, Sierra de la Neblina, Sierra Maigualida, Sierra Parima). Táchira; Colombia (Caquetá), Guyana, Ecuador (Zamora-Chinchipe), Peru (Amazonas, Huánuco, San Martín), Brazil (Amazonas, Pará, Roraima). ◆Fig. 263. Bonnetia paniculata is the most widespread species in the genus. It occurs over a wide altitudinal range, and is in the Andes as well as in the Guayana Shield.

Bonnetia rubicunda (Sastre) A.L. Weitzman & P.F. Stevens, BioLlania Edición Especiál 6: 558. 1997. —Neblinaria rubicunda Sastre, Bull. Mus. Natl. Hist. Nat., Misc. ser. 4, 9(2): 129, fig. 1. 1987. Pachycaulous shrub or small tree, leaf scars prominent on corky bark; peduncles 6– 8 cm long, pedicels 2 cm long, bracts 3–5, foliaceous; petals pink, 2.5 × 1.5 cm. Moist scrub on exposed ridges of tepui slope, 1500– 2300 m; Bolívar (expected). Guyana (slopes of Mount Ayanganna, Mount Roraima, and Mount Wokomung).

Bonnetia ptariensis Steyerm., Ann. Missouri Bot. Gard. 74: 649. 1987. Shrub 1–2.5 m tall; petals yellow. Rocky tepui summit scrub, 2300–2400 m; Bolívar (Ptari-tepui). Endemic. See note under Bonnetia bolivarensis.

Bonnetia sessilis Benth., London J. Bot. 2: 363. 1843. —Kieseria sessilis (Benth.) Kuntze, Revis. Gen. Pl. 1: 62. 1891. Bonnetia phelpsii Gleason, Brittonia, 3: 170. 1939. Bonnetia martiana Maguire, Mem. New York Bot. Gard. 23: 153. 1972. Bonnetia martiana var. venadoensis Maguire, Mem. New York Bot. Gard. 23: 153. 1972. Shrub to small tree 0.1–8 m tall; petals white-pink. Lowland white-sand savannas to tepui meadows and tepui scrub, 100–2200 m; Bolívar (Amaruay-tepui, Aprada-tepui, Auyán-tepui, Cerro Arepuchi, Cerro Akurimá, Cerro Guanococo, Cerro Guaiquinima, Cerro Ichún, Cerro Venado, Chirikayen, Gran Sabana, Icabarú, Ilú-tepui, Macizo del Chimantá, Murisipán-tepui, Río Canaracuni, Río Caruay, Río Uaiparú, Roraima-tepui, Salto Eutobarima, Serranía Marutaní, Sierra de Lema, Sororopán-tepui, Uaipántepui, Upuigma-tepui), Amazonas (Caño Pimichín, Cerro Parú, Cerro Yapacana, Cerro Autana, El Venado savanna, Río Atabapo, Río Siapa on slopes of Cerro Aracamuni, west of Río Temi, upper Río Yatúa, Sierra Parima). Colombia (Caquetá, Guainía, Vaupés), Guyana (widespread on tepuis and Mount Pakaraima), Brazil (Amazonas, Roraima). ◆Fig. 264. See note under Bonnetia crassa.

Bonnetia roraimae Oliv., Trans. Linn. Soc. London, Bot. 2: 272, pl. 37B, figs. 9–17. 1887. Densely branched shrub or tree 2–12 m tall, with a dense canopy; petals pink. Often the dominant woody species in tepuislope and summit forests, 1600–2900 m; Bolívar (Angasima-tepui, Apacará-tepui, Aparamán-tepui, Aprada-tepui, Auyántepui, Carrao-tepui, Ilú-tepui, Kamarkawarai-tepui, Karaurín-tepui, Kukenán-tepui, Macizo del Chimantá [Acopán-tepui, Chimantá-tepui, Churí-tepui], Murisipán-tepui, Ptari-tepui, Roraima-tepui, Sororopán-tepui, Tramen-tepui, Tereké-yurén-tepui, Yuruanítepui). Guyana (Mount Ayanganna, Mount Roraima), Brazil (Roraima). ◆Fig. 269. Bonnetia roraimae is one of the dominant woody species in forests of the eastern tepuis of Bolívar state. Bonnetia roseiflora Maguire, Mem. New York Bot. Gard. 23: 145. 1972. Low shrub 2–3 m tall, branched, with slender, sprawling stems; petals pink. Tepuisummit thickets and meadows, 1900–2100 m; Bolívar (Cerro Jaua). Endemic. The only apparent difference between Bonnetia roseiflora and B. steyermarkii is petal color. Sterile specimens cannot be reliably identified.

Bonnetia steyermarkii Kobuski, J. Arnold Arbor. 29: 402. 1948. Shrub 1–4 m tall; calyx green, petals yellow. Shrub islands on rocky tepui summits, 1600–2500 m; Bolívar (Aparamán-tepui,

Bonnetia 321

Auyán-tepui, Murisipán-tepui, Ptari-tepui, Tereké-yurén-tepui). Endemic. ◆Fig. 270. See note under Bonnetia roseiflora. Bonnetia tepuiensis Kobuski & Steyerm., J. Arnold Arbor. 29: 399. 1948. Bonnetia tepuiensis subsp. minor Steyerm., Ann. Missouri Bot. Gard. 71: 326. 1984. Shrub 1–3.5(–12) m tall with dense canopy; petals white-pink; leaves 1.5–3.5 × 1.5–2 cm, apex rounded, retuse. Rocky open tepui summits, tepui forests with Bonnetia roraimae, 1000–2500 m; Bolívar (Auyántepui, Caraurín-tepui, Carrao-tepui, Cerro Venamo, Ilú-tepui, Kamarkawarai-tepui, Macizo del Chimantá [Río Tírica], Murisipán-tepui, Roraima-tepui, Sierra de Lema, Tereké-yurén-tepui, Uei-tepui, Yu-

Fig. 263. Bonnetia paniculata

ruaní-tepui). Guyana (Mount Ayanganna, Mount Roraima). ◆Fig. 268. Bonnetia tristyla Gleason, Bull. Torrey Bot. Club 58: 396. 1931. Bonnetia tristyla subsp. nervosa Steyerm., Ann. Missouri Bot. Gard. 74: 649. 1987. Shrub 1–6 m tall; pedicels 3–6 cm long; petals yellow. Tepui bogs and scrub, meadows, 600–2100 m; Bolívar (Cerro Jaua, Cerro Sarisariñama), Amazonas (slopes of Cerro Aracamuni, Cerro Avispa, Cerro Duida, Cerro Marahuaca, Cerro Vinilla, Cerro Yapacana, Sierra de la Neblina, Sierra Unturán). Colombia (Vaupés), Brazil (Amazonas). ◆Fig. 271. Bonnetia wurdackii Maguire, Mem. New York Bot. Gard. 23: 147. 1972. Shrub to small tree 1–6 m tall, with a dense canopy; petals yellow. Tepui shrublands on rock outcrops, 1900–2600 m; Bolívar (Macizo del Chimantá [Abacapá-tepui, Amurí-tepui, Apacará-tepui, Chimantá-tepui, Churí-tepui, Murey-tepui, Toronó-tepui]). Endemic. ◆Fig. 273.

Fig. 264. Bonnetia sessilis

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Fig. 265. Bonnetia neblinae

Fig. 266. Bonnetia lanceifolia

Fig. 267. Bonnetia kathleenae

Bonnetia 323

Fig. 269. Bonnetia roraimae

Fig. 268. Bonnetia tepuiensis

Fig. 270. Bonnetia steyermarkii

Fig. 271. Bonnetia tristyla

Fig. 272. Bonnetia ahogadoi

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Fig. 273. Bonnetia wurdackii

Fig. 274. Bonnetia multinervia

Fig. 275. Bonnetia maguireorum

Clavija 325

THEOPHRASTACEAE by James S. Miller and Bertil Ståhl Trees or shrubs, often sparsely branched. Leaves alternate, often densely clustered at the ends of branches, simple, entire, serrate, or spiny-margined, the apex often culminating in a spine (Bonellia), often with subepidermal fibers that become evident in dried specimens, exstipulate. Inflorescence terminal or lateral, racemose, rarely corymbose, or with the flowers solitary. Flowers actinomorphic, hypogynous, bisexual or unisexual (Clavija, the plants then polygamodioecious), bracteate. Sepals (4)5, free, imbricate, persistent, gland-dotted; corolla sympetalous, fleshy, (4)5lobed, the lobes imbricate. Stamens 5, opposite the corolla lobes; filaments basally connate in a short tube or connate for their entire length (Clavija), adnate to the corolla tube at the base; anthers bithecal, extrorse, longitudinally dehiscent, sometimes with a prolonged, apiculate connective; staminodes 5, alternate with the stamens, fused with the corolla, petaloid or gibbous. Ovary superior, 1-locular, placentation free central or basal; style terminal; stigma usually discoid or capitate; ovules few to numerous, anatropous to hemitropous, immersed in mucilage. Fruit baccate, indehiscent. Seeds 1–several, with abundant, oily endosperm. U.S.A. (southern Florida), Mexico, Central America, West Indies, South America south to Paraguay; 7 genera and ca. 100 species, 2 genera and 3 species in the flora area. Key to the Genera of Theophrastaceae 1.

1.

Inflorescences lateral; leaves large and clustered at the stem apex; flowers crateriform; staminodes gibbous; plants sparsely branched or unbranched .................................................................................. 1. Clavija Inflorescences terminal; leaves small and more evenly spread out; flowers campanulate, staminodes flattened; plants much-branched .................................................................................................. 2. Jacquinia

1. CLAVIJA Ruiz & Pav., Fl. Peruv. Prodr. 142, t. 30. 1794. Understory shrubs or small trees, seldom branched. Leaves alternate, petiolate, usually clustered at the stem apex, simple, entire, serrulate, or serrate, often with linear sclereids just beneath the epidermis, usually elongate and coriaceous. Inflorescences racemose, mostly clustered among and beneath the leaves. Flowers 4- or 5-merous, often showy, unisexual or bisexual (plants polygamodioecious). Sepals connate basally, imbricate, persistent; corolla pale orange, orange-red, or sometimes red, glabrous, fleshy, crateriform, the tube short, with thickened staminodia borne in the mouth alternate with the corolla lobes. Stamens 4 or 5; filaments in pistillate flowers connate at base only, in staminate and bisexual flowers connate into a tube; anthers in staminate and bisexual flowers forming a peltate disk in the mouth of the corolla tube. Fruit baccate, coriaceous, globose, 1–several-seeded, pulp orange, sweet. Seeds large. Neotropics; ca. 50 species, 4 in Venezuela, 2 of these in the flora area.

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T HEOPHRASTACEAE

Fig. 276. Clavija imatacae

Clavija 327

Fig. 277. Clavija lancifolia subsp. chermontiana

328

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Key to the Species of Clavija 1.

1.

Young shoots glabrous or sparsely puberulous toward apices; petioles rather poorly demarcated, 4–8% of leaf length; leaf surfaces, except for veins and veinlets, smooth or very sparsely striate ................ C. imatacae Young shoots densely puberulous (appearing as a thin, light brown felt); petioles well demarcated, 10–13% of leaf length; leaf surfaces finely striate, at least on lower surface ............................................. C. lancifolia

Clavija imatacae Ståhl, Opera Bot. 107: 62. 1991. —Cola de pava. Clavija lancifolia auct. non Desf. 1832: sensu Mez in Engl., Pflanzenr. IV. 236a(15): 27. 1903, pro parte. Shrub or tree to 3 m tall, unbranched or rarely sparsely branched; leaves relatively large and thin; flowers fragrant. Deciduous or evergreen forests, 50–400 m; southern Delta Amacuro, northeastern Bolívar. Adjacent Guyana. ◆Fig. 276. Clavija imatacae has frequently been misidentified as C. longifolia (Jacq.) Mez. Clavija lancifolia Desf., Nouv. Ann. Mus. Hist. Nat. 1: 402, t. 14. 1832.

Guyana, Suriname, French Guiana, Amazonian Brazil and Bolivia; 2 subspecies, 1 in the flora area. C.

lancifolia subsp. chermontiana (Standl.) Ståhl, Opera Bot. 107: 56. 1991. —Clavija chermontiana Standl., Field Mus. Nat. Hist., Bot. Ser. 8: 148. 1930. —Cola de pava, Pavito, Yuu-kamohi. Unbranched shrub or tree 0.5–3 m tall; fruits edible. Nonflooded forests, 50–500 m; central and northwestern Bolívar, central to northern Amazonas. Guyana, Suriname, French Guiana, Amazonian Brazil and Bolivia. ◆Fig. 277.

2. JACQUINIA L., Fl. Jamaic. 27. 1759. Shrubs or small trees, much-branched. Leaves pseudoverticillate, usually rigid-coriaceous, entire, lacking or with a poorly developed mucro. Inflorescences terminal or pseudoaxillary racemes, sometimes bracteate. Flowers 5-merous, bisexual, whitish. Sepals 5, free or slightly connate at the base, imbricate, widely ovate to rounded, often ciliate or crenate; corolla white or whitish, campanulate, the 5 lobes imbricate, with 5 petaloid staminodia inserted in the mouth of the corolla tube, slightly smaller than and alternate with the corolla lobes. Stamens 5; filaments dilated basally, connate into a tube at the base and basally adnate to the corolla tube; anthers ovoid, dehiscing longitudinally, nearly basifixed. Ovary 1-locular with numerous ovules; style not clearly differentiated from the ovary; stigma capitate. Fruits baccate, few-seeded, coriaceous, globose to ovoid. Seeds immersed in mucilaginous pulp. U.S.A. (southern Florida), islands off the coast of Mexico and Honduras, West Indies, Venezuela, Colombia, Trinidad, Brazil; 13 species, 4 in Venezuela, 1 of these in the flora area. Jacquinia armillaris Jacq., Enum. Syst. Pl. 15. 1760. —Piacayure. Jacquinia revoluta Jacq., Fragm. Bot. 64. 1809. Jacquinia arborea auct. non Vahl: sensu Kunth in H.B.K., Nov. Gen. Sp. 3: 250. 1818 [1819].

Shrub to 3 m tall; leaves obovate to oblanceolate, apically rounded. Seasonally dry forests, 200–400 m; Bolívar (near El Palmar, near Upata). Aragua, Carabobo, Nueva Esparta, Sucre; Lesser Antilles, northern Colombia, Trinidad, eastern Brazil. ◆Fig. 278.

Jacquinia 329

Fig. 278. Jacquinia armillaris

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T HURNIACEAE

THURNIACEAE by Ricarda Riina and Paul E. Berry Perennial herbs with an upright rootstock, solid-pithed. Leaves tristichous, simple, parallel-veined, all basal, with sheathing base, the blades elongate, flat or V-shaped in cross section, the margin smooth or serrulate. Inflorescences of 1 or 2 large globose to ellipsoid heads, or smaller heads racemosely arranged, subtended by spreading to reflexed leafy bracts on triangular peduncles. Flowers bisexual, regular; pedicels short and swollen; tepals 6, persistent, subequal, whitish, spotted. Stamens distinct, in 2 whorls of 3; filaments adnate to the base of the tepals; anthers basifixed, tetrasporangiante, introrse, opening by longitudinal slits; pollen in tetrads. Gynoecium 3-carpellate, superior, with 3 or more erect ovules; ovary distally tapering, with 3 elongated stylodia; stigmas dry. Fruit a loculicidal capsule, usually with 3 spindle-shaped seeds hispid-spinose at both ends, 1 per locule; embryo small, cylindric, embedded in copious endosperm. Southeastern Colombia, southern Venezuela, Guyana, Suriname, French Guiana, northern Brazil; 1 genus and 3 species, 2 species in the flora area. All three species of Thurnia live in or along rivers, being completely submerged for most of the year, but then emerging from the water as the rivers recede during the dry season. They prefer rivers low in suspended sediments. Therefore, they are usually found in “black-water” rivers that drain areas overlain by whitesand soils derived from the erosion of the Guayana Shield and that leach out tannins and humic acids from the vegetation. Alternatively, they occur in “clearwater” rivers that lack the humic acids of the black-water rivers but do not carry the silt that is generated by younger mountains like the Andes. Vegetatively, Thurnia species are perennial herbs that closely resemble sedges (Cyperaceae, such as Scleria), especially since their stems are solid, generally trigonous, and two of the three species have sharp serrations on the leaf margins. However, when they flower after the river level drops, the differences from Cyperaceae are readily apparent. The flowers of Thurniaceae are bisexual and are grouped in heads subtended by spreading, leafy bracts; the 6 tepals and 6 exserted stamens are bright white, making the heads quite attractive. In fact, bees and flies have been observed visiting heads of Thurnia polycephala, suggesting that the family may be insect-pollinated, rather than wind-pollinated, as earlier workers have assumed. Other characters that distinguish the family from the Rapateaceae and Juncaceae, where Thurnia was formerly placed, include the anthers opening by longitudinal slits; an elongate ovary with 3 terminal stigmas; and a capsular fruit producing 3 unusual, elongate seeds that form spines at both the tip and the base. The third species of Thurnia, T. jenmanii Hook. f., is only known from the Potaro River in Guyana. It differs from the other two in having smooth leaf margins. It has one or two large heads that are more ellipsoid than the globose head of T. sphaerocephala. At low river levels, the hearts of the stems of both species of Thurnia in the flora area are an important food source for the broad-headed Amazonian turtle, Peltocephalus dumerilianus. 1. THURNIA Hook. f. in Hook., Icon. Pl. pl. 1407, 1408. 1883. Description same as for family. Southeastern Colombia, southern Venezuela, Guyana, Suriname, French Guyana, northern Brazil. 3 species, 2 in Venezuela, both in the flora area.

Thurnia 331

Fig. 279. Thurnia sphaerocephala

Fig. 280. Thurnia polycephala

332

T HURNIACEAE

Key to the Species of Thurnia 1. 1.

Inflorescence with a single large head 3–7 cm diameter ..... T. sphaerocephala Inflorescence racemose, with several smaller heads, each 1.2–2.5 cm diameter ................................................................................ T. polycephala

Thurnia polycephala Schnee, Bol. Soc. Venez. Ci. Nat. 8: 243. 1943. Aquatic herb rooted to the substrate; leaf margins serrate; heads numerous in a racemose inflorescence, each head 1.2–2.5 cm diameter; stamens and perianth parts white. In colonies rooted in river beds on sand bars or shallow rock surfaces, 50–300 m; Amazonas (Caño Cabeza de Manteco tributary of Río Autana, Caño Cañame, Caño Pimichín, Caño San Miguel, Caño Yagua near Cucurital, lower slopes of Cerro Sipapo, south of Puerto Ayacucho, Río Atabapo, Río Atacavi, Río Cataniapo, Río Guasacavi, Río Sipapo, Río Temi). Southeastern Colombia, northern Brazil. ◆Fig. 280. Flowering plants observed by Gerrit Davidse along the lower Río Cataniapo had heads with white stamens and perianth parts. These heads were visited by small bees, which may be pollinators of this species. Thurnia sphaerocephala (Rudge) Hook. f. in Hook., Icon. Pl. pl. 1407. 1883.

—Mnasium sphaerocephala Rudge, Pl. Guian. t. 12. 1805. —Dewaado (Yekwana), Maraca. Aquatic herb forming clumps, submerged much of the year but emergent in the dry season; flowering heads solitary, 3–7 cm diameter; stamens and perianth white. In and along streams and rivers in open or forested swampy areas, Mauritia palm swamps, 50– 1500 m; Bolívar (Caño Anacapá, El Paují, near Las Bonitas, Río Acanán, Río Apacará, Río Aparamán, Río Chicanán, Río Chizca, Río Caura, Río Icabarú, between lower Río Kamay and Río Aranac, Río Kanarakuni, Río Pacairao near Kavanayén, Río Tirica above Techiné-merú, Río Uaiparú), Amazonas (Caño Galipero, Cerro Sipapo, Coromoto, Río Cataniapo, Río Cuao, Río Cunucunuma, Río Gavilán at main road crossing, upper Río Orinoco, upper Río Yudi, Río Siapa above Raudal Gallineta, Sabana de Huachapana, Samariapo, near San Fernando de Atabapo). Southeastern Colombia, Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 279.

THYMELAEACEAE by Zachary S. Rogers, Paul E. Berry, and Julian A. Steyermark Trees, shrubs, or lianas. Leaves alternate or opposite, simple, margins entire; venation usually brochidodromous (except for Lophostoma in the flora area); stipules absent. Inflorescences axillary or terminal, capitate, umbellate, or racemose, often compound; bracts and bracteoles present or absent, foliaceous or modified. Flowers tubular (Thymelaeoideae) or nontubular (Octolepidoideae), actinomorphic, hypogynous, bisexual or unisexual; unisexual flowers often with staminodes or pistillodes. Floral tube when present long to short, usually cylindrical or urceolate, lower portion persistent around base of fruit or caducous, articulation above ovary absent (in the flora area); calyx lobes 4 or 5, usually imbricate; petals usually reduced to squamiform scales or absent. Stamens 1–many (4, 8, or 10 in the flora area), in 2 whorls at different heights (4 stamens in a single whorl in Schoenobiblus); filaments fused with tube when present, upper whorl of filaments included or exserted, lower whorl usually included; anthers 4-sporangiate, usually basifixed and introrse;

T HYMELAEACEAE 333

extragynoecial disk present or absent, composed of small free scales or fused into a cup-shaped structure. Ovary superior, usually 1- or 2-locular (Thymelaeoideae), to 12-locular (Octolepidoideae), sessile or stipitate, with 1 anatropous suspended ovule in each carpel; style terminal or lateral; stigma capitate, papilliate. Fruits drupaceous. Seeds usually with a crustaceous seed coat; endosperm absent to copious. Widespread in temperate and tropical regions of both hemispheres; 40–45 genera and ca. 700–800 species, 4 genera and 10 species in the flora area. Thymelaeaceae are easily recognized in the field by their fibrous, exstipulate leaves with entire margins, their strong fibrous bark, and their usually tubular flowers and single-seeded drupaceous fruits in the flora area. The family is now considered to be a basal member of Malvales based on molecular data (Alverson et al. 1998; Fay et al. 1998; Bayer et al. 1999; Savolainen et al. 2000), and has recently been recircumscribed by Herber (2003) to include two subfamilies: Thymelaeoideae (ca. 40 genera, 700 species) and Octolepidoideae (ca. 8 genera, 50 species). Molecular (Wurdack and Horn 2001) and morphological (Horn 2004) data now suggest that Tepuianthus, a small genus nearly endemic to the flora area (treated as a monogeneric family in this volume), is sister to Thymelaeaceae, and could be recognized as a basal subfamily within Thymelaeaceae. This interpretation clearly makes morphological sense considering the shared similarities between Tepuianthus and the paleotropical members of the ancestral subfamily Octolepidoideae (e.g., nontubular flowers, multicarpellate capsular fruits, noncrotonoid pollen grains).

Literature cited Alverson, W. S., K. G. Karol, D. A. Baum, M. W. Chase, S. M. Swensen, R. McCourt, and K. J. Sytsma. 1998. Circumscription of Malvales and relationships to other Rosidae: evidence from rbcL sequence data. Amer. J. Bot. 85: 876–887. Bayer, C., M. F. Fay, A. Y. De Bruijn, V. Savolainen, C. M. Morton, K. Kubitzki, W. S. Alverson, and M. W. Chase. 1999. Support for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid atpB and rbcL DNA sequences. Bot. J. Linn. Soc. 129: 267–303. Fay, M. F., C. Bayer, W. S. Alverson, A. Y. de Bruijn, and M. W. Chase. 1998. Plastid rbcL sequence data indicate a close affinity between Diegodendron and Bixa. Taxon 47: 43–50. Herber, B. E. 2003. Thymelaeaceae. Pages 373–396, in K. Kubitzki, ed., The Families and Genera of Vascular Plants. IV. Flowering Plants. Dicotyledons. Malvales, Capparales and Non-betalain Caryophyllales. Berlin: Springer. Horn, J. 2004. The morphology and relationships of the Sphaerosepalaceae (Malvales). Bot. J. Linn. Soc. 144: 1–40. Savolainen, V., M. F. Fay, D. C. Albach, A. Backlund, M. van der Bank, K. M. Cameron, S. A. Johnson, M. D. Lledó, J.-C. Pintaud, M. Powell, M. C. Shehan, D. E. Soltis, P. S. Soltis, P. Weston, W. M. Whitten, K. J. Wurdack, and M. W. Chase. 2000. Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences. Kew Bull. 55: 257–309. Wurdack, K. J., and J. W. Horn. 2001. A re-evaluation of the affinities of the Tepuianthaceae: molecular and morphological evidence for placement in Malvales, p. 151. Albuquerque, NM: Botany 2001, Abstracts.

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Key to the Genera of Thymelaeaceae 1.

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

Lianas or scrambling shrubs; leaves opposite (rarely alternate on fast growing shoots); venation composed of many close-together parallel veins ..................................................................................... 3. Lophostoma Upright shrubs or trees; leaves alternate; venation composed of brochidodromous vein loops .................................................................. 2 Flowers bisexual; calyx lobes 5; stamens 10 ............................... 2. Lasidenia Flowers unisexual; calyx lobes 4; stamens 8 ............................................ 3 Floral tube longer than calyx lobes; stamens 8, included or slightly exserted; leaves 2–21 × 1–7 cm ................................................ 1. Daphnopsis Flowal tube much shorter than calyx lobes; stamens 4, conspicuously exserted; leaves 30–36 × 12–15 cm .................................. 4. Schoenobiblus

1. DAPHNOPSIS Mart., Nov. Gen. Sp. Pl. 1: 65. 1824. Shrubs or trees. Leaves alternate, petiolate, venation brochidodromous. Inflorescences axillary or terminal, capitate, umbellate, axes often compound. Flowers unisexual (plants dioecious), each sex usually with staminodes or pistillodes, pedicellate or sessile. Floral tube cylindrical, funnel-shaped, or urceolate, persistent or caducous; calyx lobes 4, usually spreading, shorter or rarely as long as floral tube; petals absent or scale-like, sometimes fused to form a small, lobed, membranous ring. Stamens 8, sometimes a few abortive, in 2 whorls of different heights; filaments usually short. Subgynoecial disk fused to form a cup-shaped ring, or composed of several distinct scales. Ovary 1–locular; style terminal. Fruits drupaceous; fruit wall fleshy or dry. Seeds with a crustaceous or membranous coat; endosperm scanty or absent. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; 50–65 species, 7 in Venezuela, 6 of these in the flora area. Key to the Species of Daphnopsis 1. 1. 2(1).

2.

3(2). 3.

Inflorescences composed of sessile flowers in dense subsessile fascicles, usually borne on leafless portions of the stem .................. D. steyermarkii Inflorescences pedunculate, arising from the leaf-bearing branches ...... 2 Leaf blades acuminate to caudate-acuminate at the apex and base; petioles 5–15 mm long; inflorescences usually dichotomously branched with elongated secondary axes; petals present, in the form of a connate ring ......................................................................................... D. americana Leaf blades rounded to obtuse, or at least not acuminate at the apex; petioles 2–6 mm long; inflorescences with flowers crowded at the end of a penduncle, generally without secondary branches, or without elongated secondary axes; petals absent ..................................................... 3 Lower surface of blades glabrous; midrib on lower surface with some spreading trichomes; leaves drooping ..................................... D. dircoides Lower surface of blades usually covered with appressed pubescence; midrib on lower surface with appressed trichomes only; leaves ascending or spreading ........................................................................................... 4

Daphnopsis 335

4(3).

4.

5(4).

5.

Floral tube reddish white, lobes white; subgynoecial scales 4, with a dark reddish zone below the blunt apex; interior of floral tube with sparsely pubescent patches about 1/3 of the distance from the base of thetube .............................................................................................. D. nevlingiana Floral tube and lobes yellow or pale gray-green; subgynoecial scales 8, uniformly pale or straw-colored; interior of floral tube either uniformly densely pubescent in lower 1/2 or with elongated trichomes extending moderately from base to apex .................................................. 5 Tertiary venation of upper leaf surface consisting of a fine twisted, contorted, irregular insculpted pattern (best seen under magnification); subgynoecial disk with filiform lobes of uniform width throughout; interior of floral tube densely sericeous in lower 1/2 (at least in the staminate flowers) ............................................................ D. guaiquinimae Tertiary venation of upper leaf surface consisting of a subelevated reticulate pattern, not as above; subgynoecial disk with linear-lanceolate lobes (lobes often bifid and of unequal lengths), tapering from a broadened triangular base to a narrow, acicular apex; interior of floral tube nearly glabrous in the staminate flowers, moderately or sparsely pubescent with elongated trichomes from base to summit in the pistillate flowers ......................................................................... D. longipedunculata

Daphnopsis americana (Mill.) J.R. Johnst., Contr. Gray Herb. 34: 242. 1909. —Laurus americanus Mill., Gard. Dict. ed. 8, no. 10. 1768. Shrub or tree to 15 m tall; leaves 3–21 × 1–8 cm; inflorescences terminal or axillary, umbellate, 10–75-flowered (staminate), 10– 25-flowered (pistillate), branching dichotomously 1–9 times; peduncles 2–45 mm long; pedicels 1.5–8.5 mm long; petals in the form of a connate ring; fruits 6–15 × 3–9 mm. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador; 7 subspecies, 1 in Venezuela and the flora area. D. americana subsp. caribaea (Griseb.) Nevling, Ann. Missouri Bot. Gard. 44: 315. 1959. —Daphnopsis caribea Griseb., Fl. Brit. W. I. 278. 1860. Tree 8–15 m tall, young stems densely pubescent; leaves caducous on mature stems, blades lanceolate to elliptic or oblanceolate, 5–22 × 1.5–7 cm, subcoriaceous, upper surface glabrous, lower surface densely pubescent, apex caudate-acuminate or acuminate, base attenuate-cuneate; midrib raised below, densely pubescent; petioles 5–15 mm long, pubescent; pedicels 1.5–3.5 mm long; floral

tube obconic or suburceolate, 3–4.5 mm long (staminate); 1.5–2 mm long (pistillate); subgynoecial disk composed of 0.25 mm tall scales, divided nearly to base; pistillode glabrous; ovary 1.5–2 mm long; fruits 6–7 × 3–4 mm. Lower montane forests, forests bordering savannas, 400–500 m; Bolivar (Altiplanicie de Nuria east of Miamo). Aragua, Falcón, Nueva Esparta; Central America, West Indies, Colombia. Daphnopsis americana subsp. caribaea is recognized in the flora area by its larger leaves (attaining 22 × 7 cm), longer petioles (5–15 cm versus 5 cm or less), and by the presence of petals, which form a connate ring near the mouth of the floral tube. The species also occurs at lower elevation compared to other Daphnopsis in the area (400–500 m versus 1100 m and above). Daphnopsis dircoides Steyerm., Fieldiana, Bot. 28: 420. 1952. Shrub to 2 m tall, bark lenticellate; leaves drooping, caducous on mature stems, blades oblong-elliptic or ovate-oblong, 5.5–12.5 × 1.5–6 cm, coriaceous, upper surface glabrous and dull green, lower surface silvery–colored, apex obtuse or rounded, base obtuse; vena-

336

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tion on upper surface contorted, loosely and inconspicuously reticulate, strongly raised on lower surface, pubescent; petioles 3–6 mm long. Tepui scrub on slopes and tepui summits, 1900–2300 m; Bolivar (slopes of Ilútepui, summit of Sororopán-tepui). Endemic. According to Steyermark, the habit of Daphnopsis dircoides resembles that of Dirca palustris L. (North American leatherwood). Daphnopsis dircoides is distinguished from other species in the flora area by its drooping leaves with dull green upper and silver lower leaf surfaces with purple venation. This species may only be known from the two sterile examined collections. Daphnopsis guaiquinimae Steyerm., Ann. Missouri Bot. Gard. 74: 652, fig. 9. 1987. Shrub ca. 1.5 m tall; bark conspicuously covered with orbicular lenticels, young stems densely pubescent; leaves caducous on mature stems, blades broadly elliptic or ovateelliptic, 3–6 × 1–3 cm, subcoriaceous, upper surface glabrous, lower surface moderately pubescent, apex rounded or obtuse, base cuneate-attenuate; midrib raised on lower surface, densely pubescent; venation pubescent, higher orders reticulate; petioles 2–3 mm long, pubescent; inflorescences terminal or axillary, capitate-umbellate, 4–9-flowered; peduncles 1–2 cm long, pubescent; flowers tan-gray; pedicels 1–2 mm long; floral tube cylindrical, 5–7 mm long, pubescent externally, densely pubescent in lower 1/2 internally, otherwise glabrous; calyx lobes oblong-elliptic or suborbicular, 1–1.5 mm long, 0.5–0.8 mm wide, erect, pubescent on both surfaces, apex obtuse or rounded; ovary densely pubescent. Tepui summit scrub, 1600–1700 m; Bolívar (Cerro Guaiquinima). Endemic. Daphnopsis guaiquinimae is recognized by its fine contorted tertiary venation on the upper leaf surface, short peduncles, and subgynoecial disk with 8 long filiform lobes that resemble the teeth of a comb. It is known only from the type collection. Daphnopsis longipedunculata Gilg ex Ule & Domke, Notizbl. Bot. Gart. BerlinDahlem 12: 723. 1935. —Waramiku-yek (Pemón).

Shrub or treelet to 6 m tall; bark lenticellate, young stems glabrescent; leaves caducous on mature stems, blades broadly oblong-elliptic, 3.5–11 × 1–5 cm, chartaceous, upper surface pubescent or glabrescent, lower surface densely pubescent, apex rounded or obtuse tip, often slightly apiculate or emarginate, base rounded or cuneateattenuate; midrib raised below, densely pubescent on both sides; petioles ca. 6 mm long, pubescent; inflorescences terminal or axillary, umbellate, 5–15-flowered; peduncles 1.5–9 cm long, pubescent; floral tube greenish 4–6 mm long, moderately to sparsely pubescent externally (pistillate), nearly glabrous internally (staminate); calyx lobes ovate, 0.7–1.5 × 0.7–1.2 mm, pubescent on both surfaces, apex obtuse or rounded; ovary ovoid, 1.5–3.5 mm long, pubescent. Montane scrub, 1100–1400 m; Bolivar (between Kavanayén and Carrao-tepui, base of Roraima-tepui, Sorochoroyen in Río Caroní basin). Guyana side of Mt. Roraima. ◆Fig. 282. Daphnopsis longipedunculata is distinguished from others in the flora area by the pubescent adaxial leaf surfaces, peduncles reaching 9 cm long, and by the very narrow triangular lobes of the subgynoecial disk. Daphnopsis nevlingiana Steyerm., Ann. Missouri Bot. Gard. 74: 653. 1987. Tree ca. 3 m tall; bark not lenticellate, young stems densely pubescent; leaves caducous on mature stems, blades broadly oblongelliptic or obovate, 4–7.5 × 2–4.5 cm, coriaceous, upper surface glabrous, lower surface densely pubescent, apex rounded, base rounded or cuneate-attenuate; midrib pubescent above, raised and densely pubescent below; petioles 3–4 mm long, moderately pubescent; floral tube reddish white, cylindrical, ca. 5 mm long, densely pubescent externally, mostly glabrous internally; calyx lobes red with white apices, ovate-elliptic or suborbicular, 1 × 0.7–0.8 mm; ovary ovoid, ca. 2.4 mm long, densely pubescent; style ca. 1.5 mm long. Tepui summit scrub, 1200–1400 m; Bolívar (Cerro Sarisariñama). Endemic. Daphnopsis nevlingiana has a distinctive reddish white floral tube and a dull yellow subgynoecial disk with thickened lobes.

Daphnopsis 337

Fig. 281. Daphnopsis steyermarkii

Fig. 282. Daphnopsis longipedunculata

338

T HYMELAEACEAE

Daphnopsis steyermarkii Nevling, Mem. New York Bot. Gard. 17(1): 452, fig. 5. 1967. Small shrub or tree to 6 m tall; bark not lenticellate, usually covered by lichens, young stems glabrous, mature stems conspicuously covered with nodules of old inflorescences; leaves caducous on mature stems; blades broadly obovate or elliptic, 3–13.5 × 1– 7 cm, coriaceous, both surfaces glabrous, apex obtuse or subacute, margin often strongly revolute, base rounded or cuneate, sometimes subcordate; midrib glabrous, strongly raised below; venation strongly raised on both surfaces; petioles ca. 2 mm long, glabrous; inflorescences 2–8-flowered, usually borne on defoliated portion of stem, fasciculate or nearly so; peduncles 0–7 mm long; flowers pale green, subsessile, floral tube cylindrical or urceolate, ca. 2.5 mm long (staminate), 1.5–3.1 mm long (pistillate), glabrous externally, pubescent at least in lower half internally; calyx lobes suborbicular, 0.5–1 mm long, 0.7–1 mm wide, erect or reflexed,

glabrous on both surfaces except densely pubescent at apex, apex obtuse or rounded; ovary ovoid, ca. 1 mm long, sparsely pubescent or glabrescent, stipitate; stipe 0.5–0.7 mm long; fruits green-white, ovoid, ca. 9 × 7 mm, glabrescent with a pubescent apex, surrounded at base by remains of tube; style persistent. Streamsides and rocky outcrops on tepui slopes and summits, 1700–2600 m; Bolívar (Cerro Jaua, Macizo del Chimantá), Amazonas (Cerro Marahuaka, Sierra de la Neblina, Sierra de Maigualida). Guyana. ◆Fig. 281. Daphnopsis steyermarkii differs markedly from the other species in the flora area by the densely congested many-flowered fasciculate inflorescences borne on the defoliated portion of the stems. Older stems are often lichen encrusted and always possess characteristic nodular scars left behind from old inflorescences. The species can be distinguished vegetatively from others in the flora area by its thick, coriaceous leaf blades with strongly revolute margins.

2. LASIADENIA Benth., London J. Bot. 4. 632. 1845. Monoecious shrubs or small trees. Leaves alternate, petiolate; venation brochidodromous. Inflorescences terminal, umbellate or capitate; peduncles short. Flowers hermaphroditic. Floral tube cylindrical or urceolate, distinctly 10-ribbed, densely pubescent externally, glabrous internally, persistent, splitting longitudinally as fruit develops; calyx lobes 5, imbricate; petals absent. Stamens 10, in 2 whorls of different heights within tube, upper whorl near opening, lower whorl included; filaments short. Subgynoecial disk minute, densely pubescent. Ovary 1locular, sessile, pubescent; style terminal, shorter than ovary. Fruit wall hard and thin. Seed with crustaceous coat. Colombia, Venezuela, Brazil; 2 species, both in the flora area. Key to the Species of Lasiadenia 1. 1.

Leaves oblong-elliptic, rarely ovate; apex rounded or obtuse; flowers subsessile; floral tube 5–7 mm long ....................................... L. ottohuberi Leaves ovate; apex acute, acuminate, or apiculate; flowers distinctly pedicellate; floral tube 11–15 mm long .................................... L. rupestris

Lasiadenia ottohuberi Plowman & Nevling, Brittonia 38: 114, pl. 1. 1986. Shrub to 3 m tall, bark usually lenticellate; young stems densely to sparsely pubescent; leaf blades oblong-elliptic, rarely ovate, 3–6 × 1.2–3.2 cm, coriaceous, upper surface glabrous, lower surface densely pu-

bescent, apex rounded or obtuse, apiculate or slightly emarginate, base cuneate-attenuate; petioles ca. 3 mm long, densely pubescent; inflorescences terminal or subterminal, capitate, 6–15-flowered; peduncles 1–2 mm long, densely pubescent; flowers subsessile, tangray or gray; floral tube urceolate (rarely

Lasiadenia 339

suburceolate), 5–7 mm long; calyx lobes ovate-elliptic or suborbicular, 3–3.5 mm long, 2–2.5 mm wide, pubescent, apex obtuse or rounded; subgynoecial disk minute, cupshaped, densely pubescent. Seasonally flooded black-water riparian thickets and white-sand savannas, 100–200 m; Amazonas (base of Cerro Yapacana, Río Guasacavi, Santa Cruz). Adjacent Colombia (Guainía). Lasiadenia ottohuberi is characterized by oblong-elliptic or rarely ovate leaves, rounded or obtuse apices, and subsessile 5–7 mm long flowers. Lasiadenia rupestris Benth., London J. Bot. 4: 632. 1845. Shrub to 2.5 m tall, bark often lenticellate; young stems densely to sparsely pubescent; leaf blades ovate, (2–)3.3–9.4 × 1.5–3.3 cm, chartaceous, apex acute or acuminate, often apiculate, base rounded or cuneate, rarely subcordate; petioles 2–3 mm long, densely pubescent; inflorescences terminal, umbel-

Fig. 283. Lasiadenia rupestris

late, 5–10-flowered; peduncles 3–26 mm long, densely pubescent; flowers white or white-yellow, borne on 2–4 mm long densely pubescent pedicels; floral tube 11–15 mm long; calyx lobes narrowly ovate-triangular, pubescent on both surfaces, 7–8 × ca. 2 mm, apex obtuse or rounded; subgynoecial disk composed of ca. 4 densely pubescent fused or free scales. Sandy or rocky white-water or black-river banks, 50–200 m; Bolívar (Puerto Ordaz, Río Parhueña, San Félix), Amazonas (Pimichín, Río Orinoco above Samariapo, Río Sipapo, Santa Lucia). Anzoátegui, Apure; Colombia, Guyana, Brazil. ◆Fig. 283. Lasiadenia rupestris is distinguished from L. ottohuberi, the only other species in the genus, by its ovate leaves with acute or acuminate apices and its distinctly pedicellate flowers measuring 11–15 mm long.

340

T HYMELAEACEAE

3. LOPHOSTOMA (Meisn.) Meisn. in DC. Prodr. 14: 600. 1857. —Linostoma sect. Lophostoma Meisn. in Mart., Fl. Bras. 5 (1): 72. t. 30. 1855. Lianas or scandent shrubs, bark lenticellate, axillary branches sometimes curved or recurved for climbing. Leaves opposite (rarely alternate); secondary veins many, close together, ± parallel, terminating at margin, veins more pronounced on lower surface. Inflorescences terminal, racemose, compound panicles or thyrses, branching dichotomously 1–several times; primary peduncles 1–5 mm long, usually bearing 1 or 2 reduced bract-like leaves, rachis 3–6 mm long; secondary peduncles to 1 mm long. Flowers bisexual (plants monoecious). Floral tube cylindrical, persistent; calyx lobes 5, nearly erect, shorter than floral tube; petals 5, scale-like, deeply bifid (appearing as 10 scales), usually pubescent. Stamens 10, inserted in the floral tube in 2 whorls; filaments 1–3 mm long, filiform, exserted or one whorl included. Subgynoecial disk minute or absent, annular or lobed, glabrous. Ovary 1-locular, densely sericeous, sessile; style terminal (subterminal in L. amoenum), glabrous; stigma capitate; . Fruit enclosed by but not fused to the persistent floral tube; fruit wall smooth to ribbed and spinose-mammillate. Amazonian Colombia, Venezuela, and Brazil, 4 species, 1 in Venezuela and the flora area. Lophostoma amoenum Nevling, J. Arnold Arbor. 44: 161. 1963. Liana ca. 2.5 m long, axillary branches sometimes curved or recurved for climbing; leaves opposite (alternate on vigorous shoots); blades ovate or elliptic, 3.5–5.5 × 1–3

cm, coriaceous, both surfaces glabrous, apex acute or obtuse, base cuneate; midrib raised below; petioles 2–4 mm long, glabrous; inflorescences terminal, 10–16-flowered, compound racemose panicles, 2- or 3-branched, often dichotomously branched; primary pe-

Fig. 284. Lophostoma amoenum

Schoenobiblus 341

duncles 1–5 mm long, rachis 3–6 mm long; secondary peduncles to 1 mm long; bracteole minute, rose-colored; flowers pink, distinctly pedicellate; floral tube cylindrical, 15–16 mm long, pubescent externally, upper half glabrous internally, otherwise pubescent; calyx lobes 5, elliptic, 4–4.5 mm long, ca. 1.5 mm wide, upper surface glabrous, lower surface pubescent; petals 5, bifid to base (appearing as 10 scales), fleshy, ca. 0.5 × 0.25–0.5 mm, densely pubescent on outer surface, inner surface glabrous or pubescent; subgynoecial disk absent; ovary ellipsoid, ca. 1.5 mm long,

stigma slightly exserted. Seasonally flooded riparian forests along black-water rivers, 100–200 m; Amazonas (Caño San Miguel, Santa Cruz). Endemic (but almost certainly in adjacent Guainía in Colombia). ◆Fig. 284. Lophostoma amoenum can be distinguished from other species of Thymelaeaceae occurring in the flora area by its lianescent habit, its oppositely arranged leaves, and its characteristic closely arranged parallel venation, which terminates in a pronounced marginal vein.

4. SCHOENOBIBLUS Mart., Nov. Gen. Sp. Pl. 1: 65. 1824. Shrubs or trees. Leaves alternate, petiolate; venation brochidodromous. Inflorescences terminal, umbellate, inflorescence axis branched 1–several times. Flowers unisexual (plants dioecious), borne on long pedicels. Floral tube shorter than

Fig. 285. Schoenobiblus amazonicus

342

T HYMELAEACEAE

calyx lobes, persistent at base of fruit; calyx lobes 4, spreading; petals absent. Stamens 4, exserted; filaments long, adnate to base of calyx lobes. Subgynoecial disk absent or minute. Ovary usually glabrous; style terminal; stigma globose, usually near opening of the tube. Fruits slightly fleshy, smooth. Seed coat crustaceous to coriaceous; endosperm absent. Costa Rica, Panama, Colombia, Venezuela, Trinidad, Ecuador, Peru, Brazil, Bolivia; ca. 10 species, 4 in Venezuela, 1 of these in the flora area. Schoenobiblus amazonicus Steyerm., Ann. Missouri Bot. Gard. 74: 654, fig. 10. 1987. Tree ca. 6 m tall; leaf blades broadly obovate-oblanceolate, ca. 36 × 12.5–14 cm, apex acuminate or acute, base cuneate-attenuate; petioles 6–9 mm long; inflorescences terminal or axillary, 10–20-flowered, compoundumbellate, branching nearly dichotomous, primary peduncles 1- or 2-branched, secondary peduncles 1–3, branched; pedicels 10–18

mm long; calyx lobes 4, oblong-elliptic, ca. 6 × 1.5 mm, spreading, apex rounded; stamens 4, filaments 5–5.5 mm long, glabrous. Undergrowth of evergreen lowland forests, 100– 200 m; Amazonas (along road from Puerto Ayacucho to Samariapo). Endemic. ◆Fig. 285. Schoenobiblus amazonicus is the only species of Thymelaeaceae in the flora area with 4 stamens that are borne on long exserted filaments.

TILIACEAE and newly segregated MUNTINGIACEAE by Laurence Dorr and Paul E. Berry Previously, Muntingia has been placed in either Elaeocarpaceae or Tiliaceae, but molecular studies now place it in Muntingiaceae, a family distinct from all of the other Malvaceae sensu lato (Bayer et al. 1998; Stevens 2001 onward). Muntingiaceae are included here following Tiliaceae because that is where Cronquist (1981) placed the genus in his classification system, and we did not cover Muntingiaceae in its proper alphabetical sequence in the flora.

References Bayer, C., M. W. Chase, and M. F. Fay. 1998. Muntingiaceae, a new family of dicotyledons with Malvalean affinities. Taxon 47: 37–42. Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Columbia University Press, New York. Stevens, P. F. (2001 onward). Angiosperm Phylogeny Website. Version 5, May 2004. http://www.mobot.org/MOBOT/research/APweb/

T ILIACEAE 343

Key to Muntingiaceae and Tiliaceae 1.

1.

Leaf blades 2-ranked, toothed, with asymmetrical bases and heteromorphic stipule-like prophylls; flowers in extra-axillary fascicles; petals shortly clawed and crumpled in bud; fruit a berry ................ .............................................................................. Muntingiaceae, p. 362 Leaf blades variable, stipules usually present but prophylls lacking; inflorescences of modified cymose units, not extra-axillary; petals not clawed and crumpled in bud; fruit various .................... Tiliaceae, p. 343

TILIACEAE by Laurence J. Dorr and Willem Meijer Trees, shrubs, or herbs; indumentum often of stellate hairs, sometimes also simple and arachnoid hairs, and/or lepidote or fimbriate scales, or plants glabrescent. Leaves alternate, simple; leaf blades entire or 3–5-lobed, base cuneate, rounded, cordate, or sometimes oblique, margin entire or serrate, usually palmately or pinnipalmately veined; petioles often pulvinate apically; stipules present, often deciduous. Inflorescences axillary or terminal, sometimes leaf-opposed, mostly cymose, the cymes arranged into racemiform, paniculiform, or umbelliform systems, or flowers solitary; bracts and bracteoles usually present, sometimes united to form an involucre (i.e., epicalyx), or large and folicaeous forming a wing (Tilia). Flowers usually bisexual, sometimes unisexual (plants then monoecious, dioecious, or rarely gynodioecious), actinomorphic, usually 4- or 5-merous, rarely 3-merous. Calyx valvate in bud; sepals (3–)5, free or campanulate at base, often persistent, in some cases accrescent in fruit. Petals equal in number to sepals, or sometimes absent, free or rarely connate, contorted, imbricate or valvate, often with glands or glandular tissue basally. Stamens usually many, or less often 5 or 10, inserted above base of petals or borne on an androgynophore, often above a hairy disk-like ring (i.e., urceolus), often partly staminodial or reduced to staminodes in functionally pistillate flowers, free or united at base or united in 5 or 10 phalanges; anthers dorsifixed, basifixed, or sub-basifixed, 2-locular, opening by longitudinal slits or apical pores, or by a combination of the two, connective with or without sterile, apical appendages. Ovary superior, with 2–5(–10) syncarpous carpels or in some cases with 5(6) apocarpous carpels (Christiana, pro parte), locules equal in number to carpels, or rarely (Mollia) unilocular by failure of partitions to meet in center, sometimes with false septae, each cavity with 1–many seeds; placentation axile (or intruded parietal); ovules usually anatropous; style 1 in syncarpous species; stigma punctiform, capitate, or divided into as many branches as carpels in the ovary, rarely sessile. Fruits 2–many-locular, with 1–many seeds per locule, loculicidally or septicidally dehiscent capsules, or dry or somewhat fleshy drupes with 1 or several pyrenes, or a schizocarp with dehiscent or indehiscent mericarps; fruits sometimes winged, sometimes provided with prickles or spines. Seeds 1–many, often 1 per locule, winged or not, rarely arillate; endosperm copious to scanty or absent; embryo usually straight; cotyledons usually leaf-like, often lobed or dissected.

344

T ILIACEAE

As traditionally circumscribed, the Tiliaceae are worldwide albeit most abundant in the tropics; ca. 35 genera and ca. 700–800 species, 8 genera and 30 species in the flora area. Current opinion (see e.g., Bayer and Kubitzki, Fam. Gen. Vasc. Pls. 5: 225–311. 2003) favors submerging Tiliaceae in a more broadly circumscribed Malvaceae (which would include the Bombacaceae, Malvaceae, and Sterculiaceae of the Flora of the Venezuelan Guayana). If that classification is adopted, then all of the genera treated as Tiliaceae here, except Christiana, belong in Malvaceae subfamily Grewioideae. The exception, Christiana, is placed in Malvaceae subfamily Brownlowioideae. Key to the Genera of Tiliaceae 1. 1. 2(1).

2.

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

5.

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

7.

Herbs, subshrubs, or shrubs < 5 m tall; plants of weedy habitats and old fields ....................................................................................................... 2 Shrubs or trees > 5 m tall; plants of forests or other native vegetation ..... 3 Fruit subglobose to cylindric (silique-like), 2- or 3-valved, loculicidally dehiscent, unarmed; sepals unappendaged; leaf blades entire, base not cordate ..................................................................................... 3. Corchorus Fruit ellipsoid to spherical, 2- or 3(–5)-locular, indehiscent, covered with spines or bristles; sepals often with small, subapical appendages; leaf blades 3-lobed or entire, base broadly cordate ...................... 7. Triumfetta Fruits with spines or bristles; anther connective with a sterile, apical appendage ..................................................................................... 1. Apeiba Fruits unarmed; anther connective without a sterile, apical appendage ................................................................................................................ 4 Flowers unisexual; plants dioecious; petals small, < 1 cm long, relatively inconspicuous; fruits dehiscent or not; seeds not winged .................... 5 Flowers bisexual; plants hermaphroditic; petals large, > 1 cm long, showy; fruits dehiscent; seeds winged or not ....................................... 6 Flowers 3–5(–7)-merous, calyx campanulate, petals longer than calyx; androgynophore absent; pistillate flowers with 5(6) carpels fused at base; styles 5, free; capsules apocarpous, ca. 1 cm diameter, each maturing carpel loculicidally dehiscent; leaf blades broadly ovate, base cordate .................................................................................... 2. Christiana Flowers 4-merous, sepals free, petals shorter than sepals; androgynophore present; pistillate flowers with 4 completely fused locules; style very short; stigma divided into 4 reflexed arms; capsules syncarpous, indehiscent, ca. 4–5 cm diameter; leaf blades ovate-oblong, base not cordate ........................................................................................ 8. Vasivaea Flowers exinvolucellate; fruits 2-valved, globose or compressed perpendicular to the septum .................................................................... 6. Mollia Flowers involucellate (i.e., epicalyx present); fruits 5-valved, not globose or compressed perpendicular to the septum ......................................... 7 Epicalyx of (4–)6–9, briefly to ± entirely connate involucral bracts; anther locules free from one another above the middle and diverging; anthers opening at first by triangular pores, then by longitudinal slits to base; leaves with stellate and arachnoid hairs .............................. 5. Lueheopsis Epicalyx of 5–9, almost free involucral bracts; anther locules parallel, not diverging; anthers opening by longitudinal slits alone; leaves with stellate hairs alone ............................................................................ 4. Luehea

Apeiba 345

1. APEIBA Aubl., Hist. Pl. Guiane 537. 1775. Aubletia Schreb., Gen. Pl. ed. 8, 353. 1789, non Gaertn. 1788, nec Lour. 1790, nec Pers. 1807. Oxytandrum Neck., Elem. Bot. 2: 255. 1790. Trees; indumentum of stellate or stellate and simple hairs. Leaf blades simple, ovate to elliptic or obovate, base rounded or cordate, margin entire or serrulate, apex short acuminate; domatia present or not; petioles slightly pulvinate apically; stipules caducous. Inflorescences leaf-opposed, mostly few-flowered cymes; bracteate, bracts caducous. Flowers bisexual, (4)5-merous. Sepals triangular to lanceolate, free or slightly connate at base. Petals nearly as long as sepals, obovate to spatulate, yellow or white, without glands, glabrous. Stamens numerous, free (outer stamens sometimes joined basally into a short tube); outer stamens often sterile (i.e., staminodial); filaments short; anthers linear, basifixed, introrse, opening by longitudinal slits (and sometimes also with apical pores), connective with a sterile, apical appendage. Ovary syncarpous, globose, 5–10-locular; ovules numerous in each locule; style simple, slender; stigma minutely dentate. Fruit a woody capsule, depressed-globose, often with a sunken apex, spiny or bristly, indehiscent or with a small operculate-like opening. Seeds numerous, on elongated funiculi filling the locules, embedded in pulp, not winged; cotyledons orbicular-cordate. Southern Mexico, Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 9 species, 8 in Venezuela, all in the flora area. The common name Cabeza de negro is widely applied to species of this genus. The soft light wood of Apeiba is used for rafts, and fibers from the bark are used to make rope and cordage. Apeiba ulmifolia Kunth in H.B.K. {Nov. Gen. Sp. (quarto ed.) 5: 347. 1821 [1823]}, described from an unspecified locality on the Orinoco River, is a mixture of Luehea alternifolia (Mill.) Mabb. (leaves) and Apeiba cf. tibourbou Aubl. (flowers). Key to the Species of Apeiba 1. 1. 2(1). 2.

3(2).

3.

4(3). 4.

Leaf blades with 3 or 4 pairs of secondary veins in addition to 1 or 2 pairs of basal veins; sepals and petals 4(5)-merous ............................. A. glabra Leaf blades with 5–11 pairs of secondary veins in addition to 1–3 pairs of basal veins; sepals and petals (4)5-merous .......................................... 2 Branchlets, petioles, and inflorescences uniformly stellate-tomentose; capsules covered with 0.2–0.6 cm long, subglabrous spines ................ 3 Branchlets, petioles, and inflorescences stellate-tomentose, mixed with 2–4 mm long simple hairs; capsules covered with 0.5–4 cm long, hairy bristles .................................................................................................... 5 Lower surface of leaves with conspicuous tufts of brown hairs (i.e., domatia) in the axils of primary veins only; opening of capsule without teeth, slightly sunken ................................................................ A. petoumo Lower surface of leaves with tufts of brown hairs (domatia) in the axils of both primary and secondary veins, the domatia sometimes indistinct; opening of capsule with 5–7 teeth, slightly elevated ........................... 4 Leaf blades below with pale brown to gray-brown indumentum; capsules with 0.4–0.6 cm long spines ............................................... A. macropetala Leaf blades below with gray indumentum or subglabrous; capsules with 0.2–0.3 cm long spines ...................................................... A. membranacea

346

5(2).

5.

6(5).

6.

7(6). 7.

T ILIACEAE

Leaf blades below with densely whitish gray to pale-brown indumentum, the stellate hairs completely covering the epidermis; leaf margin minutely dentate to subentire; lower surface of leaves with distinct tufts of brown hairs (domatia) in the axils of the primary veins, the domatia less distinct in the axils of secondary veins............................... A. uittienii Leaf blades below with brown to yellowish brown indumentum of stellate hairs, the epidermis clearly visible below the hairs; leaf margin minutely serrate to serrate; lower surface of leaves with indistinct tufts of yellow hairs (domatia) in the axils of primary and secondary veins, or else lacking domatia .............................................................................. 6 Indumentum of branchlets yellowish brown; capsules with 1.5–2 cm long bristles, these covered with appressed and erect hairs, operculate-like opening ca. 10 mm diameter ............................................ A. schomburgkii Indumentum of branchlets rusty brown; capsules with 2–4 cm long bristles, these covered with appressed hairs, operculate-like opening 2–4 mm diameter ................................................................................... 7 Leaf blades ± rugose to bullate; petals white; capsules with flexible spines to 4.5 cm long .................................................................. A. albiflora Leaf blades not rugose or bullate; petals yellow; capsules with shorter, stiffer spines ............................................................................. A. tibourbou

Apeiba albiflora Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 209, t. 20. 1922. —Cabeza de mono, Majaguillo erizado. Apeiba tibourbou var. rugosa Szyszyl., Diagn. Pl. Nov. (Szyszylowicz) 2. 1894. Apeiba tibourbou var. krukofii Uittien, Recueil Trav. Bot. Néerl. 32: 248. 1935. Tree to 22 m tall, trunk buttressed; petals white; capsule bristles slender, flexible, to 2 (–3) cm long, hairy. Riparian forests, 50– 200(–400) m; Delta Amacuro (Río Toro at border with Bolívar), Bolívar (Reserva Forestal Imataca, Río Palmar), Amazonas (Río Ventuari). Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas, coastal Bahia, Maranhão, Mato Grosso, Pará, Rondônia). Apeiba albiflora often is considered a synonym of A. tibourbou, but it is distinguished readily by its ± rugose to bullate (versus smooth) upper leaf surface, white (versus yellow) petals, and smaller capsules with longer, more flexible bristles. Apeiba glabra Aubl., Hist. Pl. Guiane 541, t. 214 (plate mislabeled Apeiba levis). 1775. Apeiba aspera Aubl., Hist. Pl. Guiane 545, t. 216 (excluding fruit). 1775. Apeiba burchellii Sprague, Bull. Herb. Boissier sér. 2, 5: 703. 1905.

Tree to 30 m tall, trunk buttressed; petals yellow; capsule bristles slender, fragile, to 6 mm long, glabrous. Evergreen lowland forests, 100–200 m; Amazonas (Río Cunucunuma, base of Sierra de la Neblina). Colombia (Amazonas), Suriname, French Guiana, Brazil (Amapá, Maranhão, Pará). Apeiba macropetala Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 120. 1925. —Apeiba echinata var. macropetala (Ducke) Ducke, Arch. Inst. Biol. Veg. 4: 52. 1938. —Cabeza de negro, Guána. Tree to 30 m tall, trunk not buttressed; petals yellow; capsule spines stout, rigid, 0.4–0.6 cm long, with appressed, simple hairs. Nonflooded forests, 100–300 m; Bolívar (Río Caura), Amazonas (Río Cunucunuma). Guyana, Brazil (Amazonas, Mato Grosso, Pará, Rondônia), Bolivia (Pando). According to Steyermark et al. 125862 (US!, VEN), the fruit is used for combing hair, although there is a conflicting superstition that you will lose your hair by using the fruit in this manner. Apeiba membranacea Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 61. 1861. —Peine de mono, Penenimpuatsi (Baniva).

Apeiba 347

Fig. 286. Apeiba tibourbou

Fig. 288. Apeiba membranacea

Fig. 287. Apeiba petoumo

348

T ILIACEAE

Tree to 10(–40) m tall, trunk sometimes buttressed; petals bright yellow; capsule spines thick, pyramidal, 0.2–0.3 cm long, glabrous. Often along riversides, 100–200 m; Bolívar (Río Paragua basin, Sierra Imataca), Amazonas (Río Casiquiare above Capibara, Sierra Parima). Central America, Colombia, Trinidad (cultivated), Guyana, Ecuador, Peru, western Amazonian Brazil (Acre, Amazonas, Maranhão, Mato Grosso, Pará, Roraima), Bolivia. ◆Fig. 288. Apeiba petoumo Aubl., Hist. Pl. Guiane 543, t. 215 (flowering branch) 1775. —Onotillo. Apeiba echinata Gaertn., Fruct. Sem. Pl. 2: 189, t. 121. 1802. Apeiba hypoleuca Steud., Flora 26: 755. 1843. Tree to 30(–35) m tall, trunk sometimes buttressed; petals yellow; capsule spines stout, pyramidal, 1–3 mm long, glabrous. Lower montane forests, 200–600 m; Delta Amacuro (east of Río Grande and east-northeast of El Palmar), Bolívar (Altiplanacie de Nuria, southwest base of Amaruay-tepui, north of El Palmar, Sierra Imataca). Monagas; Guyana, Suriname, French Guiana, Brazil (Amapá, Pará). ◆Fig. 287. The name Apeiba aspera Aubl. also has been applied to this species, but A. aspera was based on a mixed collection. JansenJacobs and Meijer (Flora of the Guianas, ser. A, 17, 1995: 9, 14) typified A. aspera with the flowering (as opposed to the fruiting) element and placed it in synonymy under A. glabra. Apeiba schomburgkii Szyszyl., Diagn. Pl. Nov. (Szyszylowicz) 2. 1894. —Cabeza de negro, Erizo, Katunyo’ (Panare), Wana. Apeiba tibourbou var. membranacea Lockh. ex Griseb., Fl. Brit. W.I. 708. 1864. Apeiba surinamensis Uittien in Pulle, Fl. Suriname 3(1): 51. 1932; Uittien, Recueil

Trav. Bot. Néerl. 30: 184. 1933. Tree to 20(–25) m tall, trunk with plank buttresses; petals white; capsule bristles slender, 1.5–2 cm long, covered with appressed and a few erect hairs. Evergreen or semideciduous lowland to lower montane forests, sometimes bordering savannas, 50–600 m; Bolívar (Altiplanicie de Nuría, Anacoco, El Callao, La Paragua, Reserva Forestal Imataca, Río Ariza 55 km northeast of San Francisco de La Paragua, Río Ore, San Félix, Serranía de Maigualida), Amazonas (Puerto Ayacucho). Monagas, Táchira, Zulia; Colombia, Trinidad, Guyana, Suriname, Brazil (Amazonas, Pará, Roraima). Apeiba tibourbou Aubl., Hist. Pl. Guiane 538, t. 213. 1775. —Chapominaji. Tree to 15(–30) m tall, trunk sometimes buttressed; petals yellow; capsule bristles slender, flexible to 1 cm long, covered with ± appressed hairs. Lowland forests and secondary forests, near sea level to 300 m; Delta Amacuro (Departamento Tucupita), Bolívar (between La Paragua and El Cristo, Río Aro, Río Suapure, Río Villacoa, San Francisco de la Paragua, San Pedro de las Dos Bocas), Amazonas (Departamento Atures, Río Orinoquito, San Juan de Manapiare). Common in lower elevations of Venezuela; common in the Neotropics from Mexico to Bolivia. ◆Fig. 286. Apeiba uittienii Jans.-Jac. & Westra, Brittonia 47: 335, fig. 2. 1995. —Cabeza de negro. Tree to 20(–30) m tall, trunks not buttressed?; petals deep or bright yellow; capsule bristles stiff, to 0.5 cm long, puberulent. Riparian forests, lower montane forests, 100–600(–800) m; Bolívar (Río Canaracuni, upper Río Caura, Río Mawela, Río Nichare), Amazonas (Río Mawarinuma east of Sierra de la Neblina, Serranía Batata, Simarawochi). Brazil (Roraima: Sierra Parima).

2. CHRISTIANA DC., Prodr. 1: 516. 1824. Speirostyla Baker, J. Linn. Soc., Bot. 25: 298. 1889. Asterophorum Sprague, Bull. Misc. Inform. Kew 1908: 249. 1908. Tahitia Burret, Notizbl. Bot. Gart. Berlin-Dahlem 9: 609. 1926. Trees; indumentum of stellate hairs, sometimes glabrescent. Leaf blades simple, ovate (flora area species) to narrowly elliptic, sometimes 3-lobed, margin entire (flora area species) or irregularly dentate; stellate-hairy (flora area species)

Christiana 349

or glabrous; without domatia; stipules caducous. Inflorescences axillary or terminal panicles or cymules; bracteate. Flowers unisexual (plants then monoecious or dioecious). Calyx ± campanulate, 3–5-lobed, with an annular nectary (sometimes nectary expressed as isolated patches of tissue). Petals 5(–7), spatulate, exceeding the calyx lobes in length. Staminate flowers: stamens many, briefly connate at base, all fertile; anthers dorsifixed, dehiscing by longitudinal slits, locules spreading, connective without a sterile, apical appendage; pistillode lacking. Pistillate flowers: sterile stamens (i.e., staminodia) many, briefly connected at base, carpels (1)2–5(6), free or fused to varying degrees; styles 5, partially connate at the base; stigmas 5, flattened and irregularly lobed, reflexed; ovules 1 per carpel. Fruits capsules or fol-

Fig. 289. Christiana africana

350

T ILIACEAE

licles, syncarpous or rarely apocarpous (flora area species), loculicidally dehiscent. Seeds 1 per carpel, spherical, usually mottled; endosperm fleshy; cotyledons large and foliaceous. Southern Mexico, Central America, South America (Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil), Tahiti, tropical Africa, Madagascar; 5 species; 1 in Venezuela. Christiana africana DC., Prodr. 1: 516. 1824. —Cabeza de negro. Christiana cordifolia Hook. f., Niger Fl. 238. 1849. Carpodiptera ?schomburgkii Baill., Adansonia 10: 181. 1872. Christiana madagascariensis Baill., Bull. Mens. Soc. Linn. Paris 1: 542. 1885. Speirostyla tiliifolia Baker, J. Linn. Soc., Bot. 25: 299, t. 50. 1889. Tree to 8(–16) m tall; leaf blades entire, broadly ovate; petals white or yellow; fruit separating into 5 ovoid-ellipsoid carpels, densely stellate hairy. Riparian forests, 50– 100 m; Bolívar (Caruachi, Parque Cachamay

near Puerto Ordaz, lower Río Caroní). Zulia; Southern Mexico, Belize, Nicaragua, Costa Rica, Guyana, Ecuador, Brazil, Africa, Madagascar. ◆Fig. 289. The disjunct African-American distribution of Christiana africana is remarkable and has not been explained satisfactorily. There are three other American species of Christiana, all of which occur in South America. All three are rarely collected and have very narrow ranges, as does the species found in Tahiti. Christiana africana is the only species in the genus with a fully apocarpous gynoecium.

3. CORCHORUS L., Sp. Pl. 529. 1753; Gen. Pl. ed. 5: 234. 1754. Antichorus L., Syst. Nat. ed. 12, 2: 264. 1767; Mant. Pl. 9, 64. 1767. Nettoa Baill., Adansonia 6: 238. 1866. Oceanopapaver Guill., Bull. Soc. Bot. France 79: 226. 1932. Shrubs, subshrubs, or annual or perennial herbs (flora area species); indumentum of simple or stellate hairs. Leaf blades simple, ovate, elliptic, lanceolate, or oblong, margin serrate, crenate, or dentate, sometimes with a pair of setae or filiform appendages at base of blade; without domatia; stipules subulate, often ± as long as petioles. Inflorescences leaf-opposed (or terminal) few-flowered fascicles, or flowers solitary; bracteate, bracts minute. Flowers bisexual, (4)5-merous. Sepals often cucullate. Petals clawed, yellow, without glands. Stamens numerous, (5–)20–40, all fertile; filaments sometimes inserted on a short torus, or sometimes united at base to form a short androgynophore, androgynophore sometimes with nectar glands opposite the petal bases; anthers versatile, dorsifixed, introrse, opening by longitudinal slits, anther locules ± as long as wide, connective without sterile, apical appendage. Ovary syncarpous, 2–5(–10)-locular; ovules 2–many in each locule; style simple, short; stigma cup-shaped, lobed, or capitate-fimbriate. Fruit a capsule, subglobose to narrowly cylindric, sometimes silique-like, straight or slightly curved, rostrate or not, smooth, spiny, or muricate, loculicidally 2–5-valved, often with transverse septae between the seeds. Seeds 2–many, polygonal, glabrous; endosperm fleshy; often with curved embryo; cotyledons foliaceous. Tropics and subtropics worldwide; 40(–100?) species, 6 in Venezuela, 3 of these in the flora area. Many species of Corchorus are weedy; some are cultivated for fiber (jute) and others locally used as vegetables. The generic name means “low-quality vegetable” in Greek.

Corchorus 351

Key to the Species of Corchorus 1.

1. 2(1). 2.

Ovary 3-locular; capsules ± triangular in cross section, splitting into 3 winged valves; leaf blades ovate, often with a pair of setose or filiform appendages at the base, glabrous or with a few short hairs ..... C. aestuans Ovary 2-locular; capsules and leaves not as above .................................. 2 Branchlets ± appressed-pubescent, mixed with 2 mm long erect hairs; capsules sparsely pilose with hairs to 2 mm long ........................ C. hirtus Branchlets subglabrous with a line of pubescence; capsules sparsely appressed pubescent with hairs to 0.5 mm long ..................... C. orinocensis

Corchorus aestuans L., Syst. Nat. ed. 10, 2: 1079. 1759. —Corchorus acutangulus Lam., Encycl. 2: 104. 1786. Herbaceous annual or perennial to 1 m tall; leaf blades mostly ovate, often with a pair of basal setae, upper and lower surface sparsely appressed-pilose; capsules narrowly cylindric, winged, glabrous, apex 3-horned, each horn with 2 recurved or horizontal arms. Habitat unknown in the flora area, probably upland savannas, ca. 900 m; Bolívar (near Roraima-tepui). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Nueva Esparta, Sucre, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Trinidad, Tobago, Ecuador, Brazil; pantropics. A single, old collection is known from the flora area, Schomburgk 857 (NY), gathered on the Guyana side of Mount Roraima.

[1823]. —Corchorus hirtus var. orinocensis (Kunth) K. Schum. in Mart., Fl. Bras. 12(3): 127. 1886. Corchorus argutus Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 337. 1821 [1823]. Corchorus argutus var. benthamii K. Schum. in Mart., Fl. Bras. 12(3): 130. 1886. Herbaceous annual to 1 m tall; leaf blades narrowly ovate, subglabrous; capsules narrowly cylindric, not winged, glabrous, apex beaked, beak 8–10 mm. Open places of lowlands and hills, 50–300 m; Bolívar (Ciudad Bolívar, Paviche, Río Paragua), Amazonas (northeast of Puerto Ayachucho, Río Manapiare savannas, San Juan de Manapiare,

Corchorus hirtus L., Sp. Pl. ed. 2, 1: 747. 1762. —O’mochahcho ipeñ (Panare). Corchorus pilolobus Link, Enum. Hort. Berol. Alt. 2: 72. 1822. Herbaceous annual to 1 m tall; leaf blades narrowly ovate or ovate, both surfaces sparsely appressed-pilose; capsules narrowly cylindric, not winged, sparsely pilose with hairs to 2 mm, apex beaked, beak 1–2 mm. Open areas, 50–1000 m; Bolívar (Corozal, El Tigre near Río Cuchivero, 6 km from Maniapure toward Caicara, Río Uairén). Aragua, Barinas, Carabobo, Mérida, Táchira, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Trinidad, Tobago, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 290. Corchorus orinocensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 337. 1821

Fig. 290. Corchorus hirtus

352

T ILIACEAE

Santa Isabel). Apure, Aragua, Barinas, Carabobo, Miranda, Portuguesa, Táchira, Trujillo, Vargas, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Jansen-Jacobs and Meijer (Flora of the Guianas, ser. A, 17, 1995: 26) were the first to unite Corchorus orninocensis and C. argutus,

which were published simultaneously, and when the two names are considered synonymous, the former name must be used. Bornstein in Howard (Flora of the Lesser Antilles 5, 1989: 191–192) confounded Corchorus orinocensis with C. hirtus and a third species of Corchorus that is not found in the flora area.

4. LUEHEA Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: 410. 1801, nom. cons. Alegria DC., Prodr. 1: 516. 1824. Brotera Vell., Fl. Flumin. 322. 1825 [1829], non Cav. 1799, nec Spreng. 1801, nec Spreng. 1802, nec Willd. 1800 [1803]. Trees or shrubs; indumentum of stellate hairs. Leaf blades simple, elliptic to ovate-oblong, base acute, rounded, or cordate, often slightly oblique, margin dentate or serrate, apex acute to acuminate; without domatia; stipules caducous. Inflorescences many-flowered axillary cymes, or flowers solitary and terminal. Flowers bisexual, showy, with an epicaylx of 5–9, almost free, lanceolate, involucral bracts. Sepals 5, lanceolate. Petals 5, linear to obovate or ± orbicular, white, yellow, or violet, thickened-glandular at the base, glabrous except for appressed white hairs at the base abaxially. Androgynophore absent; stamens numerous, ± united into 5 phalanges, intergrading into outer anther-less filaments (i.e., staminodes), these sometimes lacking (not flora area species) or fused and petaloid (not flora area species), the phalanges villose basally; anthers dorsifixed, in bud introrse and straight, at anthesis extrorse and becoming hippocrepiform (horseshoe-shaped), opening by longitudinal slits, connective without sterile, apical appendage. Ovary syncarpous, 5locular, villose, ovules 10–20, ascending; style simple; stigma capitate, discoid or lobed. Fruit a woody capsule, imperfectly loculicidal from the apex, opening halfway or less by 5 valves, cylindrical to ellipsoid. Seeds numerous, winged on one side. Mexico, Central America, West Indies, South America south to Argentina; ca. 25 species, 4 in Venezuela, 3 of these in the flora area. Key to the Species of Luehea 1. 1. 2(1).

2.

Flowers solitary or occasionally inflorescences with a few large flowers; petals orbicular, 3–5.5 cm long; capsules to 6 cm long..............L. candida Inflorescences many-flowered; petals not orbicular, < 4 cm long; capsules to 4 cm long ............................................................................................ 2 Petals broadly obovate, 3–4 cm long; capsules 2.5–4 cm long, ovoid-cylindrical, dehiscent about 3/5 of their length, not markedly channeled in the middle of the valves ................................................... L. alternifolia Petals linear to oblanceolate or narrowly obovate, 0.8–1.5 cm long; capsules to 1.5 cm long, ovoid, dehiscent to half or more of their length, channeled in the middle of the valves ..................................... L. cymulosa

Luehea alternifolia (Mill.) Mabb., Gard. Bull. Singapore 54: 254. 2002. —Cedrus alternifolia Mill., Gard. Dict. ed. 8, Cedrus no. 3. 1768. —Cabeza de negro. Luehea speciosa Willd., Ges. Naturf.

Freunde Berlin Neue Schriften 3: 410, t. 5. 1801. Luehea ferruginea Turcz., Bull. Soc. Imp. Naturalistes Moscou 31: 224. 1858. Tree to 25(–30) m tall; trunk buttressed;

Luehea 353

petals white or yellow; capsules broadly cylindric to ovoid, 2.5–4 cm long, valves rounded, not channeled. Semidecidous forests, 200–500 m; scattered in northern Bolívar (Altiplanicie de Nuria, northeast of Ciudad Piar, islands in Lago Guri). Aragua, Barinas, Carabobo, Distrito Federal, Lara, Mérida, Sucre, Vargas, Yaracuy, Zulia; Mexico, Central America, Cuba, Puerto Rico, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Luehea candida (DC.) Mart., Nov. Gen. Sp. Pl. 1: 102. 1824 [1826]. —Alegria candida DC., Prodr. 1: 517. 1824. —Guácimo, Guácimo cimarrón, Ya’ra ipun yo’ (Panare). Luehea nobilis Linden & Planch., Pl. Columb. 52. 1863 [1874–1875]. Tree to 15 m tall, trunk not buttressed; petals white; capsules broadly cylindric to ovoid, to 6 cm long, valves channeled, conspicuously 5-angled. Along rivers and woods bordering savannas, 50–300 m; Bolívar (La Paragua, La Unión de Río Caura, San Pedro). Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Guárico, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; Mexico, Central America, northern Colombia. ◆Fig. 291.

Fig. 291. Luehea candida

The wood of Luehea candida serves as structural elements of Panare houses. Luehea cymulosa Spruce ex Benth., J. Linn. Soc., Bot. 5(suppl. 2): 58. 1861. —Guácimo. Luehea tessmannii Burret, Notizbl. Bot. Gart. Berlin-Dahlem 9: 836. 1926. Tree to 16 m tall, trunk buttressed; petals white; capsules ovoid to pyriform, to 1.5 cm long, valves channeled. Flooded riverbanks, flooded gallery forests, disturbed savannas, 50–200 m; Amazonas (Isla Ratón in Río Orinoco, near mouth of Río Atabapo). Apure; Colombia (Amazonas, Meta), Ecuador, Peru, Amazonian Brazil, Bolivia (Beni, Pando). Luehea cymulosa and L. seemannii Triana & Planch., which also occurs in Venezuela, are difficult to distinguish morphologically and their relationship deserves closer scrutiny. The principal differences relate to inflorescence, flower, and capsule size; for example, the former has shorter inflorescences (to 8 versus 20 cm long), shorter petals (8–13 versus 15 mm long), and smaller capsules (14–16 versus 22 mm long). The two species appear to have discrete ranges. Luehea cymulosa is Amazonian in distribution, while L. seemannii is common in Central America, northern Colombia, and western Venezuela.

354

T ILIACEAE

5. LUEHEOPSIS Burret, Notizbl. Bot. Gart. Berlin-Dahlem 9: 838. 1926. Trees; indumentum of stellate and arachnoid hairs. Leaf blades simple, elliptic to obovate, base rounded, margin entire, dentate, or crenulate, apex obtuse to acute to short acuminate; without domatia; stipules early caducous. Inflorescences axillary or terminal thryses. Flowers bisexual, showy, with an epicaylx of (4–)6–9, briefly to ± entirely connate involucral bracts. Sepals 5, lanceolate; petals 5, obovate, white, pink, or rose-violet, thick, glandular, puberulent at base abaxially. Androgynophore absent; staminal tube prominent, bearing numerous outer sterile filaments (i.e., staminodes) and inner stamens; anthers sub-basifixed, subcordate, introrse, at anthesis erect and straight, dehiscing from apical pores and at length by slits to base, anther locules free above the middle and diverging, connective without a sterile, apical appendage. Ovary syncarpous, 5-locular; ovules few, biseriate; style simple, stigma subcapitate or dilated apically. Fruit a woody capsule, cylindricovoid, imperfectly loculicidal from the apex, opening by 5 valves, valves channeled or not. Seeds small, winged. South America; 6 or 7 species, 3 in Venezuela, all in the flora area. Key to the Species of Lueheopsis 1.

1.

2(1). 2.

Involucral teeth 10–20 mm long, nearly as long as the sepals; leaf blades mostly narrowly ovate to oblong, below with a dense, ferruginous covering of matted arachnoid hairs and only a few stellate hairs on larger veins ...................................................................................... L. althaeiflora Involucral teeth 1–4 mm long, much shorter than the sepals; leaf blades broader, elliptic-ovate to suborbicular, below with many stellate hairs on larger and smaller veins (not as ferruginous as above) or else subglabrous ............................................................................................ 2 Involucral teeth 9, 3–4 m long; inflorescence to 13 cm long; capsule valves channeled ................................................................................. L. duckeana Involucral teeth 6, 1 mm long; inflorescence 1.5–2.5 cm long; capsule valves rounded, not channeled ....................................................... L. rosea

Lueheopsis althaeiflora (Spruce ex Benth.) Burret, Notizbl. Bot. Gard. Berlin-Dahlem 9: 837, 840. 1926. —Luehea althaeiflora Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 53. 1861. —Majagua, Palo de araguato. Tree to 20 m tall, trunk buttressed; petals rose-pink or violet; capsules not seen. Riparian forests, 50–200 m; Amazonas (Caño Yapacana, Río Casiquiare, Río Siapa near mouth of Caño Chimoni, San Carlos de Río Negro). Colombia (Vaupés), Brazil (Amazonas). ◆Fig. 292. Lueheopsis duckeana Burret, Notizbl. Bot. Gard. Berlin-Dahlem 9: 840. 1926. —Canoa, Ya’ra ipun yo’ (Panare) Lueheopsis violacea Standl., Trop. Woods 33: 13. 1933.

Tree to 20 m tall; trunk buttressed; petals pink or violet; capsules narrowly ovoid, to 2.5 cm long, valves channeled. Semideciduous forests on granitic slopes and savannas, 50– 400 m; Bolívar (Maniapure, northeast of Maripa); Brazil, Bolivia (Pando, Santa Cruz). Steyermark et al. 125688 (MO, US!, VEN), a sterile collection made on the Río Cunucunuma between Cerro Duida and Cerro Huachamacari (Amazonas), may also belong here. The common name Steyermark cited was Mavi-ní and he noted that the fruit was said to be eaten like an aguacate (sic!). Lueheopsis rosea (Ducke) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 9: 837, 840. 1927. —Luehea rosea Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 208. 1922.

Mollia 355

Fig. 292. Lueheopsis althaeiflora

Luehea flavescens Uittien, Recueil Trav. Bot. Néerl. 22: 363. 1925. —Lueheopsis flavescens (Uittien) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 9: 837, 842. 1926. Tree to 35(–40) m tall, trunk buttressed; petals deep rose-pink; capsules cylindrical,

3–3.5 cm long, valves rounded, not channeled. Evergreen lowland forests on flood plains, 100–300 m; Amazonas (35 km southeast of La Esmeralda). Guyana, Suriname, French Guiana, Peru, Brazil (Amazonas, Pará, Rondônia), Bolivia.

6. MOLLIA Mart., Nov. Gen. Sp. Pl. 1: 96. 1824 [1826], nom. cons. Schlechtendalia Spreng., Syst. Veg. 4(2): 295. 1827, non Willd. 1803, nom. rej., nec Less. 1830, nom. cons. Trees or shrubs; indumentum of stellate hairs and/or scales. Leaf blades simple, ovate to elliptic or oblong, margin entire or crenulate to dentate, domatia present or not; stipules rudimentary. Inflorescences axillary, few- to many-flowered cymes, sometimes on a common peduncle, or flowers solitary. Flower buds oblong, usually at least 3 times longer than wide; flowers bisexual, showy, 5-merous. Sepals linear. Petals truncate, erose apically, white or pinkish, without glands. Stamens numerous, in 2 whorls each of 5 phalanges, filaments ± united basally, outer whorl longer than the inner; anthers introrse, dimorphic, opening by longitudinal slits, outer anthers cordate basally, dorsifixed, inner anthers sagittate basally, subbasifixed, connective without a sterile, apical appendage. Ovary syncarpous, 2-locular or incompletely so (and then appearing 1-locular), with 2 placentas opposite each other and connected by pulpy tissue; ovules numerous in 2 parallel rows on each placenta; style filiform; stigma punctiform. Capsules 2-locular, loculicidal, globose or ± compressed contrary to the septum and sometimes winged along the line

356

T ILIACEAE

of dehiscence; exocarp hairy or scaly, smooth or tuberculate, locules each with 2 rows of 8–20 flattened seeds fitting between narrow membranous partitions. Seeds flat, winged or not; embryo flat, endosperm copious, oily. South America (principally Amazon Basin and Guyana); ca. 12 species, 7 in Venezuela, all of these in the flora area. Morphologically Mollia is somewhat anomalous in the Malvaceae subfamily Grewioideae, but Bayer and Kubitzki (Fam. Gen. Vasc. Pls. 5: 255. 2003) cited unpublished molecular data that support its inclusion in this subfamily. Key to the Species of Mollia 1. 1. 2(1).

2.

3(1). 3.

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

Indumentum of stellate hairs (rarely leaf surface below mostly lepidote or fimbriate scaly) .................................................................................. 2 Indumentum of lepidote or fimbriate scales ............................................. 3 Margin of leaf blades entire to undulate (rarely dentate near apex), lower surface densely white-woolly-tomentose (rarely mostly lepidote or fimbriate scaly), upper surface with scattered stellate hairs; capsules slightly compressed, margin often narrowly winged; seeds not winged ................................................................................................ M. tomentosa Margin of leaf blades conspicuously dentate, lower surface stellate pubescent, upper surface often scabrid with scattered stellate hairs; capsules globose, margin not winged; seeds winged ............................ M. ulei Inflorescences compound, both pedicels and peduncle visible; leaf blades without domatia in the axils of principal nerves below ................ M. sp. A Inflorescences simple, pedicel alone visible (peduncles absent or greatly reduced); leaf blades with domatia (sometimes inconspicuous) in the axils of principal veins below ................................................................ 4 Lower surface of leaf blade with scales ± remote, associated mostly with veins and veinlets ................................................................ M. glabrescens Lower surface of leaf blade densely covered with scales, these obscuring veins, veinlets, and areoles ................................................................... 5 Flowers small, sepals < 2.5 cm long .............................................. M. lepidota Flowers large, sepals > 3 cm long .............................................................. 6 Leaf blades broadly ovate (rarely oblanceolate), apex acute to acuminate; capsules slightly compressed, surface tuberculate ................. M. speciosa Leaf blades ovate, apex long acuminate; capsules ± globose, surface ± smooth .................................................................................. M. grandiflora

Mollia glabrescens Benth., J. Bot (Hooker) 4: 131. 1842. —Caná-acu-dajay, Cenato (Yekwana). Tree to 12(–20) m tall; margin of leaf blade dentate; domatia present, inconspicuous; capsules ± globose, covered with scales; seeds winged. Evergreen lowland forests, 100–300 m; Bolívar (El Cácaro between Río Caura and Río Paragua), Amazonas (Río Cunucunuma, Río Orinoco, Río Sipapo, San Fernando de Atabapo, Tamatama). Guyana. In the most recent revision of Mollia (Meijer, Mem. New York Bot. Gard. 29: 68–

82. 1978), the Venezuelan material of M. glabrescens was misidentified as M. gracilis Spruce ex Benth., a species with much smaller flowers that appears to be restricted to Amazonian Peru and neighboring Ecuador, Brazil, and Colombia. Mollia grandiflora Meijer, Mem. New York Bot. Gard. 29: 80. 1978. —Majagua. Tree to 10 m tall; margin of leaf blade entire (often dentate apically); domatia present, inconspicuous; capsules ± globose, surface ± smooth, covered with scales, margin

Mollia 357

winged, apex apiculate or emarginate; seeds not winged. Riparian forests, 100–200 m; Amazonas (Río Casiquiare, Río Guainía, Río Siapa, Río Yatua near Río Casiquiare, San Carlos de Río Negro). Colombia (Guainía). Mollia lepidota Spruce ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 59. 1861. Colombia (Amazonas, Vaupés), Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; 4 or 5 subspecies, 1 in Venezuela. Peruvian material of the Mollia lepidota complex often is segregated as M. williamsii Baehni. M. lepidota subsp. casiquiarensis (Baehni) Meijer, Mem. New York Bot. Gard. 29: 78. 1978. —Mollia lepidota var. casiquiarensis Baehni, Candollea 5: 417, t. 12e. 1934. Tree to 10 m tall; margin of leaf blade entire; capsules globose, slightly rimmed or winged, apex acute, not tuberculate, not winged; seeds not seen. Forested river banks, 50–200 m; Amazonas (Buena Vista, Culimacare, near the mouth of the Río Casiquiare, Solano). Brazil (Amazonas). Mollia speciosa Mart., Nov. Gen. Sp. Pl. 1: 97, t. 60. 1824 [1826]. —Cabeza de culebra, Cabeza de tamatama, Majagua orillera. Shrub or tree to 12(–25) m tall; margin of leaf blade slightly crenulate, often dentate apically; capsules slightly compressed, apiculate, sometimes emarginate, tuberculate, indistinctly winged apically, covered with scales; seeds not winged. Flooded river margins, white-sand savannas, lowland forests, 50–200 m; Amazonas (Caño Caname, Caño Guasuriapana, Caño Pasiba, Caño San Miguel, Caño Yagua, Río Atabapo near San Fernando, Río Casiquiare, Río Cunucunuma, Río Guainia, Río Guayapo, Río Orinoco, Río Sanariapo, Río Sipapo, Río Ventuari, San Carlos de Río Negro). Guyana, Brazil (Amazonas). ◆Fig. 294. Mollia tomentosa Spruce ex Benth., J. Linn. Soc., Bot. 5(suppl. 2): 60. 1861. —Coco de mono, Majagua rabipelado. Shrub or tree to 8(–15) m tall; margin of leaf blades entire to undulate (sometimes dentate); capsules slightly compressed, stellate-pubescent, margin winged; seeds not

winged. Seasonally flooded black-water riparian areas, 50–200 m; Amazonas (Río Baría, Río Guainía, Río Pasimoni, Río Yatúa). Colombia (Vaupés), Brazil (Amazonas). In some of the material we refer to Mollia tomentosa [i.e., Liesner 7341 (MO, US!), Maguire et al. 36402 (NY, US!), Ll. Williams 14305 (F, US!)] the petioles, flower buds, and fruit are stellate-hairy, but the pubescence on the lower leaf surface is more scale-like than stellate. Mollia ulei Burret, Notizbl. Bot. Gart. Berlin-Dahlem 9: 844. 1926. —Guácimo rebalsero, Vara blanca, Ya’ra ipun yo. Mollia stellaris Meijer, Mem. New York Bot. Gard. 29: 72. 1978. Mollia scabra Meijer, Mem. New York Bot. Gard. 29: 72. 1978. Shrub or tree to 12 m tall; margin of leaf blade dentate; domatia present as inconspicuous pit-like flaps of tissue not hairs; capsules globose, stellate-pubescent, margin not winged; seeds winged. Riparian forests, primary lowland and hill forests, semideciduous forests on granitic outcrops, 100– 300 m; Bolívar (Morichal El Caballo, near Panare, Reserva Forestal La Paragua, island in Río Caroní north of El Plomo, Río Caura, mouth of Río Paragua, Río Parguaza between El Carmen and Raudal Maraca), Amazonas (Puerto Ayacucho). Guyana, Peru, Brazil (Amazonas, Roraima). ◆Fig. 293. Mollia sp. A Mollia speciosa var. alata Meijer, nom. nud. Tree to 20 m tall; margin of leaf blade remotely crenulate or entire; capsules ± globose, apex emarginate, covered with scales, tuberculate, margin conspicuously winged laterally; seeds not winged. Seasonally flooded forests, 100–200 m; Amazonas (Caño San Miguel, Río Casiquiare, Río Pamoni). Brazil (Amazonas). This unnamed species is based on Aymard 9091 (MO, PORT!, US!), Colella et al. 1878 (US!), and Stergios & Stergios 11388 (US!). It resembles Mollia speciosa but differs in its branched inflorescence structure, capsules that are conspicuously winged but not compressed, entire to remotely crenulate leaf margin (not dentate apically), and lack of domatia.

358

T ILIACEAE

Fig. 293. Mollia ulei

Fig. 294. Mollia speciosa

7. TRIUMFETTA L., Sp. Pl. 444. 1753; Gen. Pl. ed. 5, 203. 1754. Bartramia L., Sp. Pl. 389. 1753, nom. rej., non Salisb. 1796, nec Hedw. 1801, nom. cons. Shrubs or herbs; indumentum of stellate or stellate and simple hairs, rarely glabrescent. Leaf blades simple, 3–5-lobed (rarely 7–9-lobed) or entire, ovate, ovateelliptic, broadly ovate, or elliptic (sometimes varying greatly within an individual by position on stem), margin irregularly serrate, often with black glands terminating the lowermost serrations of the blade or at the petiole apex; without domatia; stipules usually persistent. Inflorescences axillary and few-flowered, or terminal and paniculate with 1–12 cymes, each with (1–)3(–5) flowers per node, often with leaf-like bracts. Flowers bisexual (flora area species) or pistillate and plants

Triumfetta 359

gynodioecious. Sepals 5, free, often cucullate, often with small, subapical appendages. Petals 5 (absent in T. lappula), free, shorter than sepals, unguiculate, ciliate basally, yellow to orange, without glands. Androgynophore bearing 5 orbicular glands, very short or absent, crowned by a ciliate disk (i.e., urceolus) (absent in T. lappula) surrounding the base of the stamens, stamens 5–60, free, all fertile, or staminodial or absent in pistillate flowers; anthers versatile, dorsifixed, introrse in bisexual flowers, dehiscing by longitudinal slits, connective without a sterile, apical appendage. Ovary syncarpous, sessile on the androgynophore, ellipsoid to spherical, covered with numerous, hyaline spinules that often are incurved apically, 2- or 3 (–5)-locular, each locule 2-ovulate; style filiform; stigma capitate or shortly 2- or 3 (–5)-lobed. Fruit a capsule, ellipsoid or spherical, indehiscent (sometimes loculicidally dehiscent), 2- or 3-locular, or falsely 6–9-locular, covered with numerous spines. Seeds 1 or 2 per locule, laterally compressed, ovoid or pyriform, glabrous; embryo straight, endosperm oily, scanty. Pantropics; ca. 100–150 species, ca. 6 in Venezuela, 4 of these in the flora area. Species of Triumfetta in the flora area are weedy, usually occurring in disturbed habitats and cultivated areas. Key to the Species of Triumfetta 1. 1. 2(1). 2. 3(2). 3.

Sepals cucullate; fruit surface densely pubescent with stellate hairs, spines subglabrous ............................................................... T. rhomboidea Sepals not cucullate; fruit surface sparingly pubescent with stellate hairs or glabrous, spines sparsely retrorse-hispid to glabrous ..................... 2 Petals absent; stamens (5)10(15); urceolus wanting ..................... T. lappula Petals present; stamens > 15; urceolus present ....................................... 3 Leaf blades with stellate hairs only; petals elliptic to narrowly obovate; stamens 15–25 ....................................................................... T. semitriloba Leaf blades with both stellate and simple hairs; petals broadly obovate; stamens 25–30 ........................................................................ T. bogotensis

Triumfetta bogotensis DC., Prodr. 1: 506. 1824. Triumfetta hispida A. Rich. in Sagra, Hist. Phys. Cuba, Pl. Vasc. 10: 204. 1841 [1845]. Triumfetta valenciensis R. Knuth, Repert. Spec. Nov. Regni Veg. Beih. 43: 464. 1927, nom. nud. Triumfetta bolivariensis D.E. González, nom. nud. Shrub to 1(–2) m tall; leaf blades broadly to narrowly ovate, usually 3-lobed, indumentum of both stellate and simple hairs; petals present, yellow; sepals not cucullate; capsule spines sparsely retrorse-hispid to glabrous. Sandstone cliffs, 500–1000 m; Bolívar (Sierra de Lema, Uaipán-tepui). Aragua, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas,

Portuguesa, Sucre, Táchira, Zulia; a widespread weedy species of tropical America found in Mexico, Central America, West Indies, and South America from Colombia to Argentina and Paraguay. Triumfetta lappula L., Sp. Pl. 444. 1753, non Vell. 1827 [1831]. Shrub or herb to 2.5 m tall; leaf blades rhomboid-ovate to pandurate, sometimes slightly 3–5-lobed, indumentum of stellate hairs; petals absent; sepals not cucullate; capsule spines sparsely retrorsely hispid. Open rather dry areas, 50—300 m; Bolívar (Cerro Picacho in Altiplanicie de Nuria, Isla Santa Elena, Río Orinoco, opposite mouth of Río Parhueña). Aragua, Barinas, Cojedes, Distrito Federal, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Su-

360

T ILIACEAE

cre, Trujillo, Vargas, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Trinidad, Tobago, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 295.

fruits and simple as well as stellate hairs on the stem, often is segregated as T. rubricaulis.

Triumfetta rhomboidea Jacq., Enum. Syst. Pl. 22. 1760; Select. Stirp. Amer. Hist. 147, t. 90. 1763, non Lindl. 1821. Bartramia indica L., Sp. Pl. 389. 1753, non Triumfetta indica Lam. 1791. Triumfetta bartramia L., Syst. Nat. ed. 10. 1044. 1759, nom. illeg. Shrub or perennial herb to 3 m tall; leaf blades elliptic to broadly ovate, slightly 3lobed, indumentum of stellate hairs; petals present, yellow; sepals cucullate; capsule spines subglabrous. Open ruderal areas, ca. 50 m; Bolívar (Alcabala Casa Blanca to Isla Anacoca road, Ciudad Bolívar). Barinas, Carabobo, Mérida, Miranda; a pantropical weed found in the Americas in the U.S.A. (Florida), Belize, the West Indies, Colombia, Trinidad, Tobago, Guyana, French Guiana, Peru, Brazil. Triumfetta semitriloba Jacq., Enum. Syst. Pl. 22. 1760; Select. Stirp. Amer. Hist. 147. 1763. —Cadillo. Triumfetta rubricaulis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 342. 1821 [1823]. Shrub or herb to 2 m tall; leaf blades elliptic to broadly ovate, sometimes slightly 3(–5)lobed, indumentum of stellate hairs; petals present, yellow; sepals not cucullate; capsule spines sparsely retrorse-hispid. Open areas, slopes of granitic outcrops, 50–500 m; scattered in Bolívar (east slopes of Cerro Pijiguao, La Unión, upper Río Suapure). Anzoátegui, Apure, Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy; a pantropical weed found in the U.S.A. (Florida, Georgia) and in the Neotropics from Mexico to Argentina and Paraguay, including the West Indies. Triumfetta semitriloba is the most common weedy species in the genus. Taxonomically it has been treated either as a single, variable species or as a complex of closely related species. Our material, with (3)4-locular

Fig. 295. Triumfetta lappula

Vasivaea 361

8. VASIVAEA Baill., Adansonia 10: 193. 1872; Hist. Pl. 4: 195. 1872. Small trees or shrubs; indumentum of stellate hairs. Leaf blades simple, broadly elliptic to ovate-elliptic, sometimes obovate, base deeply to slightly cordate, oblique, margin denticulate or entire, apex acute to rounded; without domatia; stipules subulate-lanceolate, caducous. Inflorescences terminal or axillary, cymosely glomerate or leafy-spicate or racemose; bracts lanceolate, persistent. Flowers unisexual (plants dioecious), 4-merous. Staminate flowers: buds globose; sepals valvate; petals shorter than sepals, somewhat thick-cucullate, glands present basally, basal margins fimbriate; androgynophore annular, stamens ca. 30, filaments very shortly connate, anthers dorsifixed, slightly longer than wide, locules parallel, opening by longitudinal slits, pistil absent. Pistillate flowers: somewhat larger than staminate flowers, buds ellipsoid, calyx and corolla as in staminate flowers, stamens 20–30, well-developed but sterile (i.e., staminodial), exceed-

Fig. 296. Vasivaea alchorneoides

362

T ILIACEAE

ing the gynoecium in length, ovary syncarpous, ovoid, densely appressed-hairy; 4locular with thick walls, ovules 1 per locule, style very short, glabrous; stigma divided into 4 reflexed arms, umbraculiform, the arms fimbriate, divided, rather broadly lobed. Fruit ± globular, briefly rostrate, indehiscent, epicarp thin, mesocarp thick-fibrous. Seeds 4 per fruit, stony. Colombia, Venezuela, Guyana, Peru, Brazil; 2 species, 1 in Venezuela. Vasivaea alchorneoides Baill., Adansonia 10: 194. 1872. —Jiejiema, Palo bolsa. Shrub or small tree to 10 m tall; leaf blades ovate to elliptic, lower surface ± densely covered with minute stellate hairs; fruit short-stalked, obpyriform, densely yellowish brown stellate-hairy. River banks, riparian swampy forests, 50–400 m; Bolívar

(Río Cuyuní in Reserva Forestal Imataca), Amazonas (Caño Majagua, Cerro DuidaHuachamacari, Río Casiquiare, Río Cunucunuma, Río Negro). Guyana, Brazil (Pará, Roraima). ◆Fig. 296. The fruit of Vasivaea alchorneoides is adapted for dispersal by water.

MUNTINGIACEAE by Paul E. Berry Small to medium-sized trees, with pubescence of stellate, long, unbranched, and glandular trichomes. Leaves alternate, distichous, petiolate; blades asymmetrically cordate at the base, palmiveined; margins serrate; stipules lacking, but prophylls basal, heteromorphic. Flowers supra-axillary, solitary or in few-flowered clusters, bisexual, pedicellate, actinomorphic. Sepals (4)5(–7), valvate in bud, with ± spreading tips, basally connate, forming a saucer- to cup-like tube; petals (4)5(–7), free, imbricate, shortly clawed, crumpled in bud, thin, outer margins irregular, longer than the sepals; nectary on a broad disk. Stamens numerous; filaments filiform, free or almost so; anthers 2-thecal, opening by longitudinal slits. Ovary superior or inferior, syncarpous, 5–multilocular, placentation axile-laminar or bilobed and pendulous, many anatropous ovules per carpel; micropyle zigzag; style stout or almost lacking; stigma thick, conical, 5-ridged, or ± capitate. Fruit a berry. Seeds numerous, exotesta mucilaginous, endotesta crystalliferous, exotegmen shortly fibrous; endosperm present, starchy. Neotropics; 3 genera and 3 species, 1 species in Venezuela. 1. MUNTINGIA L., Sp. Pl. 509. 1753. Small to medium-sized trees with stellate pubescence. Leaves shortly petiolate, several-veined from the base, densely white or pale tomentose on the lower surface; prophylls (mistaken for stipules) linear, exceeding the petioles. Inflorescence axillary, pedunculate, 1-flowered, mainly in few-flowered extra-axillary fascicles. Sepals distinct, usually 5, sometimes 6 or 7; petals the same number as the sepals, white. Stamens shorter than the petals, inserted around an annular subperigynous disk. Ovary 5–7-locular, glandular-pilose. Fruit a red globose berry, irregularly many-locular, the stigmas persistent. Seeds minute, embedded in pulp. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 1 species.

Muntingia 363

Muntingia calabura L., Sp. Pl. 509. 1753. —Guacimillo, Majaguillo, Niguo. Tree 3–15 m tall; branches densely glandular-hairy; leaves oblong or oblong-lanceolate, the base oblique and subcordiform; fruit red, edible, very sweet. Semideciduous forests bordering savannas, 100–300 m; Bolívar (northwest of El Manteco, Represa Guri). Northern and central Venezuela, except the Andes; other countries listed under the genus. ◆Fig. 297. The bark of Muntingia calabura furnishes a tough fiber used in basket-making. Mun-

tingia has been placed in either Elaeocarpaceae or Tiliaceae, but molecular studies now place it in its own family, Muntingiaceae (see C. Bayer, M. W. Chase, and M. F. Fay. Taxon 47: 37–42. 1998). It is included here after Tiliaceae, because that is where Cronquist placed the family in his classification system.

Fig. 297. Muntingia calabura

364

T RIGONIACEAE

TRIGONIACEAE by Paul E. Berry Trees, shrubs, or vines. Branchlets terete, lenticellate. Leaves simple, entire, alternate or opposite (in the flora area), usually subcoriaceous, pinnately veined. Stipules interpetiolar, often connate, mostly deciduous. Inflorescences thryses, panicles, or racemes, sometimes reduced to cymes. Flowers bisexual, bibracteolate, obliquely zygomorphic. Sepals 5, connate, imbricate, unequal; petals 5, contorted in bud, the 2 anterior ones forming a keel, often saccate, the posterior or standard petal saccate, the 2 lateral petals spathulate. Stamens 5–12 (up to 4 staminodial), opposite the keel petals, ± united as a tube; anthers 2-locular, longitudinally dehiscent. Disk glands opposite the standard, sometimes laciniate. Ovary superior, (1–) 3(4)-locular; ovules 1–numerous, biseriate; style terminal, simple; stigma capitate. Fruit a septicidal capsule or 3-winged samara. Seed without endosperm; embryo straight; cotyledons plane, thin, with a very short radicle. Neotropics, Madagascar, Malaysia; 3 genera and 27 species, 1 genus and 7 species in the flora area. 1. TRIGONIA Aubl., Hist. Pl. Guiane 387. 1775. Treelets, shrubs, or scandent shrubs. Leaves opposite, petiolate. Inflorescences in simple or compound dichasia, cymes, or cincinni; bracts glandular or eglandular at the margins. Fruit a septicidal capsule, dehiscing from the apex to the base. Seeds 2–several per locule, pubescent. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, northern Paraguay; ca. 25 species, 12 in Venezuela, 7 of these in the flora area. Key to the Species of Trigonia 1. 1. 2(1). 2. 3(2).

3.

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

Leaf blades subrotund, length/width ratio < 3:2 .................... T. ehrendorferi Leaf blades ovate to obovate, elliptic or oblong-elliptic, length/width ratio > 3:2 ........................................................................................................ 2 Lower surface of leaves glabrous .............................................................. 3 Lower surface of leaves pubescent ............................................................ 4 Secondary veins of leaf in 4–7 pairs; pouch on standard petal with a pitcher-like base; stamens 8 or 9, staminodes present; fruit 0.6–3 cm long; seeds villous .................................................................. T. microcarpa Secondary veins of leaf in 9–11 pairs; pouch on standard petal with a globose base; stamens 6 or 7, staminodes absent; fruit 7–9 cm long; seeds spiny ........................................................................................ T. spruceana Inflorescences of thryrses ............................................................ T. reticulata Inflorescences panicles or racemes ........................................................... 5 Upper surface of leaves pubescent ............................................................ 6 Upper surface of leaves glabrous .............................................................. 7 Secondary veins of leaf in 10–16 pairs; flowers in cymules; fruits 3–7 cm long .................................................................................................. T. nivea Secondary veins of leaf in 4–12 pairs; flowers in cincinni; fruits 4.5–11 cm long ................................................................................................ T. villosa

Trigonia 365

7(5). 7. 8(7). 8.

Flowers in cymules ........................................................................... T. sericea Flowers in cincinni ..................................................................................... 8 Stamens 10 or 11; staminodes present; glands often laciniate; style 1– 2 mm long, glabrous; fruit 3–7 cm long; seeds villous .................. T. nivea Stamens 6 or 7, all fertile; glands never laciniate; style 2.8–3 mm long, villous; fruit 7–9 cm long; seeds spiny-pubescent; leaves sometimes becoming glabrous ...................................................................... T. spruceana

Trigonia ehrendorferi Lleras, Acta Amazon. 8: 379. 1978. Liana or shrub; leaf blades subrotund, 3–8 × 2–6 cm, apex mucronulate to emarginate; glands present on sepals. Margins of savannas and forests, 500–600 m, Bolívar (Canaima). Endemic.

Trigonia sericea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 4. 1821. Woody vine or small tree; fruit 3–7 cm long. Seasonally flooded forests, rocky margins of streams, 400–1000 m. Bolívar (base of

Trigonia microcarpa Sagot ex Warm. in Mart., Fl. Bras. 13(2): 131. 1875. Trigonia kaieteurensis Maguire, Bull. Torrey Bot. Club 75: 399. 1948. Liana; flowers greenish white, in groups of 3–6; petals usually 3.5–5 mm long; fruit obovate, 6–10 mm long, as wide as long; seeds 1 or 2(3) per locule. Forested slopes along streams, 50–800 m; Delta Amacuro (near Los Castillos, Río Toro), Bolívar (slopes of Amaruay-tepui, Río Botanamo, Río Toro area, Serranía de Imataca). Colombia, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 298. Trigonia nivea Cambess. in A. St.-Hil., Fl. Bras. Merid. 2: 81. 1829. Woody vine. Venezuela, Brazil, Paraguay; 3 varieties, 1 in Venezuela. T. nivea var. nivea Flowers white; petioles 4–8 mm long; peduncles, when present, to 1.5 mm long; leaves usually oblong to oblong-elliptic with dense, white pubescence; bracts usually as long as the peduncles. Edges of savanna and Mauritia palm stands, dry forests, 50–400 m; Bolívar (10 km east of Ciudad Bolívar, Repressa Guri, near mouth of Río Parguaza, east of Upata), Amazonas (Río Cataniapo). Anzoátegui, Apure, Monagas, Sucre; Guyana, Brazil, Paraguay. ◆Fig. 300. Trigonia reticulata Lleras, Fl. Neotrop. Monogr. 19: 41. 1978. Woody vine; sepals with glands. Edge of montane forests, 1300–1400 m; Bolívar (between Luepa and Cerro Venamo). Guyana.

Fig. 298. Trigonia microcarpa

366

T RIGONIACEAE

Fig. 299. Trigonia villosa var. villosa

Fig. 300. Trigonia nivea var. nivea

Trigonia 367

Fig. 301. Trigonia spruceana

Fig. 302. Trigonia sericea

368

T RIGONIACEAE

Apacará-tepui, Cerro Guaiquinima, Río Carrao, Río Cucurital), Amazonas (Sierra de la Neblina in Cañon Grande of Río Mawarinuma). Zulia; Colombia, Peru, Brazil. ◆Fig. 302. Trigonia spruceana Benth. ex Warm. in Mart., Fl. Bras. 13(2): 130. 1875. Woody vine or scandent shrub; leaves silvery on lower surface; flowers white, centrally yellow. Riparian forests, edge of savannas, 100–200 m; Amazonas (Caño Caname, Caño Pimichín, base of Cerro Moriche, lower Río Atabapo, Río Baria, Río Casiquiare, Río Emoni, Río Guianía, Río Pasimoni, Río Siapa, San Carlos de Río Negro, San Simón de Cucuy). Brazil. ◆Fig. 301.

Trigonia villosa Aubl., Hist. Pl. Guiane 338, pl. 149. 1775. Shrub or woody vine; branches densely golden brown, strigose when young, becoming glabrous with age. Venezuela, Guyana, French Guiana, Brazil; 2 varieties, 1 of these in the flora area. T. villosa var. villosa. —Bejuco negro. Secondary and gallery forests, 50–200 m; northernmost Bolívar (east of Ciudad Bolívar, La Paragua, Represa Guri area, lower Río Caura, Río Marhuanta east of Ciudad Bolívar, San Félix to Upata, Tauca to La Tigrera on Caicara–Ciudad Bolívar road). Anzoátegui, Sucre, Zulia; Guyana, French Guiana, Brazil. ◆Fig. 299.

TRIURIDACEAE by Paul J. M. Maas and Hiltje Maas Erect, saprophytic, monoecious or dioecious herbs. Rhizomes mostly well developed, creeping, subterranean, provided with many small, scale-like leaves. Roots filiform, with or without root-hairs. Stems simple, rarely branched, white, yellow, or purple. Stem leaves 0–8(–15), alternate, small and scale-like, sometimes amplexicaul, their base sometimes slightly saccate (Sciaphila). Inflorescence a terminal, bracteate raceme. Flowers unisexual or bisexual, white, yellow, or purplish, basal part tubular; pedicels distinct, often recurved in fruit. Tepals 3–6(–8), valvate, persistent, often soon reflexed, mostly triangular to deltate, their inner side often papillate (particularly when young), their apex sometimes provided with dense tufts of hairs (bearded) or with a long, tail-like appendage (inflexed in bud). Bisexual flowers: 2–6 stamens and many free ovaries. Staminate flowers: 2–6 mostly epitepalous stamens (except Triuris), free or inserted in a small or large central androphore; anthers dithecal, 4-locular, extrorsely, longitudinally or transversely dehiscent. Pistillate flowers: many free ovaries implanted on a receptacle; ovary unilocular with one basal, anatropous ovule, apical part often papillate; style one, lateral, rarely terminal (Triuris), about as long as the ovary to much exceeding it, stigmatic zone papillate, penicillate-papillate, or glabrous. Fruit an achene or follicle. Seeds globose to obovoid. Tropical (and subtropical) forested parts of the Old and New Worlds; 7 genera and ca. 80 species, 3 genera and 5 species in the flora area. Key to the Genera of Triuridaceae 1.

Plants dioecious; tepals 3, long-caudate; stamens 3, anthers inserted at the base of a fleshy, conical androphore, and alternating with the tepals; style terminal .................................................................... 3. Triuris

Sciaphila 369

1. 2(1). 2.

Plants monoecious (rarely dioecious); tepals 4–6, papillate, often bearded; stamens 2 or 3, anthers epitepalous; style lateral ................ 2 Fruit dehiscent (a follicle); seeds ± spongy on the dorsal side; tepals 4– 6; stamens 2 or 3 ...................................................................... 1. Sciaphila Fruit indehiscent (a thick-walled achene); seeds not spongy; tepals 4; stamens 2 .................................................................................. 2. Soridium

1. SCIAPHILA Blume, Bijdr. 514. 1825. Simple or rarely branched herbs. Rhizomes mostly horizontally creeping and provided with filiform, hairy to glabrous roots. Stems ± terete. Stem leaves 1–15. Inflorescence a terminal raceme, sometimes branched from the base (S. purpurea). Flowers 7–70, unisexual (plants monoecious or rarely dioecious). Tepals 4–6, inner side papillate, apex sometimes bearded. Staminate flowers 2–50, mostly concentrated in the apical part of the inflorescence; stamens 2–6, epipetalous; filaments 0– 0.8 mm long; anthers 4-locular, longitudinally (vertically) to transversely (horizontally) dehiscent; androphore very small and flat (to 1 mm tall) and cylindric. Pistillate flowers 2–35(–55), mostly concentrated in the basal part of the inflorescence; ovaries 10–many, papillate; style lateral, stigmatic zone papillate or penicellatepapillate. Fruit a follicle, ventro-dorsally dehiscent with 2 valves from apex to base. Seeds obovoid, dorsal part of testa spongy. Neotropics and paleotropics with the main center of distibution in IndoMalesia; ca. 50 species, 3 in Venezuela, all in the flora area. Sciaphila corymbosa from the Río Vaupes of Colombia and Brazil, and S. polygyna from Colombia and French Guiana may eventually be found in Venezuela. Key to the Species of Sciaphila 1. 1. 2(1).

2.

Plants dioecious; tepals 4; stamens 2; stems to 21 cm tall, sometimes basally branched; flowers 25–40 (staminate or pistillate), red .... S. rubra Plants monoecious; tepals 4–6; stamens 2 or 3; stems 9–150 cm tall; flowers white or purplish ............................................................................. 2 Stems 9–30 cm tall, simple or basally branched; flowers white, the lower 4–9 pistillate, the upper 10–50 staminate; tepals not bearded ................................................................................................... S. albescens Stems 30–150 cm tall, often repeatedly branched; flowers purplish, the lower 3–34 pistillate, the upper 2–32 staminate; tepals bearded (in young flowers) ........................................................................... S. purpurea

Sciaphila albescens Benth., Hooker’s J. Bot. Kew Gard. Misc. 7: 11. 1855. Sciaphila guianensis Sandwith, Bull. Misc. Inform. Kew 1932: 228. 1932. —Soridium guianense (Sandwith) Giesen in Engl., Pflanzenr. 4. 18: 72, fig. 16, 3–5. 1938. Saprophyte; flowers strongly fragrant; style long exceeding the ovary. Moist lowland to lower montane forests, 100–600 m; northern and eastern Bolívar, Amazonas (Piedra de Cucuy, Río Guainía, San Carlos de Río Negro, Sierra de la Neblina). Panama, Colom-

bia, Guyana, Suriname, French Guiana, Brazil (Amazonas). ◆Fig. 303. Sciaphila purpurea Benth., Hooker’s J. Bot. Kew Gard. Misc. 7: 11. 1855. Saprophyte. Lowland to montane forests, often on termite nests, 100–1200 m; Bolívar (lower slopes of Macizo del Chimantá, Gran Sabana), Amazonas (Cerro Yapacana, Salto Rojas on Río Orinoco, San Carlos de Río Negro, Sierra de la Neblina Sierra Parima). Colombia, Guyana, Peru, Brazil. ◆Fig. 304.

370

T RIURIDACEAE

Sciaphila rubra Maas, Acta Bot. Neerl. 28: 89. 1979. Saprophyte, stems densely covered with leaves. Seasonally flooded forests, 100–200 m; Amazonas (Río Autana). Brazil (Amazonas).

Fig. 303. Sciaphila albescens

This dioecious plant is very close to Sciaphila rubra, which has bisexual flowers and prior to this had only been found in the Ducke Reserve near Manaus, Brazil.

Fig. 304. Sciaphila purpurea

Fig. 305. Soridium spruceanum

Triuris 371

2. SORIDIUM Miers, Trans. Linn. Soc. London 21: 49. 1852. Unbranched or basally branched herbs. Rhizomes horizontally creeping, provided with filiform, densely hairy roots. Stems ± terete. Stem leaves 4–10. Inflorescence a terminal all-sided raceme. Flowers 10–50, unisexual, plants monoecious. Tepals 4, the inner sides papillate. Staminate flowers 7–43, concentrated in the apical part of the inflorescence; stamens 2(3), epitepalous, subsessile; anthers 2-locular when ripe, transversely (horizontally) dehiscent. Pistillate flowers 3–7, concentrated in the basal part of the inflorescence; ovaries 25–40, papillate; style lateral, stigmatic zone penicillate-papillate. Fruit an achene, with rather thick wall. Seeds globose to obovoid, without spongy layer, testa finely reticulate. Guatemala, Belize, Venezuela, Suriname, French Guiana, Brazil; 1 species. Soridium spruceanum Miers, Trans. Linn. Soc. London 21: 50. t. 7, fig. 10–28. 1852. —Sciaphila spruceana (Miers) Engler in Engler & Prantl, Nat. Pflanzenfam. 2, Abt. 1: 238. 1889. Sciaphila brevipes S.F. Blake, Proc. Biol. Soc. Wash. 38: 45. 1925.

Saprophyte; flowers fragrant; anthers 2locular. Evergreen lowland forests, 100–200 m; central and southern Amazonas. Zulia; Guatemala, Belize, Suriname, French Guiana, Brazil (Amazonas, Mato Grosso, Pará). ◆Fig. 305.

3. TRIURIS Miers, Trans. Linn. Soc. London 19: 78. 1845. Unbranched, dioecious herbs. Rhizomes short, cylindric. with filiform, glabrous to sparsely hairy roots. Stems hyaline, terete, flattened and appearing 2winged in herbarium material. Stem leaves 0–2. Inflorescence a 1–4-flowered raceme. Bracts auriculate at the base, amplexicaul. Flowers unisexual. Tepals 3, soon reflexed, glabrous, apex long-caudate, the tails in bud rolled inward like a watch spring. Staminate flowers with 3 stamens alternating with the tepals; filaments absent; androphore large, fleshy, conical to deltoid; anthers inserted at its base. Pistillate flowers with many glabrous ovaries; style terminal, glabrous. Fruit an achene. Seeds globose to ellipsoid. Guatemala, Colombia, Venezuela, Suriname, Brazil; 3 species, 1 in Venezuela. Triuris hexophthalma from the Pakaraima Mountains in Guyana may eventually be found in Venezuela. Triuris hyalina Miers, Trans. Linn. Soc. London 19: 79, t. 19. 1845. Small saprophyte. Moist montane forests, ca. 1200 m; Amazonas (Sierra de la Neblina). Guatemala, Colombia, Guyana, Suriname, Brazil. ◆Fig. 306.

Fig. 306. Triuris hyalina

372

T ROPAEOLACEAE

TROPAEOLACEAE by Paul E. Berry and Luther J. Raechal Annual or perennial herbs, often succulent, often climbing or trailing, the perennial species with a rhizome or tuber. Leaves alternate, peltate or subpeltate, entire to palmately lobed or digitate; stipules small, often caducous, or sometimes lacking; petioles mostly long and twining. Flowers axillary, solitary, showy, usually strongly zygomorphic, often long-pedicellate. Calyx 5-lobed, imbricate, the adaxial 1(3) extended into a nectariferous spur (nearly absent in Trophaeastrum); petals (2)5, distinct, often clawed, entire, serrate, dentate, ciliate, or lobed, the 3 abaxial ones (rarely absent) generally different from the other 2. Stamens 8; filaments distinct; anthers small, basifixed, longitudinally dehiscent. Ovary superior, 3-ribbed, 3-locular; ovule 1 per locule, pendulous; style simple, apical; stigma 3-lobed. Fruit a 3locular schizocarp (samaroid in Magallana). Seeds without endosperm at maturity. Central America, South America; 3 genera and 90 species, 1 species in the flora area. 1. TROPAEOLUM L., Sp. Pl. 345. 1753. Description as in the family, except fruit a schizocarp and the calyx conspicuously spurred. Central America, South America; 87 species, 11 in Venezuela, 1 of these in the flora area. Tropaeolum orinocense P.E. Berry, Novon 2: 182. 1992. Slender vine with orange-red flowers. Rare on dense growth along rivers, 300–400 m; Amazonas (upper Río Orinoco above mouth of Rio Ugueto). Endemic. ◆Fig. 307.

Fig. 307. Tropaeolum orinocense

Piriqueta 373

TURNERACEAE by María Mercedes Arbo Annual or perennial herbs, subshrubs, or shrubs, rarely trees, often with well-developed serial branches; trichomes simple, glandular, or stellate. Roots frequently gemmiferous. Leaves alternate, petioles usually present; blades serrate, crenate, or dentate, rarely subentire; venation pinnate, rarely 1-veined; stipules small or absent. Flowers actinomorphic, bisexual, frequently heterostylous, axillary, usually solitary, sometimes clustered in terminal or lateral inflorescences; peduncle free or adnate to petiole, rarely almost absent; pedicel well-developed and articulate or absent; prophylls 2, opposite or alternate, rarely absent. Sepals 5, the lobes quincuncial in aestivation, basally connate to form a tube or rarely free, deciduous with the corolla and adnate stamens, then leaving an annular scar close to the base; petals 5, contorted in aestivation, alternate with the sepals, clawed, the claw adnate to the calyx tube, rarely free, base of blade naked, provided with a ligule or a membranous corona which is continuous on the sepals and forming an annular structure. Stamens 5, episepalous, inserted at the base of the calyx tube; filaments sometimes fused by their edges with each other or to the petal claws; anthers dorsifixed or ± basifixed, dithecous, with introrse, longitudinal dehiscence. Ovary superior, 3carpellate but 1-locular with parietal placentae; styles 3, rarely bifid to biparted; stigma shortly fimbriate to penicillate; ovules anatropous. Capsules loculicidal, 3valved. Seeds albuminate, 1–5 mm long, exostome usually prominent; chalaza (end of ovule opposite micropyle) sometimes prominent; raphe linear; episperm reticulate or striate, rarely crested; aril fleshy, whitish, membranous when dry, rarely divided into hairs. Tropics and subtropics of both Old and New Worlds; 10 genera and ca. 170 species, 2 genera and 21 species in the flora area. Key to the Genera of Turneraceae 1.

1.

Trichomes of stems and leaves porrect-stellate (stellate with the central arm longer) and some simple, in some cases setiform-glandular trichomes (spreading, pluricellular, with swollen base and small head) also present; leaves generally without nectaries; floral peduncles free; pedicels present and well developed; flowers with a membranous, usually fimbriate, annular corona inserted on the throat ............ 1. Piriqueta Trichomes simple and nonglandular-setiform, without porrect-stellate hairs; leaves often with nectaries; floral peduncles partially or totally adnate to petioles, rarely free or absent; pedicels usually absent; flowers without a corona ................................................................... 2. Turnera

1. PIRIQUETA Aubl., Hist. Pl. Guiane 298. 1775. Annual or perennial herbs with simple, porrect-stellate and often setiformglandular hairs. Leaves petiolate, sometimes sessile, usually without nectaries; stipules reduced to glandular prominences or absent. Inflorescences axillary, 1– many-flowered, rarely in a leafless terminal raceme. Flowers often heterostylous; floral peduncle free; pedicel articulate, well-developed; prophylls inconspicuous or absent, located either at the articulation or at the apical portion of peduncle. Sepals

374

T URNERACEAE

basally connate to form a funnel-shaped to campanulate tube; petals yellow, pink, or white, sometimes with a dark spot at base; claw fused to calyx tube; corona membranous, annular, with the superior margin fimbriate to lacerate, inserted in the throat on the base of petal blades and on sepals. Staminal filaments inserted at the base of the calyx tube, glabrous; anthers dorsifixed, ovate, basally emarginate, the apex usually bent after dehiscence. Ovary pilose, rarely glabrous; style sometimes bifid; stigma penicillate. Capsule globose to ellipsoid, outer surface smooth to verrucose. Seeds obovoid, straight to slightly bent; episperm reticulate, areoles hexagonal to transrectangular; aril unilateral or enveloping the seed. Southeastern U.S.A., Mexico, Central America, West Indies, South America (except Chile and Uruguay), Africa; 26 species, 5 in Venezuela, 3 of these in the flora area. Key to the Species of Piriqueta 1.

1. 2(1).

2.

Plants without setiform-glandular hairs; fruits smooth; leaves linear, elliptic, or ovate, 16–70 × 1–20 mm; indument lax to tomentose; flowers yellow, sometimes whitish ......................................................... P. cistoides Plants with setiform-glandular hairs; stems and hairs sticky; fruits rugose; flowers yellow, orange, pink, or pinkish yellow ............................ 2 Leaves ovate, obtuse, rarely acute, 20–65 × 10–32 mm, with undulatecrenate margins; hairs stellate, many-rayed; flowers large, 12–22 mm long, pinkish yellow, heterostylous .......................................... P. undulata Leaves ovate or elliptic, 13–40 × 5–15 mm, acute, with serrate margins; hairs few-rayed; flowers small, 5–9 mm long, homostylous ....... P. viscosa

Piriqueta cistoides (L.) Griseb., Fl. Brit. W. I. 297. 1860. —Turnera cistoides L., Sp. Pl. ed 2, 1: 387. 1762. —Celedonia. Erect, simple or branched herb, 10–80 cm tall; flowers bright yellow, sometimes whitish. Savannas, edges of rivers or lakes, usually a pioneer plant in disturbed, open places on sandy or rocky soils, 50–300 m. Southeastern U.S.A., Mexico, Central America, West Indies, South America to northern Argentina and southern Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of P. cistoides 1. Flowers heterostylous, (8–)11–20 mm long, the difference in length between androecium and gynoecium is 2–5 mm in brevistylous flowers, 1.5–5 mm in longistylous flowers; plants frequently perennial; leaves elliptic or linear with margin often revolute ................ subsp. caroliniana 1. Flowers homostylous, small, 4–11 mm long, sometimes the androecium to 1.5 mm longer than the gynoecium or the latter slightly longer (to 1 mm longer); annual plants; leaves ovate, elliptic to linear, usually with a flat margin ........... ................................... subsp. cistoides

P. cistoides subsp. caroliniana (Walter) Arbo, Ann. Missouri Bot. Gard. 77: 351. 1990. —Waltheria caroliniana Walter, Fl. Carol. 17. 1788. —Piriqueta caroliniana (Walter) Urb., Jahrb. Königl. Bot. Gart. Berlin 2: 71. 1883. Piriqueta caroliniana var. integrifolia Urb., Jahrb. Königl. Bot. Gart. Berlin 2: 72. 1883. Bolívar (Río Cuchivero basin), Amazonas (La Esmeralda, near Puerto Ayacucho, Río Coro Coro basin). Guárico, Portuguesa, Zulia; southeastern U.S.A., West Indies, Colombia, Brazil, Bolivia. P. cistoides subsp. cistoides Piriqueta villosa Aubl., Hist. Pl. Guiane 298, t. 117. 1775. Piriqueta tomentosa Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 128. 1823. Piriqueta lanceolata Benth., J. Bot. (Hooker) 4: 117. 1842. Piriqueta foliosa Garcke, Linnaea 27: 63. 1849. Piriqueta cistoides var. genuina Urb., Jahrb. Königl. Bot. Gart. Berlin 2: 73. 1883. Turnera tomentosa Willd. ex Schult. in

Piriqueta 375

Fig. 308. Piriqueta viscosa subsp. viscosa

Fig. 309. Piriqueta cistoides subsp. cistoides

Fig. 310. Piriqueta undulata

376

T URNERACEAE

Roem. & Schult., Syst. Veg. 6: 678. 1820. Turnera aturensis D. Dietr., Syn. Pl. 2: 1008. 1840. Common in Bolívar (lower basins of Río Caroní and Parguaza, Río Cuyuní basin), Amazonas (near Puerto Ayacucho, Río Atabapo basin). Apure; Mexico, Central America, West Indies, South America south to southern Brazil, Bolivia, Paraguay, northern Argentina. ◆Fig. 309. Piriqueta undulata Urb., Repert. Spec. Nov. Regni Veg. 13: 154. 1914. Herb or subshrub 20–90 cm tall, with a xylopodium (thickened woody rootstock); stems and leaves somewhat sticky; flowers pinkish yellow. Sandy savannas, 50–400 m; Bolívar (Canaima, Río Parguaza basin), Amazonas (near Puerto Ayacucho, lower Río Ventuari basin). Anzoátegui, Guárico. ◆Fig. 310. Piriqueta viscosa Griseb., Cat. Pl. Cub. 114. 1866. Herb or shrub, 20–90 cm tall; stems and leaves sticky; flowers small, homostylous, to 10 mm long. Central America, West Indies, Colombia, Venezuela, Guyana, Suriname,

French Guiana, Brazil, Bolivia; 2 subspecies, both in the flora area. Key to the Subspecies of P. viscosa 1. Flowers yellow or orange; seeds short, 1.5–1.6 × 0.9–1 mm; episperm with large areoles .................... subsp. tovarensis 1. Flowers pink, with or without a purple spot on the throat; seeds long, 1.6–1.9 × 0.6–0.7 mm; episperm with small areoles ..................................... subsp. viscosa P. viscosa subsp. tovarensis Urb., Jahrb. Königl. Bot. Gart. Berlin 2: 69. 1883. Lowland to upland savannas and Mauritia palm swamps, 300–1000 m; Bolívar (Altiplanicie de Nuria, Gran Sabana). Aragua; Costa Rica, Panama, Colombia. P. viscosa subsp. viscosa Lowland to upland savannas, 100–800 m; Bolívar (basins of Río Caroní and Río Cuyuní), Amazonas (near Puerto Ayacucho, lower Río Ventuari). Sucre, Trujillo; Central America, West Indies, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 308.

2. TURNERA Plum. ex L., Gen. Pl. 87. 1737. Annual or perennial herbs, shrubs, or rarely trees; trichomes simple, stellate, or, when glandular, usually sessile. Leaves usually entire, sometimes pinnatifid, often with nectaries; stipules well-developed or sometimes absent. Flowers frequently heterostylous, usually solitary, sometimes clumped in lateral or terminal inflorescences; peduncle free or adnate to petiole, sometimes absent; pedicels usually absent, rarely developed; prophylls conspicuous. Sepals basally connate to form a cylindric to funnel-shaped calyx tube; petals usually yellow, rarely white, red, or purplish, sometimes with a yellow or purple spot at the base. Staminal filaments sometimes connate or adnate from the base of the calyx tube to the petal claws up to the throat of the tube and then forming a nectariferous pocket between each of the 5 sepals and the corresponding filament; anthers dorsifixed or ± basifixed. Ovary usually pilose; stigmas penicillate, sometimes shortly lobulate. Capsules smooth or verrucose. Seeds usually reticulate, sometimes longitudinally striate, exceptionally crested; aril unilateral, lobate or lacerate. Neotropics, Africa, some introduced and weedy in Asia and Oceania; 102 species, 22 in Venezuela, 18 of these in the flora area. The species of Turnera from the flora area belong in six of the nine series of the genus. Series Canaligerae, the most advanced in the genus, is represented by just one species (T. scabra). The series Salicifoliae, Annulares, and Capitatae include two species each. Series Leiocarpae is represented by six species, three of which are apparently endemic. Series Stenodictyae is represented by five species, two of which are endemics.

Turnera 377

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

3(2).

3. 4(2).

4.

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

6.

7(6).

7.

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

Floral peduncle absent or nearly so; flowers yellow, small, grouped in axillary or terminal capituliform inflorescences; fruits rugose ............... 2 Floral peduncle well developed; flowers solitary, sometimes grouped at branch apices; fruits rugose or smooth ................................................. 5 Upper surface of leaves glabrous with the exception of the middle vein; nectaries minute, marginal; calyx 2–6 mm long; seeds pilose ............ 3 Upper surface of leaves pilose; nectaries circular, 0.5 mm diameter, located between the petiole and the blade or on the margins of the blade base; calyx 6–8 mm long; seeds glabrous ............................................. 4 Leaves linear or narrowly lanceolate, 8–15 cm long, the apex acuminate; secondary veins with a divergence angle of ca. 70°; calyx 4–6 mm long ....................................................................................................... T. venosa Leaves elliptic, 4–8 cm long, the apex obtuse; secondary veins with a divergence angle of ca. 52°; calyx 2–3 mm long .................... T. steyermarkii Petioles 8–12 mm long; leaves 30–80 × 9–20 mm, tomentose, lanceolate, the base attenuate; ovary 5–12-ovulate; stigmas multibranched ....................................................................................... T. schomburgkiana Petioles 4–7 mm long; leaves 20–80 × 12–40 mm, tomentose-velutinous, ovate or ovate-elliptic, the base obtuse, sometimes cuneate; ovary 3- or 4-ovulate; stigmas few-branched ...................................... T. waltherioides Floral peduncle free; corolla yellow, orange, or reddish; styles pilose or hirsute at medial or basal zone; fruits rugose ...................................... 6 Floral peduncle adnate to petiole; flowers yellow; styles pilose or glabrous; fruits smooth or rugose ............................................................ 12 Leaves without nectaries, glabrous; flowers large, 25–30 mm long, yellow; calyx 25–28 mm long; prophylls 2, heart-shaped; staminal filaments adnate to calyx tube base and connate by their edges at various heights ...................................................................................... T. annectens Leaves bearing nectaries in petiole or blade, pilose at least on lower surface; flowers large or small, 8–27 mm long, yellow, orange, or reddish; calyx 7–24 mm long; prophylls not heart-shaped; staminal filaments not connate by their edges .................................................................... 7 Leaves usually large, to 17 cm long, without glandular hairs; floral peduncle 0.5–13 mm long; calyx 7–24 mm long, seeds obovoid, 2.5–4 mm long, longer than wide, with conspicuous longitudinal and slender transverse stripes .................................................................................. 8 Leaves smaller, to 8 cm long, with glandular hairs; floral peduncle short, 0.5–3 mm long, calyx 8–12 mm long; seeds subglobose, 1.3–1.8 mm long, almost as long as wide, reticulate .............................................. 11 Stipules minute, triangular, 0.3–0.6 mm long; floral prophylls lanceolate to ovate lanceolate, 2–9 mm wide .................................................. T. acuta Stipules small, 0.7–3 mm long; floral prophylls subulate, linear, or lanceolate, 0.3–3 mm wide ......................................................................... 9 Foliar nectaries with markedly pilose margins; foliar scars conspicuous or not; calyx 7–10 mm long; staminal filaments puberulous .............................................................................................. T. macrophylla

378

T URNERACEAE

9. 10(9).

10.

11(7). 11. 12(5). 12. 13(12).

13. 14(13).

14.

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

16. 17(16). 17.

Foliar nectaries with glabrous margins; foliar scars conspicuous; calyx 8– 23 mm long; staminal filaments puberulous or glabrous .................. 10 Stipules 0.6–0.9 mm long; leaves with 14–20 minute, sessile nectaries close to the margin indentations and sometimes at the base; calyx 17–23 mm long; corolla red-orange; staminal filaments glabrous ..................................................................................................... T. castilloi Stipules 1.5–3 mm long; leaves with a pair of nectaries, disk-shaped in young leaves; calyx 8–14 mm long; corolla yellow, sometimes whitish; staminal filaments puberulous .............................................. T. cicatricosa Leaves usually tomentose on lower surface, with both simple and glandular-capitate, sessile, yellow trichomes ........................................ T. odorata Leaves glabrescent on the lower surface; pubescence of simple and glandular, stipitate trichomes, especially on young leaves ........... T. breviflora Calyx totally glabrous, ovary and fruit glabrous or with a few hairs at apex; plant grass-like ............................................................. T. guianensis Calyx with at least some trichomes; ovary and fruit pilose or hirsute; plant not grass-like .............................................................................. 13 Calyx tube glabrous, the lobules pilose; ovary and fruit glabrous except for a few hairs at apex; seeds with prominent reticule nodes; chalaza conical, prominent ...................................................................... T. pumilea Calyx tube pilose; ovary and fruits pilose; seeds without prominent reticule nodes; chalaza rounded ................................................................ 14 Flowers 15–35 mm long; corolla conspicuously longer than the calyx; staminal filaments marginally adnate to petal claws, making 5 nectariferous pockets between the stamens and the calyx tube; prophylls inserted at the base of the calyx; fruits rugose ........................... T. scabra Flowers 6–9 mm long, corolla as long as the calyx or somewhat longer; staminal filaments adnate only basally to calyx tube; prophylls inserted at receptacle base; fruits smooth ............................................. 15 Calyx tube glabrous except for some trichomes on the distal part of the veins; nectaries with glabrous margins .................................... T. paruana Calyx tube pilose, rarely glabrescent at base; nectaries with a hairy border ......................................................................................................... 16 Leaves elliptic, obovate, or ovate, grayish green, 4–15 mm wide, length:width ratio 1.5–5(–7.5):1; margins serrate or serrulate, rarely crenate ........................................................................................... T. lineata Leaves linear or narrowly elliptic, 2–5 mm wide, length:width ratio 7– 10:1; margins entire or serrulate ........................................................ 17 Leaves silvery-green, tomentose-sericeous, obtuse at the apex, margins entire .......................................................................................... T. argentea Leaves plain green, acute at the apex, serrulate, with short and curly trichomes mixed with longer, antrorse ones ..................................... T. huberi

Turnera acuta Willd. ex Schult. in Roem. & Schult., Syst. Veg. 6: 678. 1820. —Guanaguanare. Turnera carpinifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 125. 1823. Turnera pilosula Urb., Repert. Spec. Nov.

Regni Veg. 13: 155. 1914. Shrub 0.5–2 m tall with reclining branches; leaves chartaceous, 2–10 × 1–4 cm; flowers reddish orange. Secondary forests, forest edges, rocky outcrops, 50–800 m; Bolívar (basins of Río Caroní and Río Paragua), Ama-

Turnera 379

zonas (basins of Río Casiquíare and Río Negro, upper Río Orinoco, Río Sipapo). Amazonian Colombia, Peru, and Brazil. ◆Fig. 313. Turnera annectens Arbo, Ann. Missouri Bot. Gard. 77: 340. 1990. Shrub to 50 cm tall; leaves lanceolate, elliptic, or obovate, 3–6 × 1–2.5 cm. Tepui meadows and shrublands, 1400–1900 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 316. Turnera argentea Arbo, Ann. Missouri Bot. Gard. 77: 342. 1990. Shrub to 1 m tall; leaves 10–27 × 2–10 mm; flowers yellow. Open savannas, ca. 100 m; Amazonas (near the confluence of Río Orinoco and Río Ventuari). Endemic. Turnera breviflora Moura, Phyton (Horn) 12: 267. 1970. Subshrub; flowers yellow; leaves 20–56 × 11–26 mm. Riparian forests, 100–200 m; Bolívar (lower Río Paragua). Aragua, Carabobo, Distrito Federal, Mérida, Miranda; Guyana, Amazonian Brazil. Turnera castilloi Arbo, Ann. Missouri Bot. Gard. 77: 344. 1990. Shrub 1–3 m tall with supporting branches; leaves 1.5–15 × 0.6–4 cm, elliptic or ovate. Riparian forests, 50–100 m; Amazonas (Río Cataniapo basin). Endemic. Turnera cicatricosa Arbo, Ann. Missouri Bot. Gard. 77: 346. 1990. Decumbent shrub 1.5–3 m tall; leaves elliptic, lanceolate, or sometimes obovate, 1–8.5 × 0.6–3 cm. Forest edges, savannas, riparian forests, 100–1100 m; Bolívar (upper Río Caroní basin), Amazonas (near San Fernando de Atabapo). Guyana. ◆Fig. 318. Turnera guianensis Aubl., Hist. Pl. Guiane 291, t. 114. 1775. —Sauce. Turnera humboldtii Spreng., Syst. Veg. 1: 941. 1825. Herb 10–70 cm tall; fruits smooth. Wet or humid savannas, 50–300 m; Bolívar (near Río Parguaza), Amazonas (near Puerto Ayacucho, Río Manapíare basin). Anzoátegui, Apure, Guárico, Monagas; Amazonian Colombia, Trinidad, Guyana, northern Brazil. ◆Fig. 311.

Turnera huberi Arbo, Ann. Missouri Bot. Gard. 77: 347. 1990. Herb or subshrub 15–40 cm tall; leaves 17–57 mm long; flowers yellow or sometimes whitish. Sandy savannas, secondary vegetation on sandy or rocky soils, 50–100 m; Amazonas (Cerro Moriche, near Puerto Ayacucho). Endemic. Turnera lineata Urb., Repert. Spec. Nov. Regni Veg. 13: 157. 1914. Turnera ulmifolia f. serissima Killip, J. Wash. Acad. Sci. 24: 51. 1954. Herb or subshrub 20–50 cm tall; flowers yellow. Humid savannas with sandy or rocky soils, 50–1100 m; common in Bolívar and Amazonas. Eastern Colombia, Guyana, Amazonian Brazil. ◆Fig. 312. Some specimens of Turnera lineata from Amazonas are atypical in having lax indument with androecium and gynoecium of similar length; other specimens have relatively narrower leaves. Turnera macrophylla Urb., Jahrb. Königl. Bot. Gart. Berlin 2: 95. 1883. Turnera muricata Rusby, Mem. New York Bot. Gard. 7: 308. 1927. Shrub 1–2 m tall; leaves 3–16 × 1–6 cm; flowers yellow-orange. Riparian vegetation, 100–200 m; Amazonas (near Cerro Morrocoy). Peru, Amazonian Brazil, Bolivia. ◆Fig. 315. Turnera odorata Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. —Clavo de especie amarilla, Clavo de especie de sabana. Turnera corchorifolia Willd. ex Schult. in Roem. & Schult., Syst. Veg. 6: 678. 1820. Turnera tomentosa Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 125. 1823. Turnera hexandra Spreng., Syst. Veg. 4(cur. post.): 124. 1827. Turnera parviflora Benth., J. Bot. (Hooker) 4: 16. 1842. Thin shrub 1–4 m tall; leaves aromatic, 3– 8 × 1.5–3 cm; flowers yellow, fragrant. Savannas, forests on granitic outcrops, forest margins, 50–400 m; Bolívar (basins of lower Río Caura and Río Paragua), Amazonas (near Puerto Ayacucho, lower Río Ventuari basin). Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 321.

380

T URNERACEAE

Turnera paruana Arbo, Ann. Missouri Bot. Gard. 77: 348. 1990. Shrub to 50 cm tall; leaves brownish red when dry, 15–30 × 6–11 mm; seeds 1.8 mm long. Upland tepui savannas, 700–800 m; Amazonas (Cerro Parú). Endemic.

Turnera venosa Urb., Bot. Jahrb. Syst. 25(Beibl. 40): 3. 1898. Shrub to 2 m tall; leaves subcoriaceous, discolorous. Edges of black-water rivers, ca. 100 m; Amazonas (Río Temi). Amazonian Brazil. ◆Fig. 319.

Turnera pumilea L., Syst. Nat. ed. 10, 2: 965. 1759. —Conejito. Herb 5–30 cm tall; branches spreading; leaves serrate, 20–50 × 5–20 mm, clumped rosette-like at the apex of the branches; flowers small, 5–7 mm long; fruits smooth. Savannas, shrubby slopes, 300–400 m; Bolívar (Guacaima near Upata). Anzoátegui, Aragua, Guárico, Monagas, Zulia; Mexico, Central America, West Indies, South America south to northern Argentina. ◆Fig. 322.

Turnera waltherioides Urb., Repert. Spec. Nov. Regni Veg. 13: 158. 1914. Shrub 1–1.5 m tall; flowers 6–7 mm long; leaves with deeply impressed veins on the lower side. Upland gallery forests, tepui slopes, 1200–1400 m; Bolívar (slopes of Auyán-tepui and Uaipán-tepui, Gran Sabana). Guyana (Kanuku Mountains), Brazil (Mount Roraima). ◆Fig. 320.

Turnera scabra Millsp., Publ. Field. Columbian Mus., Bot. Ser. 2: 77. 1900. —Malvavisco amarillo, Sereno. Turnera ulmifolia var. intermedia Urb., Jahrb. Königl. Bot. Gart. Berlin, 2: 140. 1883. Subshrub or shrub 0.2–1 m tall; leaves ovate to elliptic, 2–14 × 1–4 cm; corolla showy, golden yellow. Savannas, forest edges, secondary vegetation, always in sunny places, 50–300 m; Bolívar (lower basins of Río Caroní, Río Caura, Río Cuyuní, and Río Paragua), Amazonas (near Puerto Ayacucho). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Distrito Federal; Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, northern Brazil. ◆Fig. 314. Turnera schomburgkiana Urb., Jahrb. Königl. Bot. Gart. Berlin 2: 132. 1883. Shrub to 60 cm tall; flowers 5–8 mm long. Edges of rivers in forested areas, 400–500 m; Bolívar (Macizo del Chimantá). Guyana (Mount Roraima). ◆Fig. 317. Turnera steyermarkii Arbo, Ann. Missouri Bot. Gard. 77: 349. 1990. Branched shrub 1–2 m tall; leaves coriaceous; seeds striate, 3 mm long. Seasonally flooded white-sand savannas and caatinga forests along black-water rivers, 100–200 m; Amazonas (Caño San Miguel, Caño Yagua, Río Guasacavi, Río Temi). Brazil (northern Amazonas).

Fig. 311. Turnera guianensis

Turnera 381

Fig. 312. Turnera lineata

Fig. 313. Turnera acuta

Fig. 314. Turnera scabra

382

T URNERACEAE

Fig. 315. Turnera macrophylla

Fig. 316. Turnera annectens

Turnera 383

Fig. 317. Turnera schomburgkiana

Fig. 319. Turnera venosa

Fig. 318. Turnera cicatricosa

384

T URNERACEAE

Fig. 320. Turnera waltherioides

Fig. 322. Turnera pumilea

Fig. 321. Turnera odorata

Typha 385

TYPHACEAE by Luther J. Raechal Perennial, semiaquatic herbs with starchy rhizomes, often colonial. Flowering stems elongate. Leaves alternate, elongate, flat or planoconvex, with sheathing base and narrow, parallel-veined, spongy blades. Flowers unisexual (plants monoecious), crowded into a dense, elongate-cylindric, terminal spike, the staminate ones above, the pistillate below, the 2 portions of the spike contiguous or separated by a distinct articulation. Perianth of capillary bristles or slender scales. Stamens (1–)3(–8); filaments short, distinct or connate; anthers basifixed, dithecal, with a broad connective prolonged beyond the thecae; pollen in monads or tetrads. Fertile ovary elevated on a gynophore, 1-carpellate with 1 ovule; style terminal, persistent and elongating in fruit; stigma shortly decurrent, linear to spatulate. Fruit an achene borne on an elongated gynophore, subtended by capillary bristles. Seeds with endosperm. Cosmopolitan; 1 genus and 10 species, 1 species in the flora area. 1. TYPHA L., Sp. Pl. 971. 1753. Description and distribution as for family; 10 species, 1 in Venezuela. Typha domingensis Pers., Syn. Pl. 2: 532. 1807. —Enea. Staminate and pistillate spikes usually separated; pistillate spikes pale brown; leaves nearly flat. Marshes and edges of ponds and rivers, near sea level to 100 m; Delta Amacuro (mouth of Caño Güiniquina, between Punta Araguabisi and Punta Baja), Bolívar (Tumeremo–Bochinche road). Widespread elsewhere in Venezuela at low elevations, but to 2250 m in Táchira; cosmopolitan. ◆Fig. 323.

Fig. 323. Typha domingensis

386

U LMACEAE

ULMACEAE by James S. Miller and Paul E. Berry Trees or shrubs. Leaves alternate or rarely opposite, often distichous, simple, often unequal-sided; stipules paired, caducous. Inflorescences axillary, cymose, or the flowers solitary in the leaf axils. Flowers small, actinomorphic, hypogynous, unisexual (the plants monoecious), or sometimes bisexual. Sepals (2–)5(–9), free or connate and ± campanulate, imbricate, persistent. Petals absent. Stamens the same number as or more than the calyx lobes and opposite them; filaments free or basally adnate to the sepals; anthers bithecal, dehiscing longitudinally; pollen 2–numerous-porate or -colpate. Ovary superior, 2(3)-carpellate, unilocular or rarely bilocular; styles 2, free and divergent; stigmas decurrent; ovule solitary, pendulous, anatropous or amphitropous. Fruits samaroid, drupaceous, or a nut. Seeds with little or no endosperm. Widespread in temperate and tropical regions of both hemispheres; 16–18 genera and ca. 150 species, 3 genera and 4 species in the flora area. Key to the Genera of Ulmaceae 1. 1. 2(1). 2.

Plants usually armed with spines, lianas or scandent shrubs; pistillate flowers usually solitary; stigmas with 4 ultimate divisions ......... 2. Celtis Plants lacking spines, upright shrubs or trees; pistillate flowers in cymose inflorescences; stigmas with 2 ultimate divisions ...................... 2 Leaf blades entire, coriaceous, the venation mainly pinnate; fruit > 1 cm long ........................................................................................ 1. Ampelocera Leaf blades entire to serrulate, chartaceous, palmately veined; fruits < 1 cm long ........................................................................................ 3. Trema

1. AMPELOCERA Klotsch, Linnaea 20: 541. 1847. Medium to large trees. Leaves with bases nearly symmetric to slightly asymmetric; margins entire or coarsely dentate; venation pinnate or palmate. Inflorescence sparsely to densely branched, cymose or fasciculate, often with bisexual flowers toward the apex and functionally staminate flowers near the base. Tepals 4 or 5, usually basally connate. Stamens (3)8(16); anther connective prolonged beyond apex of the anther. Style branches 2, free to the base; stigma unbranched. Fruit a fleshy, globose to compressed drupe, the stigma asymmetrically placed. Seed globose. Mexico, Belize, Honduras, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 10 species, 1 in Venezuela. Ampelocera edentula Kuhlm., Anais Reunião Sul-Amer. Bot. 3(1): 75. 1938. —Juanasi (Yekwana). Tree to 15 m tall; leaves elliptic to subrhomboid, 4–20 × 2–9 cm, the petioles 5– 12 mm long and irregularly peeling; fruit yellowish, 1–1.5 cm long. Evergreen lowland

forests, riparian forests, 100–300 m; Bolívar (Río Nichare basin), Amazonas (Río Mawarinuma, between Yavita and Maroa). Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 324.

Celtis 387

Fig. 324. Ampelocera edentula

2. CELTIS L., Sp. Pl. 1043. 1753. Trees or shrubs, occasionally scrambling and becoming a liana, sometimes with recurved spines. Leaves alternate, distichous, entire or serrate, often unequal at the base; stipules lateral. Inflorescences axillary, of two types, the staminate usually cymose, the pistillate flowers solitary or in few-flowered fascicles. Flowers small, pedicellate, unisexual or bisexual (plants monoecious or polygamo-monoe-

388

U LMACEAE

cious). Sepals (4)5, imbricate, basally connate; petals absent. Stamens (4)5, opposite the sepals, inserted on the margin of a distinct disk, absent in pistillate flowers; anthers bithecal, dehiscing longitudinally. Ovary ovoid, sessile; stigmatic surface on the inner face of the two divergent styles. Fruits drupaceous, globose or ovoid. Temperate and tropical regions of both hemispheres; 60–75 species, 3 in Venezuela, 1 of these in the flora area.

Fig. 325. Celtis iguanaea

Trema 389

Celtis iguanaea (Jacq.) Sarg., Silva 7: 64. 1895. —Rhamnus iguanaeus Jacq., Enum. Syst. Pl. 16. 1760. —Maiz tostado. Scrambling shrub, sometimes becoming lianoid, or a tree to 5(–12) m tall; flowers small, green; fruits yellow, orange, or red. Disturbed forests and forest edges, riparian

forests, 0–400 m; Delta Amacuro (Caño JotaSabuca), Bolívar (Altiplanicie de Nuria, Hato La Vergareña), Amazonas (Puerto Ayacucho, Río Orinoco). Anzoátegui, Apure, Aragua, Carabobo, Distrito Federal, Lara, Mérida, Sucre, Táchira, Zulia; widespread in Mexico, Central America, West Indies, south to Argentina. ◆Fig. 325.

3. TREMA Lour., Fl. Cochinch. 562. 1790. Sponia Comm. ex Lam., Encycl. 4: 139. 1797. Trees or shrubs, the stems lacking spines. Leaves alternate, distichous, serrate or rarely entire; stipules lateral, free, often caducous. Inflorescences axillary, cymose, fasciculate, or the flowers solitary. Flowers small, bisexual or more often unisexual (plants monoecious or polygamo-dioecious). Sepals (4)5, basally connate; petals absent. Stamens (4)5, opposite the sepals; usually lacking totally in pistillate flowers; filaments basally adnate to the sepals, short; anthers oblong, bithecal, dehiscing longitudinally. Ovary sessile, reduced or absent in staminate flowers, unilocular; styles 2, persistent, basally connate; stigmas linear. Fruit drupaceous, ovoid to subglobose, capped by the persistent styles; seeds with carnose or scant endosperm. Tropical and subtropical parts of both hemispheres; ca. 20 species, 4 in Venezuela, 2 of these in the flora area. The species in this genus are extremely variable.

Fig. 326. Trema integerrima

390

U LMACEAE

Fig. 327. Trema micrantha

Key to the Species of Trema 1. 1.

Leaf margin entire; leaves not scabrous on the upper surface; fruits yellow to orange ......................................................................... T. integerrima Leaf margin serrate; leaves scabrous on the upper surface; fruits orange to red ....................................................................................... T. micrantha

Trema integerrima (Beurl.) Standl., Contr. Arnold Arbor. 5: 55. 1933. —Sponia integerrima Beurl., Kongl. Vetensk. Acad. Handl. 1854: 144. 1856. —Hubokawa (Yanamano), Mochochito, Yaraboba. Tree to 30 m tall; flowers small, green; fruits yellow to orange. Secondary and primary lowland to lower montane moist forests, 50–600 m; Amazonas (La Mantequilla, Ocamo). Barinas, Mérida, Zulia; Panama, Colombia, Guyana, Suriname, Peru, northern Brazil. ◆Fig. 326.

Trema micrantha (L.) Blume, Ann. Mus. Bot. Lugduno-Batarum 2: 58. 1853. —Rhamnus micranthus L., Syst. Nat. ed. 10. 2: 937. 1759. —Boboyo winae (Guahiba), Cuweyoja (Ye’kwana), Guámari (Baniba). Shrub or tree to 5(–20) m; flowers small, green; fruits red to orange. Secondary vegetation, thickets, gallery and riparian forests, occasional in moist and tall forests on white sand, near sea level to 900 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; widespread in the Neotropics from Mexico to Argentina. ◆Fig. 327.

U RTICACEAE 391

URTICACEAE by Amy Pool Herbs, subshrubs, or rarely small, soft-wooded trees or lianas, stinging hairs often present, sap usually watery. Leaves alternate or opposite, simple, often with calcium carbonate cystoliths; blades mostly ovate to lanceolate; margins entire, dentate, serrate, or rarely incised; venation usually palmate, sometimes pinnate; petiolate; stipules usually present. Inflorescences axillary, basically cymose; bracteate. Flowers usually unisexual (plant monoecious, dioecious, or polygamous). Staminate flowers: tepals (3)4 or 5(6), equal, valvate, usually green or white; stamens usually as many as tepals and opposite; filaments inflexed in bud, pollen dispersal explosive upon release of stamens from tepals; anthers 2-locular, dehiscence longitudinal; rudimentay ovary usually present. Pistillate flowers: tepals none or 3–5 and distinct or variously connate into a tube; sometimes with prominent staminodes; ovary 1-locular; ovule 1, basal; style 1; stigma penicillate, linear, or filiform. Fruit an achene, often partially or entirely enclosed in an accrescent perianth which sometimes becomes fleshy and berry-like. Seed 1; endosperm various or absent; embryo straight. Tropics, subtropics, and some temperate areas in both the New World and Old World; 40–45 genera and ca. 800 species, 6 genera and 13 species in the flora area. Species distribution is based upon specimens seen. As most species of Urticaceae are rarely collected, they perhaps may be found in additional localities. Key to the Genera of Urticaceae 1. 1. 2(1).

2.

3(1).

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

Leaves opposite (upper leaves sometimes alternate in Boehmeria) ........ 2 Leaves alternate ........................................................................................ 3 Stems 1–2 m; leaves toothed; inflorescence an elongated spike with widely separated clusters of flowers (often with reduced leaves at tips of spikes); pistillate flowers with perianth fused into sturdy tube enclosing ovary and later the fruit, ovate in flower to circular in fruit ................................................................................................. 1. Boehmeria Stems ≤ 0.5 m; leaves toothed or entire; inflorescences in loosely branched panicles or in short cymose clusters in axils of leaves; pistillate flowers with perianth of 3 parts, 1 larger than the other 2 and somewhat hooded ............................................................................. 4. Pilea Leaves entire; pistillate flowers with perianth fused into thin tube enclosing ovary and later fruit, elliptic, ripping at maturity to reveal fruit .................................................................................................. 5. Pouzolzia Leaves toothed; pistillate flowers with perianth absent or present and not fused into tube ....................................................................................... 4 Inflorescences in dense sessile glomerules; pistillate flowers with perianth absent; stinging hairs or spines never present .................. 3. Phenax Inflorescences branched; pistillate flowers with persistent perianth; at least some stinging hairs or spines present ......................................... 5 Herbs or subshrubs with succulent stems, without spines; perianth not becoming fleshy in fruit; monoecious ........................................ 2. Laportea

392

5.

U RTICACEAE

Shrubs or trees, with or without spines; perianth becoming fleshy and berry-like (only apparent on live material); usually dioecious ..... 6. Urera

1. BOEHMERIA Jacq., Enum. Syst. Pl. 9. 1760. Herbs, subshrubs, shrubs, or weak trees, lacking stinging hairs. Leaves opposite, sometimes alternate toward the branch apex, cystoliths punctiform and usually present in upper epidermis; margin toothed; venation usually palmate; stipules paired, usually free, deciduous. Inflorescences various but flowers always clustered in glomerules. Flowers unisexual (plant monoecious). Staminate flowers: tepals 3 or 4 (4 in the flora area); stamens as many as tepals. Pistillate flowers: perianth fused into tube enclosing ovary and toothed at apex; stigma linear. Fruit enclosed within the strongly persistent perianth tube. Mainly tropics of America and Asia but extending to temperate areas of eastern North America and eastern Asia, poorly represented in Africa, Australia, and New Zealand; 80–100 species, 5 in Venezuela, 1 of these in the flora area. Boehmeria cylindrica (L.) Sw., Prodr. 34. 1788. —Urtica cylindrica L., Sp. Pl. 984. 1753. Herb or woody subshrub. Disturbed areas, ca. 50 m; scattered in Delta Amacuro. Wide-

spread in eastern Canada, eastern and southern U.S.A., Central America, West Indies, Colombia, French Guiana(?), Brazil, Paraguay, Argentina, Uruguay. ◆Fig. 328.

2. LAPORTEA Gaudich., Voy. Uranie 498. 1826 [1830], nom. cons. Fleurya Gaudich., Voy. Uranie 497. 1826 [1830]. Annual herbs, sometimes perennial shrubs, stinging hairs present. Leaves alternate, cystoliths punctiform, present in upper epidermis; margin usually toothed; venation pinnate; stipules paired, partly connate across the petiole. Inflorescences laxly paniculate with flowers in loose glomerules, pedunculate. Flowers unisexual (plant monoecious). Staminate flowers: tepals 4 or 5, depressed in bud; stamens as many as tepals. Pistillate flowers: tepals 4, free, 2 inner larger than 2 outer; stigma elongate, usually subulate. Fruit partly covered by enlarged lateral tepals. Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, South Africa, Madagascar, Asia, Malasia, Oceania; 22 species, 1 in Venezuela. Laportea aestuans (L.) Chew, Gard. Bull. Singapore 21: 200. 1965. —Urtica aestuans L., Sp. Pl. ed. 2, 1397. 1763. —Fleurya aestuans (L.) Gaudich. ex Miq. in Mart., Fl. Bras. 4(1): 196. 1853. —Guaritoto. Herb or subshrub. Disturbed areas, ca. 50 m; Delta Amacuro (Capure). Aragua, Cojedes, Distrito Federal, Guárico, Miranda, Monagas, Portuguesa, Sucre, Zulia; Mexico,

Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, tropical Africa, Madagascar, India, Sumatra, Java, lesser Sunda Islands. ◆Fig. 329. Laportea aestuans is known from the flora area by only one collection and that was not available to the author. This species is probably more common than indicated by the lack of collections.

Laportea 393

Fig. 328. Boehmeria cylindrica

Fig. 329. Laportea aestuans

394

U RTICACEAE

3. PHENAX Wedd., Ann. Sci. Nat. Bot. sér. 4, 1: 191. 1854. Shrubs, herbs, or rarely trees, lacking stinging hairs. Leaves alternate, cystoliths usually punctiform and present in upper epidermis; margin toothed; venation usually palmate; stipules paired and free. Inflorescences of dense glomerules in axils of leaves, sessile or nearly sessile; bracts often broad and perianth-like. Flowers unisexual (plants monoecious or dioecious). Staminate flowers: perianth 4lobed, fused below middle, rounded or acuminate at apex in bud (may resemble pistillate perianth of Boehmeria); stamens 4. Pistillate flowers: perianth lacking; stigma linear to filiform. Fruit embedded in numerous persistent bracts. Southeastern U.S.A., Mexico, Central America, West Indies, southward to Argentina; ca. 12 species, 4 in Venezuela, 2 of these in the flora area. Key to the Species of Phenax 1. 1.

Bracts ovate with apex acute, margins irregularly short-ciliate; styles > 1.5 mm long, distinctly short pubescent at 10× ........................ P. hirtus Bracts circular with apex rounded, margins uniformly long-ciliate; styles < 1.5 mm, nearly glabrous ....................................................... P. sonneratii

Phenax hirtus (Sw.) Wedd. in A. DC., Prodr. 16(1): 235(38). 1869. —Boehmeria hirta Sw., Prodr. 34. 1788. Herb or shrub. Near streams, ca. 1200– 1300 m; Amazonas (slope of Sierra de la Neblina). Falcón, Lara, Zulia; Mexico, Central America, Jamaica, Hispaniola, Colombia, Ecuador, Peru, Bolivia, Argentina. Urtica hirta Sw. is cited as the basionym by Weddell (Fl. Ind. Occid. 1: 285. 1797). However, Swartz did not publish this combination. The page cited by Weddell pertains to Boehmeria hirta. Phenax sonneratii (Poir.) Wedd. in A. DC., Prodr. 16(1): 235(37). 1869. —Parietaria sonneratii Poir. in Lam., Encycl. 5: 15. 1804. —Picapica. Phenax vulgaris Wedd., Ann. Sci. Nat. Bot. sér. 4, 1: 192. 1854. Herb or subshrub. Disturbed areas, ca. 50 m; Delta Amacuro (El Palmar). Aragua, Distrito Federal, Falcón, Lara, Miranda, Monagas, Sucre, Zulia; Mexico, Central America, West Indies, Colombia, French Guiana, Ecuador, Peru, Brazil, introduced(?) in India. ◆Fig. 330.

Fig. 330. Phenax sonneratii

Pilea 395

4. PILEA Lindl., Coll. Bot. t. 4. 1821, nom. cons. Herbs or rarely subshrubs, lacking stinging hairs. Leaves opposite, cystoliths linear, fusiform, or less often punctiform and present in upper or upper and lower epidermis; margin serrate or less often entire; venation usually palmately 3-veined; stipules paired and connate across the base of the petiole. Inflorescences paniculate, spicate, or capitate. Flowers unisexual (plants monoecious or dioecious). Staminate flowers: tepals 3 or 4, fused near base, often with short, vertical protuberances on outer surface near apex; stamens as many as tepals. Pistillate flowers: tepals 3, identical, linear or usually one larger and hooded with other 2 identical and not hooded; stigma sessile and penicillate. Fruit with tightly fitting perianth persistent at base. Pantropics, a few species in temperate areas; ca. 600 species, 29 in Venezuela, 5 of these in the flora area. Key to the Species of Pilea 1.

1.

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

3.

4(2).

4.

Leaves entire, < 1 cm long, generally unequal at a node, the larger obovate, the smaller orbicular, venation pinnate with secondary veins obscure; inflorescences nearly sessile; achenes 0.3–0.5 mm long ............................................................................................... P. microphylla Leaves toothed at least above middle, from < 1–8 cm in length, nearly equal at a node (< 25% difference), ovate to obovate, venation palmate or 3-pliveined; inflorescences pedunculate or nearly sessile, achenes 0.5–1.2 mm long ..................................................................................... 2 Leaves mostly < 2 cm long; achenes 0.8–1.2 mm long ............................. 3 Leaves mostly > 2 cm long; achenes 0.5(–0.7) mm long ........................... 4 Leaves ovate to elliptic-ovate, margin serrate nearly to base; stipules triangular, opaque, 0.5–1 mm long; transverse secondary veins apparent below ...................................................................................... P. dauciodora Leaves orbicular, obovate to spatulate (rarely ovate), margin crenate with teeth only on distal half; stipules orbicular, transparent, 1.5–2 mm long; transverse secondary venation obscure below ........................... P. fendleri Leaves ovate, apex acute (obtuse), margins coarsely serrate, longer petioles 1.5–6.5 cm; inflorescences pedunculate, peduncle 1–5 cm long .................................................................................................. P. pubescens Leaves obovate, oblong, or orbicular (rarely some ovate, then upper leaves attenuate basally), apex obtuse to rounded (acute), margins bluntly crenate, longest petioles < 1 cm; inflorescences sessile or nearly sessile (peduncle < 0.5 cm) ................................................... P. involucrata

Pilea dauciodora Pav. ex Wedd., Ann. Sci. Nat. Bot. sér. 3, 18: 223. 1852. Stem decumbent. Sink holes, ca. 2600 m; Amazonas (Cerro Marahuaka). Aragua, Barinas, Distrito Federal, Mérida, Miranda, Táchira; southern Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 331.

Pilea fendleri Killip, Field Mus. Nat. Hist., Bot. Ser. 13(2): 341. 1937. Pilea dauciodora var. crenata Wedd. in A. DC., Prodr. 16(1): 139. 1869.. Pilea leptophylla Killip, Contr. U.S. Natl. Herb. 26: 387. 1936, non Urb. 1899. Stem erect to 8 cm, lower portion repent. Shaded mossy boulders, 50–100 m; Delta

396

U RTICACEAE

Fig. 331. Pilea dauciodora

Fig. 332. Pilea pubescens

Pilea 397

Fig. 333. Pilea involucrata

Fig. 334. Pilea microphylla

Amacuro (Río Amacuro). Aragua; Nicaragua, Colombia, Peru. Pilea involucrata (Sims) Urb., Symb. Antill. 1: 298. 1899. —Urtica involucrata Sims, Bot. Mag. 51: pl. 2481. 1824. —Pilea pubescens var. involucrata (Sims) Wedd., Arch. Mus. Hist. Nat. 9: 258. 1856. Pilea chrysosplenioides Wedd., Ann. Sci. Nat. Bot. sér. 3, 18: 231. 1852. Stem erect, 5–30 cm tall, lower portion often repent. Exposed rocks in cloud forests, along rivers, 50–600 m; Delta Amacuro (Río Amacuro), Bolívar (Miamo). Aragua, Distrito Federal, Falcón, Lara, Miranda, Nueva Esparta, Sucre, Táchira, Yaracuy, Zulia; Panama, West Indies, Colombia. ◆Fig. 333. The collection from Delta Amacuro is atypical in having a few ovate leaves with acute apices, characteristics normally found in Pilea pubescens. Pilea involucrata is also reported from Ecuador but these collections, while having nearly sessile inflorescences,

have the other key characteristics of P. pubescens, and here are considered to belong to that variable and widespread species. Similar collections have been made in Táchira. Pilea microphylla (L.) Liebm., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Math. Afh. ser. 5, 2: 296. 1851. —Parietaria microphylla L., Syst. Nat. ed. 10, 1308. 1759. Succulent, erect 2–30 cm tall, or procumbent and often forming mats. Exposed boulders, ca. 500 m; Bolívar (Miamo). Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Táchira, Yaracuy, Zulia; U.S.A. (southern Florida), Mexico,

398

U RTICACEAE

Central America, West Indies, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 334. The specimen of Pilea microphylla from the flora area was initially identified as P. serpyllacea (Kunth) Liebm., a species very similar in appearance but differing in having leaves equal at a node and subglobose and pedunculate pistillate inflorescences with peduncles as long or longer than the subtending leaves. Pilea pubescens Liebm., Danske Vidensk.

Selsk. Skr. Naturvidensk. Math. Afh. ser. 5, 2: 302. 1851. Stem erect, 10–50 cm tall, lower portion often repent and rooting at nodes. Streams, 500–900 m; Bolívar (Matacuchillo west of Santa Elena), Amazonas (upper Río Orinoco). Distrito Federal, Falcón, Lara, Mérida, Miranda, Táchira, Trujillo, Yaracuy; Mexico, Central America, West Indies, Colombia, Suriname, Ecuador, Peru, southeastern Brazil. ◆Fig. 332. Intermediaries exist between this species and Pilea involucrata (see above).

Fig. 335. Pouzolzia occidentalis

Urera 399

5. POUZOLZIA Gaudich., Voy. Uraine 503. 1826 [1830]. Shrubs, subshrubs, or rarely small trees, lacking stinging hairs. Leaves usually alternate, cystoliths usually punctiform and present in upper epidermis; margin mostly entire; venation often subpalmate or pinnate with 2 prominent basal secondary veins; stipules paired and free. Inflorescences of glomerules in axillary clusters (in the flora area) or glomerules arranged in spikes. Flowers unisexual (plant usually monoecious). Staminate flowers: perianth (3)4(5)-lobed, perianth parts narrowed at apex to form elongated tip or abruptly inflexed and transversely plicate on the back; stamens as many as tepals. Pistillate flowers: perianth fused into tube enclosing ovary and toothed at apex, usually with longitudinal ribs; stigma filiform. Fruit enclosed in persistent perianth, perianth thin and ripping at maturity (in the flora area). Pantropics; ca. 50 species, 3 in Venezuela, 1 of these in the flora area. Pouzolzia occidentalis (Liebm.) Wedd., Arch. Mus. Hist. Nat. 9: 410. 1857. —Leucococcus occidentalis Liebm., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Math. Afh. ser. 5, 2: 311. 1851. Shrub or tree 1.5–6 m tall; leaves ovate to

elliptic-ovate, long acuminate at apex. Rocky escarpment, 400–600 m; Bolívar (Miamo). Carabobo, Cojedes, Lara, Miranda, Monagas, Portuguesa, Sucre, Zulia; Honduras to Panama, Puerto Rico, Colombia, Ecuador. ◆Fig. 335.

6. URERA Gaudich., Voy. Uranie 496. 1826 [1830]. Large herbs, shrubs, or small trees, scattered stinging hairs usually present, sharp stinging spines often present. Leaves alternate, cystoliths punctiform, linear, present in upper epidermis or not apparent or absent; margin various; venation usually pinnate; stipules free or ± fused across base of petiole. Inflorescences axillary or cauliflorous, paniculate, cymose or simple with only a few clustered flowers. Flowers unisexual (plant most usually dioecious, occasionally monoecious). Staminate flowers: tepals 4 or 5, slightly imbricate, usually depressed in bud; stamens as many as tepals. Pistillate flowers: perianth parts 4, equal or very unequal (in the flora area), subtending or loosely enclosing ovary; stigma subsessile, usually penicillate-capitate. Fruit at least partially surrounded by perianth which becomes enlarged, fleshy, and berry-like on live material; on dried specimens perianth appears thin and bract-like appressed to fruit, enclosing or subtending. Neotropics, Africa, Asia; 35–75 species, 6 in Venezuela, 3 of these in the flora area. Key to the Species of Urera 1.

1.

2(1).

Stinging spines absent; leaf margin crenate, teeth 0.4–0.7 cm apart; fruit 0.8–1 mm long, larger perianth parts in fruit 0.6–0.8 mm, widely ovate; stigma penicillate ...................................................... U. caracasana Stinging spines present on stem and petiole; leaf margin shallowly sinuate-dentate, teeth (1–)1.5–3.5 cm apart or leaves deeply pinnately lobed; fruit 1.5 mm long or longer, larger perianth parts in fruit 1 mm long or longer, widely ovate to elliptic; stigma penicillate or elongate, curved and subterminal ........................................................................ 2 Leaf margin sinuate-dentate; fruit 2.4–3.5 mm long, larger perianth parts in fruit 1–2 mm long, widely ovate; stigma penicillate ................................................................................................... U. baccifera

400

2.

U RTICACEAE

Leaf margin deeply pinnately lobed; fruit 1.5 mm long, larger perianth parts in fruit 1–1.2 mm long, narrowly elliptic; stigma (in fruit) curved and subterminal ........................................................................ U. laciniata

Urera baccifera (L.) Gaudich. ex Wedd., Ann. Sci. Nat. Bot. sér. 3, 18: 199. 1852. —Urtica baccifera L., Sp. Pl. ed. 2, 1398. 1762 [1763]. —Guaritoto, Ortega, Picaton, Priniga-moza. Shrub 2–3 m tall, with stinging spines and hairs; fruit appearing white and berry-like; inflorescence branches purplish. Forests, 100–1300 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua,

Carabobo, Distrito Federal, Falcón, Miranda, Monagas, Lara, Portuguesa, Táchira, Yaracuy, Zulia; southern Mexico, Central America, West Indies, southward in South America to Argentina. ◆Fig. 336.

Fig. 336. Urera baccifera

Urera 401

Fig. 337. Urera caracasana

Urera caracasana (Jacq.) Gaudich. ex Griseb., Fl. Brit. W. I. 154. 1859. —Urtica caracasana Jacq., Pl. Hort. Schoenbr. 3: 71, pl. 386. 1798. —Guaritoto montanero. Shrub or tree 2–6 m tall, spineless but with irritating hairs; fruit yellow-green, turning gray; inflorescence branches green,

frequently becoming swollen and diseased(?). Margins of forests, deciduous thickets, 50– 300 m; widespread in Bolívar and Amazonas. Anzoátegui, Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Mérida, Miranda, Monagas, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Panama, Trinidad-Tobago, throughout South America. ◆Fig. 337.

402

U RTICACEAE

The name Urera caracasana is based on staminate material and therefore cannot be applied with total confidence. The earliest name for this species based upon pistillate material is probably U. capitata Wedd. var. capitata. Urera caracasana is used here in a stricter sense than that normally employed. It is more frequently used to include Urera corallina (Liebm.) Wedd., which is found primarily in Central America from Mexico through Panama, and the West Indies but also extends into Venezuela and Colombia. It is easily distinguished from U. caracasana by brilliant orange to red, 1.2–2 mm long fruiting perianths that enclose the achene. The fruiting perianths of U. caracasana on the other hand only cover about 3/4 of the achene,

are 0.6–0.8 mm long, and are yellow-green. Staminate and sterile plants can usually be differentiated by the venation. In U. caracasana the secondary veins usually extend out to the margin. In U. corallina the secondary veins usually arc upward before reaching the margin. Urera laciniata Goudot ex Wedd., Ann. Sci. Nat. Bot. sér. 3, 18: 203. 1852. Shrub 2 m tall, with stinging spines and hairs; inflorescence green, white-green, or green-yellow; sap white; fruits green-yellow. Near rivers, 100–200 m; Amazonas (Río Mawarinuma). Nicaragua, Costa Rica, Panama, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

VALERIANACEAE by Nereida Xena de Enrich Annual or perennial herbs, rarely subshrubs or shrubs, erect or climbers, generally rhizomatose, frequently with a strong odor that persists after drying, due to valepotriate compounds in the roots. Leaves opposite, decussate, simple or compound, exstipulate, petiolate or sessile, frequently sheathing at base, the basal leaves sometimes in a rosette, the cauline ones generally smaller or sometimes absent; blades entire to dentate, serrate, or crenate. Inflorescence terminal, axillary, or scapose, determinate, composed of aggregates of dichasia or compound cymes, densely flowered, terminal branches sometimes scorpioid. Flowers small, white, creamy, or pink, bisexual or unisexual (plants hermaphroditic, gynodioecious, dioecious, or polygamous). Calyx tube adnate to ovary, the limb generally reduced or involute in flower, usually persistent and variously modified in fruit, appendages varied in shape or in filiform segments; corolla gamopetalous, the tube long and narrow, funnelform or subcampanulate, sometimes rotate, either straight, gibbous, or spurred, usually widened and ± hirsute at the throat, the limb 5-lobed, rarely 3lobed, regular, oblique or bilabiate (Centranthus, Plectritis, Fedia), the lobes imbricate in bud. Stamens 1–4; filaments free, inserted in the corolla tube, alternate with corolla lobes; anthers bilocular, versatile or basifixed, introrse, longitudinally dehiscent, the thecae sometimes sulcate and then the anthers 4-lobed. Ovary inferior, 3carpellate and 3-locular, only 1 locule fertile, the sterile ones sometimes wider than the fertile one or reduced to crests or wings (Plectritis); ovule 1, pendent from apex of fertile locule, anatropous; style filiform; stigma papillose, subtruncate or more commonly 2- or 3-lobed. Fruit a cypsellate achene, crowned by the reduced calyx, or calyx developed into appendages of various forms or into plumose pappus. Seed pendulous; testa membranous; endosperm absent or scarce; embryo straight, the cotyledons oblong, larger than the radicle.

Valeriana 403

Cosmopolitan, absent only in Australia, mostly in temperate zones and tropical highlands; 8 genera and 350 species, 1 species in the flora area. 1. VALERIANA L., Sp. Pl. 31. 1753; Gen. Pl. ed. 5. 19. 1754. Oligococe Willd. ex DC., Prodr. 4: 632. 1820. Porteria Hook., Icon. Pl. 9: 864. 1852. Amblyorhinum Turcz., Bull. Soc. Nat. Moscou 25(2): 173. 1852. Perennial herbs, subshrubs, or small shrubs, rarely annual herbs or twining plants, rhizomatous or with woody, succulent, or tuberous roots. Stem subscapose or foliate, fistulose, terete or occasionally ± quadrangular. Leaves simple or compound, petiolate or sessile, frequently sheathing at the base; blades entire, serrate, or crenate, firm or membranous. Inflorescence of dichasia, thyrses, or compound cymes, dense or ± scorpioid, bracteate. Flowers bisexual or unisexual (plants gynodioecious, dioecious, or polygamous). Calyx with involute limb in flower and unfolding in fruit into a plumose-ciliate pappus formed of 5–20 sections connected by a fine, basal membrane, or reduced into a short crown both in flower and fruit; corolla white, creamy, or pink, tubular, funnelform, or campanulate, tube short or rarely elongated, frequently gibbous in the anterior basal portion, the limb with 5 equal or subequal lobes, corolla of pistillate flowers 1/3–1/2 the length of hermaphoditic flowers. Stamens 3, adnate to corolla; anthers introrse, with 2 thecae, these sometimes sulcate and then the anthers 4-lobed. Ovary with only the adaxial locule fertile, the abaxial ones vestigial; stigma terminal, 3-lobed. Fruit a cypsellate compressed achene, with calyx persistent in fruit or forming a short, irregularly dentate or lobed crown. Seed filling completely the fertile locule, without endosperm. Almost worldwide in temperate regions or mountains, except Australia; ca. 200 species, ca. 16 in Venezuela, 1 of these in the flora area. Valeriana scandens L., Sp. Pl. ed. 2. 47. 1762. Valeriana phaseoli Braun, Sem. Hort. Berol. 13(2). 1851. Twining or climbing perennial herb; root fasciculate and tuberous with several herbacous stems; leaves long-petiolate, simple or compound; inflorescence of large cymes ending in short, monochasial branches; flowers white; fruits pyriform with a plumose pappus. Subtropical and tropical moist forests, 500–2300 m; U.S.A. (Florida) to Panama, West Indies, all of South America; 3 varieties, all in Venezuela, 2 of these in the flora area.

V. scandens var. candolleana (Gardner) Müll. Arg. in Mart., Fl. Bras. 6(4): 344. 1855. —Valeriana candolleana Gardner, Lond. J. Bot. 4: 112. 1845. Valeriana alpina Vell., Fl. Flumin. 28. 1825 [1829]. Valeriana scandens var. dentata Müll. Arg. in Mart., Fl. Bras. 6(4): 344. 1855. Twining climber. Wet tropical forests, 1000–2500 m; Bolívar (Macizo del Chimantá, Río Aponguao, Roraima-tepui, Sororopántepui), Amazonas (Sierra de la Neblina). Mérida; Mexico, Central America, Cuba, Santo Domingo, Puerto Rico, Brazil. ◆Fig. 338.

Key to the Varieties of V. scandens 1. Leaves ovate-cordate, base shallowly cordate, margin serrate or dentate .............. ................................. var. candolleana 1. Leaves cordate, base deeply cordate-emarginate, margin entire or crenate ............. ................................... var. subcordata

V. scandens var. subcordata Müll. Arg. in Mart., Fl. Bras. 6(4): 344. 1855. Climbing herb. Wet and riparian forests, 900–1000 m; Bolívar (Santa Elena de Uairén). Anzoátegui; Brazil.

404

V ALERIANACEAE

Fig. 338. Valeriana scandens var. candolleana

VELLOZIACEAE by James S. Miller and Paul E. Berry Xerophytic herbs, shrubs, or sometimes tree-like, the stems simple or subdichotomously branched, the upper parts covered with persistent leaf bases, the lower parts covered with appressed, adventitious roots. Leaves clustered at stem apices, narrow and elongate, often pungent-pointed. Flowers axillary, or on short or elongate peduncles, actinomorphic, epigynous, bisexual or rarely unisexual (Barbaceniopsis). Tepals 6, in 2 whorls, petaloid, free or basally connate into a short to elongate tube. Stamens 6 in 2 whorls (Vellozia) or in 6 bundles of 2– several; filaments free or basally adnate to the tepals; anthers linear, basifixed or dorsifixed, dehiscing longitudinally; pollen often in tetrads. Ovary inferior, 3carpellate, 3-locular, the placentation lamellate-axile; ovules numerous, anatropous; style terminal, simple; stigma capitate or slightly 3-lobed. Fruit a loculicidal capsule. Seeds numerous; endosperm abundant, hard. Panama, Colombia, Venezuela, Guyana, Brazil, Bolivia, Africa, Madagascar, southern Arabia, China; 9 genera and ca. 240 species; 2 genera and 2 species in the flora area.

Barbacenia 405

Key to the Genera of Velloziaceae 1. 1.

Stigma unlobed; anthers dorsifixed, borne on elongate, terete filaments; stamens in 6 bundles of 2–several stamens each ................ 1. Barbacenia Stigma 3-lobed; anthers basifixed, borne on short, flat filaments; stamens 6, in 2 whorls ................................................................. 2. Vellozia

1. BARBACENIA Vand., Fl. Lusit. Bras. Spec. 21. 1788. Pleurostima Raf., Fl. Tellur. 2: 97. 1836 [1837]. Herbs to short shrubs, the stems simple or dichotomously branched. Leaves breaking off irregularly. Flowers axillary, bisexual. Tepals connate into a tube. Stamens in 6 bundles of 2–several stamens each; filaments short, flat, with 2 apical appendages; anthers dorsifixed. Stigma not broadly lobed. Venezuela, Brazil, Bolivia; ca. 90 species, 1 in Venezuela. Barbacenia celiae Maguire, Mem. New York Bot. Gard. 18(2): 33, fig. 6A–N. 1969. —Pleurostima celiae (Maguire) Menezes, Revista Brasil. Bot. 3: 47. 1980. Herb to 25 cm tall, in dense colonies; flowers purple. Open areas and depressions on sandstone slopes and tepui summit, 800– 2100 m; Amazonas (Cerro Yaví). Adjacent Brazil (Roraima). ◆Fig. 339. The other Barbacenia that has been reported from the Venezuelan Guayana, B. uaipanensis Maguire, is actually a member of the Bromeliaceae, Ayensua uaipanensis (Maguire) L.B. Sm. (soon to be transferred to Brocchinia).

Fig. 339. Barbacenia celiae

406

V ELLOZIACEAE

2. VELLOZIA Vand., Fl. Lusit. Brasil. Spec. 32, pl. 2. 1788. Herbs or shrubs to 4 m tall, the stems simple or dichotomously branched. Leaves deciduous along a straight, transverse line. Flowers on long peduncles from the upper leaf axils, bisexual. Tepals free or connate into a tube. Stamens 6; filaments terete, basally adnate to the tepals; anthers basifixed. Stigma 3-lobed. Panama, South America (mainly Brazil), tropical Africa, Madagascar; ca. 140 species, 1 in Venezuela. Vellozia tubiflora (A. Rich.) Kunth in H.B.K., Nov. Gen. Sp. 7: 155. 1825. —Radia tubiflora A. Rich. in Kunth, Syn. Pl. Aeq. 1: 300. 1822. Barbacenia alexandrinae R.H. Schomb. ex Hook., London J. Bot. 4: 13. 1845. —Vellozia alexandrinae (R.H. Schomb. ex Hook.) Goethart & Henrard, Blumea 2: 363. 1937. Vellozia duidae Steyerm., Fieldiana, Bot. 28: 157. 1951. Herb or low shrub to 1 m tall, the stems often crowded in clumps, fire-resistant, if

previously burned then with blackened stems; flowers with the corolla tube purple or white, the spreading lobes white. Granitic outcrops, quartz-sand savannas, exposed sandstone near streams, 50–2100 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Moriche, Gran Sabana, Luepa), Amazonas (Cerro Duida, Cerro Vinilla, Cerro Yutajé, Culebra, around Puerto Ayacucho, middle and upper Río Orinoco). Apure, Guárico; Colombia (Vichada), Brazil. ◆Fig. 340.

Fig. 340. Vellozia tubiflora

V ERBENACEAE

407

VERBENACEAE by Gerardo A. Aymard C. Herbs, shrubs, lianas, or trees, often with glandular structures; branches and branchlets often ± tetragonal, not prominently articulate. Leaves exstipulate, mostly decussate, whorled, or alternate, simple or palmately compound, petiolate, sometimes sessile; blades various in texture; margins entire or variously dentateserrate; venation pinnate. Inflorescences axillary or terminal, mostly pedunculate, in the form of spikes, thyrses, racemes, or cymes, often arranged in panicles; bracts mostly small, sometimes colored and leaf-like. Flowers pedicellate or sessile, ± zygomorphic or sometimes actinomorphic, bisexual, heterostylous (Aegiphila), or polygamous, hypogynous. Calyx gamosepalous, tubular-campanulate, mostly 4- or 5lobed or -toothed, persistent, often enlarged in fruit; corolla salverform or funnelform, the limb 4- or 5-lobed sometimes ± 2-lipped. Stamens 4(5), often didynamous, or 2 by abortion, inserted on the corolla tube, included or exserted, anthers with 2 thecae, parallel or divergent, introrse, longitudinally dehiscent; staminodes often present; disk sometimes present. Pistil 1; ovary superior, 2–4-locular, sometimes 8locular due to false partitions; each locule with 1 or 2 ovules; style 1, terminal stigma bifid, bilobed, or capitate; ovules born basally or laterally, anatropous or apically orthotropous. Fruit a drupe or schizocarp, separating into 2–4 nutlets, or a 2-valved capsule; exocarp fleshy or dry; endocarp hard. Seed with thin testa, with or without endosperm; cotyledons flat, ± thickened, parallel. Tropics and subtropics worldwide, a few species in temperate regions; about 100 genera and 2600 species, 15 genera and 74 species in the flora area. According to the Angiosperm Phylogeny Group (APG II. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linnean Soc. 141: 399–436), Avicennia belongs in the Acanthaceae, rather than in Verbencaceae, where it traditionally has been placed. Likewise, the traditional placement of genera between Lamiaceae and Verbenaceae has changed, with Aegiphila, Amasonia, Clerodendrum, Gmelina, Tectona, and Vitex now placed in the Lamiaceae. The Asian genera Gmelina and Tectona are widely cultivated in forest plantations. Verbena tenuisecta Briq., a creeping or procumbent herb with pinnatifid leaves and blue or purple flowers, may be present in the flora area in Bolívar state, but no reliable reports have been found; it is known from northern Venezuela in the Coastal Cordillera and in the Andes. Key to the Genera of Verbenaceae 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

Trees ........................................................................................................... 2 Herbs, shrubs, subshrubs, or woody lianas .............................................. 7 Leaves palmately compound ............................................................. 15. Vitex Leaves simple ............................................................................................. 3 Fruiting calyx with enlarged wing-like lobes ................................. 10. Petrea Fruiting calyx without enlarged lobes ...................................................... 4 Trees in mangrove vegetation, with pencil-like pneumatophores .................................................................................................. 3. Avicennia Trees in other habitats, without pencil-like pneumatophores ................. 5

408

V ERBENACEAE

5(4). 5. 6(5). 6. 7(1). 7. 8(7). 8. 9(8). 9. 10(9). 10. 11(8). 11. 12(11). 12. 13(12). 13. 14(13). 14. 15(14). 15. 16(14). 16. 17(16).

17. 18(11). 18. 19(18). 19. 20(19). 20.

Leaves with one or more pairs of glands at the apex of the petiole or the base of the blades ............................................................... 5. Citharexylum Leaves without glands at the apex of the petiole or the base of the blades ................................................................................................................ 6 Plants not spinose; corolla white or yellow-green; filaments exserted from the tube .................................................................................... 1. Aegiphila Plants often spinose; corolla blue or rarely white; filaments included in the tube ...................................................................................... 7. Duranta Shrubs in mangrove vegetation, with pencil-like pneumatophores .................................................................................................. 3. Avicennia Shrubs, subshrubs, herbs, or lianas in other habitats, without pencil-like pneumatophores .................................................................................... 8 Herbaceous or woody lianas ...................................................................... 9 Herbs, shrubs, or subshrubs .................................................................... 11 Filaments included in the tube; fruiting calyx with enlarged wing-like lobes .............................................................................................. 10. Petrea Filaments exserted from the tube; fruiting calyx without enlarged lobes .............................................................................................................. 10 Leaves pinnately veined; calyx 1–15 mm long, not inflated; corolla yellow or greenish white ..................................................................... 1. Aegiphila Leaves 3-veined at the base; calyx 18–20 mm long, inflated; corolla red .......................................................................................... 6. Clerodendrum Perennial or annual herbs ....................................................................... 12 Small trees, shrubs, or subshrubs ........................................................... 18 Calyx campanulate, red, 6–15 mm long (including the lobes) .... 2. Amasonia Calyx tubular-campanulate, green, 1–5 mm long .................................. 13 Flowers embedded in the rachis of the inflorescence ...... 13. Stachytarpheta Flowers not embedded in the rachis of the inflorescence ...................... 14 Fruit a fleshy drupe ................................................................................. 15 Fruit a dry schizocarp .............................................................................. 16 Branches with hooked hairs; upper pair of stamens with a glandular thickening at the apex of the filament .................................. 14. Tamonea Branches without hooked hairs; upper pair of stamens without a glandular thickening at the apex of the filament ................................ 8. Lantana Creeping or procumbent herbs.. ....................................................... 11. Phyla Erect herbs ............................................................................................... 17 Leaves glabrous or sparsely pubescent on both sides; petiole canaliculate; calyx outside without hooked hairs, fruiting calyx not accrescent .................................................................................................... 4. Bouchea Leaves pilose-pubescent on both sides; petiole not canaliculate; calyx outside with hooked hairs, fruiting calyx accrescent ....................... 12. Priva Flowers embedded in the rachis of the inflorescence ...... 13. Stachytarpheta Flowers not embedded in the rachis of the inflorescence ...................... 19 Calyx campanulate, red with dark, disk-shaped glands ........................ 20 Calyx subcampanulate or tubular-campanulate, green, without dark disk-shaped glands .............................................................................. 21 Calyx 6–15 mm long; corolla yellow, petals in a single whorl ..... 2. Amasonia Calyx 1–2 mm long; corolla white or purple, petals in a double whorl .......................................................................................... 6. Clerodendrum

Aegiphila 409

21(19). 21. 22(21). 22.

Filaments exserted from the tube ............................................... 1. Aegiphila Filaments included in the tube ............................................................... 22 Plants often spinose; inflorescences racemose; calyx 5-costate .... 7. Duranta Plants not spinose; inflorescences spicate or densely capitate; calyx not costate .................................................................................................. 23 23(22). Stigma thick; fruit a dry schizocarp; ovule 1 per locule .................. 9. Lippia 23. Stigma thin; fruit a fleshy drupe; ovules 2 per locule .................. 8. Lantana 1. AEGIPHILA Jacq., Observ. Bot. 2: 3. 1767. Manabea Aubl., Hist. Pl. Guiane 61. 1775. Trees, shrubs, or woody vines; branches and branchlets tetragonal or subterete, glabrous or variously pubescent. Leaves simple, decussate, rarely subopposite or ternate, sometimes deciduous, exstipulate, mostly petiolate, glabrous or variously pubescent, entire or dentate. Inflorescences axillary cymes, solitary, or in terminal panicles, the cymes umbelliform, glomeruliform, or head-like. Flowers pedicellate, bisexual, heterostylous, actinomorphic. Calyx campanulate or tubular, 4(5)-lobed or -toothed; corolla white or yellow, salverform or funnelform, 4(5)-lobed. Stamens 4 or 5, isomorphic, inserted somewhat above the base or near the middle of the corolla tube; filaments exserted; anthers oblong, basifixed or medifixed. Ovary 4-locular, each locule with 1 ovule; style bifurcate. Fruit a drupe with fleshy exocarp, globose or subglobose; fruiting calyx enlarged and indurated. Seeds (1–3)4, sometimes fewer by abortion, without endosperm. Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; ca. 150 species, 37 in Venezuela, 21 of these in the flora area. Key to the Species of Aegiphila 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

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

Lianas or scandent shrubs ......................................................................... 2 Small trees or erect shrubs ...................................................................... 13 Branches and branchlets densely ochraceous-villous or densely pubescent ......................................................................................................... 3 Branches and branchlets glabrous, papillose, puberulent, sparsely villous, or very minutely pubescent .......................................................... 6 Branches and branchlets densely ochraceous-villous .............................. 4 Branches and branchlets densely pubescent ............................................ 5 Branches and branchlets with hairs 1–3 mm long; leaves coriaceous, ovate, densely punctate on the lower surface; calyx 8–10 mm long; bracteoles entire ......................................................................... A. hoehnei Branches and branchlets with hairs ca. 0.5 mm long; leaves membranous, elliptic or ovate-elliptic, not punctate on the lower surface; calyx 1–1.5 mm; bracteoles trilaciniate .......................................... A. hystricina Leaves glabrous on both sides, densely punctate on the lower surface; corolla not papillose ......................................................................... A. elata Leaves strigose on the upper surface, pubescent with disk-shaped glands on the lower surface; corolla papillose .................................... A. racemosa Leaves with disk-shaped glands on both sides ......................................... 7 Leaves without disk-shaped glands ........................................................ 10

410

V ERBENACEAE

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

9. 10(6). 10. 11(10). 11. 12(11). 12. 13(1). 13. 14(13). 14. 15(14). 15. 16(15). 16. 17(15). 17. 18(17). 18. 19(14). 19. 20(19). 20. 21(20).

Leaves oblong; calyx 1–1.8 mm long ...................................... A. glandulifera Leaves elliptic or oblong-elliptic; calyx 2–9 mm long ............................... 8 Branches and branchlets with indument of erect hairs ...... A. membranacea Branches and branchlets papillose or very minutely pubescent ............. 9 Leaves acuminate; calyx 7–9 mm long, glabrous; corolla white or cream, 15–25 mm long, the outside papillose or very minutely pubescent ................................................................................................ A. macrantha Leaves caudate; calyx 2–5 mm long, puberulent, corolla yellow glabrous, 3–6 mm long ................................................................................... A. filipes Leaves oblong-lanceolate; fruiting calyx ca. 11 mm diameter ...... A. lewisiana Leaves oblong, oblong-elliptic, or ovate; fruiting calyx 5–9 mm diameter .............................................................................................................. 11 Branches and branchlets densely puberulent; leaves sparsely strigulose with white hairs on both sides; corolla white ........................... A. pendula Branches and branchlets glabrous; leaves glabrous on both sides, or appressed-pubescent on lower surface.................................................... 12 Leaf base rounded or subcordate; lower surface of leaves appressedpubescent ......................................................................................... A. elata Leaf base acuminate; leaves glabrous on both sides ........................ A. laevis Leaves serrate mostly in the upper half ............................... A. venezuelensis Leaves entire ............................................................................................ 14 Leaves without disk-shaped glands or black punctations on both surfaces .............................................................................................................. 15 Leaves with disk-shaped glands, black punctations, or both on both surfaces ...................................................................................................... 19 Leaves mostly densely appressed-pubescent or the lower surface densely short-pubescent .................................................................................... 16 Leaves glabrous or sparsely pilose on lower surface, except on primary and secondary veins ............................................................................. 17 Upper surface of leaves densely pubescent; pedicels pannose, 1–3 mm long; corolla yellow ........................................................................ A. mollis Upper surface of leaves glabrous or sparsely puberulent; pedicels pubescent, 3–6 mm long; corolla white .......................................... A. intermedia Inflorescences panicles; flowers sessile or pedicel ca. 1 mm long; calyx ca. 1 mm long ................................................................................ A. spruceana Inflorescences cymes; pedicel 2–8 mm long; calyx 2–4 mm long ........... 18 Petioles canaliculate; leaves membranaceous or subcoriaceous, the base acuminate; inflorescences 5–7 cm long ......................................... A. laevis Petioles not canaliculate; leaves coriaceous, the base rounded; inflorescences 14–16 cm long ....................................................................... A. elata Branches and branchlets densely ochraceous-sericeous, densely appressed-villous with gray, white, and yellow hairs, or velutinous .... 20 Branches and branchlets glabrous, puberulent, shortly pubescent, or sparsely appressed pubescent ............................................................. 23 Base of leaves acute, cuneate, or attenuate; corolla tube 2–5 mm long .............................................................................................................. 21 Base of leaves attenuate; corolla tube 6–12 mm long ............................ 22 Inflorescences 6–17 cm long; calyx mucronate or truncate; corolla greenish or yellow, lobes 3–4 mm long ................................................... A. mollis

Aegiphila 411

21. 22(20).

22. 23(19). 23. 24(23). 24. 25(23). 25. 26(25). 26.

Inflorescences 1–6 cm long; calyx dentate; corolla white, lobes 1.5–2 mm long .......................................................................................... A. parviflora Lower surface of leaves densely ochraceous-sericeous; calyx outside not black-punctate; corolla white, lobes ovate-elliptc, 1–1.8 mm wide .............................................................................................. A. roraimensis Lower surface of leaves strigose; calyx outside black-punctate; corolla yellowish, lobes ovate or lanceolate-ligular, 2–4 mm wide ...... A. integrifolia Leaves oblong, obovate, or oblanceolate ................................................. 24 Leaves elliptic, oblong-elliptic, or ovate-elliptic ..................................... 25 Leaves oblong, secondary veins 5–7, subcoriaceous; corolla pale yellow .................................................................................................... A. perplexa Leaves obovate or oblanceolate, secondary veins 10–13, membranaceous; corolla white .......................................................................... A. bracteolosa Branchlets glabrescent; lower surface of leaves glabrous or puberulent; corolla yellow, 3–6 mm long .......................................................... A. filipes Branchlets puberulent or appressed-villose; lower surface of leaves appressed-pubescent; corolla white, 7–12 mm long ............................... 26 Petioles 1–7 mm long; leaf base acuminate; pedicels minutely puberulent to glabrescent; filaments 9–12 mm long ................................... A. laxiflora Petioles 5–15 mm long; leaf base attenuate-acuminate; pedicels densely appressed-pubescent; filaments 2–4 mm long .................. A. venezuelensis

Aegiphila bracteolosa Moldenke, Repert. Spec. Nov. Regni Veg. 33: 115. 1933. Shrub 1–3 m tall; corolla white. Montane forests, 1000–1500 m; Amazonas (Caño Piedra near Cerro Cuao, Cerro Aracamuni, 75 km southeast of Puerto Ayacucho). Colombia, Guyana, Peru, Brazil, Bolivia. Aegiphila elata Sw., Prodr. 31. 1788. Liana or small shrub. Neotropics; 2 varieties, both in the flora area. Key to the Varieties of A. elata 1. Leaves ovate or oblong-elliptic .. var. elata 1. Leaves widely ovate ...... var. macrophylla A. elata var. elata Liana; corolla yellow. Evergreen lowland forests, 100–300 m; Bolívar (Río Asa, lower Río Paragua). Aragua, Apure, Barinas, Carabobo, Mérida, Monagas, Trujillo; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, Ecuador. A. elata var. macrophylla (Kunth) LópezPal., Pittieria 5: 15. 1973. —Aegiphila macrophylla Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 251. 1817 [1818]. Shrub or small tree; corolla yellow. Evergreen lowland forests, 100–200 m; Amazonas

(Río Mawarinuma, San Carlos de Río Negro). Apure, Barinas, Carabobo, Mérida, Yaracuy; Trinidad. Aegiphila filipes Mart. & Schauer ex Schauer in A. DC., Prodr. 11: 652. 1847. —Tabaquillo. Liana, scandent shrub, or shrub; corolla yellow or white. Semideciduous forests, Trachypogon savannas, 200–300 m; Bolívar (El Manteco, islands in Lago Guri, Isla Anacoco, mouth of Río Paragua), Amazonas (Río Cunucunuma). Trujillo; Panama, Colombia, Peru, Brazil (Acre, Amazonas, Pará), Bolivia. Aegiphila glandulifera Moldenke, Brittonia 1: 187. 1932. Liana or scandent shrub. Neotropics; 3 varieties, 1 in Venezuela. A. glandulifera var. pyramidata Moldenke, Phytologia 1: 203. 1937. —Hoja babaso, Shinnate (Yekwana). Corolla yellow or white. Evergreen lowland and disturbed forests, 100–400 m; Bolívar (southwestern base of Cerro Guaiquinima, Santa María de Erebato), Amazonas (Río Baría, Río Casiquiare, Río Cunucunuma, Río Mawarinuma, Río Ocamo, San Carlos de Río Negro, Solano). Apure, Monagas; Brazil. ◆Fig. 342.

412

V ERBENACEAE

Aegiphila hystricina Aymard & Cuello, Novon 14: 22. 2004. Liana or scandent shrub; corolla white or creamy yellow. Lower montane forests, 400– 500 m; Bolívar (Gran Sabana, Kamarata), Amazonas (upper Río Carinagua south of Puerto Ayacucho). Endemic. Aegiphila hoehnei Moldenke, Phytologia 6: 224. 1937. Liana. Neotropics; 3 varieties, 1 in Venezuela. A. hoehnei var. venezuelensis Moldenke, Phytologia 43: 193. 1979. —Bejuco de pereza, Laurel orillero, Merecure. Vine or scandent shrub with many lateral hanging branches; corolla white or light yellow. Edges of evergreen lowland forests, white-sand scrub (bana), disturbed areas, 100–200 m; Amazonas (Pimichín, Río Mawarinuma, San Carlos de Río Negro, YavitaMaroa road). Endemic. Aegiphila integrifolia (Jacq.) B.D. Jacks., Index Kewensis 1: 386. 1895. —Callicarpa integrifolia Jacq., Enum. Syst. Pl. 12. 1760; Select. Stirp. Amer. Hist. 15, t. 173, fig. 7. 1763, nom. superfl. —Humarí (Baniva), Macuderi (Bale), Tabaquero, Tabaquillo, Udidi (Piaroa). Aegiphila guianensis Moldenke, Repert Spec. Nov. Regni Veg. 33: 125. 1933. —Aegiphila integrifolia var. guianensis (Moldenke) López-Pal., Revista Fac. Farm. Univ. Andes 14: 21. 1974. Shrub or small tree; corolla white. Semideciduous to evergreen lowland and lower montane forests, disturbed areas, near sea level to 1200 m; Delta Amacuro (Caño Guara near Isla Guara, Río Orocoima), Bolívar (Canaima, El Paují, El Tigre, Isla Anacoco, Karaurín-tepui, Kilómetro 88, La Paragua, Río Aro, Río Maniapure, Santa Elena de Uairén, Tumeremo, Upata), Amazonas (Isla Ratón, La Esmaralda, Puerto Ayacucho, Río Atacavi, Río Casiquiare, Río Negro, Samariapo, San Antonio del Orinoco, Tamatama, Yavita). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Guárico, Lara, Mérida, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Acré,

Amazonas, Roraima, Pará), Bolivia. 346.

◆ Fig.

Aegiphila intermedia Moldenke, Repert. Spec. Nov. Regni Veg. 33: 128. 1933. —Tabaquillo. Liana or scandent shrub; corolla white. Evergreen lowland forests, disturbed areas, ca. 100 m; Amazonas (Isla Ratón). Brazil (Amazonas, Pará). Aegiphila laevis (Aubl.) J.F. Gmel., Syst. Nat. 259. 1791. —Manabea laevis Aubl., Hist. Pl. Guiane 66, pl. 25. 1775. Shrub; corolla yellow. Lower montane forests, 400–1300 m; Bolívar (Río Tírica near Macizo del Chimantá), Amazonas (Río Cunucunuma, Sierra de la Neblina). Cojedes, Distrito Federal, Guárico, Lara, Zulia; Colombia, Guyana, Suriname, French Guiana, Brazil. Aegiphila laxiflora Benth., Ann. Nat. Hist. 2: 449. 1839. —Tabaquillo. Shrub to small tree; corolla white. Semideciduous to wet forests, 100–300 m; Bolívar (El Manteco, Isla Anacoco, mouth of Río Paragua, Upata). Barinas, Mérida, Monagas; Trinidad, Guyana. ◆Fig. 348. Aegiphila lewisiana Moldenke, Repert. Spec. Nov. Regni Veg. 33: 131. 1933. Liana or scandent shrub; corolla white. Evergreen lowland forests, 100–200 m; Bolívar (Isla Anacoco, Río Cuyuní). Carabobo, Distrito Federal, Lara, Mérida, Portuguesa, Sucre. ◆Fig. 345. Aegiphila macrantha Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 173. 1925. —Erocuaja (Warao). Liana; corolla white or green. Evergreen lowland forests, near sea level to 300 m; Delta Amacuro (Caño Araguabisi), Bolívar (Bochinche, El Dorado, El Palmar, Tumeremo). Miranda; Trinidad, Guyana, Suriname, French Guiana, Brazil (Pará). ◆Fig. 343. Aegiphila membranacea Turcz., Bull. Soc. Imp. Naturalistes Moscou 36: 219. 1863. Liana or scandent shrub; corolla yellow. Premontane to montane forests, 200–1300 m; Bolívar (Anawaray-Parú on middle Río Caura, Cerro Venamo, El Palmar, El Paují,

Aegiphila 413

La Escalera), Amazonas (Santa Bárbara del Orinoco). Apure, Mérida, Zulia; Colombia, Suriname, French Guiana. Aegiphila mollis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 250, t 130. 1817 [1818]. Neotropics; 5 varieties, all in Venezuela, 3 of these in the flora area. Key to the Varieties of A. mollis 1. Leaves puberulent or glabrescent ............. ................................... var. intermedia 1. Leaves tomentose or pubescent ............... 2 2. Leaves densely hirsute-tomentose ............ ........................................... var. mollis 2. Leaves short-pubescent ........................... ................................... var. surfaceana A.

mollis var. intermedia Moldenke, Repert. Spec. Nov. Regni Veg. 33: 128. 1933. —Rastrojero blanco. Shrub; corolla white or cream. Deciduous to evergreen lowland forests, 100–200 m; Bolívar (Caicara del Orinoco, La Paragua), Amazonas (Santa Barbara del Orinoco). Apure, Mérida, Monagas; Colombia. A. mollis var. mollis. —Tabaquillo. Shrub; corolla yellow. Deciduous to semideciduous forests, 100–200 m; Bolívar (Ciudad Bolívar, Maripa), Amazonas (middle Río Orinoco). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Distrito Federal, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Yaracuy; Costa Rica, Nicaragua, Panama, Colombia, Ecuador(?), Peru, Brazil (Amazonas). ◆Fig. 344. A. mollis var. surfaceana (Moldenke) Moldenke, Phytologia 25: 430. 1973. —Aegiphila surfaceana Moldenke, Bull. Torrey Bot. Club 58: 462. 1931. Shrub; corolla yellow. Evergreen lowland forests, disturbed areas, 100–200 m; Bolívar (Caicara, Río Parhueña), Amazonas (Capibara, San Carlos de Río Negro). Colombia. Aegiphila parviflora Moldenke, Repert. Spec. Nov. Regni Veg. 33: 135. 1933. —Tabaquillo. Shrub; corolla white. Deciduous forests, disturbed areas, 50–100 m; Bolívar (Ciudad

Bolívar south to Camurira). Apure, Guárico; Brazil (Pará). Aegiphila pendula Moldenke, Repert. Spec. Nov. Regni Veg. 33: 135. 1933. Shrub or liana, corolla white. Wet montane forests, ca. 1500 m; Amazonas (Sierra de la Neblina). Zulia. Aegiphila perplexa Moldenke, Repert. Spec. Nov. Regni Veg. 33: 136. 1933. Shrub or small tree; corolla yellow. Evergreen lowland forests, ca. 50 m; Delta Amacuro (Santa Catalina). Trinidad, Tobago. Aegiphila racemosa Vell., Fl. Flumin. 37. 1825 [1829]. Shrub or liana; corolla pale yellow. Premontane to montane forests, 100–1700 m; Bolívar (Altiplanicie de Nuria, Cerro Venamo, Icabarú, Santa Elena de Uairén). Mérida, Táchira, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil (Amapá, Amazonas, Pará, Roraima). Aegiphila roraimensis Moldenke, Repert. Spec. Nov. Regni Veg. 33: 138. 1933. Aegiphila steyermarkii Moldenke, Fieldiana, Bot. 28: 512. 1953. Aegiphila steyermarkii var. obtusifolia Moldenke, Fieldiana, Bot. 28: 513. 1953. Shrub; corolla white. Wet lower montane to high-tepui forests, 800–2500 m; Bolívar (Auyán-tepui, El Paují, Guiaquinima-tepui, Kavanayén, Macizo del Chimantá [Chimantá-tepui], Ptari-tepui, upper Río Yuruaní, Santa Elena de Uairén, Sororopán-tepui), Amazonas (Cerro Huachamacari). Guyana. ◆Fig. 347. Aegiphila spruceana Moldenke, Repert. Spec. Nov. Regni Veg. 33: 139. 1933. Shrub; corolla white. Evergreen lowland forests, 100–200 m; Amazonas (Piedra Cocuy). Colombia, Brazil (Amazonas). Aegiphila venezuelensis Moldenke, Fieldiana, Bot. 28: 514. 1953. Aegiphila steyermarkii var. macrophylla Moldenke, Phytologia 7: 119. 1960. Aegiphila cowanii Moldenke, Phytologia 27: 63. 1973. Shrub or small tree. Venezuela; 2 varieties. Endemic.

414

V ERBENACEAE

Key to the Varieties of A. venezuelensis 1. Leaves serrate mostly in the upper half of the blade ........................... var. serrata 1. Leaves entire ........... var. venezuelensis

(Cerro Jaua, Cerro Sarisariñama, Cerro Venamo, Macizo del Chimantá [Chimantátepui], Guaiquinima-tepui, Ilú-tepui, Ptaritepui, Serranía Marutaní), Amazonas (Cerro Aracamuni, Sierra Parima). Guyana(?).

A. venezuelensis var. serrata Moldenke, Phytologia 6: 323 1958. Corolla white. Wet montane forests, 1500– 1700 m; Bolívar (Cerro Jaua), Amazonas (Sierra de la Neblina). Aragua, Yaracuy. ◆Fig. 341. A. venezuelensis var. venezuelensis Corolla white. Wet montane and hightepui Bonnetia forests, 1400–2600 m; Bolívar

Fig. 341. Aegiphila venezuelensis var. serrata

Fig. 342. Aegiphila glandulifera var. pyramidata

Aegiphila 415

Fig. 343. Aegiphila macrantha

Fig. 344. Aegiphila mollis var. mollis

416

V ERBENACEAE

Fig. 345. Aegiphila lewisiana

Fig. 346. Aegiphila integrifolia

Amasonia 417

Fig. 347. Aegiphila roraimensis

Fig. 348. Aegiphila laxiflora

2. AMASONIA L. f., Suppl. Pl. 48. 1781 [1782], nom. cons. Herbs or subshrubs, mostly woody at the base, usually branched only at the base. Leaves alternate or opposite, mostly ± clustered below the inflorescences, simple, sometimes deciduous, exstipulate; blades membranaceous, glabrous becoming pubescent-hirsute. Inflorescences terminal, racemiform, composed of shortly pedunculate cymes subtended by large leaf-like, colored bracts. Flowers pedicellate, bisexual, slightly zygomorphhic. Calyx campanulate, 5-lobed subequal; corolla yellow or white, salverform, the limb 5-lobed, slightly 2-lipped. Stamens 4, didynamous, inserted somewhat above the base of the corolla tube; anthers ovoid, dorsifixed. Ovary 4-locular, each locule with 1 ovule; style shortly bifurcate. Fruit a drupe with fleshy exocarp; fruiting calyx somewhat enlarged. Neotropics; ca. 4 species, 2 in Venezuela, both in the flora area. Key to the Species of Amasonia 1. 1.

Bracts of the inflorescences oblong-lanceolate, ovate, lanceolate, oblong, or elliptic ................................................................................ A. campestris Bracts of the inflorescences obovate .............................................. A. obovata

418

V ERBENACEAE

Amasonia campestris (Aubl.) Moldenke, Torreya 34: 8. 1934. —Taligalea campestris Aubl., Hist. Pl. Guiane. 625. 1775. —Adarro-Itipi (Curripaco), Candela, Farfara roja, Rabo de guaca, Rabo de zorro. Amasonia arborea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 253. 1817 [1818]. Amasonia lasiocaulos Mart. & Schauer ex Schauer in DC., Prodr. 11: 678. 1847. Amasonia spruceana Moldenke, Repert. Spec. Nov. Regni Veg. 37: 213. 1934. Herb or subshrub; bracts red; corolla yellow or white. Edges of forests, savannas, disturbed areas, 50–1800 m; Delta Amacuro (Los Castillos de Guayana), widespread in Bolívar and Amazonas. Anzoátegui, Falcón, Guárico, Monagas, Sucre; Colombia, Guy-

ana, Suriname, French Guiana, Brazil (Bahia, Ceará, Distrito Federal, Goiás, Maranhão, Minas Gerais, Pará). ◆Fig. 349. Amasonia obovata Gleason, Bull. Torrey Bot. Club. 58: 463. 1931. Herb or subshrub; bracts red; corolla yellow. Edges of forests and savannas, 100–1100 m; Amazonas (Cerro Duida). Endemic.

Fig. 349. Amasonia campestris

Avicennia 419

3. AVICENNIA L., Sp. Pl. 110. 1753. Shrubs or trees of maritime regions, with pencil-like pneumatophores; branches and branchlets terete, prominently articulate. Leaves decussate, simple, petiolate, exstipulate; blades entire. Inflorescences axillary and terminal spikes; bracts and bracteoles scale-like and closely subtending the flower. Flowers sessile, actinomorphic, bisexual. Calyx with 5 nearly free lobes; corolla campanulate-rotate, the limb 4-lobed, the petals connate basally. Stamens 4, inserted in the throat of the corolla tube, equal or subdidynamous. Ovary incompletely 4-locular, each locule with 1 ovule; style with a bifid stigma; ovules born apically, orthotropous. Fruit a compressed ellipsoid or ovoid capsule, often with narrow persistent stylar beak, 1–4 cm long, dehiscent by 2 valves; exocarp juicy and fleshy. Seed usually 1; embryo viviparous, radicle pubescent. Pantropics; ca. 8 species, 2 in Venezuela, both in the flora area. As mentioned following the family description, Avicennia is now placed in the Acanthaceae. Key to the Species of Avicennia 1. 1.

Leaves usually acute at apex; corolla lobes oval to oblong, tomentose on both sides; styles 4–5 mm long ............................................. A. germinans Leaves usually rounded at apex; corolla lobes obovate, glabrous above; styles 1–2 mm long or sessile .............................................. A. schaueriana

Avicennia germinans (L.) L., Sp. Pl. ed. 3, 2: 891. 1764. —Bontia germinans L., Syst. Nat. ed. 10, 2: 1122. 1759. Avicennia nitida Jacq., Enum. Syst. Pl. 25. 1760. Avicennia tomentosa var. cumanensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 283. 1817 [1818]. —Avicennia germinans var. cumanensis (Kunth) Moldenke, Phytologia 30: 15. 1975. Avicennia tomentosa var. guayaquilensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 284. 1817 [1818]. —Avicennia

Fig. 350. Avicennia germinans

420

V ERBENACEAE

germinans var. guayaquilensis (Kunth) Moldenke, Phytologia 29: 75. 1974. Avicennia germinans f. venezuelensis Moldenke, Phytologia 47: 222. 1980. Tree with abundant erect pneumatophores; flowers white. Frequently forms associations with Laguncularia racemosa (L.) Gaertn. f. and Rhizophora mangle L. in saline or mangrove areas, near sea level to 50 m; Delta Amacuro (Caño Angosturita east of Pedernales, Caño Guiniquina). Aragua, Anzoátegui, Carabobo, Dependencias Federales, Falcón, Miranda, Monagas, Nueva Esparta, Sucre, Zulia; U.S.A (Florida), Mexico, Central America, Antilles, Colombia, Guy-

ana, Suriname, French Guiana, Ecuador, Peru, Brazil (Pará). ◆Fig. 350. Avicennia schaueriana Stapf & Leechm. ex Moldenke, Lilloa 4: 336. 1939. Avicennia schaueriana f. candicans Moldenke, Phytologia 35: 13. 1976. Tree with abundant erect pneumatophores; flowers white. Frequently forms associations with Laguncularia racemosa (L.) Gaertn. f. and Rhizophora mangle L. in saline or mangrove areas, near sea level to 50 m; expected in Delta Amacuro. Trinidad, Guyana, Suriname, French Guiana, Brazil (Ceará, Marahnão, Pará, Paraná, Santa Catarina).

4. BOUCHEA Cham., Linnaea 7: 252. 1832. nom. cons. Annual or perennial herbs; branches and branchlets terete. Leaves decussate, exstipulate, petiolate or subsessile; blades crenate, rarely entire. Inflorescences terminal or axillary, racemose or spicate. Flowers pedicellate. Calyx tubular, 5-ribbed, teeth setaceous, equal; corolla funnelform, the tube straight or curved, lobes unequal, blue, rose, or purple. Stamens 4, didynamous; filaments inserted medially or higher; anthers with parallel cells. Ovary 2-locular; ovule 1 per locule; style filiform; stigma bilobed, clavate with a posterior tooth or tubercle. Fruit dry, oblong or linear, rostrate, included in the calyx, separating into 2 cocci, one of these commonly sterile. Pantropics, most common in the Neotropics; ca. 10 species, 1 in Venezuela. Bouchea prismatica (L.) Kuntze, Revis. Gen. Pl. 2: 502. 1891. —Verbena prismatica L., Sp. Pl. 19. 1753. Annual or perennnial herb. Neotropics; 3 varieties, all in Venezuela, 1 of these in the flora area. B. prismatica var. longirostra Grenzeb.,

Ann. Missouri Bot. Gard. 13: 81. 1926. Open areas, edges of towns, 50–100 m; Bolívar (Ciudad Bolívar). Anzoátegui, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo; Mexico, Central America, Antilles, Colombia, Ecuador. ◆Fig. 351.

5. CITHAREXYLUM L., Sp. Pl. 625. 1753. Trees or shrubs, sometimes spiny; branches and branchlets usually tetragonal. Leaves decussate or in whorls of 3, simple, subopposite, entire or dentate, exstipulate, deciduous; usually with one or more pairs of glands at the apex of the petiole or the base of the blade. Inflorescences axillary and terminal racemes or panicles composed of racemes. Flowers pedicellate or sessile, bisexual, actinomorphic or somewhat zygomorphic. Calyx cupuliform, truncate, or weakly 5-lobed; corolla white or yellow, ± hairy in the throat, salverform, 5-lobed. Stamens 4 or 5, or the 5th staminodial, inserted near the middle of the corolla tube, included; anthers oblongellipsoid, medifixed. Ovary perfectly or imperfectly 4-locular, each locule with 1 ovule; style with a capitate or 2-lobed stigma. Fruit a drupe separating into 2 pyrenes by a median fissure, fruiting calyx somewhat enlarged; exocarp fleshy. U.S.A. (Florida, Texas), Mexico, Central America, Bermuda, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia,

Citharexylum 421

Fig. 351. Bouchea prismatica var. longirostra

Paraguay, Argentina, Uruguay; 125 species, 10 in Venezuela, 5 of these in the flora area. Key to the Species of Citharexylum 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Leaves glabrous on the upper surface, pubescent on the lower surface mostly on the veins ................................................................................ 2 Leaves glabrous on both sides ................................................................... 3 Leaves elliptic; petiole 2.5–4 cm long; corolla 15–18 mm long ..... C. poeppigii Leaves oblong or elliptic; petiole 5–8 mm long; corolla 9–13 mm long ............................................................................................. C. venezuelense Shrubs or small trees to 16 m tall; glands at the apex of the petiole flat or impressed ................................................................................. C. spinosum Trees 20–30 m tall; glands at the apex of the petiole cup-shaped ........... 4 Leaves oblong-elliptic or subovate, 8–28 cm long, the apex acute-acuminate ................................................................................... C. macrophyllum Leaves obovate-elliptic, 6–7 cm long, the apex rounded ....................C. sp. A

422

V ERBENACEAE

Fig. 352. Citharexylum macrophyllum

Fig. 353. Citharexylum spinosum

Citharexylum macrophyllum Poir. in Lam., Encycl. suppl. 2: 367. 1811. —Totumillo blanco. Citharexylum amazonicum Moldenke, Repert. Spec. Nov. Regni Veg. 37: 216. 1934. Tree 20–30 m tall; corolla white. Evergreen lowland forests, 100–200 m; Delta Amacuro (Caño Arature, Río Grande). Aragua, Falcón; Guyana, Suriname, French Guiana, Ecuador, Brazil (Amazonas, Pará, Roraima). ◆Fig. 352.

Citharexylum poeppigii Walp., Repert. Bot. Syst. 4: 76. 1845. —Cerezillo, Totumillo negro. Citharexylum poeppigii f. anomalum Moldenke, Phytologia 8: 549. 1963. Tree to 25 m tall; corolla white. Evergreen lowland to lower montane forests, 100–900 m; Delta Amacuro (Río Amacuro), Bolívar (El Dorado, El Palmar, Santa Elena de Uairén). Apure, Barinas, Mérida, Táchira; Colombia, Ecuador, Peru, Brazil (Amazonas).

Clerodendrum 423

Citharexylum spinosum L., Sp. Pl. 625. 1753. —Totumillo, Totumillo blanco, Totumillo sabanero. Citharexylum fruticosum L., Syst. Nat. ed. 10, 2: 1115. 1759. Citharexylum fruticosum var. brittonii Moldenke, Lilloa 4: 311. 1939. Shrub or small tree to 16 m tall; corolla white. Deciduous to evergreen lowland to premontane forests, 100–500 m; Delta Amacuro (Pedernales), Bolívar (Altiplanicie de Nuria, El Manteco, El Miamo, Guri, Río Asa, mouth of Río Paragua). Anzoátegui, Aragua, Distrito Federal, Falcón, Nueva Esparta, Sucre; Panama, West Indies, Trinidad, Guyana, Suriname, French Guiana. ◆Fig. 353.

Citharexylum venezuelense Moldenke, Repert. Spec. Nov. Regni Veg. 37: 237. 1934. Small tree to 20 m tall; corolla white. Evergreen lowland forests, ca. 200 m; Bolívar (El Dorado). Apure, Barinas, Distrito Federal, Mérida, Portuguesa, Táchira. Citharexylum sp. A. Tree to 20 m tall. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma). Endemic. This distinct taxon is known from a single collection: Steyermark & Luteyn 129770 (NY, VEN).

6. CLERODENDRUM L., Sp. Pl. 637. 1753, spelling variant: Clerodendron. Large herbs, shrubs, trees, or woody lianas, usually unarmed or sometimes spiny at the base of the petiole. Leaves decussate or in whorls, simple, entire or variously dentate, deciduous. Inflorescences of axillary and/or terminal pedunculate cymes, or the cymes arranged in a terminal panicle, the cymes mostly loose-flowered, sometimes head-like. Flowers pedicellate, bisexual, slightly zygomorphic. Calyx campanulate, regular, 5-lobed or -toothed; corolla simple or double, white, pink, blue-violet, or red, salverform, the tube straight or slightly curved, the limb 5-lobed, lobes subequal or the lower lobe somewhat larger. Stamens 4, didynamous, inserted on the upper part of the corolla tube, long-exserted, involute in bud; anthers oblongovoid, medifixed, 2-locular with parallel thecae, opening by longitudinal slits. Ovary imperfectly 4-locular, each locule with 1 ovule; style with a shortly bifid stigma. Fruit a drupe, globose or obovoid, often 4-sulcate or -lobed, separating into 4 stones at maturity, fruiting calyx somewhat enlarged; exocarp fleshy. Seeds oblong, without endosperm. Pantropics, most abundant in Asia and Africa; ca. 560 species, 9 in Venezuela, 3 of these in the flora area. Key to the Species of Clerodendrum 1. 1. 2(1). 2.

Vines or scandent subshrubs; calyx inflated; corolla red ......... C. thomsonae Subshrubs or shrubs to small trees, 5–7 m tall; calyx not inflated, corolla white, purple, or pink ............................................................................ 2 Subshrubs; petioles 3–25 cm long; leaves widely ovate, 5–28 cm wide, margins dentate; corolla double ......................................... C. philippinum Shrubs to small trees 5–7 m tall; petioles 4–12 cm long; leaves oblongelliptic, 4–7 cm wide, margins entire; corolla simple .......... C. ternifolium

Clerodendrum philippinum Schauer in A. DC., Prodr. 11: 667. 1847. —Melino. Clerodendrum fragrans (Vent.) Willd., Enum. Pl. 659. 1809, as “clerodendron.” —Volkameria fragrans Vent., Jard. Malmaison 2: pl. 70. 1804.

Clerodendrum fragrans var. pleniflora Schauer in A. DC., Prodr. 11: 666. 1847, as “clerodendron.” Shrub 1–3 m tall. Edges of towns, disturbed areas, 100–200 m; Bolívar (El Manteco), Amazonas (Misión San Francisco de

424

V ERBENACEAE

Fig. 354. Clerodendrum philippinum

Guayo). Aragua, Distrito Federal, Falcón, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy; southern China to southern tropical Asia, widely cultivated and naturalized as an ornamental shrub from southern U.S.A. to Chile. ◆Fig. 354. Clerodendrum ternifolium Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 244. 1817. Shrub to small tree. Edges of towns, 100– 200 m; Bolívar (Ciudad Bolívar). Anzoátegui, Falcón, Guárico, Sucre; Colombia.

Clerodendrum thomsonae Balf., Edinburgh New Philos. J. ser. 2, 15: 253. 1862. Vine or climbing shrub; calyx greenish white. Edges of towns, disturbed areas, near sea level to 900 m; Delta Amacuro (Tucupita), Bolívar (El Paují, middle Río Caura), Amazonas (Maroa). Aragua, Distrito Federal, Mérida; tropical West Africa, widely cultivated and naturalized as an ornamental shrub from Central America to northern South America.

7. DURANTA L., Sp. Pl. 637. 1753. Shrubs, often spinose. Leaves decussate or in whorls, simple, entire or dentate-serrate, deciduous. Inflorescences axillary and terminal racemes. Flowers pedicellate, bisexual, ± zygomorphic. Calyx tubular, 5-ribbed, each rib terminating in a tooth; corolla salverform, the tube straight or somewhat curved, the limb oblique, slightly unequally 5-lobed. Stamens 4(5), or the 5th staminodial, didynamous,

Duranta 425

inserted above the middle of the corolla tube, included; anthers ovoid, medifixed or dorsifixed, erect, the thecae divergent. Ovary normally 8-locular, each locule with 1 ovule; style with a capitate stigma. Fruit a drupe, 4-stoned, each stone 2-seeded, fruiting calyx accrescent, flask-shaped, enclosing the fruit. U.S.A. (Florida, Texas), Mexico, Central America, Bermuda, Antilles, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; ca. 40 species, 8 in Venezuela, 4 of these in the flora area. Key to the Species of Duranta 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Leaves glabrous on both sides ................................................................... 2 Leaves sparsely appressed-pubescent on both sides ................................ 3 Leaves coriaceous, entire; calyx glabrous to glabrescent outside ................................................................................................ D. obtusifolia Leaves membranous or chartaceous, serrate mostly in the uppper half of blades; calyx pubescent outside .................................................... D. erecta Spinose vine; leaf apex cuspidate ....................................................... D. sp. A Unarmed shrub; leaf apex acute or acuminate ........................................ 4 Margin of leaf blades flat, not revolute; calyx tube 3–5 mm long ..... D. erecta Margin of leaf blades revolute; calyx tube 7–8 mm long .................. D. sp. B

Duranta erecta L., Sp. Pl. 637. 1753. —Melero. Duranta repens L., Sp. Pl. 637. 1753. Duranta plumieri var. alba Mast., Gard. Chron. ser. 3, 63: 44. 1888. Duranta repens var. canescens Moldenke, Phytologia 1: 436. 1940. Duranta repens var. lopez-palacii Moldenke, Phytologia 26: 177. 1973. Shrub or small tree, often spinose; flowers blue, rarely white. Edges of towns, deciduous to evergreen lowland forests, near sea level to 200 m; Delta Amacuro (Pedernales, Tucupita), Bolívar (Ciudad Bolívar, El Palmar, Puerto Ordaz), Amazonas (Río Cataniapo). Aragua, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, Zulia; U.S.A. (Florida), Mexico, Central America, Antilles, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Brazil, Bolivia, Argentina, Uruguay. ◆Fig. 355. Duranta obtusifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 254. 1817 [1818]. Duranta coriacea Hayek, Repert. Spec. Nov. Regni Veg. 2: 88. 1906. Shrub or small tree, often spinose; flowers blue, rarely white. Semideciduous to lower montane forests, 100–300 m; Bolívar (Altiplanicie de Nuria). Mérida; Colombia, Ecuador, Peru.

Fig. 355. Duranta erecta

426

V ERBENACEAE

Duranta sp. A Spinose vine. Wet tepui summit forest, ca. 2000 m; Bolívar (Macizo del Chimantá [Chimantá-tepui]). Endemic. This taxon is based on Steyermark et al. 128050 (MO, VEN).

Duranta sp. B Shrub to 4 m tall; corolla white with blue. Wet tepui forests, 1800–1900 m; Amazonas (Sierra de la Neblina). Endemic. This taxon is based on Nee 30691 (NY, VEN).

8. LANTANA L., Sp. Pl. 626. 1753; Gen. Pl. ed. 5, 275. 1754. Shrubs or erect herbs, sometimes subscandent, scandent, or prostrate; branches and branchlets ± scabrous and hirtous-pubescent or tomentose with simple trichomes. Leaves opposite or ternate, simple, dentate, often rugose. Inflorescences axillary, solitary, fasciculate, spicate, or capitate, densely flowered racemes; bracts conspicuous. Flowers sessile, bisexual, ± zygomorphic. Calyx tubular, truncate, 2-lobed or 4- or 5-dentate; corolla salverform, the tube straight or somewhat curved, the limb oblique, slightly unequally 4- or 5-lobed. Stamens 4, didynamous inserted about the middle of the corolla tube, included; anthers ovoid, medifixed. Ovary 2-locular, each locule with 1 ovule; style with an oblique, capitate stigma. Fruit a fleshy 2-seeded drupe; exocarp usually ± fleshy (rarely dry); endosperm hard; fruiting calyx not enlarged. Seeds without endosperm. Pantropics, most common in the Neotropics; ca. 240 species, 16 in Venezuela, 9 of these in the flora area. Key to the Species of Lantana 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(3). 5. 6(5). 6. 7(6). 7. 8(6).

8.

Branches, leaves, and peduncles glutinous (sticky to the touch even in herbarium specimens), with trichomes with apical glands ................. 2 Branches, leaves, and peduncles not glutinous, with or without glandular trichomes ................................................................................................ 3 Peduncles 3–15 cm long; corolla yellow, orange, or red .............. L. glutinosa Peduncles 2–2.7 cm long; corolla violet, white, or pink .......... L. ruiz-teranii Corolla yellow, orange, or red .................................................................... 4 Corolla white, violet, or pink ..................................................................... 5 Lower surface of leaves canescent-pubescent; calyx glabrous or sparsely pilose ....................................................................................... L. cujabensis Lower surface of leaves without canescent trichomes, sometimes glabrescent; calyx pubescent, mixed with glandular globules ............ L. camara Peduncle of inflorescence 12–25 cm long .......................... L. achyranthifolia Peduncle of inflorescence 3–10 cm long .................................................... 6 Leaves bullate ............................................................................................ 7 Leaves not bullate ...................................................................................... 8 Branches scabrous-hirsute; leaves ovate ......................................... L. radula Branches sparsely pilose; leaves oblong or lanceolate-triangular ...... L. fucata Leaves with 3 leaflets, the outside sericeous-strigose, the inside subglabrous; lower surface of leaves, outside of bracts, and outside of corolla with orange glandular globules ...................................... L. trifolia Leaves simple; bracts acute or acuminate, appressed-pubescent on both sides; without glandular globules ........................................... L. canescens

Lantana achyranthifolia Desf., École Bot. ed. 3, 392. 1829.

Tabl.

Subshrub or herb; corolla white or violet. Evergreen lowland forests, 100–200 m;

Lantana 427

Bolívar (Río Parguaza), Amazonas (Río Manaviche). Aragua, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Guatemala, Honduras, Colombia. ◆Fig. 356. Lantana camara L., Sp. Pl. 627. 1753. —Cariaquito, Cariaquito colorado, Fiikcki (Yanomami), Toqui-voquiris (Yekwana). Lantana aculeata L., Sp. Pl. 627. 1753. —Lantana camara var. aculeata (L.) Moldenke, Torreya 34: 9. 1934. Lantana mista L., Syst. Veg. ed. 2, 417. 1767. —Lantana camara var. mista (L.) L.H. Bailey, Cycl. Amer. Hort. 2: 884. 1900. Lantana moritziana Otto & A. Dietr., Allg. Gartenzeit 9: 372. 1841. —Lantana camara var. moritziana (Otto & A. Dietr.) López-Pal., Revista Fac. Farm. Univ. Andes 1421. 1974. Lantana armata Schauer in A. DC., Prodr. 11: 597. 1847. Lantana armata var. guianensis Moldenke, Phytologia 51: 244.1982. Aromatic shrub or subshrub to 3 m tall; corolla yellow, orange, or red. Edges of towns, disturbed areas, 50–1000 m; Delta Amacuro (Río Cuyubini basin, Tucupita), Bolívar (Ciudad Bolívar, Gran Sabana, Isla Anacoco, Río Canaracuni, Río Pao, Río Parguaza, Río Yuruaní, Santa Elena de Uairén, Upata), Amazonas (Caño Caname, Isla Ratón, Río Cunucunuma, upper Río Siapa, Río Ventuari, San Antonio, San Carlos de Río Negro, San Fernando de Atabapo, Santa Bárbara del Orinoco). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Acré, Minas Gerais, Pará, Paraná), Bolivia, Paraguay, Argentina. ◆Fig. 358. The branches of Lantana camara are used to make a tea to treat fever. The plant is also used as an ornamental shrub. This is a very variable species in indument, inflorescence type, and the color of flowers. Lantana canescens Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 259. 1817 [1818].

—Cariaquito blanco. Subshrub or herb; corolla white or pale violet. Edges of gallery forests and savannas, disturbed areas, near sea level to 900 m; Delta Amacuro (Tucupita), Bolívar (Altiplanicie de Nuria, Cerro Bolívar, Ciudad Bolívar, El Manteco, El Palmar and south at Mina de El Manganeso, Isla Anacoco, Santa Elena de Uarién, Upata). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Sucre, Táchira, Zulia; Mexico, Colombia, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia, Argentina. Lantana cujabensis Schauer in A. DC., Prodr. 111: 599. 1847. Subshrub; corolla yellow or orange. Edges of forests and towns, 100–400 m; Bolívar (Santa María de Erebato), Amazonas (San Carlos de Río Negro, San Fernando de Atabapo). Barinas, Mérida; Colombia, Ecuador, Peru, Brazil, Bolivia. Lantana fucata Lindl., Bot. Reg. 10: 798. 1824. Subshrub; corolla violet. Edges of towns, ca. 50 m; Bolívar (Ciudad Bolívar). Aragua, Carabobo, Distrito Federal, Lara, Mérida, Miranda, Nueva Esparta, Táchira, Trujillo, Yaracuy; Mexico, Jamaica, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Lantana glutinosa Poepp., Allg. Gartenzeitung 10: 315. 1842. —Cariaquito. Subshrub or shrub, sometimes armed; corolla yellow or red. Edges of towns, savannas, forests, near sea level to 800 m; Delta Amacuro (Caño Guiniquina), Bolívar (Ciudad Bolívar, Gran Sabana, Maripa, middle Río Orinoco). Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Portuguesa, Trujillo, Zulia; Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Lantana radula Sw., Prodr. 92. 1788. Shrub; corolla white or pink with yellow throat. Upland savannas, 1000–1300 m; Bolívar (Kavanayén, Santa Elena de Uairén, Uarama-tepuí in northeastern Gran Sabana). Antilles, Guyana, Brazil (Bahia). ◆Fig. 359.

428

V ERBENACEAE

Lantana ruiz-teranii López-Pal. & Steyerm., Acta Bot. Venez. 14(1): 163. 1983. Armed subshrub; corolla violet, white, or pink. Upland savannas, 900–1200 m; Bolívar (Kavanayén, San Ignacio de Yuruaní, Santa Elena de Uairén). Endemic. ◆Fig. 357. Lantana trifolia L., Sp. Pl. 626. 1753. —Cariaquito morado. Lantana trifolia var. rigiduscula Briq., Annuaire Conserv. Jard. Bot. Genève 7– 8: 303. 1904. Lantana trifolia f. hirsuta Moldenke, Phytologia 3: 113. 1949. Lantana trifolia f. oppositifolia Moldenke, Phytologia 4: 179. 1953. Lantana trifolia f. rubierensis Moldenke, Phytologia 9: 187. 1963. Subshrub or herb; corolla blue, pink, or violet. Savannas, granitic outcrops, disturbed areas, 100–900 m; Bolívar (El Manteco, Río Paragua, Santa Elena de Uairén, Santa María de Erebato), Amazonas (Isla Ratón, Puerto Ayacucho). Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Miranda, Portuguesa, Sucre, Táchira, Trujillo; Mexico, Central America, Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 360. The branches of Lantana trifolia are used to make a tea to treat fever.

Fig. 357. Lantana ruiz-teranii

Fig. 356. Lantana achyranthifolia

Fig. 358. Lantana camara

Lippia 429

Fig. 359. Lantana radula

Fig. 360. Lantana trifolia

9. LIPPIA L., Sp. Pl. 633. 1753. Erect bushes, shrubs, subshrubs, or herbs, rarely trees, glabrous or variously pubescent with simple trichomes, often hirsute or tomentose. Leaves decussate or ternate, simple, entire, dentate, or lobed, exstipulate, petiolate or sessile, thinmembranous to heavy-coriaceous, the upper surface rugose or not; venation mostly pinnate. Inflorescences axillary or terminal, solitary or fasciculate, spicate or capitate, densely flowered racemes, the racemes sometimes arranged in panicles; bracts conspicuous. Flowers small, sessile, usually bisexual, ± zygomorphic. Calyx small, membranous tubular-campanulate, sometimes 2-lipped or 4-toothed; corolla salverform, the tube straight or somewhat curved, the limb oblique, unequally 4- or 5lobed. Stamens 4, didynamous, inserted near the middle of the corolla tube, included or slightly exserted; anthers ovoid, medifixed unappendaged, the thecae parallel, often aborted or absent on pistillate plants. Ovary globose, 2-locular, each locule with 1 ovule, basal and erect or affixed laterally near the base; style often short; stigma capitate. Fruit a small, dry, 2-parted schizocarp, fruiting calyx not enlarged. Seeds without endosperm. Pantropics, mostly Neotropics; ca. 250 species, 7 in Venezuela, 3 of these in the flora area. Key to the Species of Lippia 1. 1.

Calyx ca. 2 mm long; corolla outside without glandular globules ...... L. alba Calyx ca. 1 mm long; corolla outside with glandular globules ................ 2

430

2(1). 2.

V ERBENACEAE

Petioles 0–1 mm long; leaves narrowly obovate; bracts of inflorescences narrowly elliptic, apex obtuse; fruit cordiform ..................... L. micromera Petioles 1–10 mm long; leaves ovate to narrowly ovate or elliptic; bracts of inflorescences ovate, the apex acute-acuminate; fruit subglobose ............................................................................................... L. origanoides

Lippia alba (Mill.) N.E. Br. in Britton and P. Wilson, Bot. Porto Rico 6: 141. 1925. —Lantana alba Mill., Gard. Dict. ed. 8. 1768. Lippia geminata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 266. 1817 [1818]. —Lantana geminata (Kunth) Spreng., Syst. Veg. 2: 763. 1825. Aromatic subshrub or herb; flowers pink, violet, blue, or white, the throat usually yellow. Edges of forests and savannas, disturbed areas, on granitic outcrops, near sea level to 200 m; Delta Amacuro (Caño Mánamo, Tucupita), Bolívar (Ciudad Bolívar, Moitaco, Río Orinoco at mouth of Río Cinaruco), Amazonas (Puerto Ayacucho, San Fernando de Atabapo). Apure, Aragua, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, Antillas, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Lippia alba is widely cultivated as an ornamental or as a treatment for stomach illnesses. Lippia micromera Schauer in A. DC., Prodr. 11: 587. 1847. —Oregano. Aromatic subshrub to 2 m tall; flowers white. Edges of towns and disturbed areas, near sea level to 200 m; Delta Amacuro (between Tucupita and Pedernales), Bolívar (Ciudad Bolívar), Amazonas (Río Cuao). Aragua, Distrito Federal, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Sucre, Táchira, Trujillo; Antilles, Guyana. ◆Fig. 361. Lippia micromera is sometimes cultivated as a food condiment or as a medicine for stomach illnesses. Lippia origanoides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 267. 1817 [1818]. —Orégano. Lippia schomburgkiana Schauer in A. DC., Prodr. 11: 577. 1847. Aromatic shrub or subshrub to 3.5 m tall; flowers white, the throat yellow. Open areas, savannas, edges of towns with Curatella

Fig. 361. Lippia micromera

Fig. 362. Lippia origanoides

Petrea 431

americana forests, 100–1500 m; Bolívar (Auyán-tepui, Cerro Altamira east of Ciudad Piar, Cerro Bolívar, Cerro Cuchivero, Ciudad Bolívar, islands in Lago Guri, Kavanayén, La Vergareña, Puerto Ordaz, Río Parguaza, Río Suapure, upper Río Túriba north of Túriba, Río Yuruaní, base of Roraima-tepui, San Pedro de las Bocas, Santa Elena de Uairén).

Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Táchira, Zulia; Trinidad, Colombia, Guyana, Brazil. ◆Fig. 362. Lippia origanoides is sometimes cultivated as a food condiment or a medicine for stomach illnesses.

10. PETREA L., Sp. Pl. 626. 1753. Shrubs, trees, or woody lianas. Leaves decussate or in whorls, deciduous, exstipulate, simple. Inflorescences axillary and terminal racemes; bracts mostly inconspicuous. Flowers pedicellate, bisexual, actinomorphic or sometimes zygomorphic; torus distinct, mostly swollen like a ring. Calyx violet, blue, white, or pale green, campanulate, 5-lobed, the lobes mostly equal and longer than the tube, with a corona inside at the base of the lobes, 5-lobed or -sinuate; corolla salverform, 5lobed, the lobes subequal, rounded. Stamens 4, inserted near the middle of the corolla tube, included and equal in size; filaments short and slender; anthers oblong or ovoid, medifixed; sometimes 1 staminode present. Ovary subglobose or oblong, borne on a ± conspicuous disk, ± completely 2-locular, the ovules 1 per locule; style with a small capitate stigma. Fruit a drupe, 2-stoned, enclosed in the calyx tube, fruiting calyx accrescent, the lobes stiffened, acting as wing for the fruit, calycinal corona curved inward and closing the mouth of the calyx. Seeds laterally or apically attached, lacking endosperm. Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 14 species, 6 in Venezuela, 5 of these in the flora area. Key to the Species of Petrea 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

Erect shrubs or trees to 20 m tall; branchlets densely pubescent; filaments glabrous; style pubescent ............................................. P. pubescens Vining shrubs or woody lianas; branchlets glabrous, puberulent or sparsely pilose; filaments sparsely pilose; style glabrous ................... 2 Lower surface of leaves ± pitted due to enlarged veins ....... P. macrostachya Lower surface of leaves not pitted and without enlarged veins .............. 3 Lower surface of leaves bullate when mature; ovary obovate ..... P. bracteata Lower surface of leaves not bullate when mature; ovary oblong ............. 4 Leaves membranous, smooth to the touch or very slightly roughened; calyx lobes obovate ............................................................. P. blanchetiana Leaves subcoriaceous or coriaceous, rough to the touch; calyx lobes oblong ............................................................................................. P. volubilis

Petrea blanchetiana Schauer in A. DC., Prod. 11: 617. 1847. —Tostadito. Woody liana; corolla fragrant, blue. Evergreen lowland forests, 100–200 m; Bolívar (Salto Pará), Amazonas (Río Mawarinuma). Panama, Colombia, French Guiana, Ecuador, Peru, Brazil.

Petrea bracteata Steud., Flora 26: 764. 1843. Woody liana; corolla blue, the throat white. Evergreen lowland forests, 100–200 m; Bolívar (Río Botanamo, Río Venamo), Amazonas (Río Cunucunuma, Río Padamo, Río Siapa, Río Ventuari, Tamatama).

432

V ERBENACEAE

Fig. 363. Petrea volubilis

Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 364. Petrea macrostachya Benth., Ann. Nat. Hist. 2: 448. 1839. Woody liana; corolla blue to violet-purple. Evergreen lowland forests, 100–300 m; Amazonas (Río Jenita on Río Ocamo, Río Matacuni, Río Mavaca, Río Padamo). Guyana, Suriname, French Guiana, Brazil. Petrea pubescens Turcz., Bull. Soc. Imp. Naturalistes Moscou 36: 211. 1863. —Tostadito. Petrea glandulosa Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 70. 1926. Petrea pubescens var. albicalyx Moldenke, Phytologia 18: 72. 1969. Tree to 20 m tall; corolla white to purple. Deciduous or semideciduous forests, 50–200 m; Bolívar (Caicara to El Burro), Amazonas (Puerto Ayacucho). Apure, Aragua, Barinas, Carabobo, Lara, Mérida, Portuguesa, Táchira, Trujillo; Colombia, Ecuador, Peru, Brazil, Bolivia. Petrea volubilis L., Sp. Pl. 626. 1753. —Bejuco mayo, Moradito, Nazareno, Tostadito. Petrea arborea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 282. 1817 [1818].

Petrea kohautiana C. Presl, Abh. Königl. Böhm. Ges. Wiss. ser. 5, 3: 529. 1845. Petrea aspera Turcz., Bull. Soc. Imp. Naturalistes Moscou 36: 211. 1863. Woody liana or shrub; flowers blue or white. Evergreen lowland forests, 50–400 m; Delta Amacuro (Los Castillos de Guayana,

Petrea 433

Fig. 364. Petrea bracteata

Río Toro), Bolívar (Ciudad Bolívar, El Palmar, Isla Anacoco, Río Botanamo, Tumeremo, Upata), Amazonas (Puerto Ayacucho to Gavilán, Quebrada El Grillo between Sama-

riapo and San Fernando de Atabapo). Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Mexico, Central America, Antilles, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 363. Petrea volubilis is widely cultivated as an ornamental.

434

V ERBENACEAE

11. PHYLA Lour., Fl. Cochinch. 66. 1790. Creeping or procumbent herbs, sometimes subshrubs, glabrous often hirsute or tomentose with malpighiaceous trichomes. Leaves opposite, simple, entire or dentate, exstipulate, petiolate or sessile. Inflorescences spicate, axillary, spikes short, cylindric, never arranged in corymbs or panicles. Flowers small, sessile, embedded in the axil of small cuneate-ovate bracts, usually bisexual, ± zygomorphic. Calyx small, membranous, tubular-campanulate, sometimes 2-lipped or 4-toothed; corolla salverform, the tube straight or somewhat curved, the limb oblique, unequally 4-lobed. Stamens 4, didynamous, inserted near the middle of the corolla tube, included or slightly exserted; anthers ovoid, medifixed unappendaged, the thecae parallel, often aborted or absent on pistillate plants. Ovary globose, 2-locular, each locule with 1 ovule, basal and erect or affixed laterally near the base; style often short; stigma 2-lobed. Fruit a small, dry, 2-parted schizocarp when mature, fruiting calyx enlarged and covering the fruit. Seeds 2, without endosperm. Canada, U.S.A., New World tropics and subtropics; ca 10 species, 6 in Venezuela, 1 of these in the flora area. Phyla betulifolia (Kunth) Greene, Pittonia 4: 48. 1899, “betulaefolia.” —Lippia betulifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 264. 1817 [1818]. —Hierbabuena de laguna. Creeping herb to 50 cm tall, rooting at the nodes; flowers white or purple. Edges of wet areas, savannas, disturbed areas, near sea level to 200 m; Delta Amacuro (Boca Macareo, Caño Macareo, Curiapo, Río Cuyubini basin, Tucupita), Bolívar (Cerro Baraguán along Río Orinoco, Ciudad Bolívar, El Tigre, Río Cuchivero, Río Parguaza, Upata), Amazonas (Atures, San Fernando de Atabapo). Apure, Barinas, Carabobo, Cojedes, Guárico, Monagas, Portuguesa, Zulia; Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 365.

Fig. 365. Phyla betulifolia

12. PRIVA Adans., Fam. Pl. 2: 505. 1763. Perennial herbs, stems, branches and branchlets ± tetragonal, often decumbent. Leaves decussate or subopposite, simple, exstipulate, sessile or petiolate, thinmembranous, mostly dentate-serrate. Inflorescences racemes, mostly terminal, sometimes axillary; bracts not conspicuous. Flowers pedicellate or subsessile, bisexual, ± zygomorphic. Calyx tubular, 5-toothed; corolla white, blue, rose, or violet, salverform or funnelform, the tube straight or somewhat curved, the limb 5-lobed, 2-lipped, the upper lip 3-lobed with one large central lobe and 2 medium-sized lateral lobes, the lower lip 2-lobed with 2 equal small lobes. Stamens 4, didynamous,

Priva 435

inserted near the middle of the corolla tube, included; anthers oblong or ovoid, dorsifixed, staminode sometimes present. Ovary (2–)4-locular, each locule with 1 ovule; style with an unequally 2-lobed stigma. Fruit a dry, sometimes woody, 2parted schizocarp, echinate or ridged outside, fruiting calyx accrescent, enclosing the fruit. Seeds lacking endosperm. Pantropics; ca. 25 species, 1 in Venezuela. Priva lappulacea (L.) Pers., Syn. Pl. 2: 139. 1806. —Priva lappulacea L., Sp. Pl. 19. 1753. Priva lappulacea f. albiflora Moldenke, Phytologia 17: 114. 1968. Perennial herb, sometimes decumbent; flowers white, blue, lavender, violet, or pink. Edges of towns, disturbed areas, near sea level to 500 m; Delta Amacuro (Pedernales, Sacupana), Bolívar (Altiplanicie de Nuria, Ciudad Bolívar, San Felíx). Anzoátegui, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Monagas, Nueva Esparta, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 366.

Fig. 366. Priva lappulacea

436

V ERBENACEAE

13. STACHYTARPHETA Vahl, Enum. Pl. 1: 205. 1804, nom. cons. Annual or perennial herbs or shrubs, glabrous, sparsely pilose to hirsute-pubescent. Leaves opposite, alternate, or sometimes in whorls, simple, dentate, often rugose. Inflorescences terminal, spicate, the spikes elongate or abbreviated, densely or loosely many-flowered, occasionally few-flowered; bracts mostly not conspicuous. Flowers sessile, ± embedded in the rachis, bisexual, ± zygomorphic. Calyx tubular, 5-ribbed and unequally 5-toothed; corolla white, blue, purple, pink, or red, salverform, the tube straight or curved, the limb slightly unequally 5-lobed. Stamens 2, inserted above the middle of the corolla tube, included; anthers ellipsoid, dorsifixed, the thecae divaricate, one above the other, staminodes 2. Ovary 2-locular, each locule with 1 ovule; style with a minute capitate stigma. Fruit a dry 2-parted schizocarp, oblong-linear, fruiting calyx enclosing the fruit. Seeds erect, linear, without endosperm. Mexico, Central America, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, naturalized in tropical Asia, Africa, and Oceania; ca. 140 species, 8 in Venezuela, 6 of these in the flora area. Key to the Species of Stachytarpheta 1. 1.

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

5.

Branches, leaves, and peduncles of inflorescences densely white-pubescent; bracts linear-lanceolate, ca. 1 mm wide ................... S. lopez-palacii Branches, leaves, and peduncles of inflorescences glabrous to tomentosehirsute, without white pubescence; bracts lanceolate, ovate-elliptic, or narrowly ovate to subulate, 2–12 mm wide ......................................... 2 Branches tomentose-hirsute; corolla with glandular hairs outside; filaments and staminodes ca. 3 mm long ................................................... 3 Branches glabrous or sparsely tomentose-hirsute; corolla glabrous outside; filaments and staminodes 0.5–1 mm long ................................... 4 Bracts sparsely hirsute and puberulent with disk-shaped glands near the apex outside; corolla blue ............................................................ S. sprucei Bracts pubescent-hirsute without disk-shaped glands near the apex outside; corolla red or pink ........................................................... S. mutabilis Leaves linear, narrowly elliptic, 0.2–3 cm wide ..................... S. angustifolia Leaves ovate or elliptic, 2.5–6 cm wide ..................................................... 5 Upper surface of leaves sparsely strigillose, the lower surface strigose; calyx 4–5 mm long, hirsute-pubescent outside; filaments glabrous or sparsely pilose, ca. 0.5 mm long .......................................... S. cayennensis Upper surface of leaves glabrous or subglabrous, the lower surface sparsely strigillose; calyx 6–7 mm long, glabrous outside, pubescent only near the apex; filaments pubescent, ca. 1 mm long .... S. jamaicensis

Stachytarpheta angustifolia (Mill.) Vahl, Enum. Pl. 1: 205. 1804. —Verbena angustifolia Mill., Gard. Dict. ed. 8, no. 15. 1758. —Verbena. Stachytarpheta angustifolia var. elatior Schrad. ex Schult., Mant. 1: 172. 1822. —Stachytarpheta angustifolia f. elatior (Schrad. ex Schult.) López-Pal., Revista Fac. Farm. Univ. Andes 15: 79. 1974.

Stachytarpheta surinamensis Miq. ex Pulle, Enum. Vasc. Pl. Surinam 402. 1906, nom. nud. Stachytarpheta elatior var. jenmanii Moldenke, Phytologia 1: 472. 1940. Herb or small shrub; corolla blue. Edges of savannas, wet areas, swamps and marshes, 50–300 m; Bolívar (Altiplanicie de Nuria, Ciudad Bolívar, Moitaco, middle Río Orinoco,

Stachytarpheta 437

Fig. 367. Stachytarpheta angustifolia

Upata). Anzoátegui, Apure, Aragua, Carabobo, Cojedes, Falcón, Guárico, Mérida, Monagas, Portuguesa; Honduras, Jamaica, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 367. Stachytarpheta cayennensis (Rich.) Vahl, Enum. Pl. 1: 208. 1804. —Verbena cayennensis Rich., Actes Soc. Hist. Nat. Paris 1: 105. 1792. —Verbena. Verbena dichotoma Ruiz & Pav., Fl. Peruv. 1: 23. 1798. —Stachytarpheta dichotoma

Fig. 368. Stachytarpheta sprucei

(Ruiz & Pav.) Vahl, Enum. Pl. 1: 207. 1804. Stachytarpheta cayennensis f. albiflora Moldenke, Bot. Gaz. 106: 163. 1933. —Stachytarpheta dichotoma f. albiflora (Moldenke) Moldenke, Phytologia 28: 426. 1974. Herb or small shrub; corolla white, pale blue, or blue. Edges of towns, savannas, disturbed areas, 50–200 m; Delta Amacuro

438

V ERBENACEAE

(Caño Mánamo, Pedernales, Sacupana), Bolívar (Ciudad Bolívar, El Dorado, La Paragua, La Prisión, Río Caura, Río Cuchivero, Río Venamo), Amazonas (Río Caname, San Fernando de Atabapo, Yavita). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Stachytarpheta jamaicensis (L.) Vahl, Enum. Pl. 1: 206. 1804. —Verbena jamaicensis L., Sp. Pl. 19. 1753. —Verbena. Stachytarpheta pilosiuscula Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 279. 1817 [1818]. Herb or small shrub; corolla blue, violet, or purple. Edges of towns, savannas, disturbed areas, 50–200 m; Delta Amacuro (Misión San Francisco de Guayo, Pedernales, Sacupana, Tucupita), Bolívar (Ciudad Bolívar, Tumeremo), Amazonas (Río Guainía). Monagas, Nueva Esparta, Sucre; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador. Stachytarpheta lopez-palacii Moldenke, Phytologia 31: 27. 1975.

Herb or subshrub; corolla pale blue. Upland savannas, disturbed areas, 100–900 m; Bolívar (Villa Lola). Sucre. Stachytarpheta mutabilis (Jacq.) Vahl, Enum. Pl. 1: 208. 1804. —Verbena mutabilis Jacq., Collectanea 2: 335. 1788. Herb or small shrub; corolla red or pink. Edges of rivers, disturbed areas, 100–200 m; Bolívar (La Prisión). Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Lara, Mérida, Miranda, Portuguesa, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana. Stachytarpheta sprucei Moldenke, Phytologia 1: 474. 1940. —Adéketón-yek (Pemón), Ehchipipin (Panare). Stachytarpheta roraimensis Moldenke, Phytologia 1: 458. 1940. Stachytarpheta roraimensis var. pubescens Moldenke, Phytologia 3: 117. 1949. Shrub to 4 m tall; corolla blue. Open areas, savannas, igneous outcrops, 50–1000 m; Bolívar (Caicara del Orinoco, Caño Canaima, Gran Sabana, Kamarata, Río Parguaza, Santa Elena de Uairén), Amazonas (Puerto Ayacucho). Colombia, Guyana, Brazil (Roraima). ◆Fig. 368.

14. TAMONEA Aubl., Hist. Pl. Guiane 659. 1775. Ghinia Schreb., Gen. Pl. 1: 19. 1789. Herbs or subshrubs. Leaves decussate, simple, entire or dentate-serrate. Infloresences axillary or terminal racemes; bracts not conspicuous. Flowers pedicellate or sessile, bisexual, ± zygomorphic. Calyx tubular-campanulate, 5-toothed; corolla blue, violet, or white, funnelform or salverform, the tube straight, the limb unequally (4)5-lobed. Stamens 4, didynamous, inserted near the middle of the corolla tube, included, the 2 upper stamens with a glandular thickening; anthers ovoid, dorsifixed. Ovary 4-locular, each locule with 1 ovule; style with an oblique, oblong stigma. Fruit a drupe, fruiting calyx accrescent, cyathiform; exocarp thin, fleshy; endocarp hard. Neotropics; ca 10 species, 1 in Venezuela. Tamonea spicata Aubl., Hist. Pl. Guiane 659. 1775. —Ghinia spicata (Aubl.) Moldenke, Phytologia 1: 169. 1935. —Borrachera. Herb or subshrub; flowers blue or violet, the throat yellow inside. Edges of towns, Trachypogon savannas, disturbed areas, 50– 1400 m; Bolívar (Altiplanicie de Nuria, El

Dorado, Gran Sabana, Isla Anacoco, Puerto Ordaz, San Félix, Santa Elena de Uairén, Tumeremo, Upata). Anzoátegui, Monagas; Mexico, Central America, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 369. Tamonea spicata is reportedly poisonous to cattle.

Vitex 439

Fig. 369. Tamonea spicata

15. VITEX L., Sp. Pl. 638. 1753. Trees or shrubs. Leaves glabrous, tomentose, or villous, decussate or ternate, palmately compound, 3–7-foliolate, rarely 1-foliolate; leaflets chartaceous or membranous, sometimes coriaceous, mostly petiolate, entire or dentate. Inflorescences axillary and/or terminal cymes, solitary or arranged in panicles, the cymes divaricate, umbelliform, or head-like; bracts mostly not conspicuous. Flowers pedicellate or sessile, bisexual, zygomorphic. Calyx campanulate to tubular, 5-lobed or -toothed; corolla white, violet, or blue, bearded or villous on the largest lobe near the mouth, salverform, the tube straight, the limb obliquely 5-lobed, the 2 upper and 2 lateral lobes subequal, the lower lobe largest. Stamens 4, didynamous, inserted somewhat above the base of the corolla tube and sericeous around the insertion; anthers ovoid, dorsifixed. Ovary (2)4-locular, each locule with 1 ovule; style with a short bifid stigma. Fruit a drupe; exocarp fleshy, fruiting calyx somewhat enlarged, often accrescent. Seeds obovate or oblong, erect, without endosperm. Tropical and subtropical regions, with a few species in the temperate regions of Europe and Asia; ca. 350 species, 11 in Venezuela, 9 of these in the flora area.

440

V ERBENACEAE

Key to the Species of Vitex 1.

1.

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

4.

5(2). 5. 6(5). 6. 7(6). 7. 8(7). 8.

Branchlets, petioles, and inflorescences with golden brown pubescence; calyx 15–25 mm long; corolla exterior densely appressed golden brown villous ........................................................................................... V. triflora Branchlets, petioles, and inflorescences without golden brown pubescence, sparsely pubescent to glabrous; calyx 1–7 mm long; corolla sparsely pubescent, if densely appressed not golden brown villous outside ......................................................................................................... 2 Inflorescences capituliform or umbelloid cymes ....................................... 3 Inflorescences thyrsoid, paniculate, or cymes divaricate ......................... 5 Trees 30–40 m tall; leaves absent or nearly so during the flowering period; cymes not dense .............................................................. V. stahellii Shrub to small trees 5–15 m tall; mature leaves present during the flowering period; cymes compact ................................................................. 4 Branches subglabrous; leaves caudate, subglabrescent except on the veins on the lower surface; calyx outside sparsely appressed-pubescent; fruit ca. 8 mm long ............................................................. V. capitata Branches shortly pubescent; leaves acuminate or long acuminate, the lower surface velutinous-tomentose; calyx densely pubescent outside; fruit ca. 13 mm long ...................................................... V. schomburgkiana Corolla white, white-sericeous outside, 4–6 mm long .................... V. sprucei Corolla blue or violet, glabrescent, pilosulous to densely appressed-pubescent, not white-sericeous outside, 8–13 mm long ........................... 6 Petiole disk-shaped, enlarged at the apex.. ............................... V. compressa Petiole without disk, not enlarged at the apex ......................................... 7 Leaflets coriaceous; inflorescences a thyrsoid- panicle, 20–36 cm long ................................................................................................. V. calothyrsa Leaflets chartaceous or membranous; inflorescences loose spreading cymes, 4–20 cm long .............................................................................. 8 Petioles strigulose to glabrescent, slighty flabelliform at the apex; inflorescences 4–9 cm long; calyx 4–5 mm long ................................... V. klugii Petioles puberulent, entire at the apex; inflorescences 12–20 cm long; calyx 1–3 mm long ................................................................. V. orinocensis

Vitex calothyrsa Sandwith, Bull. Misc. Inform. 1930: 157. 1930. —Cedrillo. Vitex pacimonensis Spruce ex Sandwith, Bull. Misc. Inform. 1930: 157. 1930, as synonym. Tree to 25 m tall; corolla blue or blue-violet. Evergreen lowland forests, sand savannas, 100–200 m; Amazonas (Caño Culebra, Caño San Miguel, Isla Ratón, Río Baría, Río Casiquiare, Río Pasimoni, Río Siapa, Río Yatúa, San Carlos de Río Negro, Trapichote). Colombia, Brazil. ◆Fig. 371.

Vitex capitata Vahl, Eclog. Amer. 2: 50, pl. 18. 1798. —Escobillo, Guarataro, Piedrero, Piqueguaro, Serae-serae (Panare), Totumillo morado. Small tree to 15 m tall; corolla blue or purple. Semideciduous to evergreen forests, sometimes in rocky places, 50–500 m; Bolívar (Caicara del Orinoco, Canaima, Guri, La Paragua, Maripa, Puerto Ordaz, Río Asa, middle Río Caura, Río Orocopiche near Ciudad Piar, Río Paragua near Raudal Auraima), Amazonas (Puerto Ayacucho, Río Coro Coro west of Cerro Yutajé). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Guárico,

Vitex 441

Lara, Monagas, Portuguesa, Sucre; Trinidad, Suriname, Colombia, Brazil. ◆Fig. 373. Vitex compressa Turcz., Bull. Soc. Imp. Naturalistes Moscou 36: 224. 1863. —Cenicero, Guarataro, Guarataro lechoso, Totumillo. Tree to 30 m tall; corolla blue or violet. Semideciduous to evergreen forests, 50–600 m; Bolívar (Altiplanicie de Nuria, El Dorado, El Manteco, El Pao, Guri, Puerto Ordaz, Río Asa, Río Botanamo, Río Cuyuní, Río Oris in Río Paragua basin, Río Orocopiche near Ciudad Piar, Río Supamo, Río Venamo), Amazonas (Macabana in Río Ventuari basin, upper Río Cataniapo basin). Anzoátegui, Aragua, Barinas, Carabobo, Falcón, Guárico, Lara, Miranda, Monagas, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana. ◆Fig. 372. Vitex klugii Moldenke, Phytologia 1: 101. 1934. —Conejemuro (Bale), Palo de arco morado, Palo de hambre. Tree to 20 m tall; corolla blue. Evergreen lowland forests, 100–700 m; Amazonas (Río Atacavi, upper Río Yatúa on Sierra de la Neblina, San Carlos de Río Negro, San Fernando de Atabapo, San Juan de Manapiare, 2.5 km west of Yavita). Peru. ◆Fig. 375. Vitex orinocensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 247. 1817 [1818]. Neotropics; 2 varieties, both in the flora area. Key to the Varieties of V. orinocensis 1. Apex acuminate, acute, or obtuse .......... .................................... var. multiflora 1. Apex long-acuminate or caudate ............ ................................... var. orinocensis V.

orinocensis var. multiflora (Miq.) Huber, Bol. Mus. Paraense Hist. Nat. 5: 217. 1909. —Vitex multiflora Miq., Linnaea 18: 739. 1844. —Guarataro, Palo de arco, Sesejudi (Yekwana), Totumillo, Waka-dame (Arekuna). Shrub or tree to 20 m tall; corolla blue or purple. Semideciduous to evergreen lowland forests, 50–900 m; Bolívar (Caicara del

Orinoco, Maripa, Río Caura, Río Hacha in Río Icabarú basin, Río Paramichí, Río Suapure), Amazonas (Caño Marieta, Cariche, Puerto Ayacucho, Río Casiquire). Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 374. V. orinocensis var. orinocensis Small tree; corolla blue or purple. Semideciduous forests, 100–200 m; Amazonas (near confluence of Río Meta with Río Orinoco). Guárico; Colombia. Vitex schomburgkiana Schauer, Linnaea 20: 483. 1847. Vitex appunii Moldenke, Phytologia 1: 484. 1941. Shrub or tree to 20 m tall; corolla blue or purple. Expected in eastern Bolívar state. Apure, Guárico; Guyana (Cerro Roraima base, Rupununi Savanna), Brazil (Roraima). Outside the flora area, Vitex schomburgkiana is known from riparian forests and edges of savannas at 100 to 1000 m. Vitex sprucei Briq., Bull. Herb. Boissier sér. 1, 4: 346. 1896. —Marfíl. Vitex spongiocarpa Ducke, Trop. Woods 31: 20. 1932. Vitex spongiocarpa var. longidentata Moldenke, Phytologia 2: 31. 1941. Vitex sprucei var. longidentata Moldenke, Phytologia 2: 31. 1941. Vitex sprucei var. vaupesensis Moldenke, Phytologia 6(6): 332. 1958. Tree to 30 m tall; corolla white. Evergreen lowland forests, shrubby white-sand savannas, 100–200 m; Amazonas (Río Pasimoni). Colombia, Brazil (Amazonas). Vitex stahelii Moldenke in Pulle, Fl. Suriname 4(22): 310. 1940. —Totumillo, Totumillo morado. Tree to 40 m tall; corolla blue. Evergreen lowland forests, 100–400 m; Delta Amacuro (Los Castillos de Guayana, Serranía de Imataca), Bolívar (El Palmar, mouth of Río Paragua, Río Toro). Anzoátegui; Guyana, Suriname, French Guiana. ◆Fig. 376.

442

V ERBENACEAE

Fig. 370. Vitex triflora

Fig. 371. Vitex calothyrsa

Vitex 443

Fig. 372. Vitex compressa

Fig. 373. Vitex capitata

444

V ERBENACEAE

Fig. 374. Vitex orinocensis var. multiflora

Fig. 375. Vitex klugii

V IOLACEAE 445

Fig. 376. Vitex stahelii

Vitex triflora Vahl, Eclog. Amer. 2: 49. 1798. —Guarataro, Palo de arco, Palo de arco morado, Waranshikuai (Yekwana). Vitex triflora var. quinquefoliolata Moldenke, Phytologia 1: 104. 1934. —Vitex triflora f. quinquefoliolata (Moldenke) Moldenke, Phytologia 44: 384. 1979. Small tree to 15 m tall; corolla blue, violet, or purple. Evergreen lowland forests, 100– 300 m; Amazonas (Culebra, Mavaca, Río

Atacavi, Río Casiquiare, Río Cataniapo, Río Mawarinuma, Río Yureba, Santa Bárbara del Orinoco). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Acré, Amapá, Amazonas, Pará). ◆Fig. 370. The scraped bark of Vitex triflora is placed into alcoholic beverages such as rum or red wines and is used to treat rheumatism and other body pains.

VIOLACEAE by Ángel Fernández del Valle Herbs, subshrubs, shrubs, climbers, or trees. Leaves alternate, opposite, or rosulate (not in the flora area), sometimes distichous, the blades simple, entire or toothed, sometimes lobulate, pinnativeined, rarely basiveined, without indument or with simple hairs; stipules persistent or caducous. Inflorescences axillary or terminal, sometimes cauliflorous, in spikes, racemes, or panicles, sometimes flowers solitary.

445

Fig. 376. Vitex stahelii

Vitex triflora Vahl, Eclog. Amer. 2: 49. 1798. —Guarataro, Palo de arco, Palo de arco morado, Waranshikuai (Yekwana). Vitex triflora var. quinquefoliolata Moldenke, Phytologia 1: 104. 1934. —Vitex triflora f. quinquefoliolata (Moldenke) Moldenke, Phytologia 44: 384. 1979. Small tree to 15 m tall; corolla blue, violet, or purple. Evergreen lowland forests, 100– 300 m; Amazonas (Culebra, Mavaca, Río

Atacavi, Río Casiquiare, Río Cataniapo, Río Mawarinuma, Río Yureba, Santa Bárbara del Orinoco). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Acré, Amapá, Amazonas, Pará). ◆Fig. 370. The scraped bark of Vitex triflora is placed into alcoholic beverages such as rum or red wines and is used to treat rheumatism and other body pains.

VIOLACEAE by Ángel Fernández Herbs, subshrubs, shrubs, climbers, or trees. Leaves alternate, opposite, or rosulate (not in the flora area), sometimes distichous, the blades simple, entire or toothed, sometimes lobulate, pinnativeined, rarely basiveined, without indument or with simple hairs; stipules persistent or caducous. Inflorescences axillary or terminal, sometimes cauliflorous, in spikes, racemes, or panicles, sometimes flowers solitary.

446

V IOLACEAE

Flowers bisexual, actinomorphic to strongly zygomorphic, aestivation imbricate or open, pedicels generally articulate with 2 bracteoles. Sepals 5, quincuncial, equal or not, free or connate at the base, mostly persistent; corolla hypogynous to ± perigynous; petals 5, free, convolute, quincuncial, or non-overlapping, equal or not; in zygomorphic flowers the anterior petal spurred or gibbous to just emarginate. Stamens 5(3), alternate, hypogynous or mostly so, in zygomorphic flowers the 2 anterior petals with appendices extended toward the base of the spurred petal; filaments short, free or connivent around the ovary; anthers generally introrse, sometimes connivent in a ring or tube surrounding the ovary, longitudinally dehiscing and usually with connectives, dorsal scales very variable in size and shape; pollen grains binucleate, 3–5-colporate. Ovary superior, free, 1-locular, with 2–5 carpels, most commonly 3; ovules 1–many per carpel, anatropous; placentation parietal; style usually simple, filiform, erect or curved; stigma variable in shape, erect or curved to the anterior petal or deflexed, truncate, capitate, or occasionally pulvinate. Fruits capsular, valvately loculicidal, sometimes a berry, or nut-like and indehiscent. Seeds with albumen, glabrous or tomentose to pilosulous, sometimes winged; endosperm fleshy; embryo straight, sometimes slightly curved. Cosmopolitan, but most diverse in the Neotropics; 23 or 24 genera and 850– 900 species, 8 genera and 29 species in the flora area. The monotypic genus Noisettia Kunth has been recorded from Suriname, French Guiana, Peru, Brazil, and Bolivia, and may eventually be found in the flora area. Key to the Genera of Violaceae 1. 1.

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

Flowers zygomorphic, the anterior petal spurred or saccate; 2 anterior stamens with dorsal nectariferous glands into the anterior petal ...... 2 Flowers actinomorphic or slightly zygomorphic, in the latter case the anterior petal just wider than the others; all stamens equal in shape and size .......................................................................................................... 3 Lianas or scandent shrubs; flowers 3–6.8 cm long, anterior petal spurred .............................................................................................. 2. Corynostylis Herbs; flowers 0.8–2 cm long (in the flora area), anterior petal saccate ................................................................................................ 4. Hybanthus Flowers slightly zygomorphic, with one petal longer and wider than the others ...................................................................................................... 4 Flowers completely actinomorphic ............................................................ 5 Inflorescences cymose, the anterior petal emarginate; filaments fused only at the base; capsule papillate ...................................... 1. Amphirrhox Inflorescences racemose, the anterior petal rounded; filaments completely fused into a tube; capsule smooth ............................. 6. Paypayrola Dorsal connective scales of thecae absent; placentae 4 or 5, ovules > 22 per placenta; fruit a nut, indehiscent ................................................. 5. Leonia Dorsal connective scales of thecae present; placentae 3, ovules 1–22 per placenta; fruit a capsule, dehiscing or not ............................................ 6 Leaves alternate or opposite; inflorescence racemose or thyrsoid; petals usually whitish; connective scales apical and/or lateral, well developed, wider than half the diameter of the anther, commonly orangebrown; ovules 1–4 per placenta .................................................. 7. Rinorea

Amphirrhox 447

6. 7(6).

7.

Leaves alternate; inflorescence cymose; petals variously colored; connective scales apical, transparent; ovules 6–22 per placenta ................... 7 Petals whitish, yellowish, or reddish; connective scales 1/ 2 as or almost equaling diameter of the anthers; ovules 7–22 per placenta .......................................................................................... 3. Gloeospermum Petals salmon-colored; connective scales minute, < 1/2 the diameter of the anther; ovules 6–10 per placenta ............................. 8. Rinoreocarpus

1. AMPHIRRHOX Spreng., Syst. Veg. 4: 51. 1827. Spathularia A. St.-Hil., Mém. Mus. Hist. Nat. 11: 51, 491. 1824. Braddleya Vell., Fl. Flumin. 93. 1825 [1829]. Spatellaria Rchb., Consp. Regn. Veg. 189. 1828. Bradleya Kuntze, Revis. Gen. Pl. 1: 40. 1891. Treelets or shrubs. Leaves alternate, oblong-lanceolate or oblanceolate, base cuneate or attenuate, apex acuminate, erect or falcate, margin remotely repand-serrate, generally with dark mucros, principal and secondary veins prominent on both sides, 9–14 lateral veins, pinnate, arched, tertiary venation reticulate; petiole slightly swollen at base, glabrous, dark-colored; stipules caducous, herbaceous, strongly ovate-deltoid, venulate. Inflorescence terminal or axillary, peduncle terete, striate, with sparse squamiform hairs, unicellular, pale, < 0.5 mm long; pedicels ar-

Fig. 377. Amphirrhox longifolia

448

V IOLACEAE

ticulate, with bracteoles < 1 mm long. Flowers bisexual, slightly zygomorphic, glabrous. Sepals 5, unequal, aestivation quincuncial; petals 5, free, non-overlapping, somewhat fleshy, clawed, the anterior slightly bilobed or emarginate, the others oblong. Stamens equal; filaments fused only at the base, dorsal connective scales prolonged. Ovary superior, sessile, ovoid, glabrous, 1-locular, 3 parietal placentae with many ovules; style erect; stigma slightly swollen. Fruit a loculicidal capsule, warty, glabrous. Seeds ovoid, glabrous, dark. Neotropics; 1 species sensu lato (4 or 5 described). Amphirrhox longifolia (A. St.-Hil.) Spreng., Syst. Veg. 4(2): 99. 1827. —Spathularia longifolia A. St.-Hil., Hist. Pl. Remarq. Brésil 318, t. 28. 1825. —Café de venado, Gaspadillo, Gaspadillo blanco, Hicaquillo, Palo de cuchara, Palo de troja, Pate grulla, Ihkodï tawï, Tiaebo hiEi hele y Tiebo here (Höti), Fedejodo y Hede-hede (Ye’kwana), Wesemonama (Yanomami). Treelet 4–7 m tall; leaves alternate, 16–28 × 5–9 cm, petiole 0.6–1.2 cm long; stipules 1.2–2.6 mm long; inflorescence in long pedunculate dichasia, flowers 2.4–3.6 cm long, whitish to yellowish; capsule verruculose,

2.7–3.6 cm long; seeds 5–7 mm long. Evergreen lowland to lower montane forests, near rivers and streams, 50–800 m; common in Delta Amacuro, Bolívar, and Amazonas. Sucre; Central America, Colombia, Guayana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 377. Amphirrhox longifolia (sensu lato) is a species with great morphological plasticity and is widely distributed. Further studies could demonstrate the existence of an additional species, which would be differentiated mainly by the type of inflorescence, the floral morphology, and the size and consistency of the leaves.

2. CORYNOSTYLIS Mart., Nov. Gen. Sp. Pl. 1: 25, t. 17. 1824 [1823]. Calyptrion Ging. ex DC., Prod. 1: 288. 1824. Scandent shrubs or lianas, branchlets puberulous. Leaves alternate, petiolate, entire or serrulate, stipules deciduous. Inflorescences racemes, terminal or axillary, or flower solitary and axillary. Flowers zygomorphic, on long, articulate peduncles; bracts 2, caducous. Calyx quincuncial, sepals subequal, free; petals non-overlapping, whitish to yellowish, anterior petal spurred and longer than the others. Stamens 5, 4 fused in pairs plus one free, the 2 anterior ones longer than the others, pubescent and prolonged; anthers with apical, squamiform connective. Ovary with 3 placentae; ovules many; style claviform, curved; stigma embedded at the style apex. Capsule woody, 3-valved. Seeds flattened, ± square. Neotropics; 4 species, all in the flora area. Key to the Species of Corynostylis 1. 1. 2(1). 2. 3(2). 3.

Liana; leaves palmativeined; spur to 5 cm long ........................... C. volubilis Scandent shrub or liana; leaves pinnativeined; spur 1–3 cm long .......... 2 Climbing shrub; leaves 6–9 × 2–4.3 cm, glabrous, sometimes falcate; petiole 5–13 mm long ........................................................................ C. arborea Liana; leaves 2–6.3 × 1–2.4 cm, pubescent to glabrescent; petiole 3–8 mm long ......................................................................................................... 3 Leaves and petioles darkly pubescent ....................................... C. pubescens Leaves and petioles lightly glabrescent ............................. C. carthagenensis

Corynostylis 449

Fig. 378. Corynostylis volubilis

Corynostylis arborea (L.) S.F. Blake, Contr. U.S. Natl. Herb. 23: 837. 1923. —Viola arborea L., Syst. Nat. ed. 10, 2: 1239. 1759. —Bejuco de seguera, Verga de toro. Viola hybanthus L., Sp. Pl. ed. 2. 1328. 1753. —Corynostylis hybanthus (L.) Mart., Nov. Gen. Sp. Pl. 1: 26, t. 17, 18. 1823. Scandent shrub; blade elliptic, 6–9 × 2–4.3 cm, ovate to ovate-elliptic, entire, acuminate at the apex, acute to rounded at the base, penniveined, somewhat coriaceous; sepal margin ciliate, corolla whitish, spur subcylindric, to 3 cm long; capsule to 4–5 cm diameter. Gallery forests, on wet soils near rivers and streamlets, near sea level to 500 m; Delta Amacuro (Caño Atoiba, Caño Jota-

Sabuca, Orocoima in front of Isla El Toro), Bolívar (Río Botanamo, Río Caroní, Río Caura, Río Cuyuní, Río Nichare, Río Oris), Amazonas (Río Casiquiare, Río Cunucunuma, Río Ocamo, Río Orinoco between Río Atabapo and La Esmeralda, Raudal Monserrat in upper Río Orinoco). Apure, Guárico, Sucre, Zulia; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, introduced into Trinidad. Corynostylis carthagenensis H. Karst., Fl. Columb. 2: 53, t. 127, figs. 9–14. 1858 [1859]. Low-growing liana; leaves grouped at branchlet tips; blade elliptic to ovate, acuminate, 2–6.3 × 1–2.4 cm, serrate, base some-

450

V IOLACEAE

what attenuate to the petiole; petiole 3–8 mm long, puberulous; flowers white, peduncles 2.5–4.7 cm long; sepals ciliate along the margin; spur 1.5–3.1 cm long, subcylindrical to conical; capsule ovoid, ca. 3 cm diameter. Seasonally flooded river bank forests, 50–100 m; Bolívar (Mereyal on the banks of Río Orinoco, Río Cuyuní). Anzoátegui, Cojedes, Guárico; Colombia. Corynostylis pubescens S. Moore, Trans. Linn. Soc. London, Bot. ser. 2, 4: 306. 1895. Liana, branchlets pubescent; leaf blade elliptic to obovate, 5 × 1.5 cm, pubescent on the lower surface and costa on the upper surface; petiole 0.5 cm long, pubescent; peduncle 5 cm long; sepals triangular, pubescent along the costa, spur 1.8–2.2 cm long. Riparian forests, 100–200 m; Amazonas (Río Orinoco below mouth of Río Atabapo). Brazil, Bolivia. Corynostylis pubescens is distinguished from other species in the genus by the pubescence on the lower surface of the leaves and on the costa of the sepals. The only collection from the flora area (Wurdack & Adderley 42784, NY, VEN) seems to agree with the

original description of C. pubescens, a species from Pando in Bolivia and Mato Grosso do Sul in Brazil. More material is needed to establish the correct identification of the flora area specimen, which could also prove to be a new species. Corynostylis volubilis L.B. Sm. & A. Fernández, Caldasia 6: 143. 1954. —Bejuco de seguera. Liana to 15 m tall, glabrous; leaf blade lanceolate-ovate, sometimes falcate, glabrous, chartaceous, 7–9.5 × 3–4.3 cm, acuminate, palmativeined with 3 well marked secondary veins, margin serrate; petiole 9–11 mm long; peduncle 6–7 cm long; flowers white; spur claviculate, 5 cm long; ovary ovoid, capsule 2–4 cm long. Riparian forests mostly along black-water rivers, 100–200 m; Amazonas (Río Baría, Río Casiquiare, Río Cuao, San Carlos de Río Negro). Colombia, Peru, Brazil. ◆Fig. 378. Corynostylis volubilis is easily distinguished from other species of Corynostylis in having palmately veined leaves, by the longer spur of its anterior petal, and by its tall liana habit.

3. GLOEOSPERMUM Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 128. 1862. Trees or treelets. Leaves alternate, distichous, toothed, rounded to cuneate at the base; stipules caducous. Inflorescence axillary, cymose. Flowers actinomorphic, subsessile. Sepals quincuncial, subequal, ± persistent, minutely pilosulous or ciliolate; petals quincuncial, rarely non-overlapping. Filaments flattened, free or fused into a short tube; anthers appendiculate, appendix almost equaling the anthers. Ovary ovoid; style erct; stigma truncate or claviform. Fruit globose, indehiscent. Seeds mucilaginous. Neotropics; ca. 15 species, 2 in Venezuela, 1 of these in the flora area. Gloeospermum sphaerocarpum Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 129. 1862. Tree or treelet to 9 m tall; branchlets pilosulous, glabrescent when older; leaf blade elliptic to ovate, 5–14 × 3–4.8 cm, acuminate, glabrous, lateral veins 8–11, margin usually serrate; petioles to 10–12 mm long; sepals basally connate, 2 outer ones smaller than the other inner 3, margin ciliolate; petals equal, fleshy, connective scales apical, transparent; ovary subglobose to pyriform, glabrous; style erect; capsule globose, to 4 cm diameter; seeds

10–12. 100–200 m; Amazonas. Miranda, Sucre, Yaracuy; Panama, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 379. Gloeospermum sphaerocarpum is known from the flora area by a single collection (Maguire et al. 41956, NY). This was collected at 100 m in Amazonas state, but was not seen for the writing of this treatment. The other species of Gloeospermum in Venezuela, G. andinum (Tul.) Melch., is found scattered in the central part of the Cordillera de la Costa, in forests between 900 and 1350 m elevation.

Hybanthus 451

Fig. 379. Gloeospermum sphaerocarpum

4. HYBANTHUS Jacq., Enum. Syst. Pl. 2. 1760. Herbs or shrubs. Leaves alternate or opposite, entire to serrulate, stipules generally persistent. Flowers solitary and axillary, or in fascicles or terminal racemes, zygomorphic, pedicels articulate. Sepals free, unequal, persistent; anterior petal clawed or sacciform at the base. Stamens free or fused, the 2 anterior ones longer than the others, with nectariferous dorsal appendices, connective elongated and membranous. Ovary 1-locular, with 3 placentae and 2–many ovules. Capsule 3-valved, elastically dehiscent. Seeds globose. Pantropics; ca. 100 species, 9 in Venezuela, 3 of these in the flora area. Key to the Species of Hybanthus 1. 1. 2(1). 2.

Shrub; leaves distichous, pedicels < 5 mm long ................ H. phyllanthoides Herb or subshrub; leaves spirally or decussately arranged, pedicels 5– 11 mm long ............................................................................................. 2 Erect, glabrous; leaves opposite, lanceolate; anterior petal heart-shaped; peduncles 6–7 mm long .................................................... H. oppositifolius Commonly prostrate, pubescent to puberulous; leaves alternate, elliptic; anterior petal inversely trullate; peduncles to 12 mm long ... H. calceolaria

Hybanthus calceolaria (L.) Schulze, Notizbl. Bot. Gart. Berlin-Dahlem 12: 114. 1934. —Viola calceolaria L., Sp. Pl. ed. 2, 2: 1327. 1763. —Ionidium calceolaria (L.) Vent., Jard. Malmaison 1: sub t. 27. 1803. —Conejilla. Viola ipecacuanha L., Mant. Pl. 2: 484. 1771. —Ionidium ipecacuanha (L.) Vent., Jard. Malmaison 1: sub t. 27. 1803. —Hybanthus ipecacuanha (L.) Baill., Hist. Pl. 4: 344. 1873. Herb, prostrate or suberect, pubescent to hirsutulous; leaves alternate, elliptic, acumi-

nate, serrate, subsessile, 2–4.5 × 1.3–2.6 cm; flowers solitary, axillary, peduncles 7–11 mm long, bractlets linear, 5–7 mm long, villous; sepals unequal, ciliolate, anterior petal strongly widened in its distal median part, apex villous; ovary ovoid, hirsutulous; style erect; capsule hirsute. Savannas, wet soil near rivers, along roads, sometimes a weed in gardens and farms, 50–400 m; Bolívar (near Ciudad Bolívar, Maripa, Reserva Forestal Imataca, Río Paragua, Upata). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Falcón, Guárico, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre,

452

V IOLACEAE

Trujillo, Zulia; Mexico, Costa Rica, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 380.

with the original description and are tentatively assigned to H. phyllanthoides pending further studies.

Hybanthus oppositifolius (L.) Taub. in Engl. & Prantl, Nat. Pflanzenfam. III. 6: 333. 1895. —Viola oppositifolia L., Sp. Pl. ed. 2, 2: 1327. 1763. —Ionidium oppositifolium (L.) Roem. & Schult., Syst. Veg. 5: 395. 1819. Ionidium circaeoides Kunth in H.B.K., Nov. Gen. Sp. 5: 379. 1821 [1823]. —Hybanthus circaeoides (Kunth) Baill., Traité Bot. Méd. Phan. 841. 1884. Erect herb, glabrous, to 1.5 m tall; leaves opposite, petiole 1–4 mm long, lanceolate to linear-lanceolate, 0.4–0.9 × 3–5.3 cm, slightly serrulate margin; flowers solitary, peduncles 6–14 mm long with 2 minute linear, glabrous bracteoles, anterior petal obdeltoid or heartshaped; ovary globose, glabrous, style slightly curved; capsule glabrous with subpersistent style. Seasonally flooded shrubby savannas, banks of rivers and streamlets, mattorales, near sea level to 300 m; Delta Amacuro (Los Cocos near Río Orinoco), Bolívar (Isla Anacoco on Río Cuyuní, Moitaco, Raudal Guanaguaña and Raudal Ceiato on Río Caura, Río Erebato, near Río Orinoco, lower Río Pao basin, San Martín de Turumbán southwest of Tumeremo), Amazonas (San Juan de Manapiare). Mexico, Guatemala, Honduras, Nicaragua, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Hybanthus phyllanthoides (Planch. & Linden) L.B. Sm. & A. Fernández, Caldasia 6: 130. 1954. —Ionidium phyllanthoides Planch. & Linden, Ann. Sci. Nat. Bot. sér 6, 17: 125. 1862. —Cafecillo, Escoba. Deciduous shrub to 2(–4) m tall, branchlets puberulous; leaves distichous, blade rhomboid to ovate, glabrous, repand-crenate to serrate, 0.6—1.1 × 1.5–2.1 cm; petiole 1–3 mm long; flowers solitary, peduncles 5–7 mm long. Understory of semideciduous forests, 100–500 m; Bolívar (northeastern part between Upata and El Palmar). During the dry season, almost all individuals of Hybanthus phyllanthoides lose their leaves. This species was formerly known only from the type collection near Pamplona, Colombia. Collections from Bolívar state agree

Fig. 380. Hybanthus calceolaria

Leonia 453

5. LEONIA Ruiz & Pav., Fl. Peruv. 2: 69, t. 222. 1799. Steudelia Mart., Nov. Gen. Sp. Pl. 3: 168. 1829 [1832], non Spreng. 1822. Shrubs or small trees. Leaves with translucid dots, alternate, entire, toothed to remotely serrate. Inflorescence cymose or racemose, axillary or cauliflorous in L. glycycarpa. Flowers actinomorphic, with quincuncial aestivation; pedicels articulate. Sepals unequal; petals free or slightly fused at the base, oblong. Stamens 5 (3 in Andean L. triandra), fused into a tube, without dorsal connective scales, thecae sessile, placed horizontal over the staminal tube. Ovary globose, ovoid, 1-locular, with 4 or 5 placentae; ovules numerous. Fruit indehiscent, nuciform, pericarp subligneous. Seeds embedded in mucilaginous pulp, oblong to obovate. Nicaragua, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 6 or 7 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Leonia 1.

1.

Leaves subsessile, the blade base decurrent onto the petiole, 6–10 pairs of secondary veins, margin toothed; inflorescences axillary or ramiferous, 2–2.4 cm long; fruit ca. 2 cm diameter, yellow at maturity (drying dark) ...................................................................................................... L. cymosa Leaves petiolate, the blade base cuneate, rounded to obtuse, 3–5 pairs of secondary veins, arched toward the apex, margin entire; inflorescences cauliflorous, 10–20 cm long; fruit ca. 5 cm diameter, grayish light brown at maturity ............................................................................. L. glycycarpa

Leonia cymosa Mart., Nov. Gen. Sp. Pl. 2: 88, t. 169. 1826 [1827]. —Anakasha (Yekwana), Guakabakadeupadek, Kúno-to-wo-ku-deu, Mamoncillo, Mutuiyek (Pemón), Uri tote hyEi (Höti), Yarakaru nishesahri. Small tree 3–10 m tall, lenticellate; leaves chartaceous, subsessile, obovate to elliptic, blade 5.8–17.5 × 3.7–7.3 cm, glabrous, secondary veins anastomosed, base attenuate to cuneate to the petiole, margin slightly serrulate and toothed; inflorescences cymes or fascicles, axillary or ramiferous; filaments fused into a short tube; fruit to 2.5 cm diameter, yellow. Evergreen lowland to lower montane forests and riparian/seasonally flooded forests, 100–700 m; Bolívar (Cerro Maigualida, Río Caroní basin, middle and lower Río Caura, Río Cucurital, Río Paragua basin, Supamo-Parapoy mines), Amazonas (Culebra, road from Maroa to Yavita, Mavaca, Raudal Xiquitapora on Río Putaco, Río Casiquiare, San Carlos de Río Negro, Serranía Batata on Cuao-Sipapo massif, Sierra Tapirapecó).

Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. The pulp of the fruits of Leonia cymosa has a sweet taste and is eaten by local people. Leonia glycycarpa Ruiz & Pav., Fl. Peruv. 2: 69, t. 222. 1799. —Anakasha, Merecurillo, Nomimadi (Yekwana). Tree 5–20 m tall, glabrous, lenticellate; leaves coriaceous, blade elliptic, entire, 9–18 × 3.5–7 cm, glabrous, secondary veins in 3–5 pairs, arched to the apex; petioles 1.6–2.3 cm long; inflorescences commonly cauliflorous, peduncles 7–18 cm long; petals whitish to light yellow; capsule 2–4.8 cm diameter. Evergreen lowland to lower montane forests, 200–800 m; Bolívar (Río Akanán, Río Ariza, Río Erebato, Río Icabaru, Río Paragua, Río Tabaro tributary of Río Nichare, Salto Para vicinity of Río Caura), Amazonas (Caño Baría, Río Siapa, Simarawochi at Sierra Parima). Nicaragua, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 381.

454

V IOLACEAE

Fig. 381. Leonia glycycarpa

6. PAYPAYROLA Aubl., Hist. Pl. Guiane 249, t. 99. 1775. Lignonia Scop., Intr. Hist. Nat. 292. 1777. Payrola Juss., Gen. Pl. 427. 1789. Wibelia Pers., Syn. Pl. 1: 21. 1805, non P.G. Gaertn., B. Mey. & Scherb. 1801, nec Bernh. 1801. Periclista Benth., J. Bot. (Hooker) 4: 108. 1842. Shrubs or trees. Leaves alternate, ovate, obovate, or long-lanceolate to sublanceolate-oblong, base acute, cuneate, or attenuate, apex acute or acuminate to short-cuspidate, margin repand-serrate, principal and secondary veins prominent on both sides, lateral veins pinnate, arched, blade glabrous, petiole glabrous, sulcate on the upper side; stipules caducous. Inflorescence terminal, axillary, or ramulose, racemose; pedicels articulate, with bracteoles. Flowers bisexual, slightly irregular, glabrous, subsessile. Sepals 5, subequal, sometimes patent, persistent, margin sometimes ciliolate, aestivation quincuncial; petals tubular, lobes 5, oblong, unequal, the anterior ones wider than the others, fleshy, non-overlapping. Filaments 5, completely fused into a long tube; anthers sessile, dorsal connective scales minute or wanting. Ovary glabrous or pubescent, trigonous-ovate, oblong-acute to conic, 3 placentae with numerous ovules; style erect. Fruit a 3-valved capsule, smooth, glabrous. Seeds light brown. Neotropics; 9 or 10 species (3 of them still unnamed), 3 in Venezuelan, all in the flora area. Paypayrola is badly in need of further study. More collections are needed since material in herbaria is scanty and the genus is poorly known.

Paypayrola 455

Key to the Species of Paypayrola 1.

1.

2(1). 2.

Leaves clearly lanceolate to elliptic-lanceolate, leaf base long-attenuate, petiole 2.5–3.5 cm long; corolla linear-cylindric, 0.9–1.3 mm wide; sepals ovate acute ........................................................................ P. longifolia Leaves ovate, obovate, or oblong, leaf base short-attenuate, cuneate, acute, or rounded, petiole 0.5–2 cm long; corolla swollen at base, 7– 10 mm wide; sepals ovate-obtuse to ovate-suborbiculate .................... 2 Petiole thick, 1–2 cm long; raceme laxiflorous, flowers 15–20 mm long, appendix of the connective minute .......................................... P. grandiflora Petiole slender, 0.5–1.5 cm long; raceme densiflorous, flowers 8–14 mm long, appendix of the connective lacking ............................... P. guianensis

Paypayrola grandiflora Tul., Ann. Sci. Nat. Bot. sér. 3, 7: 371. 1847. Paypayrola ventricosa Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 153. 1849. Shrub or small tree to 10 m tall; leaves alternate, glabrous on both sides, elliptic to obovate-oblong, acuminate, rounded to cuneate at base, margin entire; inflorescence terminal, axillary, or ramulose, racemes 6–11 cm long, pedicels 1–3 mm long; flowers fleshy, cupuliform, corolla light orange to yellowish; anthers semi-ovate, with minute apendix; ovary as long as the filament tube, conical, shortpuberulous at the base of the style. Evergreen lowland forests, 100–200 m; Amazonas (along road between Yavita and Maroa). Colombia, Guyana, French Guiana, Peru, Brazil. Paypayrola guianensis Aubl., Hist. Pl. Guiane 249, t. 99. 1775. Shrub or tree 3–7 m tall, glabrous; leaves glabrous, ovate-lanceolate to obovate-oblong, acuminate, 13–20 × 7–10, cm, sometimes bicolored; inflorescence commonly a terminal raceme, 4–7 cm long, densiflorous; pedicels 1–2 mm long; flowers to 1.4 cm long, whitish; sepals ovate-suborbiculate, unequal; petals longlinear to oblong-lanceolate, 8–10 mm long, pistil twice as large as the filament tube; seeds light brown, 9–11 mm long. Riparian forests, 200–300 m; Amazonas (Río Cunucunuma). Colombia, Guyana, Suriname, French Guiana, Brazil. Paypayrola longifolia Tul., Ann. Sci. Nat. Bot. sér. 3, 7: 372. 1847. —Gaspadillo, Gaspadillo negro, Jabón, Pulvillo mágico. Tree or treelet to 4–12 m tall, glabrous; leaves long-lanceolate, acute, long-apiculate

Fig. 382. Paypayrola longifolia

456

V IOLACEAE

to long-acuminate, entire, base long-attenuate to the petiole, 16–30 × 4–8 cm, clearly bicolored when dry; petiole 2–3.5 cm, curved at base; inflorescence cauliflorous to ramiflorous, rarely terminal, 4–6 cm long, pauciflorous; flowers 1–1.3 cm long, solitary, sepals oblong, connective appendage equaling the anther length; ovary shorter than the filament tube, conical-obtuse, pistil erect. Evergreen lowland forests on seasonally flooded soils, terra firme evergreen to semideciduous forests, 50–900 m; Delta Amacuro (northeast of El Palmar, Los

Castillos de Guayana, Río Amacuro, Río Cuyuní), Bolívar (Cerro Abismo, El Dorado to Santa Elena de Uairén road, Río Acanán at Guadeqen, Río Apacará, Río Caroní at Eutobarima, Río Hacha, mouth of Río Paragua, Serranía de Imataca, Serranía Supamo region, Sierra de Lema), Amazonas (Capibara, Cerro Huachamacari, Río Siapa). Monagas; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 382. Paypayrola longifolia is the most common species of Venezuelan Paypayrola.

7. RINOREA Aubl., Hist. Pl. Guiane 235, t. 93. 1775, nom. cons. Conohoria Aubl., Hist. Pl. Guiane 239. 1775. Passoura Aubl., Hist. Pl. Guiane 2(suppl.): 21. 1775. Riana Aubl., Hist. Pl. Guiane 237. 1775. Alsodeia Thouars, Hist. Vég. Îsles Austra. Afrique 55. 1805. Shrubs or trees. Leaves alternate or opposite, stipules caducous, domatia sometimes present at the axils of the secondary venation. Inflorescences thyrsoid, (pseudo)racemose or rarely cymose, terminal or axillary. Flowers actinomorphic, solitary or in cymules, with articulate pedicels. Sepals 5, free, with quincuncial aestivation; petals 5, free, generally non-overlapping. Stamens 5; filaments varying from free to completely fused into a tube; anthers introrse, with lateral dehiscence, provided with dorsal connective, laminar scales conspicuous. Ovary superior, glabrous or hairy, 1-locular, divided into 3 carpels with a single filiform style and stigma, placentae parietal, 1–4 ovules per placenta. Capsule loculicidal into 3 valves, coriaceous to subwoody, sometimes with indument. Seeds 1–4, subglobose to pyriform, glabrous or pilosulous; endosperm copious. Pantropics; 160 species, 17 in Venezuela, 14 of these in the flora area. Rinorea neglecta Sandwith was mentioned for the flora area (Amazonas state) based on a misidentification of Huber & Cerda 1450 (NY, VEN), which belongs to R. ovalifolia. However, R. neglecta is expected in Bolívar state, since it has been collected in Guyana. Another species erroneously noted for Amazonas state is R. vaupesana L.B. Sm. & A. Fernández, based on Guánchez 90 (TFAV). The specimen corresponds to R. paniculata, but R. vaupesana comes from the upper Río Negro in Brazil and from the Río Vaupes in Colombia, so future collections in the Orinoco Venezuelan border south of Puerto Ayacucho could add this species to the flora area. Finally, another expected species is R. amapensis, which has been found on the Colombian bank of the Río Guainía, opposite Maroa in Amazonas state. Key to the Species of Rinorea 1. 1. 2(1). 2.

Leaves alternate; inflorescence thyrsoid; filaments fused at base into a tube ......................................................................................... R. paniculata Leaves opposite; inflorescence thyrsoid, racemose, or pseudoracemose; filaments free or fused ........................................................................... 2 Inflorescence thyrsoid, cymules lateral with 1–9 flowers ........................ 3 Inflorescence racemose or pseudoracemose, flowers solitary or 1–3 in small lateral basal cymules ................................................................... 4

Rinorea 457

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

5(4).

5.

6(5). 6. 7(6). 7. 8(7). 8. 9(7). 9. 10(6). 10. 11(10).

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

Ovary subglobose, sometimes with 1 or 2 pilose hairs, ovules 2 per placenta; style filiform, 2–3 mm long; stamens 2.2–3 mm long .... R. sprucei Ovary trapezoid, glabrous, ovules generally 1 per placenta; style ± zigzag at the base, to 1 mm long; stamens 1.2–1.8 mm long ............ R. racemosa Ovary conical, style tapering to the ovary; flowers subsessile; stamens clearly curved to outside; capsule 4–7 cm long .................. R. macrocarpa Ovary subglobose, conical, pyriform, or trapezoid; style filiform, stipitiform, to subclavate; flowers pedicellate 2–7.5 mm long, stamens oriented upward and straight; capsule commonly 1.7–5 cm long ............ 5 Leaves subsessile, petioles 0.5–2 mm long, ovate or elliptic; sepals strongly 5–7-ribbed when dried, longest sepals almost equaling size of petals ...................................................................................... R. endotricha Leaves clearly petiolate, petioles 3–15 mm long, ovate, obovate, or elliptic; sepals smooth to striate when dried, if ribbed then the seeds hairy, sepals about half size of petals .............................................................. 6 Margin of leaves serrate, the teeth evident .............................................. 7 Margin of leaves subserrate, crenate, subcrenate, to ± entire .............. 10 Domatia formed by hairy tufts in the axils of the principal and secondary veins on the lower surface of the blade ................................................. 8 Domatia of hairy tufts on the lower surface of blade completely absent .... 9 Margin of leaves ciliolate, blade pubescent; 2 anterior stamens at least partially fused with the dorsal glands .................................... R. ovalifolia Margin of leaves not ciliolate, blade glabrous, only costa and secondary veins pilose or glabrescent; stamens completely free .............. R. brevipes Ovary golden- to ferruginous-strigose; leaf elliptic, noticeably narrowing at the base and at the apex ............................................... R. melanodonta Ovary brown and spiny-hispid; leaf ovate to ellipitic, the base rounded to cuneate, the apex acute to acuminate ........................................... R. riana Leaf base symmetric or slightly unequal, rounded to cuneate .............. 11 Leaf base asymmetric, oblique, and subauriculate at the petiole ......... 12 Domatia of hairy tufts present in the axils of veins on the lower surface of the blade; sepals and petals glabrous, sometimes golden-pilosulous or -ciliolate ................................................................................... R. flavescens Domatia of hairy tufts in the axils of veins on the lower surface of the blade absent; sepals and petals feruginous-hispid to -strigose ....... R. pubiflora Domatia of hairy tufts in the axils of veins on the lower surface of the blade; seeds 1 per valve ................................................................ R. camptoneura Domatia of hairy tufts in the axils of veins on the lower surface of the blade absent; seeds 2 per valve ................................................. 13 Seeds pilosulous; petals 2–3 mm long; style stipitiform to subclavate, curved at the base, 1.5 mm long .......................................... R. lindeniana Seeds glabrous; petals 4–5 mm long; style filiform, erect, 2.2–2.8 mm long ........................................................................................ R. falcata

Rinorea brevipes (Benth.) S.F. Blake, Contr. U.S. Natl. Herb. 20: 512. 1924. —Alsodeia brevipes Benth., J. Bot. (Hooker) 4: 106. 1842. Treelet with numerous small whitish callose lenticels when dried; leaves opposite, the

blade elliptic to obovate, glabrous, rachis puberulous on both sides, domatia present at the axils of the secondary veins, margin serrulate; axis of the inflorescence densely puberulous; flowers pendent; petals without marginal cilia; filaments free; ovary subglob-

458

V IOLACEAE

ose, pilosulous, style filiform, conical at the base; capsule pilosulous, veined, style often persistent at apex of one valve; seed 1 per valve, glabrous. Lower montane forests, 400– 500 m; Amazonas (upper Río Orinoco). Guyana, Suriname, French Guiana, Peru, Brazil. The presence of Rinorea brevipes in the flora area is based on a single specimen (Chaviel 31, PORT). This collection is only in fruit, so flowering material is needed to confirm the existence of the species in the flora area. Rinorea camptoneura (Radlk.) Melch. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 352. 1925. —Alsodeia camptoneura Radlk., Sitzungsber. Math. Phys. Cl. Königl. Bayer. Akad. Wiss. München 20: 186. 1891 [1890]. —Dawicú blanco. Treelet 3–10 m tall; leaves opposite, blade glabrous, commonly falcate, domatia present in axils of the secondary venation, lateral veins 8–13 pairs; filaments free, connective scales ovate, brownish; ovary densely pilose; capsule golden- to ferruginous-pilosulous; seeds 2 per valve, globose, glabrous. Flooded and nonflooded evergreen lowland forests, 100–300 m; Amazonas (Río Casiquiare, confluence of Río Sipapo and Río Cuao, Río Ventuari delta, San Carlos de Río Negro). Peru, Brazil. Rinorea endotricha Sandwith, Bull. Misc. Inform. Kew 1933: 324. 1933. —Molenillo. Shrub or treelet 1.5–3 m tall; leaves opposite, subsessile, the blade ovate to obovate, subserrate, glabrous on both sides, margin subserrate; sepals almost reaching the size of the petals, strongly veined; stamens subsessile, filaments free, connective scales ovate, acuminate, orange-brown; ovary conical, golden-strigose, style filiform; capsule acuminate, reddish-punctate when dried; seeds 2 per valve, globose, spotted, glabrous. Understory of semideciduous or evergreen lowland forests, 50–300 m; Delta Amacuro (northwest of El Palmar, southeast of Los Castillos), Bolívar (near Guri dam, El Palmar south to area of El Dorado). Guyana. Rinorea endotricha is an endemic to the eastern Venezuelan Guayana region and Mabura Hills in adjacent Guyana. It is sepa-

rated from the closely related and more widely distributed R. melanodonta by its subpetiolate leaves, leaf base rounded (versus strongly decurrent on the petiole), and the size of the sepals almost equaling the petals. Rinorea falcata (Mart. ex Eichler) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Alsodeia falcata Mart. ex Eichler in Mart., Fl. Bras. 13(1): 386. 1871. Tree or treelet to 10 m tall; leaves opposite, elliptic to irregularly ovate, acuminate, blade glabrous on both sides, rachis pilosulous above, base rounded to cuneate or obtuse, falcate toward the apex; axis of the inflorescence golden-hispidulous, pedicels articulate; filaments free; ovary subglobose, whitishto ferruginous-strigose, style filiform and taller than the stamens; capsule acuminate, pilosulous, veined; seeds 2 per valve, glabrous. Understories of evergreen lowland seasonally flooded and nonflooded forests, 50–600 m; Bolívar (Salto Pará on Río Caura, Serranía de los Pijiguaos), Amazonas (Cerro Huachamacari, upper Río Orinoco to La Esmeralda, Río Siapa). Táchira; Colombia, Suriname, French Guiana, Peru, Brazil. Rinorea falcata is closely related to R. camptoneura and R. flavescens, from which it is differentiated by its intermediate characters, such as pilosity, leaf base form, the number of sepal veins and shape, petals, ovary, and style pilosity. Rinorea flavescens (Aubl.) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Conohoria flavescens Aubl., Hist. Pl. Guiane 239, t. 95. 1775. —Alsodeia flavescens (Aubl.) Spreng., Syst. Veg. 1: 806. 1825 [1824]. —Gaspadillo, Icaco, Jonojodedö (Yekwana), Kurunmota-yek (Pemón), Molenillo. Small tree 3–12 m tall; leaves opposite, blade glabrescent to glabrous, base rounded to cuneate, axis of the inflorescence commonly purplish and shining when dried; flowers pendulous, sepals unequal; filaments free, connective scales orange-brown; ovary ovoid to conical, commonly glabrous, style filiform, surpassing the stamens; capsule acuminate, reddish brown to purplish when dried, glabrous, veined; seeds 2 per valve, glabrous, flower parts persistent at the base of the capsule. Evergreen lowland to montane forests,

Rinorea 459

50–1000 m; widespread in Bolívar and Amazonas. Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 387. Rinorea lindeniana (Tul.) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Alsodeia lindeniana Tul., Ann. Sci. Nat. Bot. sér 3, 7: 364. 1847. —Alsodeia guianensis var. lindeniana (Tul.) Eichler in Mart., Fl. Bras. 13(1): 387. 1871. Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia; 2 varieties, 1 in Venezuela. R. lindeniana var. lindeniana. —Gaspadillo, Hoglashdi (Yanomami), Molinillo. Small tree 4–8 m tall; leaves opposite, base asymmetric, blade glabrous on both sides, rachis hirtellous above, central axis of the inflorescence sparsely hirtellous; flowers pendent; sepals ovate, mucronulate; filaments free, connective scales golden brownish; ovary globose, hirsute, style curved at base; capsule hispidulous, ± asymmetric with one valve larger than the other two and the style sometimes subpersistent at the apex of one of them; seeds 1 per valve, globose, golden brownishpilosulous. Seasonally flooded and nonflooded lowland to lower montane forests, 100–700 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Falcón, Lara, Merida, Miranda, Portuguesa, Sucre, Tachira, Vargas, Yaracuy; Panama, Colombia, Trinidad, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. Rinorea lindeniana var. lindeniana is one of the most common species of Rinorea found in Venezuela. Variety fernandeziana is endemic to the border region of Colombia and Panama. Rinorea macrocarpa (Mart. ex Eichler) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Alsodeia macrocarpa Mart. ex Eichler in Mart., Fl. Bras. 13(1): 185, t. 78, fig. 1. 1871. —Akaymota-yek (Arekuna). Small tree to 10 m tall; leaves opposite, elliptic to obovate, glabrous, rachis and secondary veins pilosulous to glabrescent on both sides, domatia sometimes present in axils of the lateral veins; inflorescence a many-flowered raceme, the flowers almost sessile; petals usually completely glabrous; filaments

curved to the outside and fused basally to a short carnose tube, connective scales transparent to yellowish, fringed at margin; ovary conical and tapering into the style; capsule symmetric, acuminate, slightly hispidulous; seeds 2 or 3 per valve, globose, commonly glabrous. Understory of humid lowland forests, 100–500 m; widespread in Bolívar and Amazonas. Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 385. Rinorea macrocarpa is easily distinguished from other species of Rinorea by its subsessile flowers and fruits, its filaments oriented outside and basally fused into a tube, its transparent and marginally fringed connective scales, and by the conical shape of the ovary, which is tapered to a conical style. Rinorea melanodonta S.F. Blake, Contr. U.S. Natl. Herb. 20: 511, pl. 35. 1924. —Molenillo. Shrub or treelet to 6 m tall; leaves opposite, blade elliptic to ovate, apex acute, margin subcrenate to subserrate, glabrous on both sides; flowers pendent; filaments free, connective scales orange-brown; ovary subglobose or subconical, hairy, style filiform, not completely erect; capsule pilosulous, veined; seeds 2 per valve, pilose. Understories of semideciduous to evergreen lowland forests, 100–400 m; Bolívar (lower Río Paragua and Río Caroní to El Callao and the western slopes of Altiplanicie de Nuria). Falcón, Miranda, Nueva Esparta, Portuguesa, Sucre, Yaracuy; Colombia, Trinidad, Guyana. Rinorea melanodonta is differentiated from the closely related species R. endotricha by the shape of the leaf bases, which is symmetric and tapering into the petiole in R. melanodonta versus asymmetric and subcordate or subauriculate at the petiole in R. endotricha. Rinorea ovalifolia (Britton) S.F. Blake, Contr. U.S. Natl. Herb. 20: 513, pl. 36. 1924. —Alsodeia ovalifolia Britton, Bull. Torrey Bot. Club 16: 18. 1889. —Arunillo (Panare), Cacho de venado. Shrub or treelet, branchlets puberulous; leaves opposite, hispidulous on both sides, blade usually elliptic to ovate, veins goldenor ferruginous-hispidulous, domatia sometimes present, margin commonly subserrate; axis of the inflorescence densely hispidulous;

460

V IOLACEAE

anterior filaments embodied in fused dorsal glands, posterior filaments and dorsal glands free, connective scales brownish; ovary pilosulous, style sometimes slightly curved; capsule hispid, veined, with subpersistent style at the apex; seeds 2 per valve, globose, glabrous. Understories of semideciduous forests, shrubby savannas, granitic outcrops, 50–400 m; Bolívar (Caicara, Río Parguaza, Serranía de los Pijiguaos), Amazonas (road between Puerto Ayacucho and El Burro, between Puerto Ayacucho and Samariapo). Brazil, Bolivia, expected in Colombian Orinoquia. ◆Fig. 384. Some introgressive hybridization occurs between populations of Rinorea ovalifolia and R. pubiflora var. grandifolia f. grandifolia. Rinorea paniculata (Mart.) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Alsodeia paniculata Mart., Nov. Gen. Sp. Pl. 1: 30, t. 21. 1824 [1823]. Tree or treelet; leaves alternate, petioles to 2.5 cm long, blade elliptic to ovate, 6.5–23 × 4–10 cm, sometimes ± coriaceous, glabrous, rachis pilosulous, margin subcrenate, commonly darker colored when dry; inflorescence 5–20 × 3–6 cm, axis puberulous; filaments basally fused to a short tube; ovary trapezoid, pilosulous, style curved at base; capsule asymmetric, floral parts persistent, puberulous; seeds 1 per valve, globose, glabrous. Understories of evergreen lowland to lower montane forests, near rivers or in seasonally flooded soils, 100–700 m; Bolívar (upper Río Caura basin), Amazonas (widespread). Colombia, French Guiana, Brazil. ◆Fig. 383. Rinorea pubiflora (Benth.) Sprague & Sandwith, Bull. Misc. Inform. Kew 1931: 171. 1931. —Alsodeia pubiflora Benth., J. Bot. (Hooker) 4: 106. 1842. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 varieties, both in the flora area. Key to the Varieties of R. pubiflora 1. Anthers reduced to 0–1.5 × 0–0.75 mm or absent, connective scales wider or linear; floral parts subpersistent at base of the capsule; seeds pilosulous or glabrous ................................... var. grandifolia

1. Anthers well developed, 1–1.8 × 0.7–1 mm, connective scales ovate; no persistent floral parts at the base of the capsule; seeds pilosulous ........... var. pubiflora R. pubiflora var. grandifolia (Eichler) Hekking, Fl. Neotrop. Mongr. 46: 109. 1988. —Alsodeia falcata var. grandifolia Eichler in Mart., Fl. Bras. 13(1): 386. 1871. —Gaspadillo, Gaspadillo blanco, Jona joreri (Yekwana), Jööna jodedü (Yekwana), Molinillo. Tree or treelet; leaves opposite, blade more coriaceous than in var. pubiflora; stamens reduced or staminodes, dorsal glands reduced or absent, connective scales wider than in var. pubiflora; style 2–3.25 mm long, hispidulous toward the base; capsule smaller than in var. pubiflora; and subtended by subpersistent floral parts; seeds 2 per valve, rarely 1 or 3, glabrous or pilosulous. Understories of humid and mesic primary or disturbed forests, granitic outcrops, 50–900 m; Delta Amacuro (Los Castillos de Guayana), Bolívar (Icabarú, Reserva Forestal La Paragua, Río Caura and Río Paragua basin, Río Karún, Urimán), Amazonas (Culebra, south of Limon de Parhueña, Puerto Ayacucho). Two forms have been differentiated in Rinorea pubiflora var. grandifolia. Both are found in the flora area: f. grandifolia has glabrous seeds, f. andersonii (Sandwith ex Hekking) Hekking has pilosulous seeds. R. pubiflora var. pubiflora. —Pata de paují, Tuliwaedi hele (Höti). Alsodeia pubiflora Benth., J. Bot. (Hooker) 4: 106. 1842. Passoura guianensis Aubl., Hist. Pl. Guiane 2(suppl.) 21, t. 38. 1775. —Alsodeia guianensis (Aubl.) Eichler in Mart., Fl. Bras. 13(1): 387, t. 78, fig. 2. 1871. Tree or treelet; leaves opposite, glabrous, rachis strigose on the lower surface, domatia present in the axils of the secondary venation; flowers pendent; stamens developed, filaments free, dorsal glands equaling or slightly longer than the filaments; style 2–2.8 mm long, usually completely glabrous; capsule 1.7–3 × 0.5–1 cm long, without persistent floral parts in its base; seeds 2 or 3(4) per valve, brownish-puberulous. Common as under-

Rinorea 461

growth in primary and disturbed, evergreen or semideciduous forests, 100–400 m; Bolívar (Río Caura and Río Paragua basin, road between Caicara and San Juan de Manapiare), Amazonas (Caño Iguana, Culebra, Mavaca, Sierra Parima, Yutajé). Mérida, Táchira, Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 386.

Rinorea racemosa (Mart.) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Alsodeia racemosa Mart., Nov. Gen. Sp. Pl. 1: 29, t. 20. 1824 [1823]. Treelet; leaves opposite, glabrous on the upper surface, pilose on the lower surface, domatia sometimes present, lateral veins 11– 21 pairs, acumen 0.2–2.5 mm long; inflorescence thyrsoid; sepals and petals commonly

Fig. 383. Rinorea paniculata

Fig. 384. Rinorea ovalifolia

462

V IOLACEAE

Fig. 385. Rinorea macrocarpa

Fig. 387. Rinorea flavescens

Fig. 386. Rinorea pubiflora var. pubiflora

Fig. 388. Rinorea sprucei

Rinoreocarpus 463

pilosulous over the costa; filaments fused into a tube; ovary subglobose to trapezoid, glabrous, style ± sigmoid at the base, equaling the stamens; capsule with floral parts subpersistent, glabrous; seeds 1(2) per valve. Evergeen lowland forests, 100–200 m; Amazonas (San Pedro de Cataniapo). Colombia, Peru, Brazil. Rinorea riana Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Conohoria riana DC. ex Ging. in DC., Prodr. 1: 312. 1824, nom. illeg. —Rinorea riana (DC.) Kuntze, Revis. Gen. Pl. 1: 42. 1891, nom. illeg. —Gaspadillo, Gaspadillo blanco, Jö’nö jodedü (Yekwana), Mamoncilla, Molenillo. Riana guianensis Aubl., Hist. Pl. Guiane 237, t. 94. 1775. Treelet; branchlets with whitish callose lenticels; leaves opposite, blade narrowly ovate to elliptic, wider from the middle to the apex, coriaceous, glabrous, base rounded to cuneate, margin usually subserrate; filaments free, connective scales brown; ovary subglobose, covered by erect, hispid, brown hairs, style filiform, sometimes slightly curved at base, exceeding the stamens; capsule velutinous, veined; seeds 1 or 2 per valve, hirtellous. In undergrowth of evergreen lowland to lower montane forests, sea-

sonally flooded or not, sea level to 700 m; Delta Amacuro (Eleanor Creek, Río Grande, Río Toro, Sierra de Imataca), Bolívar (Maripa, Reserva Forestal Imataca, widespread in Río Caura and Río Paragua basins, Río Chicanán, Río Venamo, Sierra Maigualida, Supamo mines), Amazonas (35 km south of Puerto Ayacucho). Trinidad, Guyana, Suriname, French Guiana, Brazil. Rinorea sprucei (Eichler) Kuntze, Revis. Gen. Pl. 1: 42. 1891. —Alsodeia sprucei Eichler in Mart., Fl. Bras. 13(1): 385. 1871. Small tree; leaves opposite, blade narrowly ovate to elliptic, acuminate, pilosulous on the lower surface, lateral veins 9–15, margin entire or slightly subcrenate, acumen 1–6 cm long; flowers pendent; sepal exterior densely pilose; petals golden-strigillose on both sides; filaments nearly fused into a tube; ovary glabrous, sometimes with 1 or 2 long hairs, style erect, exceeding the stamens, stigma deflexed; capsule asymmetric, with subpersistent floral parts at base, commonly glabrous; seeds 2 per valve, globose, shining. Forests on white-sand soils, caatinga and seasonally flooded margins of rivers, 100–300 m; widespread in Amazonas. Colombia, Brazil. ◆Fig. 388.

8. RINOREOCARPUS Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 144. 1925. Trees. Leaves alternate, glabrous. Inflorescences axillary, cymose. Flowers actinomorphic, salmon-colored, aestivation of sepal quincuncial, petals apotact. Filaments free, dorsal appendix of the connective apical, minute, narrower than the anthers and colorless. Ovary conical-ovoid, glabrous, placentae 3, each with 6–10 ovules; style filiform, 2 times the length of the stamens. Capsule 3-valved, veined. Seeds 6–12, pyriform, glabrous, dark-colored, and shining. Neotropics (Amazonian and Guayanan region); 1 species. Rinoreocarpus ulei (Melch.) Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 173. 1930. —Gloeospermum ulei Melch., Notizbl. Bot. Gart. Berlin-Dahlem 9: 164. 1924. —Cacaguillo. Rinoreocarpus salmoneus Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 144. 1925. Treelet to 10 m tall. Understory forests over granitic or lateritic hills, 200–600 m; Bolívar (La Vergareña, Río Caura, Serranía de los Pijiguaos). Colombia, Suriname,

French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 389. Rinoreocarpus ulei is differentiated from the related genus Rinorea by the shape of the inflorescence, which is cymose in Rinoreocapus versus thyrsoid to (pseudo)racemose in Rinorea, and by larger orangish flowers. It is distinguished from Gloeospermum by the dehiscent capsular type of fruit in Rinoreocarpus versus an indehiscent berry in Gloeospermum.

464

V IOLACEAE

Fig. 389. Rinoreocarpus ulei

VISCACEAE by Job Kuijt Leafy or scale-leaved parasitic plants on branches of trees or shrubs. Stems brittle, terete, angled, or flattened, lateral ones sometimes with one or more pairs of scalelike basal cataphylls; epicortical roots absent, the haustorial attachment single. Leaves opposite, often coriaceous or succulent, sometimes reduced to scales, sessile or petiolate, exstipulate; blade entire, venation various but often obscure. Inflores-

V ISCACEAE 465

cence axillary or terminal or both, indeterminate, a pedunculate spike with flowers along one or more expanding fertile internodes. Flowers unisexual (plants monoecious or dioecious), with a single perianth whorl, 2–4-merous, small, sessile. Anthers sessile, 1- or 2-locular; pollen spinulose, tricolpate. Ovary inferior, 1-locular, the single basal placental papilla containing 2 embryo sacs; stigma capitate, sessile or nearly so. Fruit a 1-seeded berry, petals persistent, part of the exocarp becoming a mass of viscin attached to the seed. Endosperm and embryo green, the latter dicotylous. Worldwide (mostly tropical and subtropical); 7 genera and ca. 500 species, 2 genera and 53 species in flora area. Viscaceae have often been regarded as a subfamily, Viscoideae, of Loranthaceae sensu lato, but are different in being monochlamydeous, in having basically determinate inflorescences, and having a different basic chromosome number. All genera except Arceuthobium (Northern Hemisphere) are restricted to either the Old World or the New World. Key to the Genera of Viscaceae 1. 1.

Anthers 1-locular; flowers in 1–3 series above each fertile bract, series sometimes reduced to 1 flower; mostly above 1000 m .. 1. Dendrophthora Anthers 2-locular; flowers in 2 or 3 series above each fertile bract, mostly with > 1 flower per series at least in the lower inflorescence; usually lower elevations (1000 m or less) ..................................... 2. Phoradendron

1. DENDROPHTHORA Eichler in Mart., Fl. Bras. 5(2): 102. 1868. Parasitic mostly on dicotyledonous shrubs and trees, often somewhat succulent and olive to brownish green, brittle, the stem with inconspicuous nodal constrictions. Dioecious or monoecious, in the latter case the sexes in various distributional patterns on inflorescences or individuals. Inflorescences mostly axillary or clustered at the nodes, consisting of a peduncle of mostly one internode followed by 1–several fertile internodes. Flowers 1–3-seriate, (2)3(4)-merous, often sunken in the swollen inflorescence axis. Anthers 1-locular, sessile, dehiscing via an irregular, transverse slit. Fruit frequently white, fleshy, with copious viscin; endosperm and embryo bright green. Southern Mexico, Central America, much of the West Indies, southward to Peru and Bolivia; ca. 120 species, ca. 25 in Venezuela, 15 of these in the flora area. The distinction between Dendrophthora and Phoradendron continues to be a troublesome one, being based primarily on the very small 1-locular versus 2-locular anthers, respectively. There is a strong tendency for the former genus to be limited to higher elevations and the latter one to low elevations, but several exceptions are known in the Venezuelan area. For example, D. elliptica is known from low elevations, while P. aphyllum, P. crassifolium, and P. ptarianum are reported from high elevations, while D. crispula and D. elliptica may occur as low as 100 m. Uniseriate flowers in the flora area are limited to Dendrophthora. The anther criterion makes it technically impossible at this time to make a reliable generic assignment for several species, where no anther material is available, these species here being treated under Dendrophthora [D. densifrons, D. fendleriana, D. oligantha, and D. tenuiflora].

466

V ISCACEAE

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

3. 4(3). 4.

5(2). 5. 6(5). 6. 7(6). 7. 8(7). 8. 9(7). 9. 10(9). 10. 11(5). 11. 12(11). 12. 13(12). 13. 14(13).

Leafy unforked shoots with several pairs of intercalary cataphylls ............................................................................................... D. fendleriana Unforked shoots lacking intercalary cataphylls (rarely with 1 pair), or stems dichotomous ................................................................................. 2 At least some branches terminating in an inflorescence ......................... 3 All inflorescences lateral ........................................................................... 5 Inflorescences with 3 or more fertile internodes, flowers mostly 3 per fertile bract; larger lateral inflorescences with at least 1 pair of sterile bracts; leaves to 7 cm long ......................................................... D. crispula All inflorescences lacking sterile bracts, fertile internodes 1–3; flower 1(2 or 3) per fertile bract; leaves < 2 cm long ....................................... 4 Leaves orbicular; inflorescences with mostly 1 fertile internode; flower 1 per bract only ............................................................................ D. jauana Leaves obovate to elliptic; inflorescences with mostly 2 or 3 fertile internodes; flowers 2- or 3-seriate when more than 1 flower per bract .................................................................................................. D. oligantha Inflorescence < 8 mm long; leaves ≤ 2.5 cm long ...................................... 6 Inflorescence ≥ 10 mm long; leaves > 2.5 cm long .................................. 11 Fertile internode slender, the basal portion stalk-like, flowerless; 1 flower per fertile bract ....................................................................... D. tenuiflora Fertile internode short, not slender, without stalk-like base; 1–several flowers per fertile bract ......................................................................... 7 Flowers 2 or more per fertile bract, 2- or 3-seriate .................................. 8 Flowers 1 per fertile bract and/or 1-seriate .............................................. 9 Leaves linear to very narrowly lanceolate, with very acute apex; fruit ovoid, smooth when fresh ........................................................ D. decipiens Leaves broadly obovate to elliptcal, apex rounded; fruit globose, often warty when fresh ....................................................................... D. elliptica Leaves 1–2 mm long .................................................................. D. microsoma Leaves to 25 mm long .............................................................................. 10 Leaves broadly elliptic to orbicular; inflorescences 2–3 mm long; flower 1 per fertile bract ......................................................................... D. jauana Leaves narrowly lanceolate; inflorescence to 8 mm long; flowers 3 or 4 per fertile bract .............................................................................. D. roraimae Monoecious, the basal internode of the inflorescence exclusively staminate, others pistillate ............................................................. D. macbridei Dioecious or monoecious, but in the latter case fertile internodes mixed .............................................................................................................. 12 Flower one above each fertile bract, the lower part of the fertile internode stalk-like ................................................................................. D. tenuiflora Flowers several to many per bract, l–3-seriate, the base of each fertile internode not stalk-like .......................................................................... 13 Flowers 3-seriate, or both 2- and 3-seriate on the same inflorescence ..... 14 Flowers 1- or 2-seriate only ..................................................................... 16 Dichotomous, i.e., forked, the new sprouts each with one pair of expanded leaves, and the apex aborting; basal cataphylls 2 or 3(–6) pairs, flowers 20–40 per fertile bract ................................................................ D. obliqua

Dendrophthora 467

14.

Unforked, the new sprouts with several pairs of leaves, apex not aborting; basal cataphylls 1(2) pairs; flowers mostly < 20 per fertile bract .............................................................................................................. 15 15(14). Monoecious; inflorescence to 2 cm long, absent or with 1 pair of sterile bracts and 2(3) fertile internodes; flowers ≤ 7 per fertile bract ................................................................................................ D. densifrons 15. Dioecious; inflorescences to 4 cm long, with mostly 1–3 pairs of sterile bracts and 4 or 5 fertile internodes; flowers (staminate plant) > 8 per fertile bract ............................................................................ D. intermedia 16(13). Leaf apex acute; inflorescence to 1.5 cm long ............................... D. nitidula 16. Leaf apex rounded; inflorescence 2–4 cm long ....................................... 17 17(16). Plants not yellow-dotted; inflorescence with 1–4 pairs of sterile cataphylls ...................................................................................... D. intermedia 17. Plants usually irregularly yellow-dotted at least when dry; inflorescence with 0(1) pair of sterile bracts .................................................. D. tepuiana Dendrophthora crispula (Rizzini) Kuijt, Bot. Jahrb. Syst. 122: 176. 2000. —Phoradendron crispulum Rizzini, Revista Fac. Agron. (Maracay) 8(3): 86. 1975. Dendrophthora lacryma-jobi E.A. Kellogg, Novon, 6: 35, fig. 2. 1996. Plant monoecious, stems terete, basal cataphylls 3–5 pairs, on smaller twigs followed by one pair of leaves, terminal and 2 axillary inflorescences all lacking sterile internodes; leaves to 7 × 2.5 cm, elliptic to narrowly obovate, apex rounded; inflorescence ≤ 2.3 cm long, the longer ones axillary with 1–3 short sterile internodes; flowers 3 per fertile internode, probably 2-seriate, nearly all flowers pistillate. Savanna edges to montane forests, 100–1300 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Macizo del Chimantá, Río Tonoro, Salto Uraima), Amazonas (Caño Caname, Cerro Marahuaka, Cerro Yutajé). Endemic. Dendrophthora decipiens Kuijt, Proc. Kon. Ned. Akad. Wetensch. 93: 131. 1990. Plant yellowish to deep green, diffusely branched, with one very low and inconspicuous pair of cataphylls on lateral branches; inflorescence very small, to 4 mm long, and usually with only one fertile internode bearing 1–3 flowers above each bract in 2 or 3 series; fruits ovoid, conical when dry. Tepui summits, 1300–2800 m; Bolívar (Acopántepui, Auyán-tepui, Ilú-tepui, Kukenán-tepui), Amazonas (Cerro Coro Coro, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Río Manapiare). Endemic.

Dendrophthora densifrons (Ule) Kuijt, Novon 6: 33. 1996. —Phoradendron densifrons Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 292. 1915. Plant monoecious, tawny, dark when dry, with 1 or 2 pairs of basal cataphylls, densely leafy; leaves ovate-elliptic, 3–6 cm long, petiole distinct, 4–7 mm long; inflorescence to 2.5 cm long, with one pair of sterile cataphylls and 3 fertile internodes; flowers 4–6 per bract, 3-seriate. Open forested slopes, 2100– 2500 m; Bolívar (Macizo del Chimantá). Guyana (Mount Roraima). ◆Fig. 391. Dendrophthora elliptica (Gardner) Krug & Urb., Ber. Deutsch. Bot. Ges. 14: 285. 1896. —Viscum ellipticum Gardner, London J. Bot. 4: 106. 1845. —Phoradendron ellipticum (Gardner) Eichler in Mart., Fl. Bras. 5(2): 119. 1868. Dendrophthora elliptica var. stenophylla Krug & Urb., Bot. Jahrb. Syst. 24: 69. 1897. Dendrophthora rubicunda Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 289. 1915. Phoradendron dendrophthora Rizzini, Revista Fac. Agron. (Maracay) 8: 87. 1975. Dendrophthora hylaeana Rizzini, Rodriguésia 41: 8. 1989. Small, often bright or yellowish green, much-branched, with one pair of small, low basal cataphylls; leaves tending to be ellipticovate, but sometimes rather narrow, to 2.5 cm long; inflorescences usually < 3 mm long, with 1(2) fertile internodes, flowers 1–3 per fertile bract, 2- or 3-seriate; fruit mostly

468

V ISCACEAE

warty, especially at the base. Open scrub and savanna margins to dense, forested slopes, 1000–2300 m, but occasionally much lower; Bolívar (fairly common), Amazonas (common). Aragua, Falcón, Miranda, Portuguesa, Trujillo, Yaracuy; Colombia, Ecuador, Peru, Brazil. ◆Fig. 392. Dendrophthora fendleriana (Eichler) Kuijt, Novon 6: 38. 1996. —Phoradendron fendlerianum Eichler in Mart., Fl. Bras. 5(2): 129. 1868. Phoradendron duidanum Trel., Bull. Torrey Bot. Club 58: 358. 1931. ?Phoradendron tatei Trel., Bull. Torrey Bot. Club 58: 359. 1931. Phoradendron spectabile Rizzini, Revista Brasil. Biol. 31(2): 200. 1971. Phoradendron spectabile var. altimontanum Rizzini, Revista Brasil. Biol. 31(2): 201. 1971. Plant apparently dioecious, stems unforked, terete, stout, with 3 pairs of prominent intercalary cataphylls between successive pairs of foliage leaves; basal cataphylls about 4 pairs, the upper ones more than halfway to the next foliage leaves, these fleshy, to 7 × 4 cm, elliptic, venation obscure; inflorescences axillary, with 3 or 4 pairs of sterile bracts and 5 fertile internodes; flowers to 7 per fertile bract, 3-seriate. Rock outcrops and tepui summits, 1400–2500 m; Bolívar (Cerro Jaua), Amazonas (Cerro Aracamuni, Cerro Marahuaka, below Salto los Monos, Sierra de la Neblina). Mérida. ◆Fig. 394. Dendrophthora intermedia (Rizzini) Kuijt, Novon 6: 35. 1996. —Phoradendron intermedium Rizzini, Rodriguésia 18–19: 190. 1956. Rather large plants, glabrous, unforked, internodes to 6 cm long, terete, basal cataphylls one pair 3–6 mm above axil, sometimes followed by a second pair 1.5 cm higher, rather small; leaves to 12 × 6 cm, oblong-elliptic to obovate, apex rounded, base tapering into indistinct petiole 5–10 mm long; venation palmate or nearly so; dioecious; staminate inflorescences apparently only lateral, with 1–4 pairs of sterile basal cataphylls, to 4 cm long, the sterile internodes together to 8 mm long, followed by 4 or 5 fertile internodes; flowers ca. 8–15 or more

per fertile bract, often 3-seriate above, at other times entirely 2-seriate. Evergreen lowland to semideciduous lower montane forests, 100–700 m; Bolívar (Cerro Altamira), Amazonas (Yavita). Endemic. Dendrophthora intermedia looks somewhat like D. tepuiana, but lacks the latter’s yellowish cast and, more importantly, has longer and often 2-seriate inflorescences. Dendrophthora jauana Rizzini, Rodriguésia 41: 15. 1976. Succulent plant to 50 cm diameter, nodes swollen when older; leaves elliptic-orbicular, basal cataphylls absent or one pair at ca. 5 mm above axil; leaves to 10 mm long; inflorescence 2–3 mm long, in leaf axils, but also terminally on small lateral shoots, in some of the latter cases resulting in dichotomies; fertile internodes 1(2); flowers 1 per fertile bract. Tepui summits, 1200–2400 m; Bolívar (Auyán-tepui, Cerro Jaua), Amazonas (Cerro Sipapo, west of Río Coro Coro). Endemic. Dendrophthora macbridei (Standl. ex J.F. Macbr.) Kuijt, Bot. Jahrb. Syst. 122: 177. 2000. —Phoradendron macbridei Standl. ex J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13(2.2): 387. 1937. Dendrophthora basiandra Kuijt, Wentia 6: 32. 1961. Plant rather coarse, yellowish or brownish where exposed; leaves to 11 cm long, the single pair of basal cataphylls inserted rather high (ca. 1.5 cm) above the leaf axil. In montane shrublands on Schefflera hitchcockii (Lasser & Maguire) Maguire, Steyerm. & Frodin, ca. 2200 m; Amazonas (Cerro Coro Coro). Distrito Federal; occasional from northern South America to Bolivia. Dendrophthora microsoma Rizzini, Mem. New York Bot. Gard. 23: 858. 1972. Dendrophthora parvispicata Rizzini, Bol. Soc. Venez. Ci. Nat. 22: 324. 1976. A dwarf species, no more than 15 cm diameter, internodes < 1 cm long, lateral branches with one very low pair of basal cataphylls; leaves probably persistent; inflorescence 2–3 mm long, with 1 fertile internode, each bract with 2 flowers, 1-seriate. Brushy tepui slopes and summits, 1800– 2100 m; Bolívar (Cerro Jaua). Endemic.

Dendrophthora 469

Dendrophthora nitidula (Rizzini) Kuijt, Proc. Kon. Ned. Akad. Wetensch. 93: 135. 1990. —Phoradendron nitidulum Rizzini, Rodriguésia 28: 23. 1976. Rather slender and sparsely branched, erect, olive green, dioecious, internodes slightly keeled when young; leaves to 7 × 3 cm, lanceolate to somewhat falcate, acute at both ends; inflorescences slender, fertile internodes 3, apparently 2-seriate. Tepui cliffs and headwaters, 1700–2200 m; Bolívar (Macizo del Chimantá [Chimantá-tepui]). Endemic. ◆Fig. 390. Dendrophthora obliqua (C. Presl) Wiens, Taxon 20: 326. 1971. —Viscum obliquum C. Presl, Epimel. Bot. 255. 1849 [1851]. —Phoradendron obliquum (C. Presl) Eichler in Mart., Fl. Bras. 5(2): 134. 1868. Phoradendron guascanum Trel., Bull. Torrey Bot. Club 54: 474. 1927. Phoradendron duidanum var. hymenaeifolium Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 237. 1982. Coarse and stout plants, nodes becoming swollen; lateral branches with 2 or 3(–5) pairs of large basal cataphylls, the upper ones usually at least halfway to the next foliage leaves; leaves ovate to very broadly lanceolate, sometimes asymmetrical, very coriaceous; inflorescences to 3(–4) cm long; flowers numerous, 3-seriate. Montane forests on tepuis, 1400–1800 m; Bolívar (Aparamántepui, Cerro Jaua), Amazonas (Cerro Sipapo, Cerro Yutajé, Río Coro Coro, Río Iguapo, Sierra de la Neblina). Distrito Federal, Miranda, Táchira; Panama, Colombia, Ecuador, Peru. Dendrophthora oligantha Kuijt, Novon 6: 38, fig. 3. 1996. Plant monoecious(?), profusely branched, small-leaved, apparently succulent; basal cataphylls one pair on lateral branches, to 3 mm above the base; leaves to 12 × 7 mm, obovate-elliptic, apex rounded, base tapering into petiole ca. 1 mm long; inflorescence to 7 mm long, peduncle 1 mm long, fertile internodes 1–3; flowers 1 or 3 per fertile bract, 2- or 3-seriate; inflorescence commonly terminal on short branches bearing one pair of expanded leaves. Open mossy woods in montane forest, 1200–1300 m; Bolívar (Kavanayén). Endemic.

Dendrophthora oligantha is distinctive in that the smaller new sprouts have 3 inflorescences at the tip and probably are to be shed following flowering or fruiting. Dendrophthora roraimae (Oliv.) Ule, Bot. Jahrb. Syst. 52(Beibl. 115): 49. 1914. —Phoradendron roraimae Oliv., Timehri 5: 201. 1886. Plant often very small, rarely to 25 cm large, tawny or yellowish; leaves acuminate, rather fleshy; inflorescences mostly with 1 fertile internode; flowers 1-seriate. On trees and shrubs on tepui escarpments and summits, 1800–2600 m; Bolívar (Auyán-tepui, Cerro Jaua, Ilú-tepui, Macizo del Chimantá [Chimantá-tepui, Murey-tepui], Roraimatepui), Amazonas (Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 393. Dendrophthora tenuiflora (Steyerm. & Maguire) Kuijt, Proc. Kon. Ned. Akad. Wetensch. 93: 138. 1990. —Phoradendron tenuiflorum Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 442. 1967. Stems slightly quadrangular when young; leaves to 6 × 2 cm, subcoriaceous, olive green, narrowly oblanceolate, apex rounded; inflorescence slender, to 1 cm long, fertile internodes to 5, 1 flower above each fertile bract, placed at the far end of the internode. Forested slopes, gallery forests, 500–1700 m; Bolívar (Cerro Papelón north of Cerro Guaiquinima, Macizo del Chimantá [Chimantá-tepui]). Endemic. Dendrophthora tepuiana (Steyerm.) Kuijt, Proc. Kon. Ned. Akad. Wetensch. 93: 138. 1990. —Phoradendron tepuianum Steyerm., Fieldiana, Bot. 28: 223. 1951. Plant dioecious, olive-tawny, usually minutely yellow-dotted throughout; stems slightly compressed, becoming terete, lateral branches with 1 pair of basal cataphylls; leaves to 10 × 5 cm, obovate, apex rounded; inflorescence with 1–3 pairs of basal cataphylls, fertile internodes mostly 3; flowers to 3–9 per fertile bract, 2-seriate. Gallery forests, elfin forests on tepui summits, 900– 1800 m; Bolívar (Amaruay-tepui, 12 km north of Aparamán-tepui, Kukenán-tepui, Ptari-tepui), Amazonas (Cerro Yutajé). Adjacent Guyana. ◆Fig. 395.

470

V ISCACEAE

Fig. 390. Dendrophthora nitidula

Fig. 392. Dendrophthora elliptica

Fig. 391. Dendrophthora densifrons

Dendrophthora 471

Fig. 393. Dendrophthora roraimae

Fig. 394. Dendrophthora fendleriana

Fig. 395. Dendrophthora tepuiana

472

V ISCACEAE

2. PHORADENDRON Nutt., J. Acad. Nat. Sci. Philad. ser. 2, 1: 185. 1848. Spiciviscum H. Karst., Fl. Columb. 1(2): 73. 1858 [1860].

Small parasitic shrubs on the branches of woody plants (dicotyledons in the flora area), mostly single-stemmed, internodes terete to quadrangular, lateral branches subtended by 1 or more fused scale-leaves (cataphylls); some species with intercalary cataphylls between successive pairs of expanded leaves; rarely (two species), all leaves squamate. Leaves and scale leaves paired throughout. Branching dichotomous (either through terminal abortion or terminal inflorescences) or percurrent. Inflorescence an unbranched spike with one or more sterile basal internodes followed by 2 or more fertile internodes bearing one or more, recessed flowers in (1)2 or 3 longitudinal series above each fertile bract. Flowers unisexual, in various patterns of distribution including dioecy, small, each mostly tripartite. Anther bilocular. Fruit a 1-seeded berry, embryo and endosperm chlorophyllaceous, surrounded by viscin. Northeastern U.S.A., Mexico, Central America, West Indies, southward in South America to Bolivia and central Argentina; 230 species, 67 in Venezuela, 38 of these in the flora area. Phoradendron is closely related to, and sometimes difficult to distinguish from, the other neotropical genus of Viscaceae, Dendrophthora. The main consistent distinction lies in the anther, which is unilocular in Dendrophthora and bilocular in Phoradendron. Key to the Species of Phoradendron 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(6). 7. 8(7).

8.

Plants squamate ........................................................................................ 2 Plants foliaceous ........................................................................................ 3 Internodes terete, basal cataphylls absent; fruiting petals closed ................................................................................................... P. aphyllum Internodes flattened; basal cataphylls 1 pair, tubular; fruiting petals erect ........................................................................................ P. tunaeforme Plants dichotomous (through terminal abortion or terminal inflorescences and/or with intercalary cataphylls ............................................ 4 Plants consistently percurrent; intercalary cataphylls (largely)absent .............................................................................................................. 18 Innovations bearing 1 pair of leaves ......................................................... 5 Innovations bearing 2 or more pairs of leaves, or plants percurrent .... 13 Flowers 1-seriate ............................................................................ P. kelloggii Flowers 2- or 3-seriate ............................................................................... 6 Flowers 3 per fertile bract, the apical (median) one staminate, lateral ones pistillate; internodes quadrangular ................................ P. triflorum Flowers > 3 per fertile bract and/or sex distribution otherwise; internodes mostly terete .......................................................................................... 7 Basal cataphylls 2–7 pairs; fruiting petals ± erect .................................. 8 Basal cataphylls 1 pair; fruiting petals closed ......................................... 9 Leaves yellowish green, delicate, thin, to 2 cm wide; basal cataphylls 1 or 2 pairs; staminate flowers placed below pistillate ones on each fertile internode; fruit to 4 mm long ............................................. P. dimunitivum Leaves somewhat leathery, reddish brown, to 6 m wide; basal cataphylls 2–7 pairs; staminate flowers apparently concentrated on lowest fertile internodes; fruit to 6 mm long .................................... P. inaequidentatum

Phoradendron 473

9(7). 9. 10(9). 10. 11(10). 11. 12(9). 12. 13(4). 13. 14(13). 14. 15(13). 15. 16(15). 16. 17(16). 17. 18(3). 18. 19(18).

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

21. 22(21). 22. 23(22). 23.

Venation pinnate to obscure .................................................................... 10 Venation palmate, evident ....................................................................... 12 Leaves lanceolate, coriaceous, venation obscure ..................... P. morsicatum Leaves thin or very slightly coriaceous, venation basal or pinnate ...... 11 Leaves to 13 × 10 cm, thin, venation pinnate, evident ............ P. racemosum Leaves to 6 × 3.5 cm, no more than slightly coriaceous, with 2 basal but often obscure lateral veins ................................................... P. northropiae Flowers > 10 per fertile bract, 2- and 3-seriate even on the same plant; basal cataphylls not white-margined ....................................... P. haughtii Flowers 1–3(–5) per fertile bract, basal cataphylls white-margined when dry ............................................................................................ P. scariosum Plants partially percurrent, innovations with (1)2 pairs of leaves before a terminal inflorescence is formed; intercalary cataphylls 1 pair ........ 14 Plants normally percurrent, neither dichotomous nor with terminal inflorescences; intercalary cataphylls present or absent .......................... 15 Flowers 3 per fertile bract, the median (apical) one staminate, lateral ones pistillate; leaves lanceolate to ovate...................... P. strongyloclados Flowers 3–6(–9) per fertile bract, staminate flowers generally below pistillate ones; leaves narrowly (ob)lanceolate ...................... P. dimunitivum Internodes conspicuously flattened when young; leaves narrowly oblong, < 1 cm wide, flowers 3 per fertile bract ............................... P. platycaulon Internodes not flattened; leaves lanceolate, > 3 cm wide; flowers 3–18 per fertile bract .......................................................................................... 16 Intercalary cataphylls several pairs, at least the uppermost ones fertile (subtending inflorescences) ................................................. P. crassifolium Intercalary cataphylls one pair, not fertile ............................................. 17 Venation pinnate; leaves dark green; flowers mostly 2-seriate .... P. piperoides Venation palmate; leaves yellowish green; flowers 3-seriate ............... ............................................................................................. P. chrysocladon Fruiting petals closed or nearly so .......................................................... 19 Fruiting petals erect to spreading ........................................................... 36 Leaves shiny, often clasping the stem, with leathery margin; fertile internodes 4–7, the largest ones usually with > 30 flowers per fertile bract; (perhaps exclusively) hyperparasitic on other mistletoes ...... P. dipterum Not the above combination of characters; not normally hyperparasitic; flowers < 30 per fertile bract ............................................................... 20 Leaves widest at or below the middle ..................................................... 21 Leaves widest above the middle .............................................................. 30 Infructescence to 15 mm long; fruit 4 mm diameter, fertile internodes 2 or 3; flowers mostly 3 per fertile bract; if more, 3-seriate ....................................................................................... P. membranifolium Infructescence > 15 mm long (except P. microstachyum); fruit < 4 mm diameter; if more than 3 flowers per fertile bract, 2- or 3-seriate ........ 22 Inflorescences 3-seriate ........................................................................... 23 All or nearly all inflorescences 2-seriate ................................................ 25 Flowers 1–3 per fertile bract; young leaves often translucent red .............................................................................................P. pellucidulum Flowers > 9 per fertile bract; young leaves not translucent red .............................................................................................................. 24

474

V ISCACEAE

24(23). Leaf apex tapering; petiole distinct, ca. 10 mm; blade symmetrical; fruit globose, 3 mm diameter ........................................................... P. nervosum 24. Leaf apex rounded; petiole 3–5 mm, blade often falcate or curved; fruit ellipsoid, 2 mm thick ............................................................... P. perrottetii 25(22). Basal cataphylls mostly 1 pair ................................................................ 26 25. Basal cataphylls 2 or more pairs ............................................................. 29 26(25). Mature internodes terete; dioecious ................................... P. herbert-smithii 26. Mature internodes keeled or quadrangular; monoecious ...................... 27 27(26). Internodes keeled but not quadrangular; leaves to 10 cm long, apex ± acute; inflorescence to 25 mm long ....................................... P. bilineatum 27. Internodes sharply quadrangular to slightly keeled; leaves to 8 cm long, apex obtuse to ± rounded (sometimes minutely apiculate); inflorescence 7–25 mm long ............................................................................. 28 28(27). Leaf apex obtuse; inflorescence 15–25 mm long ................................ P. berryi 28. Leaf apex ± rounded but often minutely apiculate; inflorescence 7–15 mm ......................................................................................... P. microstachyum 29(25). Upper internodes flattened and keeled; leaf apex acute; petiole not flanged; Bolívar, Amazonas ................................................... P. undulatum 29. Internodes quadrangular becoming terete; leaf apex rounded; petiole broadly flanged distally; known only from the type (Bolívar) .............................................................................................. P. steyermarkii 30(20). Inflorescence biseriate ......................................................... P. quadrangulare 30. Inflorescence triseriate ............................................................................ 31 31(30). Flowers < 7 per fertile bract (lowest fertile internode) .......................... 32 31. Flowers > 7 per fertile bract (lowest fertile internode) .......................... 33 32(31). Young leaves often translucent red when dry, to 3.5 cm wide, base mostly long-tapering; mostly dioecious .........................................P. pellucidulum 32. Leaves blackish when dry, to 5 m wide, base contracted to distinct petiole; apparently monoecious ......................................................... P. pteroneuron 33(31). Abaxial midvein, brown-furfuraceous; venation evident, pinnate; mature fruit globose, wider than subtending internode ................. P. hexastichum 33. Abaxial midvein not furfuraceous; venation usually obscure and/or basal; fruit ovoid, no wider than subtending internode ............................... 34 34(33). Leaf base long-tapering to 1–2 m petiole; leaf apex obtuse but not usually rounded .......................................................................... P. longipetiolatum 34. Leaf base short-tapering to petiole < 5 mm long; leaf apex rounded .... 35 35(34). Dioecious; internodes terete; basal cataphylls 1–3 pairs ............ P. schultesii 35. Monoecious (very rarely dioecious); leaves tending to be falcate-asymmetrical; internodes quadrangular to keeled; basal cataphylls mostly 1 pair ........................................................................................ P. perrottetii 36(18). Flowers 9 or more per fertile bract on the longest fertile internode ..... 37 36. Flowers 9 or fewer per fertile bract on the longest internodes .............. 38 37(36). Peduncle with 3 or 4 pairs of sterile bracts; leaves to 9 × 3 cm, obovate to spathulate ............................................................................... P. pulleanum 37. Peduncle with (1)2 pairs of basal bracts; leaves to 7 × 0.7 cm, narrowly lanceolate ................................................................................... P. exiguum 38(36). Peduncle with 3 or 4 pairs of sterile bracts; flowers 7 or fewer per fertile bract ........................................................................................ P. pulleanum

Phoradendron 475

38. 39(38).

39.

40(39). 40. 41(40). 41. 42(41). 42.

Peduncle with 1(2) pairs of sterile bracts; flowers mostly 3–7 per fertile bract ..................................................................................................... 39 Flowers 3 per fertile bract, the apical (median) one staminate, the lateral ones pistillate; fruit tubercular; internodes sharply quadrangular ............................................................................................. P. mucronatum Flowers (1–)3–7 per fertile bract; sex distribution not as above; fruit smooth; internodes terete, keeled, or at most very weakly quadrangular ......................................................................................................... 40 Plants yellowish when dry; leaves coriaceous; internodes terete; basal cataphylls 2–4 pairs ............................................................. P. mairaryense Plants not yellowish when dry; leaves thin; internodes weakly quadrangular or keeled at least above; basal cataphylls 1 pair ...................... 41 Most internodes clavately expanded distally; fruit 7–8 × < 4 mm; staminate flowers below pistillate ones .................................... P. obtusissimum Internodes not clavately expanded distally; fruit < 6 × 6 mm; sex distribution otherwise ...................................................................................... 42 Fruit ± spherical, ca. 6 mm diameter; petals nearly closed, obtuse; terminal fertile internode with 3 flowers per fertile bract ......... P. membranifolium Fruit ovate, to 4 × 2.5 mm; petals erect, acute; terminal fertile internode usually with 1 flower per bract .............................................. P. trinervium

Phoradendron aphyllum Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 412. 1966. Plant monoecious, small, leafless, extremely fragile, internodes terete, basal and intercalary cataphylls lacking;. inflorescences clustered at the nodes, to 1.5 cm long in fruit, peduncle mostly simple; fertile internodes 1–several, flowers 3 or 4 per fertile bract, 2- or 3-seriate; fruit to 6 × 4 mm, ellipsoid; petals inconspicuous, closed. Lower montane forests, 500–1000 m; Bolívar (Río Cuyuní, Uei-tepui). Guyana. ◆Fig. 396. Phoradendron aphyllum is an extremely rare and inconspicuous, small species. It is presently known from only four collections. Phoradendron berryi Rizzini, Rodriguésia 41: 18. 1976. Plant percurrent, with sharply keeled to quadrangular internodes; basal cataphylls 1 pair; leaves to 7 × 2.5 m, thin, lanceolate, apex obtuse, base cuneate, venation rather obscure; inflorescence 1.5–2.5 cm, peduncle mostly simple; fertile internodes 3 or 4; flowers 5–9 per fertile bract, 2-seriate; fruit 3 × 2 mm, ovoid; petals closed. Lowland forests, near sea level to 100 m; Delta Amacuro (Jotajana), Amazonas (south of Macuruco). Endemic. ◆Fig. 398. Phoradendron bilineatum Urb., Bot. Jahrb. Syst. 23(Beibl. 57): 5. 1897.

Phoradendron carinatum Trel., Phoradendron 139. 1916. Percurrent plant, probably monoecious, internodes keeled; basal cataphylls 1 pair, sometimes with a second, slightly higher pair; leaves to 10 × 2.5 m, thin, lanceolate, apex ± acute, base cuneate, petiole 5–10 mm; inflorescence to 2.5 cm, peduncle simple or double; fertile internodes 4 or 5, flowers 3–9 per fertile bract, 2-seriate; fruit ca. 3.5 × 2.5 mm, ellipsoid, petals closed. Evergreen lowland forests, 100–400 m; Amazonas (Laguna El Sillón west of Cerro Camani, San Carlos de Río Negro). Aragua, Barinas; Colombia, Guyana, Suriname, French Guiana, adjacent Brazil. Phoradendron chrysocladon A. Gray, U.S. Explor. Exped., Phan. 15(1):743. 1854. Phoradendron knoopii Warb. ex Trel., Phoradendron 152. 1916. Phoradendron pachyphyllum Trel. Phoradendron 151. 1916. Phoradendron flavens var. longispicum Rizzini, Rodriguésia 30: 63. 1978. Plant monoecious, percurrent, golden green; internodes sharply keeled when young, becoming terete; intercalary and basal cataphylls 1 pair, often acute; leaves to

476

V ISCACEAE

7.5 × 4 cm, ovate, apex acute, base obtuse to truncate; venation palmate, with 3 or 5 strong veins reaching the apex; inflorescence to 5 m long, peduncle 4–7 mm, simple or double, fertile internodes mostly 4, flowers to 18 per fertile bract 3-seriate; fruit 5 × 3 mm, ovoid, petals closed. Lowland to montane forests, 50–2500 m; Bolívar (Amaruay-tepui, Auyán-tepui, La Escalera, Macizo del Chimantá [Amurí-tepui], Santa Elena de Uairén). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Sucre, Yaracuy; Mexico (Chiapas), Central America, Jamaica, Haiti, Lesser Antilles, Colombia, Trinidad, Guyana, Ecuador, Peru, Brazil, northern Bolivia. ◆Fig. 410. Phoradendron crassifolium (Pohl ex DC.) Eichler in Mart., Fl. Bras. 5(2): 125. 1868. —Viscum crassifolium Pohl ex DC., Prodr. 4: 280. 1830. Viscum martinicensis DC., Prodr. 4: 280. 1830. —Phoradendron martinicensis (DC.) Griseb., Fl. Brit. W. I. 314. 1864. Large, stiff-branched plant, percurrent, monoecious, internodes terete; percurrent and lateral innovations with 1 and 1 or 2 pairs of sterile cataphylls, respectively, followed by 2–5 pairs of fertile cataphylls, each of which subtends 1–3 inflorescences; inflorescences also present at foliar nodes; leaves mostly to 17 × 10 cm, coriaeous, ovate to broadly lanceolate, with 3–7 prominent, palmate veins; inflorescences 2–3.5 cm long (in fruit), with 2–6 or more crowded sterile basal internodes followed by 5–9 fertile ones; flowers 3–7 per fertile bract, 2-seriate, the terminal staminate, the others pistillate; fruit ca. 3 mm diameter, globose, petals small, erect. Lowland to montane forests, 50–2000 m; widespread in Bolívar and Amazonas. Apure, Aragua, Distrito Federal, Miranda, Sucre, Táchira, Yaracuy, Zulia; Central America, Lesser Antilles, South America except Chile, Argentina, and Uruguay. ◆Fig. 400. Phoradendron crassifolium is one of the most widely distributed species in the genus. Phoradendron diminutivum E.A. Kellogg, Novon 6: 41. 1996. Rather small and delicate, muchbranched plant, monoecious, both percurrent and with terminal inflorescences, yellowish to olive green when dry; internodes terete, short-papillate; intercalary cataphylls (1

pair) present or absent; basal cataphylls 1 or 2 pairs; leaves 2.5–5 × 1–2 cm, thin, narrowly (ob)lanceolate, apex acute or obtuse, base cuneate, venation obscure, with 3 basal veins; inflorescence to 2.5 m long, peduncle 2–3 mm, simple; fertile internodes 2–4(5), flowers 3–9 per fertile bract, 2-seriate, staminate and pistillate flowers intermixed but staminate flowers more commonly borne proximally; fruit 4 × 3 mm, ovoid, petals erect to spreading somewhat. Evergreen lowland to lower montane forests, 100–800 m; Amazonas (Río Coro Coro, Río Pasimoni). Brazil (Amazonas). Phoradendron dipterum Eichler in Mart., Fl. Bras. 5(2): 109. 1868. Phoradendron tovarense Urb., Bot. Jahrb. Syst. 23(Beibl. 57): 8. 1897. Phoradendron demerarae Trel., Phoradendron 73. 1916. Phoradendron longiarticulatum Rizzini, Revista Fac. Agron. (Maracay) 8(3): 88. 1975. Sparsely branched, percurrent plant, mostly or exclusively hyperparasitic on other mistletoes, dioecious or perhaps also monoecious, sex distribution variable; internodes strongly 4-ridged or 4-winged or somewhat dipterous; basal cataphylls 1 pair, conspicuous, tubular; leaves extremely variable broadly obovate to spathulate or somewhat falcate, apex rounded, base flattened, often clasping the node somewhat or even auriculate, veins basal; inflorescences to 10 cm, peduncle simple; fertile internodes 3–6, flowers ca. 30 or more per fertile bract, in 3–5 series, the pistillate mostly 3-seriate; fruit very small, ca. 2 mm diameter, ovoid to nearly globose, petals closed or nearly so. Evergreen lowland forests, 100–200 m; Bolívar (Río Aparamán). Aragua, Falcón, Miranda; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Paraguay, Argentina. Phoradendron dipterum is widely distributed, but seems to exist in very low numbers at any one locality. It is curiously absent from most of the Amazon Basin. Phoradendron exiguum Trel., Phoradendron 81. 1916. Slender, percurrent, monoecious plant, much branched, internodes terete to slightly quadrangular; basal cataphylls mostly 1

Phoradendron 477

pair; leaves to 7 × 0.7 cm, narrowly lanceolate or even lorate, apex ultimately rounded, base cuneate; venation with 3 basal veins but often obscure; inflorescence to 4 cm, delicate, peduncle double or simple; fertile internodes 4 or 5, flowers 9–11 per fertile bract, 2-seriate; fruit 2.5 × 2 mm, ovoid, petals small, erect or nearly so. Semideciduous forests, 50–100 m; Bolívar (Guri). Falcón, Zulia; northeastern Colombia. Phoradendron exiguum is an inconspicuous species. Phoradendron haughtii Kuijt, Syst. Bot. Monogr. 66: 226. 2003. Plant dioecious, mostly dichotomous through the formation of terminal inflorescences; occasionally, a percurrent shoot develops, characterized by 1 pair of intercalary cataphylls; basal cataphylls 1 pair; leaves to 5.5 × 3 m, thin, (ob)ovate to broadly elliptic, apex mostly rounded, base contracted to a very short petiole; venation palmate, sometimes somewhat obscure; inflorescences 4–7 cm, peduncles simple and very short (2 mm), followed by 6 or 7 fertile internodes; flowers 15–22 (staminate) or fewer (pistillate) per fertile bract, 2- or 3-seriate even on the same inflorescence; fruit 2.5 × 2 mm, globose, petals closed. Evergreen lowland forests, 100– 200 m; Amazonas (San Carlos de Río Negro). Colombia. Phoradendron herbert-smithii Trel., Phoradendron 80. 1916. Phoradendron leptarthrum Rizzini, Rodriguésia 30/31: 188. 1956. Phoradendron crenato-callosum Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 224. 1982. Plant dioecious, to 1.5 m diameter, percurrent, internodes terete when mature; basal cataphylls 1 pair; leaves 4–11 × 2–4 cm, thin, ovate to elliptic, apex rounded to slightly notched, venation palmate; inflorescences mostly 5–10 cm long, peduncle simple or double; fertile internodes to 7, flowers 5– 13(–19) per fertile bract, 2-seriate; fruit 4 × 3 mm, ovoid, tending to keep its shape when dried, petals closed. Semideciduous lowland forests, 100–200 m; Delta Amacuro (Tucupita), Bolívar (Ciudad Bolívar). Distrito Federal, Falcón, Miranda, Sucre, Zulia; Southern Mexico, Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia.

Phoradendron hexastichum (DC.) Griseb., Fl. Brit. W. I. 313. 1860. —Viscum hexastichum DC., Prodr. 4: 282. 1830. Phoradendron bolivianum Trel., Phoradendron 138. 1916. Rather large plant, monoecious, sex distribution diverse, young internodes keeled to quadrangular becoming terete; basal cataphylls mostly 1 pair, sometimes with a second pair above; leaves mostly 7–15 × 3–5 m, (narrowly) obovate, apex rounded, base longtapering into slender, indistinct petiole; venation pinnate, the lower midvein usually strikingly light brown, slightly furfuraceous; inflorescences 3–4(–6.5) cm long, peduncle simple or double, fertile internodes 3–5, flowers 6–10 or more per fertile bract, 3-seriate; fruit 3 × 3 mm, ± globose, petals closed. Lower montane to montane forests, 200– 1300 m; Bolívar (near Campamento Karún, road between El Dorado and Santa Elena, Macizo del Chimantá, Río Canaracuni, Serranía Marutaní, Sierra de Maigualida, Túriba), Amazonas (Cerro Marahuaca, Río Coro Coro, Yutajé). Aragua, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Táchira; southern Mexico, Belize, Honduras, Costa Rica, Panama, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 402. Phoradendron hexastichum is apparently absent from the the Amazon Basin. In the Caribbean the leaves tend to be broadly ovate rather than obovate. Phoradendron inaequidentatum Rusby, Bull. Torrey Bot. Club 27: 137. 1900. —Dendrophthora inaequidentata (Rusby) Trel., Phoradendron 218. 1916. Phoradendron jenmanii Trel., Phoradendron 85. 1916. Phoradendron prancei Rizzini, Revista Fac. Agron. (Maracay) 8(3): 89. 1975. Rather small monoecious plant, profusely branched, mostly dichotomous and bearing terminal inflorescences, rarely percurrent; internodes terete, nodes expanding in age; lateral shoots with 2–7 pairs of basal cataphylls, each innovation with one pair of leaves; leaves to 9 × 6 cm, elliptic, obovate, or somewhat spathulate, apex rounded, base acute, venation palmate but obscure; axillary inflorescences mostly with 1–3 basal sterile internodes, terminal ones apparently with

478

V ISCACEAE

simple peduncles; infructescence to 3 cm, fertile internodes 4 or 5, flowers mostly 1 pistillate and 2 or 3 staminate per fertile bract, the latter apparently limited to proximal fertile internodes; flowers 2- or 3-seriate to nearly 1-seriate; fruit nearly 6 × 4 mm, ovoid, petals slightly parted to erect. Evergreen lowland to montane forests, 50–2000 m; Delta Amacuro (eastern Sierra Imataca), Bolívar (north of Aparamán-tepui, Macizo del Chimantá), Amazonas (Sierra Parima). Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, northwestern Brazil, Bolivia. ◆Fig. 412. Phoradendron inaequidentatum is often recognizable by its reddish brown coloration. Phoradendron kelloggii Kuijt, Novon 6: 42. 1996. Plant much branched, at least partly dichotomous through the formation of terminal inflorescences, internodes terete; basal cataphylls 1(–3), the most distal ones sometimes subtending inflorescences; leaves to 5 × 1.5 cm, lanceolate, apex tapering but ultimately rounded, base acute, venation obscure; inflorescences 1 per axil and also terminal, to 2 cm long, peduncle simple or double; fertile internodes 3 or 4, flowers 1–3 per fertile bract, uniseriate, staminate flowers usually placed below pistillate ones; fruit 4.5 × 4 mm, ovoid, petals erect or even somewhat reflexed. Montane forest and tepui scrub, ca. 1200 m; Amazonas (Cerro Duida). Endemic. Phoradendron longipetiolatum Urb., Bot. Jahrb. Syst. 23(Beibl. 57): 6. 1897. Phoradendron ptarianum Steyerm., Fieldiana, Bot. 28: 222. 1951. Plant monoecious, percurrent, internodes compressed to keeled distally but becoming essentially terete; basal cataphylls 1, 2, or more pairs; leaves to 10(–14) × 3 cm, thin, narrowly obovate to lanceolate, often curved, apex obtuse, base long-tapering to slender, indistinct petiole 1–2 cm long; venation ± pinnate but obscure except for the midvein; staminate and pistillate inflorescences possibly separate, to 2 cm long; peduncle mostly simple, rarely double; fertile internodes 3 or 4, flowers 6–12 per fertile bract, 3-seriate; fruit 3 mm diameter, ovoid, petals closed. Rocky tepui slopes and summits, 1800–2400 m; Bolívar (Macizo del Chimantá [Apacarátepui], Ptari-tepui). Aragua.

Phoradendron mairaryense Ule, Notizbl. Bot. Gart. Berlin-Dahlem 6: 291. 1920. Phoradendron sulfuratum Rizzini, Revista Fac. Agron. (Maracay) 8(3): 89. 1975. Yellowish, rather large plant, monoecious(?), both percurrent and (lateral branches) dichotomous with terminal inflorescences; internodes terete, older nodes swollen; basal cataphylls 2–4 pairs, the upper ones placed high on the stem; leaves to 14 × 3 cm, narrowly oblong, coriaceous, apex rounded to obtuse, base tapering cuneately to indistinct petiole 4 mm long; venation ± palmate, with 3–5 prominent veins; inflorescence to 4.5 cm long, with 1 or more sterile basal internodes; fertile internodes 4 or 5, flowers 3–5(–7) per fertile bract, mostly 2seriate; fruiting petals erect. Evergreen lowland to montane forests, 200–1400 m; Bolívar (Gran Sabana), Amazonas (Río Mawarinuma). Guyana, Brazil. Phoradendron membranifolium Kuijt, Syst. Bot. Monogr. 66: 295. 2003. Dioecious(?), percurrent plant, internodes compressed and keeled above, quadrangular below and when older; basal cataphylls one pair, very small; leaves to 7 × 3 m, ovate to elliptic, thin, apex rounded, base tapering to thin petiole ca. 4 mm long; venation evident, with prominent midvein and 2 strong basal lateral veins reaching far toward the apex; infructescence to 1.5 m long, peduncle simple or double, fertile internodes 2 or 3, flowers 3(–5) per fertile bract, 3-seriate; fruit 6 mm diameter, globose, petals ± closed but not meeting. Lower montane forests, 500–600 m; Bolívar (southeast of Los Pijiguaos). Endemic. Phoradendron microstachyum Kuijt, Syst. Bot. Monogr. 66: 301. 2003. Densely leafy, monoecious, percurrent plant, internodes sharply quadrangular; basal cataphylls 1 pair, very small; leaves to 8 × 2.5 cm, lance-elliptic, apex obtuse to slightly apiculate, base tapering acutely to 3 mm petiole; venation pinnate but obscure; inflorescence 7– 15 mm long, peduncle very short, simple; fertile internodes 3 or 4, flowers 5–7 per fertile bract, 2-seriate, the staminate ones few and borne on distal parts of fertile internodes; fruit 2 × 1.5 mm, globose, petals small, ± closed. Shrubs in savannas, 100–200 m; Amazonas (base of Cerro Moriche). Endemic.

Phoradendron 479

Phoradendron microstachyum is a small, inconspicuous plant with one of the smallest inflorescences in South American Phoradendron. Phoradendron morsicatum Rizzini, Mem. New York Bot. Gard. 29: 33. 1978. Plants dichotomous, rarely with a percurrent shoot bearing 1 pair of intercalary cataphylls, orange-brown to yellowish; internodes terete, stout; basal cataphylls mostly 1(2) pairs; leaves to 9 × 5 cm, elliptic to lanceolate, coriaceous, apex and base ± acute; venation obscure; inflorescences terminal and lateral on older nodes, peduncle simple, fertile internodes 2 or 3, flowers 5–9 per fertile bract, 2-seriate; fruit 5 × 3 mm, ovoid, petals closed. Low open scrub on tepui slopes, elevation uncertain; Bolívar (Ilú-tepui). Guyana. Phoradendron mucronatum (DC.) Krug & Urb., Bot. Jahrb. Syst. 24: 352. 1897. —Viscum mucronatum DC., Prodr. 4: 282. 1830. Phoradendron emarginatum Mart. ex Eichler in Mart., Fl. Bras. 5(2): 118. 1868. Phoradendron ottonis Eichler in Mart., Fl. Bras. 5(2): 119. 1868. Phoradendron caracasanum Urb., Bot. Jahrb. Syst. 23(Beibl. 57): 4. 1897. Phoradendron lyonii Trel., Phoradendron 116. 1916. Phoradendron caerulesens Trel., Bull. Torrey Bot. Club 48: 330. 1921. Phoradendron chaguaramasarum Trel., Bull. Torrey Bot. Club 48: 330. 1921. Phoradendron caracasanum var. parvifolium Rizzini, Rodriguésia 30/31: 182. 1956. Phoradendron ottonis var. oblongifolium Rizzini, Rodriguésia 30/31: 182. 1956. Phoradendron emarginatum var. truncatoretusum Rizzini, Rodriguésia 46: 72. 1978. Plant monoecious, densely leafy, percurrent, often bluish gray; internodes usually quadrangular (sometimes compressed distally); basal cataphylls one small pair, acute; leaves to 4.5 × 3 cm, thin, elliptic to obovate, apex obtuse, rounded, and often notched, base acute to obtuse; venation palmate, with 3 prominent veins, the midvein reaching the apex, the 2 lateral ones nearly so; inflorescences to 2 cm long, 1–3 per leaf

axil, peduncle mostly simple, fertile internodes 2–4, flowers 3 per fertile bract, the apical (median) one staminate, the lateral ones pistillate; fruit 2.5 mm diameter, globose, commonly tuberculate, petals erect. Savannas and associated scrub, 50–1100 m; Bolívar (Cerro El Medano south of Caicara, between Guanaraparo and Cerro El Cume, Río Paragua, Serranía Cerbatana), Amazonas (Raudal de Atures). Anzoátegui, Apure, Aragua, Carabobo, Distrito Federal, Falón, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Trujillo, Zulia; Mexico, Guatemala, Belize, Panama, Lesser Antilles, Colombia, Trinidad, Guyana, eastern Brazil, Bolivia, Paraguay, northern Argentina. Phoradendron mucronatum is apparently absent from the Amazon Basin. It is an extremely variable species in the Venezuelan area, in inflorescence composition, leaf shape, and degree of internodal flattening, if any. Phoradendron nervosum Oliv., Vidensk. Meddel. Dansk Naturhist. Føren. Kjøbenhavn 1964: 175. 1864. Phoradendron tubulosum Urb., Bot. Jahrb. Syst. 23(Beibl. 57): 5. 1897. Phoradendron granaticola Trel., Phoradendron 70. 1916. Phoradendron taeniicaule Rizzini, Rodriguésia 30/31: 191. 1956. —Phoradendron granaticola var. taeniicaule (Rizzini) Rizzini, Rodriguésia 46: 65. 1978. Large monoecious, percurrent, often yellowish plant; internodes keeled and compressed; basal cataphylls usually 1 pair, fused and conspicuous, often light-margined; leaves 5–20 × 3–7 m, thin, ovate, apex somewhat tapering but ultimately rounded, base contracted to a distinct petiole ca. 1 cm long; venation palmate, with 5–7 major veins; inflorescence to 4(–6) cm long, peduncle simple, fertile internodes 3–5, flowers 9 or more per fertile bract, mostly 3-seriate in the flora area, sex distribution irregular, staminate flowers rare; fruits 3 mm diameter, globose, partially embedded in swollen axis, petals inconspicuous, closed. Not recorded from the flora area, but almost certainly present. Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Mérida, Miranda, Sucre, Táchira, Trujillo, Yaracuy; central Mexico, Central America, Colombia, Ecuador, Peru, Bolivia. ◆Fig. 403.

480

V ISCACEAE

Outside the flora area, the elevation is mostly 50–2000 m. Phoradendron nervosum is a variable species known from three apparently distinct populations. Phoradendron northropiae Urb., Mem. Torrey Bot. Club 12: 33. 1902. Phoradendron excisum Rizzini, Rodriguésia 30/31: 192. 1956. Rather small, erect, monoecious plant, dichotomous but often seemingly percurrent through abortion of one axillary innovation; internodes terete but slightly keeled directly below young nodes; basal cataphylls 1 pair, somewhat tubular, blunt; leaves to 6 × 3.5 cm, obovate, coriaceous, apex rounded to notched, base contracted to petiole ca. 3 mm long; venation pinnate but obscure; inflorescence < 2 cm, slender, peduncle double; fertile internodes 2 or 3, flowers 5–9, 2-seriate, the apical 1–3 flowers above each fertile bract staminate, others pistillate; fruit 3 × 2 mm, ellipsoid, petals closed. Forested tepui slopes, 100–1200 m; Amazonas (east branch of Caño Negro). Bahamas, Guyana, Peru, Brazil, Bolivia. Phoradendron northropiae is a species of wide distribution but very sparse occurrence. It strongly resembles the much larger P. racemosum, which has thin leaves with evident venation and different shape. Phoradendron obtusissimum (Miq.) Eichler, Fl. Bras. 5(2): 134m. 1868. —Viscum obtusissimum Miq., Linnaea 18: 602. 1844. Phoradendron acinacifolium var. surinamense Rizzini, Mem. New York Bot. Gard. 29: 32. 1978. Phoradendron ichythystoma Rizzini, Ernstia 24: 9. 1984. Monoecious, percurrent plant, internodes terete or slightly keeled, often strikingly clavate distally; basal cataphylls 1 pair, very small and inconspicuous; leaves to 10 × 3 cm, elliptic to (ob)lanceolate, often somewhat curved; apex rounded, base acutely tapering into indistinct petiole; venation palmate, with 3–5 basal veins; inflorescence to 2 cm long, slender; peduncle generally double, fertile internodes 3, flowers (1–)3–6 per fertile bract, 2-seriate; staminate flowers (where present) placed below pistillate ones; fruit to 7–8 × 4 mm, ellipsoid-fusiform, petals erect to spreading. Evergreen lowland to montane

forests, 50–1500 m; Bolívar (Altiplanicie de Nuria, Macizo del Chimantá [Abacapátepui], Río Suapure), Amazonas (Caño Piedra near Cerro Cuao, between Caño San Miguel and Maroa, San Carlos de Río Negro, San Fernando de Atabapo). Mérida, Miranda, Táchira; Costa Rica, Panama, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 406. Phoradendron pellucidulum Eichler in Mart., Fl. Bras. 5(2): 112. 1868. Phoradendron semivenosum Rizzini, Rodriguésia 30/31: 189. 1956. Phoradendron williamsii Rizzini, Rodriguésia 30/31: 188. 1956. Phoradendron insigne Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 415. 1966. Phoradendron pseudomucronatum Rizzini, Rodriguésia 41: 18. 1976. Phoradendron semivenosum Rizzini, Rodriguésia 41: 17. 1976, non Phoradendron semivenosum Rizzini 1956. Phoradendron semivenosum var. agostinii Rizzini, Rodriguésia 46: 86. 1978. Phoradendron semivenosum var. longipes Rizzini, Rodriguésia 46: 86. 1978. Moderate-sized mostly dioecious plant, usually percurrent, often with lacquered appearance, internodes terete to somewhat compressed-keeled, especially just below the nodes; basal cataphylls 1 pair, small and inconspicuous; leaves to 7.5 × 3.5 m, often translucent reddish when young, broadly (ob)ovate to elliptic, apex rounded, base longtapering to slender petiole; venation usually evident and with 2 strong basal lateral veins, midvein reaching the apex or nearly so; inflorescences to 8–20 mm long, peduncle simple or double, fertile internodes 2 or 3(–5), flowers 1–4 pistillate and to 21 staminate ones per fertile bract; fruit to 4 × 3 mm, ellipsoid, petals closed but not meeting. Evergreen lowland to montane forests, 100–1300 m; Bolívar (widespread), Amazonas (Caño Caname, La Esmeralda, Río Coro Coro, Río Guayapo, Río Mawarinuma, Río Pasimoni, Yavita). Apure; Colombia, Guyana, Peru, Brazil, Paraguay. ◆Fig. 401. Phoradendron perrottetii (DC.) Eichler in Mart., Fl. Bras. 5(2): 112. 1868. —Viscum perrottetii D.C., Prodr. 4: 280. 1830.

Phoradendron 481

Phoradendron dimidiatum (Miq.) Eichler in Mart., Fl. Bras. 5(2): 134m. 1868. —Viscum dimidiatum Miq., Linnaea 18: 58. 1844. Phoradendron dunstervillorum Rizzini, Rodriguésia 41: 20. 1976. Phoradendron scariosum Rizzini, Rodriguésia 41: 17. 1976, non Phoradendron scariosum Rizzini 1956. Rather large plant, monoecious or dioecious, percurrent; internodes somewhat quadrangular to keeled; basal cataphylls 1(2) pairs; leaves to 11 × 5 m, lanceolate to (ob)ovate, thin; larger ones tending to be asymmetrical or even falcate, venation palmate or nearly so; inflorescence 3–4 cm long, peduncle simple or compound, fertile internodes 3 or more, flowers to 13–19 per fertile bract; fruit 3 × 2 mm, ellipsoid, petals closed. Evergreen lowland to lower montane forests, 100–900 m; Bolívar (southeastern base of Auyán-tepui, Canaima, base of Cerro Abismo, Río Asa, Río Karaurín, Santa Elena de Uairén), Amazonas (San Carlos de Río Negro). Anzoátegui, Distrito Federal, Lara, Mérida, Miranda, Monagas, Táchira; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 404. Phoradendron piperoides (Kunth) Trel., Phoradendron 145. 1916. —Loranthus piperoides Kunth in H.B.K., Nov. Gen. Sp. 3: 443. 181. 1818 [1820]. —Viscum piperoides (Kunth) D.C., Prodr. 4: 281. 1830. Viscum dichotomum Bert. ex Spreng., Syst. Veg. 1: 488. 1825 [1824], non Viscum dichotomum D. Don 1825. Viscum fockeanum Miq., Linnaea 18: 60. 1844. Phoradendron glauco-lutescens Rizzini, Rodriguésia 41: 23. 1976. Monoecious, percurrent plant, internodes terete, with 1 pair of intercalary cataphylls; basal cataphylls several pairs; leaves to 14 × 7 m, lanceolate, apex and base mostly acute, margin often undulate; venation pinnate, evident, the midvein reaching the apex; inflorescence to 8 m long, peduncle simple with several sterile internodes, fertile internodes 5–8, flowers to 15 per fertile bract, mostly 2seriate; staminate flowers few, in various positions; fruit 3–4 × 2.5 mm, globose to ovoid, petals erect. Evergreen lowland to montane forests and scrub, near sea level to 1500 m;

Delta Amacuro (widespread), Bolívar (widespread), Amazonas (Isla Ratón, Puerto Ayacucho, Río Atabapo, Río Negro). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; widespread and common from central Mexico, Central America, the Caribbean, South America except Chile and Uruguay. ◆Fig. 407. Phoradendron platycaulon Eichler in Mart., Fl. Bras. 5(2): 108. 1868. Monoecious, pendent, percurrent plant, internodes conspicuously keeled and flattened in the same plane, becoming terete in age; intercalary and basal cataphylls 1 pair, tubular; leaves 1.5–5 cm × 2–3 mm, thin, narrowly lanceolate to elliptic, apex acute or obtuse, base acute, venation obscure; inflorescence to 2 m long, peduncle double, fertile internodes 2 or 3(4), flowers 3 per fertile bract, the apical (median) one staminate, lateral ones pistillate; fruit to 6 × 4 mm, ovoid, smooth, petals closed to erect. Evergreen lowland forests, 50–200 m; Amazonas (Caño Cotúa near base of Cerro Yapacana, Río Ventuari, San Carlos de Río Negro, Yavita). Apure, Táchira; Colombia, Peru, southern Brazil, Bolivia. ◆Fig. 405. Phoradendron platycaulon is an unmistakable species known especially from the Amazonian basin. Phoradendron pteroneuron Eichler in Mart., Fl. Bras. 5(2): 127. 1868. Phoradendron rugulosum Urb., Bot. Jahrb. Syst. 23(Beibl. 57): 13. 1897. Monoecious(?), percurrent plant, often of varnished appearance, internodes ± keeled to quadrangular, becoming terete or nearly so; basal cataphylls (1)2–4 pairs; leaves to 9 × 5 cm, thin, elliptic to obovate, apex commonly rounded, base obtuse to nearly acute; venation pinnate, mostly distinct, the adaxial midvein often sharply ridged; inflorescence 3–5 m, peduncle commonly with 2 or more sterile internodes, fertile internodes 3–7, flowers 3 or 5(–7) per fertile bract, nearly always 2-seriate; fruit 4 × 3 mm, ovoid, petals closed. Semideciduous to evergreen lowland to montane forests, 100–1400 m; Bolívar (Cerro Altamira east of Ciudad Piar, Cerro Picacho, Río Túriba, Santa Bárbara). Lara, Táchira; western Colombia, Guyana, Suriname, Brazil, Bolivia.

482

V ISCACEAE

Phoradendron pteroneuron is scarce in the Amazon basin. Phoradendron pulleanum E.H.L. Krause, Recueil Trav. Bot. Néerl. 22: 346. 1925. Dioecious(?) plant, percurrent, internodes terete, rather stout; basal cataphylls several pairs; leaves to 9 × 3 m, coriaceous, obovate to spathulate, apex rounded, base cuneate; only the midvein evident; pistillate inflorescence 2–3 cm long, peduncle mostly with ca. 4 sterile internodes, fertile internodes 3 or more, flowers 7–9 per fertile bract, 2-seriate; fruit 2.5 × 2 mm, petals open to erect. Edge of disturbed cloud forest; Amazonas (locality lost). Distrito Federal; Guyana, Suriname, French Guiana. Phoradendron quadrangulare (Kunth) Griseb., Fl. Brit. W. I. 711. 1864. —Loranthus quadrangulare Kunth in H.B.K., Nov. Gen. Sp. 3: 444. 1818 [1820]. —Viscum quadrangulare (Kunth) DC., Prodr. 4: 283. 1830. Phoradendron venezuelense Trel., Phoradendron 111. 1916. Phoradendron huricola Trel., Bull. Torrey Bot. Club 54: 471. 1927. Phoradendron venezuelense var. filispicum Rizzini, Ernstia 24: 3. 1984. Phoradendron rubrum var. longipedunculatum Rizzini, Ernstia 32: 9. 1985. Monoecious plant, percurrent, muchbranched, internodes quadrangular; basal cataphylls 1 pair, small, acute; leaves to 6.5 × 2.5 cm, oblanceolate or elliptic to nearly spatulate, thin, apex rounded to somewhat acute, base acute to cuneate, venation with 3 slender veins in basal pattern; inflorescence slender, to 3 cm long, peduncle mostly simple, fertile internodes 3–5, flowers 3–7 per fertile bract, 2-seriate, staminate flowers few and irregularly distributed; fruit 3 mm diameter, globose, petals closed. Savannas, second growth forest, cultivated trees, near sea level to 1500 m; Delta Amacuro (Tucupita), Bolívar (Ciudad Bolívar, east of Kavanayén, La Paragua, Las Trincheras, between San Félix and Puerto Ordáz, slopes of Serranía de los Pijiguaos), Amazonas (Río Cataniapo, San Juan de Manapiare). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Guárico, Mérida, Miranda, Portuguesa, Táchira, Zulia; Mex-

ico, Central America, West Indies, South America except Chile and Uruguay. ◆Fig. 399. Phoradendron quadrangulare is the most widely distributed species of the genus. It is not satisfactorily separated from one or two extraterritorial species, or from P. trinervium within the flora area, and has a complex nomenclatural history, most of which lies outside the flora area. Phoradendron racemosum (Aubl.) Krug & Urb., Bot. Jahrb. Syst.24: 46. 1897. —Viscum racemosum Aubl., Hist. Pl. Guiane 895. 1775. —Viscum flavens Sw., Prodr. 32. 1788, nom. superfl. —Phoradendron flavens (Sw.) Griseb., Fl. Brit. W. I. 313. 1860. Viscum latifolium Lam., Encycl. 3: 57. 1789, non Viscum latifolium Sw. 1797, nec Viscum latifolium Buch.-Ham. ex D. Don 1825. Viscum glandulosum Miq., Linnaea 18: 60. 1844. Phoradendron cayennense Eichler in Mart., Fl. Bras. 5(2): 129. 1868. Monoecious plant, dichotomous, large, internodes to 13 m long, terete to slightly keeled; basal cataphylls mostly 2 or 3 pairs; leaves to 13 × 10 cm, mostly broadly ovate or lanceolate to elliptic, thin, apex obtuse to rounded, base contracted, mostly obtuse; venation pinnate, evident; inflorescences often densely clustered at the nodes, each to 6 cm long, peduncle mostly with 2 sterile internodes; fertile internodes 4 or 5, flowers 5–9, 2-seriate, the apical one staminate, others pistillate; fruit 3.5 × 2.5 mm, ovoid, petals closed. Scrub and forests, near sea level to 1200 m; Delta Amacuro (Río Orocoima), Bolívar (Altiplanicie de Nuria, Amaruaytepui, Cerro Ichún, Entrerios, northwest of Kavanayén, Río Aparamán, Río Canaracuni, Río Erebato, San Ignacio, Santa Elena de Uairén), Amazonas (Yavita). Anzoátegui, Aragua, Zulia; Costa Rica, Bahamas, Greater Antilles, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 409. Phoradendron scariosum Rizzini, Rodriguésia 30/31: 189. 1956, non Phoradendron scariosum Rizzini 1976. Plant dioecious(?), dichotomous, shoot apex aborting after formation of one pair of foliage leaves; internodes terete, basal cataphylls 1 or 2 pairs, conspicuously white-mar-

Phoradendron 483

gined when dry; leaves to 11.5 × 5 cm, somewhat coriaceous, obovate to elliptic, apex rounded, base tapering ± acutely; venation basal, with 3–5 faintly evident veins; pistillate inflorescence to 1.5 cm long, peduncle simple or double, fertile internodes ca. 3, flowers 1–3(–5) per fertile bract, 2-seriate; fruit 2.5 × 2 mm, ovoid, petals closed. Gallery forests on Xylopia, 100–300 m; Bolívar (La Paragua). Miranda. Phoradendron schultesii Kuijt, Syst. Bot. Monogr. 66: 408. 2003. Dioecious plant, mostly percurrent (some smaller branches with terminal inflorescences and therefore dichotomous), internodes terete; basal cataphylls 1–3 pairs; leaves to 9 × 4 cm, coriaceous, (ob)ovate-elliptic, apex rounded, base acute; venation obscure, possibly pinnate; inflorescences 2.5– 3.5 cm long, peduncle simple or double, fertile internodes 3 or 4, flowers 7–16 per fertile bract, 3-seriate; fruit 3.5 × 2.5 mm, ellipsoid to ovoid, petals nearly closed. Evergreen lowland forest edges, ca. 100 m; Amazonas (San Carlos de Río Negro). Endemic. Phoradendron steyermarkii Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 295. 1982. Monoecious(?) plant, percurrent, internodes somewhat quadrangular, becoming terete; basal cataphylls 2 or 3 pairs; leaves to 9 × 4 cm, broadly ovate to elliptic, apex rounded, base contracted to a cuneate, flanged petiole; venation pinnate, especially evident abaxially; inflorescence to 4.5 m long, peduncle simple or sometimes double, fertile internodes 5 or 6, flowers 5–7 per fertile bract, 2-seriate; petals closed in fruit. Montane gallery forests, ca. 1100 m; Bolívar (Río Pacairao near Kavanayén). Endemic. Phoradendron strongyloclados Eichler in Mart., Fl. Bras. 5(2): 109. 1868. Phoradendron surinamense Pulle, Enum. Vasc. Pl. Surinam 155. 1906. Phoradendron essequibense Trel., Phoradendron 149. 1916. Phoradendron johnstonii Trel., Phoradendron 149. 1916. Monoecious plant with innovations bearing 2 or 3 pairs of leaves, after which a terminal inflorescence, followed by a dichotomy; internodes terete; basal and intercalary

cataphylls 1 pair, tubular, yellow-margined when dry; leaves to 4 × 2 cm, coriaeous, lanceolate to ovate-elliptic, apex acute or rounded, base acute or nearly so, venation obscure except for 3 major basal veins; inflorescence 1–3 m long, peduncle simple or double; fertile internodes mostly (2)3–5, flowers 3 per fertile bract, the apical (median) one staminate, lateral ones pistillate; fruit 4 × 3 mm, ovoid, petals erect to somewhat spreading. Evergreen lowland to montane forests, 50–1300 m; widespread in Bolívar and Amazonas. Guyana, Suriname, Brazil, Bolivia. ◆Fig. 397. Phoradendron strongyloclados is apparently absent from the main Amazonian region. Phoradendron triflorum E.A. Kellogg, Novon 6: 44. 1996. Monoecious plants, dichotomous through abortion of the shoot apex; internodes strongly quadrangular; basal cataphylls 1 pair, inconspicuous; leaves to 5 × 3 cm, obovate to oblanceolate, thin, apex truncate to emarginate, base acute, venation pinnate but obscure; inflorescence to 1.5 cm, doubling in length in fruit; sterile internodes 2, fertile internodes 4–7, flowers 3 per fertile bract, the apical (median) one staminate, lateral ones pistillate; fruit 3 × 2 mm, petals erect. Whitesand savanna and scrub, 100–900 m; Bolívar (Río Hacha), Amazonas (Santa Bárbara del Orinoco). Colombia. Phoradendron trinervium (Lam.) Griseb., Fl. Brit. W. I. 314. 1860. —Viscum trinervium Lam., Encycl. 3: 57. 1789. —Phoradendron verticillatum Fawc. & Rendle, Fl. Jamaica 3: 96. 1914, nom. superfl., non Phoradendron verticillatum (L.) Druce 1914. Phoradendron apertum Trel., Phoradendron 104. 1916. Phoradendron appunii Trel., Phoradendron 104. 1916. Phoradendron sanctae-martae Trel., Phoradendron 105. 1916. Phoradendron zuloagae Trel., Phoradendron 105. 1916. Phoradendron baileyae Trel., Bull. Torrey Bot. Club 54: 476. 1927. Phoradendron exile Rizzini, Rodriguésia 30/31: 182. 1956. —Phoradendron baileyae var. exile (Rizzini) Rizzini,

484

V ISCACEAE

Rodriguésia 46: 51. 1978. Phoradendron treleasei Rizzini, Rodriguésia 30/31: 183. 1956. Phoradendron theloneuron Rizzini, Rodriguésia 41: 16. 1976. Monoecious, much-branched, percurrent plant, internodes somewhat quadrangular but only slightly so; basal cataphylls mostly 1 pair, deeply bifid; leaves commonly to 4.5 × 2 cm, (narrowly) obovate to nearly elliptic, apex rounded, slightly notched or nearly acute, base tapering, venation with 3 or 5 basal veins; inflorescence 1–1.5 cm long, peduncle simple or double, fertile internodes 3– 6, flowers mostly 3–7 per fertile bract, 2seriate; fruit 4 × 2.5 mm, petals small, ± acute, erect. Evergreen lowland to lower montane forests, near sea level to 1000 m; Bolívar (widespread), Amazonas (Puerto Ayacucho, Río Cataniapo). Apure, Carabobo, Distrito Federal, Falón, Guárico, Miranda, Monagas, Portuguesa, Sucre, Zulia; two isolated populations in Mesoamerica, Guyana, Suriname, French Guiana, coastal portions of Brazil. ◆Fig. 408. Phoradendron trinervium is not always easily separated from P. quadrangulare except for its erect fruiting petals.

Phoradendron ayliffei Philcox, Fl. Trinidad 2: 605. 1978. Plant monoecious, large, percurrent, internodes carinate to strongly compressed when young, especially distally; basal cataphylls variable in number, mostly 2–4 pairs; leaves commonly to 11(–17) × 4(–7) cm, (broadly) lanceolate to ovate, apex acute or even acuminate, base generally acute, venation pinnate, the midvein abaxially prominent and reaching the apex; inflorescences often numerous, mostly 4–7 cm long, peduncle with as many as 5 sterile basal internodes, fertile internodes 6–10, flowers 5–7 per fertile bract, 2-seriate, apical flower above a fertile bract mostly staminate, others pistillate, or sexes intermixed irregularly; fruit 3 × 2 mm, ovoid, petals closed. Lower montane to high montane tepui forests, 600–2500 m; Bolívar (Auyán-tepuí, Cerro Guaiquinima, Cerro Jaua, Chimantá Massif, Gran Sabana), Amazonas (Sierra Parima). Southern Mexico (Chiapas), Central America, Lesser Antilles, Colombia, Ecuador, Peru, Brazil (southeastern Atlantic area), Bolivia. ◆Fig. 411.

Phoradendron tunaeforme (DC.) Eichler in Mart., Fl. Bras. 5(2): 108. 1868. —Viscum tunaeforme DC., Prodr. 4: 284. 1830. Monoecious plant, squamate, percurrent, internodes conspicuously flattened and keeled; intercalary cataphylls absent; basal cataphylls 1 pair, tubular; inflorescence mostly 3–7 mm long, peduncle simple, fertile internodes 1(2). flowers 3 per fertile bract, the apical (median) one staminate, lateral ones pistillate; fruit 6 × 5 mm, globose, smooth or nearly so, petals minute, erect. Lower montane forests, ca. 400 m; Bolívar (Río Túriba). Brazil (mostly eastern). Phoradendron undulatum (Pohl ex DC.) Eichler in Mart., Fl. Bras. 5(2): 122. 1868. —Viscum undulatum Pohl ex DC., Prodr. 4: 282. 1830. Phoradendron herminieri Trel., Phoradendron 131. 1916. Phoradendron chrysocarpum var. stehlei Trel. & Stehlé, Bull. Bot. Soc. France 92: 262. 1945.

Fig. 396. Phoradendron aphyllum

Phoradendron 485

Fig. 397. Phoradendron strongyloclados

Fig. 398. Phoradendron berryi

486

V ISCACEAE

Fig. 399. Phoradendron quadrangulare

Fig. 400. Phoradendron crassifolium

Phoradendron 487

Fig. 401. Phoradendron pellucidulum

Fig. 402. Phoradendron hexastichum

488

V ISCACEAE

Fig. 403 Phoradendron nervosum

Fig. 404. Phoradendron perrottetii

Phoradendron 489

Fig. 405. Phoradendron platycaulon

Fig. 406. Phoradendron obtusissimum

Fig. 407. Phoradendron piperoides

490

V ISCACEAE

Fig. 408. Phoradendron trinervium

Fig. 409. Phoradendron racemosum

Phoradendron 491

Fig. 410. Phoradendron chrysocladon

Fig. 411. Phoradendron undulatum

492

V ISCACEAE

Fig. 412. Phoradendron inaequidentatum

VITACEAE by Francisco Delascio Chitty and Julian A. Steyermark Climbing shrubs and vines (in Venezuela), rarely erect shrubs or trees, usually unarmed, often with watery sap, the stems often swollen or jointed and bearing tendrils, occasionally winged. Leaves mainly alternate, usually 2-ranked, often pellucid-punctate, simple, often palmately lobed, palmately 3–5- or bipinnately compound; petiole articulate at the base of the stem and thickened; stipules usually present. Inflorescence terminal or axillary, cymose, racemose, spicate, or paniculate, usually opposite a leaf. Flowers minute, numerous, bisexual or unisexual, actinomorphic. Calyx entire or 4- or 5-toothed or lobed; sepals (3)4 or 5(–7), distinct or connate at the base; petals equal in number to the sepals, valvate in bud, minute or obsolete, spreading-recurved in anthesis, distinct or apically connate, separating at the base and falling as an early deciduous cap. Stamens as many as the petals and opposite them, somewhat perigynous, arising from the base of the disk; anthers free or connate, usually introrse, 2-locular, dehiscing longitudinally. Pollen tricolporate. Disk intrastaminal, often annular or cupulate, sometimes with 5 distinct glands. Pistil 1, the ovary superior but usually somewhat sunken in the disk, mainly 2(3–8)locular; ovules 1 or 2 in each locule, anatropous, ascending; style 1, short or none; stigma discoid, capitate, or rarely 4-fid. Fruit a berry, usually 1- or 2(3–6)-locular. Seeds 1–4(–6), erect with a bony testa, 2 deep adaxial grooves and a conspicuous abaxial chalazal knot; endosperm cartilaginous. Chiefly from tropical regions of the world, but also in temperate portions; 18 genera and 800 species, 1 genus and 10 species in the flora area. Vitis vinifera L. is sometimes cultivated in the flora area, for example along fences in San Carlos de Río Negro.

Cissus 493

1. CISSUS L., Sp. Pl. 117. 1753. Mostly climbing plants, herbaceous or usually woody, sometimes fleshy. Roots often tuberous. Stems usually bearing tendrils opposite the leaves, terete or angulate, glabrous to pubescent. Leaves 3–5-foliolate or simple, toothed or often lobed; stipules in pairs, usually caducous. Inflorescence opposite the leaves, consisting of corymbiform or umbelliform cymes; bracts and bracteoles present. Flowers 4merous, pedicellate. Calyx cupular, entire or 4-lobed, valvate in bud; corolla ovoid or campanulate, petals valvate, cucullate. Stamens inserted at the base of the disk. Disk 4-lobed. Ovary 2-locular with 2 ovules. Fruit a 1(2)-seeded berry. Seeds obovoid, generally trigonous. Pantropics; ca. 350 species, 17 in Venezuela, 10 of these in the flora area. Key to to the Species of Cissus 1. 1. 2(1).

2.

3(2).

3.

4(1).

4.

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

6.

Leaves simple ............................................................................................. 2 Leaves compound, 3-foliolate or digitate .................................................. 4 Leaves coriaceous, 3-veined, glabrous, cuneate at base, abruptly caudate at apex, the acumen 7–11 mm long; leaf margins remotely dentate and revolute; inflorescence falsely terminal; flowers white; fruits ellipsoid ............................................................................................ C. venezuelensis Leaves not coriaceous, 5-veined, canescent, villous, hispid, or glabrous, base generally cordate, rounded, or truncate, apex acute to subacuminate, leaf margins densely denticulate or dentate, not revolute; inflorescence leaf opposed; flowers greenish yellow; fruits not ellipsoid ..................................................................................................... 3 Leaves chartaceous, cordiform, glabrescent or puberulous-sericeous; tendrils simple, glabrous; stipules 1–4 mm long, triangular, subovate to rhombic; flower buds subfusiform, 1–2(–3) mm long, pedicels 1–4 mm long, sericeous or glabrous; fruits lenticellate and bottle-shaped ................................................................................................ C. descoingsii Leaves fleshy or chartaceous, variable in shape, sometimes laciniate or lobed, glabrous or densely pubescent; tendrils simple or bifid, puberulous or glabrous; stipules 1–2 mm long, falcate or ovate; flower buds ellipsoid, 0.5–1 mm long, pedicels 1–7 mm long, glabrous; fruits globose, smooth ...................................................................... C. verticillata Leaves digitately 5- or 6-foliolate, rarely 4- or 7-foliolate, leaflets sessile, normally lanceolate-oblong or elliptic, 0.5–2 cm wide, margins aristate-serrate or crenulate; inflorescence red ....................... C. palmata Leaves 3-foliolate, leaflets sessile or petiolulate, oblong, ovate-lanceolate, obovate, elliptic, rhombic, or lobulate, > 2 cm wide, margins denticulate, dentate, crenulate, or softly undulate; inflorescence orange, greenish yellow, green, pink, reddish orange, or red ........................... 5 Stems cylindrical ........................................................................................ 6 Stems not cylindrical ................................................................................. 7 Stems spinose, canescent or sparsely pubescent; terminal leaflet petiolate, 3–20 mm long, rhombic to suborbicular; peduncle 6.7–12 cm long, reddish; flowers red; fruit purple ...................................... C. spinosa Stem not spinose, sericeous or puberulous; lower outer margin of the lateral terminal leaflet sessile or subsessile, 0–1.3 mm long, obovate to

494

7(5).

7. 8(7).

8.

9(8).

9.

V ITACEAE

elliptic; peduncle 0.5–2.3 cm long, green; flowers green; fruit green or whitish-translucid ....................................................................... C. trianae Stems 4–6-winged, the epidermis exfoliating; terminal leaflet elliptic to obovate, with numerous marginal teeth; peduncle 6–12 cm long, reddish pink; fruit orbicular or subpyriform, 5–6 mm diameter ....... C. erosa Stems generally angled; without the above characters ........................... 8 Plants glabrous or glabrescent; terminal leaflet ovate, subsessile, scabrous on the veins on the lower surface, occasionally the lateral leaflets absent or much reduced; stipules deltoid or falcate, 2–9 mm long, basally subgibbose; peduncle 1–2.7 cm long, reddish orange; fruit bottle-shaped ...................................................................... C. haematantha Plants pubescent; terminal leaflet rhombic, subelliptic, or subovate, petiolate, scabrous or not on the lower surface; stipules not as above; peduncle > 2.7 cm long, greenish; fruit spherical .................................... 9 Leaflets membranous-fleshy, not scabrous, irregularly lobed; petiole puberulent, 6.5–15(–24) cm long; stipules deltoid, basally auriculate, 5–19(–27) mm long; fruit 15 mm diam, lenticellate or smooth ................................................................................................ C. gongylodes Leaflets chartaceous-coriaceous, scabrous on the upper surface, not lobed; petiole tomentose, 1–9(–12) cm long; stipules spatulate or falcate, basally gibbose, 1–2.5 mm long; fruit 6–9 mm diameter, smooth ......................................................................................................... C. alata

Cissus alata Jacq., Select. Stirp. Amer. Hist. 23. 1763. —Vitis alata (Jacq.) Kuntze, Revis. Gen. Pl. 3: 40. 1898. —Bejuco parcha morada, Mano de mato, Mojo– mera (Warao), Picatón. Cissus rhombifolia Vahl, Eclog. Amer. 2: 10. 1798. —Vitis rhombifolia (Vahl) Baker in Mart., Fl.Bras. 14: 207. 1871. Cissus pubescens Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) (quarto ed.) 5: 226. 1821 [1822]. Woody vine with angled, striate, densely pubescent stems, narrowly or inconspicuously winged; terminal leaflet rhombic, subacute to acuminate, cuneately narrowed to a sessile or shortly petiolate base, shortly serrulate on the margins, to 18 cm long and 8– 10 cm wide; lateral leaflets obliquely rhombic to ovate, to 15 cm long and 7 cm wide; inflorescence reddish or greenish white or greenish cream; peduncle 3–5 cm long; fruit purple, edible. Along streams, savannas, forested margins of savannas, open thickets and disturbed areas, 50–700 m; Delta Amacuro (Caño Curiapito, Sierra Imataca), Bolívar (El Manteco, east of La Paragua, south of Los Castillos, Río Asa, Río Orinoco near Ciudad Bolívar, Serranía La Cerbatana), Amazonas (Río Mawarinuma). Common in the llanos of

Guárico, Cojedes, Barinas, and Portuguesa and elsewhere in Venezuela north of the Orinoco; widespread from Mexico, Central America, West Indies, and South America. ◆Fig. 414. Cissus descoingsii J.A. Lombardi, Candollea 51: 370. 1996. Climbing plant with terete or 4-angled stems, puberulous with lenticels; leaf normally cordiform or triangular, ovate or oblong, 5–18 cm × 2–16 cm, apex acute or acuminate, base rounded to deeply cordate at base; petiole 1–14(–19) cm long. Lower montane moist forests, 300–700 m; Bolívar (summit of Altiplanicie de Nuria, 85 km south of El Dorado). Costa Rica, Panama, Colombia, Guyana, Ecuador, Brazil. ◆Fig. 416. Cissus erosa Rich., Actes Soc. Hist. Nat. Hist. Paris 1: 106. 1792. —Bejuco de judío, Bejuco de raya, Bejuco de sapo, Eroguata (Warao), Ma-eri (Yanomami), Mawaya-yeha (Yekwana), Paraguita, Takrum-yek (Pemón). Cissus lucida Poir., Encycl. suppl. 1: 106. 1810. Cissus quadrialata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 225. 1821 [1822]. Cissus salutaris Kunth in H.B.K., Nov.

Cissus 495

Gen. Sp. (quarto ed.) 5: 225. 1821 [1822]. —Cissus erosa var. salutaris (Kunth) Planch. in A. DC. & C. DC., Monogr. Phan. 5: 548. 1887. Cissus sexangularis Ernst, J. Bot. 8: 374. 1870. Cissus guaranitica Chodat & Hassl., Bull. Herb. Boissier sér. 2, 3: 545. 1903. Vine, often woody, the stems acutely 4angled, frequently winged, mainly glabrous or sparsely pilose, greenish or reddish violet; terminal leaflet elliptic or obovate, rounded or broadly obtuse at apex, cuneately narrowed at base, 4–16 cm long, sessile or petiolulate (0–2.6 cm long); lateral leaflets obliquely ovate-elliptic to narrowly obovate, the base cuneate or rounded, 3–12 cm long, sessile or with petiolules 0–7 mm long; inflorescence red or reddish orange; berries greenish black. Riparian forests, evergreen lowland forests, savannas, edges of Mauritia palm swamps, disturbed areas, near sea level to 800 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Common throughout Venezuela; southern Mexico, Central America, West Indies, and South America. ◆Fig. 415. This species, like Cissus descoingsii and C. verticillata, is often parasitized by a rust fungus (Mycosyrinx cissi). The fruits are edible, and a water extract of the stems is used as a rubber latex coagulant. All parts of the plant have diuretic and antirheumatism properties. Cissus gongylodes (Baker) Planch. in A. DC. & C. DC., Monogr. Phan. 5: 550. 1887. —Vitis gongylodes Baker in Mart., Fl. Bras. 14(2): 210. 1871. Vitis tricuspis Baker in Mart., Fl. Bras. 14(2): 210. 1871. —Cissus tricuspis (Baker) Planch. in A. DC. & C. DC., Monogr. Phan. 5: 551. 1887. Cissus cervii Dunaiski, Acta Biol. Paran. 21: 135. 1992. Liana, stem 4-angled, succulent, hirsuteglandulose, with scattered lenticels; terminal leaflet rhomboidal, apex acuminate, base cuneate and trilobed, lobes asymmetric, subsessile, margin serrate, 8–20 cm × 6-15 cm, sessile or petiolulate (0–5 cm long); lateral leaflets trapezoidal, base truncate, bilobed, 6–15 cm × 4–11 cm, sessile or petiolulate (0– 2.5 cm long); inflorescences greenish yellow; peduncle 2–5 cm long; fruit violet, ellipsoid. Along streams and disturbed vegetation, 100–200 m; Amazonas (Río Mawarinuma).

Colombia, Peru, Brazil, Bolivia, Paraguay, Argentina. Cissus haematantha Miq., Linnaea 26: 220. 1853. —Bejuco de fuente, Bejuco de sapo, Tagua-tagua. High-climbing woody vine or sometimes with creeping stems; petiole 0.4–14 cm long; terminal leaflet ovate or elliptic-lanceolate, 3–12 × 3–7 cm, sessile or petiolulate (0–01.7 cm long); lateral leaflets obliquely ovate, 0.7– 10 × 0.4–5 cm, sessile or petiolulate (0–1 cm long); inflorescence red or orange; fruit purple, lenticellate, 1–2 cm long. Flooded forests along streams, borders of seasonally dry evergreen forests, along trails, 50–300 m; Delta Amacuro (Río Amacuro, Río Yakarita, Sierra Imataca), Bolívar (Bochinche, 34–38 km south of El Dorado in drainage of Río Cuyuni, Río Asa, Río Botanamo, Río Paragua). Monagas; Trinidad, Suriname, French Guyana, Brazil. Cissus palmata Poir., Encycl. suppl. 1: 107. 1810. —Vitis palmata (Poir.) Baker in Mart., Fl. Bras. 14: 216. 1871. Cissus bonariensis Hook, & Arnott, Bot. Misc. 3(8): 159. 1833. Vitis gibertii Baker in Mart., Fl. Bras. 14: 211. 1871. —Cissus gibertii (Baker) Planch. in A. DC. & C. DC., Monogr. Phan. 5: 543. 1887. Cissus palmata var. balansaeana Planch. in A. DC. & C. DC., Monogr. Phan. 5: 553. 1887. Cissus paraguayensis Planch. in A. DC. & C. DC., Monogr. Phan. 5: 554. 1887. Vitis bakeri Herter, Revista Sudamer. Bot. 3(4–6): 168. 1936. Liana with long adventitious roots; stem cylindridal or 4-angled, striate, glabrous or sparsely pubescent; petiole 1–7 cm long; leaflets with apex obtuse to rounded, mucronate, base narrow-cuneate, subcoriaceous or papyraceous; inflorescence red or greenish orange; fruit purple, spherical, smooth, 5–10 mm long. Seasonally flooded forests, low thickets along streams, along sand banks, 50–100 m; Delta Amacuro (Caño Araguao). Apure; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Paraguay, Argentina, Uruguay. ◆Fig. 413. Cissus spinosa Cambess. in A. St.-Hil., Fl. Bras. Merid. 1: 345. 1825 [1828]. —Vitis spinosa Baker in Mart., Fl. Bras. 14: 209. 1871.

496

V ITACEAE

Cissus parkeri Planch. in A. DC. & C. DC., Monogr. Phan. 5: 550. 1870. Cissus hassleriana Chodat, Bull. Herb. Boissier 7, app. 1: 73. 1899. Pubescent to velutinous liana; stem generally with turbinate or recurved prickles; petiole 1–11 cm long; terminal leaflet rhombic, apex acute, base cuneate, to 15 × 8 cm, petiolule 1–20 mm long; lateral leaflets suborbicular or obovate, base oblique, subequal to the terminal leaflet, petiolules 1–13 mm long; inflorescences red or yellowish red; fruit bottle-shaped, smooth. Seasonally flooded riparian forests, near sea level to 50 m; Delta Amacuro (Caño Corisal, Varadero de El Limón). Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Cissus trianae Planch. in DC., Monogr., Phan. 5: 555. 1887. Cissus martiniana Woods. & Seib., Ann. Missouri Bot. Gard. 24: 191. 1937. —Cissus trianae var. martiniana (Woods. & Seib.) Steyerm., Fieldiana, Bot. 28: 356. 1952. Liana, occasionally hemiepiphytic; petiole 1–3 cm long; leaflets obovate or elliptic, apex acute or obtuse, base cuneate, 2–6 × 1–2.5 (–3.5) cm, margin denticulate, dentate, or crenulate; inflorescence greenish yellow; fruit green to white. Montane wet forests, 800–1800 m; Amazonas (Cerro Huachamacari, Sierra de la Neblina). Anzoátegui, Aragua, Mérida, Monagas, Táchira; Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. Cissus venezuelensis Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 427, fig. 4. 1966. Liana, stem cylindrical and sparsely puberulent; leaves ovate-oblong or elliptic-oblong, caudate or mucronate, 9–14 × 4.5–7.5 cm, glabrous; petiole 1–2.5 cm long; inflorescence falsely terminal, greenish yellow or whitish cream; peduncle 3–4 cm long; fruit green, ellipsoid, 9–10 mm long. Wet montane forests, 700–1100 m; Bolívar (affluent of Río Cuyuní, basin of Río Venamo). Guyana (Mount Ayanganna, Mazaruni River). ◆Fig. 417. Cissus verticillata (L.) Nicolson & C.E. Jarvis, Taxon 33: 727. 1984. —Viscum verticillatum L., Sp. Pl. 1023. 1753.

—Phoradendron verticillatum (L.) Druce, Bot. Exch. Club Soc. Brit. Isles 3: 422. 1914. Liana, commonly with adventitious roots. Southern U.S.A., Mexico, West Indies, Central America, South America; 4 subspecies, 2 in Venezuela, 1 of these in the flora area. C. verticillata subsp. verticillata —Bejuco de caro, Bejuco de mono, Ero-Kuaja (Warao), Juichamo (Yekwana), Ma-eri hasubi (Yanomamo), Manman (Panare), Picatón, Uva, Zarza. Cissus sicyoides L., Syst. Nat. ed. 10, 2: 897. 1759. —Vitis sicyoides (L.) Morales in Poey, Repert. Fis.-Nat. Isla Cuba 1: 206. 1865, “sycioides.” Cissus cordifolia L., Sp. Pl. ed. 2: 170. 1763. Cissus ovata Lam., Tabl. Encycl. 1: 331. 1791 [1792]. Cissus smilacina Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 224. 1821 [1822]. —Vitis sicyoides var. smilacina (Kunth) Baker. in Mart., Fl. Bras. 14: 203. 1871. Cissus tamoides Cambess. in A. St.-Hil., Fl. Bras. Merid. 1: 342. 1828. Vitis sicyoides f. laciniata Baker in Mart., Fl. Bras. 14(2): 2023. 1871. —Cissus verticillata subsp. laciniata (Baker) J.A. Lombardi, Taxon 44: 204. 1995. Sprawling or climbing plant to 20 m tall, the stems terete, striate, glabrous to tomentose or villous; leaves very variable in shape, ovate, elliptic, oblong, to subrotund, 2.5–18 × 1.7–13 cm, acute to shortly acuminate at apex, base cordate, rounded or truncate, margin finely dentate or teeth acute, petiole 1–8.5 cm long; inflorescence usually greenish white, white, or yellowish, rarely reddish; fruit green turning red to black. Deciduous riparian forests, evergreen lowland forests, along streams, on rocks along rapids, disturbed places near human habitation, near sea level to 900 m; Delta Amacuro (common along caños and larger streams), Bolívar (Río Botanamo, Río Caura, Río Cuyuní, Río Grande, Río Orinoco near Ciudad Bolívar), Amazonas (Río Casiquiare, Río Matacuni, Río Ocamo, San Carlos de Río Negro, Sierra Parima). Common in Venezuela north of Río Orinoco; widespread from southern U.S.A., Mexico, West Indies, Central America, and South America.

Cissus 497

Fig. 413. Cissus palmata

Fig. 414. Cissus alata

498

V ITACEAE

Fig. 415. Cissus erosa

Fig. 416. Cissus descoingsii

Cissus 499

Fig. 417. Cissus venezuelensis

500

V OCHYSIACEAE

VOCHYSIACEAE by Luis Marcano-Berti Trees or shrubs, rarely suffruticose, unarmed; indumentum generally consisting of simple, malpighioid, or stellate hairs. Leaves simple, opposite or in whorls, rarely alternate or subalternate, generally petiolate; stipules normally developed, small or represented by glands; blades epunctate, with margins entire or subundulate, penniveined; main lateral veins on either side of the midvein 6–30 or very numerous (to 50 per cm); submarginal vein absent or present. Inflorescence consisting of a panicle of cincinni, or rarely of cymes and sometimes axillary cincinni. Flowers zygomorphic, perigynous or epigynous. Calyx imbricate with 5 subequal or very unequal lobes or sepals inserted on a very short calyx tube or hypanthium, the posterior sepal always spurred, generally convolute, the spur exserted or hidden in bud; corolla of 1 open or convolute petal, or of 3 or 5 imbricate, free petals, sometimes absent. Stamen 1, either a pseudolaminar stamen where the connective is indistinguishable from the filament, the thecae introrse, glabrous or pubescent, opening lengthwise on the adaxial side of the pseudolamina, or a more typical stamen, where the connective is easily distinguishable from the filament, with anther dorsifixed, bithecate, opening lengthwise; staminodes 0–4. Ovary superior or inferior, 1- or 3-locular, with 1–many ovules; style 1, simple; stigma 1. Fruit 3-locular and dehiscent or 1-locular, indehiscent, and generally winged. Seeds often winged. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, 2 genera in western Africa; 7 genera and ca. 222 species, 4 genera and 65 species in the flora area. Key to the Genera of Vochysiaceae 1. 1. 2(1). 2.

3(2). 3.

Branchlets or leaves with stellate hairs; ovary inferior or subinferior; fruit indehiscent, generally winged, 1-locular ........................... 1. Erisma Branchlets or leaves with simple or malpighioid hairs; ovary superior; fruit dehiscent, 3-locular; seeds winged ............................................... 2 Spur hidden in bud; stamen with a connective easily distinguishable from the filament, the anther dorsifixed; petal 1, convolute in bud ..... 2. Qualea Spur exserted in bud; stamen pseudolaminar, the connective indistinguishable from the filament; petals 0–3, convolute, imbricate, or open in bud ..................................................................................................... 3 Ovules 7 in each locule; petal 1, convolute in bud ................... 3. Ruizterania Ovules 2 in each locule; petals 0–3, imbricate or open in bud ..... 4. Vochysia

1. ERISMA Rudge, Pl. Guian. 1: 7. 1805. Debraea Roem. & Schult., Syst. Veg. 1: 4, 34. 1817. Ditmaria Spreng., Anleit. Kenntn. Gew. ed. 2, 2: 704. 1818, non Lühnem. 1809. Trees; branchlets with stellate pubescence. Leaves opposite or 3- or 4-verticillate, glabrous or stellate-pubescent; stipules absent or small; main lateral veins 6– 18 per side. Inflorescence generally a panicle or panicle of cincinni. Flowers epigynous or subepigynous, bracteate, generally bracteolate. Calyx imbricate, 5-merous, the posterior sepal or lobe deciduous, ≤ 2 times as long as the others, convolute in bud, enveloping the inner flower parts; the 4 smaller ones unequal, persistent and

Erisma 501

accrescent in fruit; spur exserted in bud; petal 1, white, yellow, or violaceous, convolute in bud, enveloping the inner flower parts, generally obcordate, 2-lobed at the apex, clawed at the base, rudimentary petals glabrous or pubescent when present. Stamen typical, the connective easily distinguishable from the filament, the anther dorsifixed; staminodes 0–4, glabrous or pubescent. Ovary inferior or nearly so, pubescent at the top, 1-locular with 2 ovules; style 1, terminal; stigma terminal or lateral-terminal. Fruit indehiscent, 1-locular, generally winged, 1-seeded. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; ca. 19 species, 8 in Venezuela, all in the flora area. Key to the Species of Erisma 1.

1.

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

3.

4(3).

4.

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

Bracts persistent, overlapping the subtended flower buds, 7.5–14 × 5.7– 10 mm, ovate to subovate; bracteoles persistent, stalked, lanceolate to oblanceolate, 2.6–4.5 × 0.5–1.8 mm; petals yellowish; fruit winged ... 2 Bracts deciduous or subpersistent, not overlapping the subtended flower buds, 2.8–11 × 2–4.3 mm oblong-eliptic to ovate; bracteoles absent or deciduous, not stalked, linear to subulate, 0.4–1.7 × 0.1–0.3; petals white, yellowish, yellow, lilac, or blue to violet; fruit winged or not ................................................................................................................ 3 Spur erect in bud, incurvate, angle 0°-10° with the spurred sepal, at anthesis pendulous, slightly incurvate to incurvate ............. E. floribundum Spur patent to subpatent in bud, straight to slightly recurved, angle 80°–100°, at anthesis pendulous, straight to recurvate ...... E. laurifolium Fruit not winged, ca. 7 × 3 cm, rugose; bracteoles present; spurred sepal (including the calyx tube or hypanthium) 10–14 mm long at anthesis; spur straight or slightly recurved, pendent, 7–8.5 mm long; petal blue to violet; anther 3.3–3.7 mm long ........................................ E. calcaratum Fruit winged, to 15 cm long; bracteoles absent or present; spurred sepal (including the hypanthium or calyx tube) 3.5–14 mm long at anthesis; spur straight to uncinate-recurved, 1.2–6 mm long; petal white, yellowish, yellow, lilac, or blue to violet; anther 1.2–3.5 mm long ........... 4 Bracteoles absent; bracts 9.1–11 × 4–6.5 mm, subpersistent; spurred sepal (including the hypanthium or calyx tube) 1.2–1.4 cm long at anthesis; spur somewhat bag-shaped, 3.5 × 4.3 mm at anthesis; petal yellow; anther 3.5 mm long; leaves 3- or 4-verticillate; petioles 2.4–4 cm long ....................................................................................................... E. japura Bracteoles present, deciduous; bracts ca. 3 × 3 mm, deciduous; spurred sepal (including the hypanthium or calyx tube) 3.5–11 mm long at anthesis; spur straight to uncinate-recurved, 1.2–6 mm long; petal white, yellowish, yellow, lilac, or blue to violet; anther 1.2–3.1 mm long; leaves opposite or verticillate; petioles to 2 cm long ............................ 5 Spur uncinate-recurved, 3.2–4 × 2.5–4 mm; petal blue to violet; anther 1.2–1.5 mm long; leaves opposite or 3- or 4-verticillate ....... E. uncinatum Spur straight, pendent, 1.2–2.5 × 1.2–1.8 mm; petal yellowish, yellow, or white to pale lilac; anther 1.2–3.1 mm long; leaves opposite .............. 6 Spurred sepal (including the calyx tube or hypanthium) 4–4.5 mm long at anthesis; spur 1.2–1.7 × 1.2–1.6 mm; petal yellow, densely pubescent; anther ca. 1.2 mm long ....................................................... E. micranthum

502

6.

7(6).

7.

V OCHYSIACEAE

Spurred sepal (including the calyx tube or hypanthium) 6–11 mm long at anthesis; spur 2–2.5 × 1.6–1.8 mm; petal yellowish or white to pale lilac; anther 1.8–3.1 mm long .................................................................. 7 Petiole 10–20 mm long; base of leaf blade cuneate; spurred sepal (including the calyx tube or hypanthium) 6–6.5 mm long at anthesis; anther 1.6–1.8 mm long; petal white to pale lilac .............................. E. splendens Petiole ca. 4.5 mm long; base of leaf blade cordate; spurred sepal (including the calyx tube or hypanthium) ca. 11 mm long at anthesis; anther ca. 3.1 mm long; petal yellowish ................................................ E. blancoa

Erisma blancoa Marc.-Berti, Pittieria 13: 12. 1986. —Yuri. Tree to 22 m tall; trunk to 80 cm diameter; petal yellowish, ca. 11.5 mm long. Riparian forests, 100–200 m; Amazonas (Río Cuao, Río Sipapo). Panama, Brazil. ◆Fig. 420. Erisma calcaratum (Link) Warm. in Mart., Fl. Bras. 13(2): 111. 1875. —Qualea calcarata Link. in Spreng. et al., Jahrb. Gewächsk. 1(3): 24. 1820. —Chimaco. Erisma violaceum Mart., Nov. Gen. Sp. Pl. 1: 137, pl. 82. 1824 [1826]. —Debraea violacea (Mart.) Steud., Nomencl. Bot. ed. 2: 486. 1841. —Ditmaria violacea (Mart.) Spreng., Syst. Veg 4(cur. post.): 8. 1827. Tree to 30 m tall; petal ca. 18 × 18 mm, mauve to blue; fruit not winged, water-dispersed. Seasonally flooded riparian forests, 100–200 m; Amazonas (Caño Yagua, Río Autana, Río Casiquiare, Río Sipapo, between Solano and San Carlos de Río Negro). Guyana, Peru, Brazil. ◆Fig. 418.

Fig. 418. Erisma calcaratum

Erisma floribundum Rudge, Pl. Guian. 7– 8, t. 1. 1805. —Debraea floribundum (Rudge) Roem. & Schult., Syst. Veg. 1: 34. 1817. —Ditmaria floribunda (Rudge) Spreng., Syst. Veg. 1: 16. 1825 [1824]. Erisma parvifolium Gleason, Bull. Torrey Bot. Club 60: 362. 1933. Erisma pallidiflorum Ducke, Arq. Inst. Biol. Veg. 2: 54. 1935. —Erisma parvifolium var. pallidiflorum (Ducke) Ducke, Arq. Inst. Biol. Veg. 4: 43. 1938. Venezuela, French Guiana, Brazil; 2 varieties, 1 in Venezuela. Variety tomentosum is known only from Brazil. E.

floribundum var. floribundum. —Jabua. Tree to 30 m tall; petal mainly yellowish, ca. 15 × 15 mm. Evergreen lowland forests, ca. 100 m; Amazonas (Yavita). French Guiana, Brazil (Amazonas, Matto Grosso). Erisma japura Spruce ex Warm. in Mart., Fl. Bras. 13(2): 109, t. 21, fig. 1. 1875. —Jabua. Tree to 35 m tall; petal 2–2.5 × 2.5–3.5 cm, yellow. Nonflooded evergreen lowland for-

Erisma 503

Fig. 419. Erisma uncinatum

Fig. 420. Erisma blancoa

ests, 50–200 m; Amazonas (Río Guainía, 2 km southeast of San Carlos de Río Negro, between San Carlos de Río Negro and Solano, between Yavita and Maroa). Southeastern Colombia, northern Brazil. In years when abundant seeds were produced, local inhabitants around Maroa and Yavita used to gather the seeds to prepare a “queso de Jabua,” or Jabua cheese. This was done by leaving the seeds to soak in streams

for about a week. They were then pounded into a pulp, tightly wrapped and pressed in leaves, and then resoaked as before. This produces a somewhat foul-smelling, cheeselike product. Erisma laurifolium Spruce ex Warm. in Mart., Fl. Bras. 13(2): 109, t. 20, 1875. Tree to 18 m tall, trunk to 130 cm circumference; petal white with yellow. Evergreen

504

V OCHYSIACEAE

lowland forests, 50–200 m; Amazonas (sector El Pozo near San Fernando de Atabapo, 126 km southwest of Santa Bárbara del Orinoco). Brazil. Erisma micranthum Spruce ex Warm. in Mart., Fl. Bras. 13(2): 112. 1875. Tree to 25 m tall; petals mainly yellow, red near the base, ca. 4.2 × 5.2 mm. Riparian forests, forest patches, lower montane forests, 100–200 m; Amazonas (near San Carlos de Río Negro, above San Juan de Ucata). Brazil. Erisma splendens Stafleu, Acta Bot. Neerl. 3: 476, fig. 4, 1954. Tree to 20 m tall; petals yellow, 10–12 × 10–12 mm. Riparian forests, ca. 100 m; Amazonas (Caño Caname). Brazil. Erisma uncinatum Warm. in Mart., Fl. Bras. 13(2): 110. 1875. —Daujen, Mo-

reillo, Mureillo, Naiyek (Arekuna), Salado, Salado morado. Erisma pulverulentum Poepp. ex Warm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1889: 28. 1889, nom. nud. Tree to 40 m tall; trunk to 1.6 m diameter; petal violaceous, ca. 10 × 10 mm. Evergreen forests at 200–500 m; Delta Amacuro (Río Toro, Serranía de Imataca); Bolívar (widespread in north, near Urimán), Amazonas (Caño Yureba off lower Río Ventuari, Cerro Duida, Cerro Moriche, Galipero ca. 30 km north of Puerto Ayacucho, Guara in Río Manapiare valley, Pintado, Río Cunucunuma). Guyana, Suriname, French Guiana, Colombia, Peru, Brazil. ◆Fig. 419. Erisma uncinatum is a commercially very important lumber-producing tree in northeastern Bolívar state.

2. QUALEA Aubl., Hist. Pl. Guiane 5. 1775. Amphilochia Mart., Nov. Gen. Sp. Pl. 1: 127. 1824 [1826]. Agardhia Spreng., Syst. Veg. 1: 4. 1825 [1824]. Schuechia Endl., Gen. Pl. 1177. 1840. Trees or shrubs. Leaves generally opposite, sometimes in whorls, or rarely alternate, glabrous or with simple hairs; stipules generally represented by concave or subplane, elliptic to rounded, subprominent glands 0.3–2.5 mm wide and < 0.3 mm long to prominent, cylindrical or urceolate glands to 3 mm long, sometimes normally developed, then triangular, 0.5–2 mm long and often with axillary glands as described above; petiole usually present; main lateral veins about 8 on either half of the blade, or very numerous (to 30 per cm). Inflorescence generally of panicles of cincinni, or rarely panicles of regular cymes. Flowers perigynous. Calyx imbricate, 5-merous, the 4 smaller sepals (2 lateral and 2 anterior) ones unequal, the posterior one spurred dorsally at the base, generally ≤ 2 times longer than the others, convolute and enveloping the inner parts; spur appressed against the back of the spurred sepal and hidden in bud by the lateral sepals at least basally, exserted or hidden at anthesis; petal 1, white, yellowish, yellow, pink, or blue, deciduous, often obcordate, clawed, convolute, enveloping the inner flower parts; rudimentary petals, if present, 0.5–1 mm long. Stamen with the connective easily distinguishable from the filament; staminal filament glabrous or pubescent; anther dorsifixed, glabrous or pubescent; staminodes, if present, glabrous, 0.5–1 mm long. Ovary superior, pubescent, 3-locular, with axile placentation; ovules 4–24 per locule, inserted in 2 rows. Fruit a 3-locular capsule. Seeds unilaterally winged. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay; ca. 48 species, 17 in Venezuela, 16 of these in the flora area. Key to the Species of Qualea 1.

Spurred sepal (including the calyx tube or hypanthium) ≤ 12 mm long at anthesis .................................................................................................. 2

Qualea 505

1. 2(1).

2.

3(2).

3.

4(3). 4. 5(1). 5. 6(5). 6. 7(6). 7. 8(7). 8. 9(8). 9. 10(5). 10. 11(10). 11. 12(11). 12. 13(12). 13. 14(10). 14. 15(14).

Spurred sepal (including the hypanthium or calyx tube) > 14 mm long at anthesis .................................................................................................. 5 Main and minor lateral veins 23–32 per cm; main lateral veins 10–16 per cm; spurred sepal (including the calyx tube or hypanthium) 10–12 mm long at anthesis ............................................................................ Q. sprucei Main and minor lateral veins to 14 per cm; main lateral veins not > 7 per cm and 16 on each half of the leaf blade; spurred sepal (including the calyx tube or hypanthium) 5.5–9.8 mm long at anthesis .................... 3 Leaf blade 8–18 × 3.5–7.5 cm; main lateral veins ca. 16 on each 1/2 of the leaf blade, 1–3.5 cm apart; vegetative buds 4–4.5 mm long; spurred sepal (including the hypanthium or calyx tube) 8–9.8 mm long at anthesis ............................................................................................. Q. psidiifolia Leaf blade 3.2–8 × 2–4 cm; main lateral veins 5–7 per cm, 1–2.5(–3) mm apart; vegetative buds 1.2–2.5 mm long; spurred sepal (including the hypanthium or calyx tube) 5.5–7.5 mm long at anthesis .................... 4 Base of leaf blade obtuse to rounded; vegetative buds 1.8–2.5 mm long ....................................................................................................... Q. dinizii Base of leaf blade cordate; vegetative buds 1.2–1.5 mm long ...... Q. rupicola Spur exserted at anthesis .......................................................................... 6 Spur hidden at anthesis .......................................................................... 10 Anther 3–4.5 mm long ............................................................ Q. pulcherrima Anther 6–12 mm long ................................................................................ 7 Anther with the connective projecting beyond the thecae with an emarginate apicule ............................................................................ Q.paraensis Anther with the connective projecting beyond the thecae with an acute apicule .................................................................................................... 8 Anther pubescent ...................................................................... Q. decorticans Anther glabrous ......................................................................................... 9 Base of leaf blade rounded to cordate; midvein pubescent on both surfaces ................................................................................................ Q. acuminata Base of leaf blade acute to obtuse; midvein glabrous on both surfaces ...................................................................................... Q. schomburgkiana Anther pubescent on the back ................................................................. 11 Anther glabrous ....................................................................................... 14 Petal white; staminal filament pubescent ........................... Q. themistoclesii Petal mostly blue or pink; staminal filament pubescent or or glabrous .............................................................................................................. 12 Staminal filament glabrous; petal blue with a yellow central streak ...................................................................................................... Q. cyanea Staminal filament pubescent; petal pink ................................................ 13 Pedicel 10–13 mm long; stipular ridge present; anther 5.7–6.2 mm long .. ............................................................................................ Q. polychroma Pedicel 6.5–8 mm long; stipular ridge absent; anther 3.6–5 mm long .............................................................................................. Q. homosepala Petals mainly blue ........................................................................ Q. suprema Petals mainly white ................................................................................. 15 Younger and older branchlets glabrous; spurred sepal (including the hypanthium or calyx tube) 1.7–2 cm long at anthesis; anther 7–8.30 × ca.

506

15.

V OCHYSIACEAE

3.2 mm; connective produced beyond the thecae with an apicule of 3.2– 3.5 mm long ........................................................................... Q. tuberculata Younger and older branchlets pubescent; spurred sepal (including the hypanthium or calyx tube) ca. 2.5 cm long at anthesis; anther ca. 12 mm long; connective produced beyond the thecae with an apicule of ca. 2 mm long ................................................................................ Q. wurdackii

Qualea acuminata Spruce ex Warm. in Mart., Fl. Bras. 13(2): 40. 1875. —Salado de rebalse, Salado rebalsero. Qualea speciosa Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr. 3: 425. 1902. Tree to 20 m tall; petal 3.5–5 × 3.5–5 cm, white with yellow and red spots. Seasonally flooded riparian forests, montane forests, 100–2000 m; Bolívar (Churí-tepui, Río Sarvén on Macizo del Chimantá), Amazonas (Caño Yagua, Cerro Sipapo, near Puerto Ayacucho, Río Baría, Río Casiquiare, Río Cuao, Río Pasimoni, Río Sipapo, Río Yatúa, Santa Bárbara). Colombia, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 422. Qualea cyanea Ducke, Arq. Inst. Biol. Veg. 2: 54. 1935. —Chimaco. Tree to 36 m tall; petal blue with a yellow central streak, 3–3.5 × 3.5–4.2 cm. Riparian forests, 100–200 m; Amazonas (Río Jénita on Río Ocamo, Río Siapa). Brazil. Qualea decorticans Ducke, Arq. Inst. Biol. Veg. 4: 39. 1938. Tree; petal pink, 2.5–3 × 3–3.5 cm. Evergreen lowland forests, ca. 200 m; expected in Amazonas. Brazil (Amazonas: near Cocui on upper Rio Negro near Venezuelan border). Qualea dinizii Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 49, t. 17, 19E. 1915. —Congrio morado, Guarai blanco, Guarapo, Guarapo amarillo. Deciduous tree to 40 m tall; trunk to 1.5 m diameter; petal lilac to purple, 10–13 × 13–18 mm. Semideciduous to evergreen lowland and lower montane forests, 50–500 m; Delta Amacuro (east of Los Castillos, Piacoa toward Río San José, Río Toro, Serranía de Imataca), northern Bolívar (widespread), Amazonas (Caño Marieta, from Galipero south to Samariapo, mouth of Río Cuao). Monagas; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 424. Qualea homosepala Ducke, Arq. Inst. Biol. Veg. 4: 38. 1938.

Tree to 30 m tall; petal blue or mauve with yellow near the middle. Riparian forests, 100–300 m; Bolívar (Río Caura near Guiñuna). Colombia, Brazil. Qualea paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 48, t. 16. 1915. Tree to 30 m tall; petal mainly white with red at the base, 3.5–4 × 3.5–4 cm. Evergreen lowland to lower montane forests, 100–1000 m; Bolívar (Serranía de los Pijiguaos), Amazonas (Cerro Duida, Culebra, Río Sipapo, Tobogán de la Selva southeast of Puerto Ayacucho). Colombia, Guyana, Peru, Brazil. Qualea polychroma Stafleu, Acta Bot. Neerl. 2: 182, fig., 11a–c. 1953. —Tukuinanwuyek (Arekuna). Tree to 40 m tall; petal pink, 3.5–4.5 × 3.5–4.5 cm. Upland evergreen forests, 800– 900 m; Bolívar (La Escalera, between Santa Elena de Uairén and Ikabarú). Guyana. Qualea psidiifolia Spruce ex Warm. in Mart., Fl. Bras. 13(2): 46, t. 8, fig 1. 1875. Tree to 35 m tall; petal lavender or pink, ca. 2.5 × 2.5 cm. Riparian forests, 100–200 m; Amazonas (Río Casiquiare, Río Guainía, Río Pasimoni, San Carlos de Río Negro). Amazonian Colombia, Brazil (Amazonas: Rio Negro basin). Qualea pulcherrima Spruce ex Warm. in Mart., Fl. Bras. 13(2): 37. 1875. Tree to 25 m tall; petal rich rose or rose pink, basally yellow, ca. 3 × 4 cm. Riparian forests, 100–200 m; Amazonas (Canaripó, Río Casiquiare, Río Pasimoni). Amazonian Colombia, Brazil (Amazonas: Rio Negro basin). ◆Fig. 423. Qualea rupicola Ducke, Arq. Inst. Biol. Veg. 4: 41. 1938. Tree to 7 m tall; petal pink, ca. 1.5 × 1.8 cm. Riparian forests, 100–300 m; Amazonas (8 km below Raudal Gallineta on the Río Siapa). Brazil (Amazonas: Rio Negro basin).

Qualea 507

Qualea schomburgkiana Warm. in Mart., Fl. Bras. 13(2): 39. 1875. Shrub to tree ca. 15 m tall; petal ca. 4 × 6 cm, white with orange-red spots near the base. Upland gallery forests, 400–1400 m; Bolívar (widespread in Gran Sabana and bases of tepuis). Guyana, Brazil. ◆Fig. 421.

Qualea suprema Ducke, Arq. Inst. Biol. Veg. 2: 53. 1935. Small tree ca. 4 m tall; petals blue. Seasonally flooded riparian forests, 100–200 m; Amazonas (Caño San Miguel between Las Tinajas and Caño Iqueven). Brazil (Amazonas: Río Curicuriary, upper Río Negro).

Qualea sprucei Warm. in Mart., Fl. Bras. 13(2): 38. 1875. —Palo de guabina, Rabo de Maquire, Traira-miri. Tree to 22 m tall; petal pink with yellow streaks, ca. 3 × 3 cm. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro, Yavita). Brazil (Amazonas: Rio Negro basin).

Qualea themistoclesii Ducke, Arq. Inst. Biol. Veg. 4: 38. 1938. —Congrio blanco, Traira-mira. Tree to 20 m tall; petal mainly white, ca. 3.5 × 4 cm. Evergreen lowland forests, riparian forests, 50–200 m; Amazonas (Río Baría, San Carlos de Río Negro). Brazil (Amazonas: Rio Negro basin). Qualea tuberculata Stafleu, Acta Bot. Neerl. 2: 189, fig 13a–d. 1953. Tree to 30 m tall; petal white, ca. 5 × 5 cm. Evergreen lowland forests, 100–200 m; Amazonas (between Maroa and Victorino along Río Guainía). Endemic.

Fig. 421. Qualea schomburgkiana

Fig. 422. Qualea acuminata

508

V OCHYSIACEAE

Fig. 423. Qualea pulcherrima

Fig. 424. Qualea dinizii

Qualea wurdackii Marc.-Berti, Pittieria 13: 7. 1986. Shrub or tree 2.5–10 m tall; petal white,

basally yellow. Along streams in savannas, ca. 100 m; Amazonas (Caño Cotúa, Cerro Moriche, Cerro Yapacana, Río Guayapo). Endemic.

3. RUIZTERANIA Marc.-Berti, Pittieria 2: 6. 1969. Qualea ser. I callophylloidea Warm.in Mart., Fl. Bras. 13(2): 30. 1875, pro parte. Qualea sect. Trichanthera Stafleu, Acta Bot. Neerl 2: 153. 1953. Trees or shrubs. Leaves opposite, rarely subopposite, glabrous or only pubescent with simple hairs on lower surface; stipules represented by crateriform,

Ruizterania 509

subprominent glands, < 0.3 mm long and 0.3–2.5 mm wide, rounded to elliptic; midvein narrowly winged on lower surface; lateral veins to 50 per cm, at 70–90° angle with the midvein; submarginal vein present. Inflorescences axillary 1–3-flowered cincinni and/or terminal or axillary panicles. Flowers perigynous. Calyx imbricate, 5-merous, the 4 smaller ones subequal, the posterior one spurred dorsally at the base, 3–4 times longer than the others, convolute and enveloping the inner flower parts; spur exserted in the bud; petal 1, convolute, enveloping the inner flower parts; rudimentary petals sometimes present, glabrous, 1–6 mm long. Stamen pseudolaminar, the connective indistinguishable from the filament, thecae with dense, simple, elongated, erect hairs arranged like the bristles of a brush, opening lengthwise, introrse, on the adaxial side of the pseudolamina, the connective not projected beyond the thecae into a hooded apex; filament glabrous; staminodes often present, glabrous, 0.5–2 × 0.05–0.15 mm. Ovary superior, pubescent, 3-locular; ovules 7 per locule, inserted in 2 rows; style terminal. Capsule 3locular, 3-valved. Seeds winged. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; ca. 13 species, 7 in Venezuela, all in the flora area. Ruizterania nitida (Stafleu) Marc.-Berti is not included here. It was described from the slopes of Cerro Duida, Amazonas, based on sterile material and could be a branchlet of a young plant of a Qualea species or a previously described Ruizterania species. Key to the Species of Ruizterania 1. 1. 2(1). 2. 3(2). 3. 4(1).

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

6.

Ovary white to grayish-pubescent ............................................................ 2 Ovary ferruginous-pubescent .................................................................... 4 Flowers arranged in cincinni, generally 2-flowered in the axils of the leaves ............................................................................................. R. retusa Flowers arranged in panicles of cincinni .................................................. 3 Apex of the leaf blade acute or obtuse to rounded and often retuse or emarginate ................................................................................ R. obtusata Apex of the leaf blade acute to acuminate ............................. R. trichanthera Flower buds nearly 7.5 mm wide above the smaller sepals; anther with a glabrous theca and the other densely covered with erect to suberect and straight hairs .......................................................... R. cassiquiarensis Flower buds < 5.6 mm wide above the smaller sepals; both thecae covered with hairs ............................................................................................... 5 Spurred sepal (including the hypanthium or calyx tube) at anthesis < 2 cm long ................................................................................................ 6 Spurred sepal (including the hypanthium or calyx tube) at anthesis > 2 cm ........................................................................................................ 7 Spurred sepal (including the hypanthium or calyx tube) 1.2–1.8(–2) cm long at anthesis; lower surface of the leaf blade glabrous to densely ferruginous-pubescent; base of leaf blade obtuse to subrounded, sometimes rounded; anther at anthesis (2.7–)5–9(–10) mm long ................................................................................................. R. ferruginea Spurred sepal (including the hypanthium or calyx tube) 2–2.6 cm long at anthesis; lower surface of the leaf blade glabrous to glabrescent; base of leaf blade rounded to cordate on the same branch; anther at anthesis (9–)10–12.5 mm long .....................................................................R. rigida

510

7(5).

7.

V OCHYSIACEAE

Spurred sepal (including the calyx tube or hypanthium) 2.5–3 cm long at anthesis; anther at anthesis 1.3–1.9 cm long; several minor lateral veins seemingly not arising from the midvein .................... R. esmeraldae Spurred sepal (including the hypanthium or calyx tube) 2–2.6 cm long at anthesis; anther at anthesis 0.9–1.25 cm long; minor lateral veins arising from the midvein ...............................................................R. rigida

Ruizterania cassiquiarensis (Spruce ex Warm.) Marc.-Berti, Pittieria 2: 10. 1969. —Qualea cassiquiarensis Spruce ex Warm. in Mart., Fl. Bras. 13(2): 34. 1875. Tree to 25 m tall, petal white with yellow from middle to base. Evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, San Carlos de Río Negro). Brazil. Ruizterania esmeraldae (Standl.) Marc.Berti, Pittieria 2: 11. 1969. —Qualea esmeraldae Standl., Bull. Torrey Bot. Club 2: 164, fig. 8. 1953. Qualea verruculosa Stafleu, Acta Bot. Neerl. 2: 164, fig. 8. 1953. —Ruizterania verruculosa (Stafleu) Marc.-Berti, Pittieria 2: 16. 1969. Shrub or tree 2–7 m tall, petal white with yellow to deep yellow streaks. Savannas, forests on banks of rivers and streams, Río Negro caatinga, 100–300 m; widespread in central and western Amazonas. Colombia, Brazil. ◆Fig. 425. Ruizterania ferruginea (Steyerm.) Marc.Berti, Pitteria 2: 11. 1969. —Qualea ferruginea Steyerm., Fieldiana, Bot. 28: 295. 1952. Qualea rubiginosa Stafleu, Acta Bot. Neerl. 2: 154, fig. 4. 1953. —Ruizterania rubiginosa (Stafleu) Marc.-Berti, Pittieria 2: 14. 1969. Qualea rubiginosa var. angustior Steyerm., Acta Bot. Venez. 2(5–8): 239. 1967. —Ruizterania rubiginosa var. angustior (Steyerm.) Marc.-Berti, Pittieria 2: 15. 1969. Qualea apodocarpa Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 473. 1966. —Ruizterania apodocarpa (Steyerm.) Marc.Berti, Pittieria 2: 10. 1969. Shrub or tree ca. 30 m tall, petal white with yellow to reddish from middle to base. Mostly riparian habitats in montane forests,

400–1700 m; widespread in Bolívar and Amazonas. Guyana, Brazil. ◆Fig. 426. Ruizterania obtusata (Briq.) Marc.-Berti, Pittieria 2: 13. 1969. —Qualea obtusata Briq., Annuaire Conserv. Jard. Bot. Genéve 20: 383. 1919. Qualea retusa var. coriacea Ducke, Arq. Inst. Biol. Veg. 4: 37. 1938. Venezuela, Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of R. obtusata 1. Lateral veins 38–44 per cm ....................... ............................. subsp. multivenosa 1. Lateral veins 16–24 per cm ....................... ................................... subsp. obtusata R. obtusata subsp. multivenosa Marc.Berti, Pittieria 18: 6. 1989. —Palo piapoco, Seje piapoco. Tree 12–30 m tall, petal pink. White-sand savannas, ca. 100 m; Amazonas (near Pimichín, between Solano and San Carlos de Río Negro). Endemic. R. obtusata subsp. obtusata Tree 4–25 m tall, petal white with yellowish base. Riparian forests, flooded black-water swamps, 100–200 m; western Amazonas. Brazil (Amazonas: Rio Negro basin). Ruizterania retusa (Spruce ex Warm.) Marc.-Berti, Pittieria 2: 13. 1969. —Qualea retusa Spruce ex Warm. in Mart., Fl. Bras. 13(2): 34, t. 4, fig. 1. 1875. Shrub or tree to 30 m tall, petal white with yellow to orange spotted toward the base. Savannas, riparian forests, 50–200 m; widespread in western Amazonas. Brazil, Bolivia. ◆Fig. 427. Ruizterania rigida (Stafleu) Marc.-Berti, Pittieria 2: 14. 1969. —Qualea rigida

Ruizterania 511

Fig. 425. Ruizterania esmeraldae

Fig. 426. Ruizterania ferruginea

Fig. 427. Ruizterania retusa

512

V OCHYSIACEAE

Stafleu, Acta Bot. Neerl. 2: 162, fig 7. 1953. Shrub or tree to 13 m tall. Upland riparian forests, 400–900 m; Bolívar (Gran Sabana). Guyana. Ruizterania trichanthera (Spruce ex Warm.) Marc.-Berti, Pittieria 2: 15.

1969. —Qualea trichanthera Spruce ex Warm. in Mart., Fl. Bras. 13(2): 35. 1875. Tree to 36 m tall; petals white streaked with red. Lower montane evergreen forests, Río Negro caatinga, 100–300 m; Amazonas (Maroa to Yavita, Río Putaco 19 km above its mouth in Río Ocamo). Brazil.

4. VOCHYSIA Aubl., Hist Pl. Guiane 18. 1775; corr. Poir. in Lam., Encycl. 8: 681. 1808. Vochy Aubl., Hist. Pl. Guiane 18. 1775. Salmonia Scop., Intr. Hist. Nat. 209. 1777. Vochya Vell. ex Vand., Fl. Lusit. Bras. Spec. 1. 1788. Cucullaria Schreb., Gen. Pl. 6. 1789. Vochisia A.L. Juss., Gen. Pl. 424. 1789. Strukeria Vell., Fl. Flumin. 8. 1825 [1829]. Trees, shrubs, or rarely subshrubs. Leaves opposite, whorled, sometimes scattered, glabrous or pubescent with simple or malpighioid hairs; stipules small, generally triangular. Flowers yellow, perigynous, in panicles of 1–10-flowered cincinni. Calyx 5-merous, imbricate, the posterior sepal or lobe spurred, 3–4 times longer than the others, convolute in bud, enveloping the inner flower parts. Petals generally 3, imbricate, sometimes 1 open in bud or absent, very rarely 2. Stamen pseudolaminar, the connective indistinguishable from the filament, the thecae introrse, glabrous or pubescent, opening lengthwise on the adaxial side of the pseudolamina, the connective projected beyond the thecae into a hooded apex; staminodes 2, opposite the lateral petals, small, glabrous or pubescent. Ovary superior, 3-locular, with 2 ovules per locule; style 1, terminal; stigma 1, terminal to lateral. Fruit 3-locular, dehiscent. Seeds winged. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 132 species, 39 in Venezuela, 34 of these in the flora area. Key to the Species of Vochysia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4).

Ovary pubescent ........................................................................................ 2 Ovary glabrous ........................................................................................... 8 Corolla present ........................................................................................... 3 Corolla absent ............................................................................................ 6 Petals ciliate at the apex; central petal 1/3 the length of the stamen in the bud; cincinni 1-flowered ............................................................. V. expansa Petals not ciliate at the apex; central petal 1/2–3/4 the length of the stamen in bud; cincinni (1)2–4-flowered on the same inflorescence ................ 4 Spur straight, erect, at an angle of 30°–70° with the rest of the sepal ............................................................................................ V. surinamensis Spur slightly to strongly incurved, at an angle of 110°–150° with the rest of the sepal ............................................................................................. 5 Spurred sepal (including the hypanthium or calyx tube) 1–1.1 cm long at anthesis; cincinni 1.6–1.8 cm long; style glabrous; stigma (lateral part) 0.3 × 0.4–0.5 mm ......................................................................... V. obscura

Vochysia 513

5.

Spurred sepal (including the hypanthium or calyx tube) 1.4–1.6 cm long at anthesis; cincinni 3–3.5 cm long; style sparsely pubescent in the lower 1/3; stigma (lateral part) 1.2 × 0.8 mm ...................... V. venezuelana 6(2). Leaves 4–9-verticillate; lower surface of leaf blade sparsely pubescent; apex rounded; cincinni 6–7 cm long; smaller sepals (including the hypanthium or calyx tube) ca. 9.5 mm long, conspicuously unequal ....................................................................................................... V. jonkeri 6. Leaves opposite or 3-verticillate; lower surface of leaf blade densely pubescent; apex acuminate to cuspidate; cincinni 2.2–4 mm long; smaller sepals (including the hypanthium or calyx tube) 3.2–5 long, subequal ................................................................................................................ 7 7(6). Lower surface of leaf blade densely malpighioid, appressed-pubescent; cincinni ca. 4 cm long; spurred sepal (including the hypanthium or calyx tube) 2.5–2.9 cm long at anthesis; anther ca. 10 times longer than filament ....................................................................................... V. pinkusii 7. Lower surface of leaf blade pubescent on the veins with malpighioid hairs, each with an erect or suberect end much longer than the other, nearly obsolete end; cincinni 2.2–2.4 cm long; spurred sepal (including the hypanthium or calyx tube) 1.3–1.6 cm long at anthesis; anther 4– 5 times longer than filament ......................................... V. steyermarkiana 8(1). Corolla absent ............................................................................................ 9 8. Corolla present ......................................................................................... 13 9(8). Leaves opposite ........................................................................................ 10 9. Leaves 3–5-verticillate ............................................................................ 11 10(9). Leaf blades 3.5–6.2 cm; petiole ca. 1 cm long; inflorescence ≤ 6.5 cm long; spur straight to slightly recurved, 5.5–10 × 1.9–2 mm; staminodes glabrous .................................................................................... V. apopetala 10. Leaf blades 6.3–12 cm long; petiole 1.7–2 cm long, malpighioid-pubescent; inflorescence 20–53 cm long; spur subinflated, 4.7–7.5 × 2.5– 3.8 mm, generally recurved, touching the pedicel; staminodes ciliate .................................................................................................... V. punctata 11(9). Leaves (3)4- or 5-verticillate, blade 11–13.5 × 6.7–7.2 cm, main lateral veins 10–12 on each half of the blade, 1.3–1.5 cm apart; inflorescence ca. 30 cm long; the ensemble of spurred sepal and spur is C-shaped; anther 12–13 times longer than the filament .............................. V. tilletii 11. Leaves 3-verticillate, blade 6.5–12.5 × 2.7–5.4 cm, main lateral veins 12– 20 on each half of the blade, 4–8 mm apart; inflorescence 13–19 cm long; spur 6–10 mm long, recurved; the ensemble of spurred sepal and spur, at least in the flower bud, S-shaped; anther 3–4 times longer than the filament ................................................................................. 12 12(11). Petiole 1.2–1.4 cm long; blade 6.5–8.6 × 2.7–4.8 cm, glabrous on both sides, concolorous, main lateral veins 12–13 on each half of the blade, 6–8 mm apart ............................................................................ V. bautistae 12. Petiole 2–2.2 cm long; blade 10.5–12.5 × 4–5.4 cm, the upper surface glabrous, the lower surface densely ferruginous-pubescent, main lateral veins 17–20 on each half of the blade, 4–7.5 mm apart ............. V. ortegae 13(8). Stamen glabrous ...................................................................................... 14 13. Stamen pubescent on the concave (anterior) surface and/or on the convex (posterior) side ..................................................................................... 20

514

V OCHYSIACEAE

14(13). Lower surface of leaf blade densely appressed-pubescent; inflorescence ≤ 6 cm long; cincinni 1-flowered; spurred sepal (including the hypanthium or calyx tube) 1.4–1.7 cm long at anthesis; spur slightly recurved ...................................................................................... V. calophylla 14. Lower surface of leaf blade glabrous or sometimes glabrescent on the midvein; inflorescence 12–30 cm long; cincinni 1–10-flowered ......... 15 15(14). Stigma lateral, 2-branched at the base; the shorter branch 1.2–1.5 mm long, the longer branch 1.5–2.5 mm long ........................................... 16 15. Stigma terminal or lateral, to 1 mm long, not 2-branched at the base .............................................................................................................. 17 16(15). Anther as long as or slightly longer than the filament; inflorescence 18– 23 cm long; cincinni 3–10-flowered in the same inflorescence; spurred sepal (including the hypanthium or calyx tube) 1.2–1.4 cm long at anthesis; spur incurved .................................................................. V. grandis 16. Anther 2–2.5 times longer than the filament; inflorescence 12–16 cm long; cincinni (1)2- or 3-flowered; spurred sepal (including the hypanthium or calyx tube) 1.1–1.3 cm long at anthesis; spur straight to recurved .................................................................................. V. pauciflora 17(15). Flower bud completely pubescent outside; spur incurved-hooked, the spurred sepal (including the hypanthium or calyx tube) 1.5–1.7 cm long at anthesis ....................................................................... V. crassifolia 17. Flower bud glabrous to glabrescent; spur straight, S-shaped, or recurved, the spurred sepal (including the hypanthium or calyx tube) 1.2–3 cm long at anthesis .................................................................................... 18 18(17). Leaves opposite; spurred sepal (including the hypanthium or calyx tube) 1.2–1.4 cm long at anthesis ................................................... V. glaberrima 18. Leaves (3)4-verticillate; spurred sepal (including the hypanthium or calyx tube) 1.2–3 cm long at anthesis .................................................... 19 19(18). Petiole 1.5–5 cm long; apex of the leaf blade rounded to rounded-emarginate; main lateral veins ca. 30 on each side of the midvein; spurred sepal (including the hypanthium or calyx tube) 2.2–3 cm long at anthesis; spur straight to slightly S-shaped, at an angle of 50°–100° with the pedicel ........................................................................................ V. catingae 19. Petiole 0.3–0.9 cm long; apex of the leaf blade acuminate; main lateral veins 14–20 on each side of the midvein; spurred sepal (including the hypanthium or calyx tube) 1.2–1.5 cm long at anthesis; spur straight to slightly recurved at an angle of 0°–20° with the pedicel ..... V. tetraphylla 20(13). Spurred sepal (including the hypanthium or calyx tube) 6–10.8 mm long at anthesis ............................................................................................ 21 20. Spurred sepal (including the hypanthium or calyx tube) 13–20 mm long at anthesis ............................................................................................ 27 21(20). Spur incurved-hooked; lower surface of leaf blade densely pubescent along the venation with malpighioid hairs which have an erect to suberect end much longer than the other, sometimes inconspicuous end; main lateral veins ca. 12 on each half of the leaf blade ................................................................................................. V. ferruginea 21. Spur straight to moderately curved; lower surface of leaf blade glabrescent to densely pubescent with appressed-malpighioid hairs; main lateral veins 15–20 on each half of the leaf blade .................................. 22

Vochysia 515

22(21). Petals glabrous; leaf blade with 2 submarginal veins ................... V. elegans 22. Petals, at least the central one, pubescent on the back; leaf blade with 1 submarginal vein .............................................................................. 23 23(22). Central petal ca. 2/3 the length of the stamen ......................................... 24 23. Central petal as long as or longer than the stamen ............................... 25 24(23). Leaf blade 8–12 × 3–4.4 cm; inflorescence to 10 cm long .......... V. vismiifolia 24. Leaf blade 12–18 × 5–7.5 cm; inflorescence to 15 cm long .................... ......................................................................................... V. cassiquiarensis 25(23). Petiole 15–20 mm long; leaf blade 10–19 × 4.3–11 cm; inflorescence 30– 43 cm long; central petal hooded, covering the apex of the anther .................................................................................................. V. splendens 25. Petiole 3–10 mm long; leaf blade 4.5–12 × 2–3.6 cm; inflorescence 6– 25 cm long; central petal not hooded at apex ..................................... 26 26(25). Lower surface of old leaves glabrous to glabrescent; apex obtuse and retuse; main lateral veins 4–6 per cm .................................... V. angustifolia 26. Lower surface of old leaves densely pubescent; apex acute to acuminate; main lateral veins 9–13 on each half of the leaf blade; submarginal vein absent or present only near the apical 1/3 ...................... V. tomentosa 27(20). Stamen 6–8 times longer than the central petal .................................... 28 27. Stamen ≤ 2 times longer than the central petal ..................................... 29 28(27). Petiole 2.5–3 cm long; leaf blade 12–18 × 5.5–7.3 cm; main lateral veins ca. 40 on each half of the blade; spur 6.2–6.9 mm long, straight, slightly incurved, or S-shaped, appressed to the pedicel or nearly so, at least near the base, at an angle of 0–20° with the pedicel ........... V. complicata 28. Petiole 0.8–1.6 cm long; leaf blade 3.5–9 × 1.5–4 cm; main lateral veins about 13 on each half of the blade; spur 8.5–10 mm long straight, erect, at an angle of 130°–160° with the pedicel ................................. V. liscanoi 29(27). Leaves 4- or 5-verticillate; spur incurved ............................................... 30 29. Leaves opposite; spur straight, slightly incurved, or recurved .............. 31 30(29). Main lateral veins 20–25 on each half of the leave blade; spur incurvedhooked ................................................................................. V. spathiphylla 30. Main lateral veins 10–12 on each half of the blade; spur recurved .............................................................................................. V. aff. laxiflora 31(29). Main lateral veins 7–10 per cm, at an angle of 70°–80° with the midvein; spur straight or slightly recurved, pendent at an angle of 160°–180° with the spurred sepal; anther glabrous on the concave (anterior) side, slightly longer than the filament .......................................... V. rubiginosa 31. Main lateral veins 1–3 per cm, at an angle of 45°–60° with the midvein; spur straight, slightly S-shaped, recurved, or incurved; anther pubescent on the concave side and > 2 times longer than the filament ..... 32 32(31). Spur recurved, the ensemble of spur and spurred sepal S-shaped at or near anthesis ....................................................................................... 33 32. Spur straight, slightly S-shaped, or incurved, never recurved, the ensemble of spur and spurred sepal U-shaped to C-shaped at or near anthesis .................................................................................................... 34 33(32). Central petal hooded, longer than the stamen, at least at or near anthesis; inflorescence ≤ 20 cm long .............................................. V. julianensis 33. Central petal not hooded, as long as the stamen, inflorescence 30–60 cm long ............................................................................................... V. saccata

516

V OCHYSIACEAE

34(32). Stipules ca. 0.8 mm long; central petal 1/3 the length of the stamen ................................................................................................. V. vismiifolia 34. Stipules 2–4.5 mm long, central petal as long as the stamen ............... 35 35(34). Petiole 10–15 mm long; leaf blade 4–6 cm wide; staminal filament and staminodes glabrous ..................................................................... V. costata 35. Petiole 3–8 mm long; leaf blade 2–3.6 cm wide; staminal filament pubescent at least on the apex ventrally; staminodes pubescent ..... V. tomentosa Vochysia angustifolia Ducke, Bull. Mus. Hist. Nat. (Paris) sér 2, 4: 738. 1932. —Palo de conoto. Vochysia javitensis Stafleu, Recueil Trav. Bot. Néerl. 41: 509. 1948. Tree 3–20 m tall. Riparian forests, 100– 200 m; Amazonas (base of Cerro Yapacana, between Pimichín and Yavita, Río Temi). Brazil (Amazonas: Rio Negro basin). Vochysia apopetala Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 311. 1915. Tree 4–20 m tall. Riparian forests, 1100– 2000 m; Bolívar (Ilú-tepui, Macizo de Chimantá, Roraima-tepui). Endemic. ◆Fig. 428. Vochysia bautistae Marc.-Berti, Pittieria 10: 12. 1982. Tree to 15 m tall. Tepui slope forests, ca. 500 m; Amazonas (Cerro Sipapo). Endemic. Vochysia calophylla Spruce ex Warm. in Mart., Fl. Bras. 13(2): 98, t. 18, fig 1. 1875. Tree 3–15 m tall. Seasonally flooded forests along black-water rivers, 50–200 m; western Amazonas. Brazil (Amazonas: Rio Negro basin). Vochysia cassiquiarensis Stafleu, Acta Bot. Neerl. 3: 405, fig. 1a–b. 1954. Tree. Riparian forests, 100–200 m; Amazonas (San José de Casiquiare). Endemic. Vochysia catingae Ducke, Arq. Inst. Biol. Veg. 4: 33. 1938. Shrub or tree 3–10 m tall. Seasonally flooded margins of black-water rivers, scrub savannas on granitic outcrops, 50–200 m; western Amazonas). Amazonian Colombia, Brazil (Amazonas: Rio Negro basin). ◆Fig. 429. Vochysia complicata Ducke, Bull. Mus. Hist. Nat. (Paris) sér 2, 4: 738. 1932. Tree 15–30 m tall. Riparian forests along

black-water rivers, 100–200 m; Amazonas (Río Guainía, Río Negro). Brazil (Amazonas: Rio Negro basin). Vochysia costata Warm. in Mart., Fl. Bras. 13(2): 100. 1875. Tree to 30 m tall. Montane to upper montane forests, 1100–1700 m; Bolívar (Gran Sabana), Amazonas (basin of Río Coro Coro west of Cerro Yutajé). Guyana, Brazil. ◆Fig. 430. Vochysia crassifolia Warm. in Mart., Fl. Bras. 13(2): 77. 1875. Vochysia curvata Klotzsch ex R.M. Schomb., Reis. Br.-Guiana 3: 1099. 1848, nom. nud. Tree 4–20 m tall. Riparian forests, lowland to upland savannas, 50–1500 m; widespread in Bolívar, Amazonas (Samariapo north toward El Burro). Guyana, Brazil (Amazonas: Rio Branco). ◆Fig. 432. Vochysia elegans Stafleu, Acta Bot. Neerl. 3: 405, fig 1c–d. 1954. Tree to 20 m tall. Evergreen lowland forests, 100–200 m; Amazonas (between San Carlos de Río Negro and Solano). Brazil (Amazonas: Rio Negro basin). Vochysia expansa Ducke, Arq. Inst. Biol. Veg. 4(1): 32. 1938. Tree to 18 m tall. Riparian forests, 100– 200 m; Amazonas (San Carlos de Río Negro). Brazil (Amazonas: Rio Negro basin). Vochysia ferruginea Mart., Nov. Gen. Sp. Pl. 1: 151, t. 92. 1824 [1826]. —Cucullaria ferruginea (Mart.) Spreng., Syst Veg. 4(cur. post.): 9. 1827. —Vochya ferruginea (Mart.) Standl. in Britton & Rose, N. Amer. Fl. 25: 302. 1924. —Caniba, Pese, Saladillo. Tree to 22 m tall. Riparian forests, savannas, 50–100 m; Bolívar (Río Suapure), cen-

Vochysia 517

tral and western Amazonas. Apure, Táchira, Zulia; Nicaragua, Costa Rica, Panama, Colombia, Guyana, Ecuador, Peru, Brazil. ◆Fig. 435. Vochysia glaberrima Warm. in Mart., Fl. Bras. 13(2): 78. 1875. —Salao. Vochysia lucida Klotzsch ex R.M. Schomb., Reis. Br.-Guiana 3: 1099. 1848, nom. nud. Tree 8–30 m tall. Riparian forests, moist savannas, 50–900 m; Bolívar (Río Parguaza, near Santa Elena de Uairén, 14 km east of Túriba), Amazonas (between Puerto Ayacucho and Samariapo, Río Sipapo). Guyana, Brazil (Amazonas, Roraima). ◆Fig. 433. Vochysia grandis Mart., Nov. Gen. Sp. Pl. 1: 146, t. 88. 1824 [1826]. —Cucullaria grandis (Mart.) Spreng., Syst. Veg. 4(cur. post.) 9. 1827. Vochysia grandis var. genuina Briq., Ann. Conserv. Jard. Bot. Genève 20: 386. 1919. Colombia, Venezuela, Brazil; 3 varieties, 1 of these in Venezuela. V. grandis var. uaupensis Warm., Symbolae 13(2): 75. 1875. Tree to 35 m tall. Evergreen lowland forests, 100–200 m; Amazonas (between Solano and San Carlos de Río Negro). Colombia, Brazil. Vochysia jonkeri Marc.-Berti, Pittieria 10: 3, fig. 1. 1982. Tree to 15 m tall. Montane tepui summit forests, ca. 1200 m; Bolívar (Cerro Guaiquinima). Endemic. Vochysia julianensis Marc.-Berti, Pittieria 18: 8. 1989. Tree to 25 m tall. Riparian forests, 100– 200 m; Amazonas (Río Yatúa at base of Sierra de la Neblina and at Laja Catipán). Endemic. Vochysia aff. laxiflora Stafleu, Acta Bot. Neerl. 3: 407. 1954. Tree ca. 25 m tall. Evergreen lowland forests, 100—200 m; Amazonas (Río Negro). Suriname. Vochysia liscanoi Marc.-Berti, Pittieria 10: 6, fig 2. 1982.

Tree 20–35 m tall. Montane forests, 1300– 1400 m; Bolívar (La Escalera). Endemic. Vochysia obscura Warm. in Mart., Fl. Bras. 13(2): 73, t. 13. 1875. —Salado, Sanaparo fino. Tree 5–25 m tall. Riparian forests, 50–200 m; western Amazonas. Colombia, Peru, Brazil. ◆Fig. 437. Vochysia ortegae Marc.-Berti & Bautista, Pittieria 24: 63–67, fig 1. 1996. Tree to 28 m tall and 60 cm diameter. Montane forests, ca. 1500 m; Amazonas (Caño Piedra on Cerro Sipapo). Endemic. Vochysia pauciflora Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 424. 1966. Tree 25–30 m tall. Upland forests, ca. 1200 m; Bolívar (La Escalera). Endemic. Vochysia pinkusii A.C. Sm., Bull. Torrey Bot. Club 67: 288. 1940. Tree 20 m tall. Montane forests, 1300– 1400 m; Bolívar (expected). Brazil. Vochysia pinkusii is probably in Bolívar. It has been collected just across the Venezuelan border in Estado Roraima, Brazil, along Maurukow Creek, in the headwaters of the Rio Cotinga. Vochysia punctata Spruce ex Warm. in Mart., Fl. Bras. 13(2): 102. 1875. Tree 15–30 m tall. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro to Solano, Yavita to Pimichín trail). Brazil. Vochysia rubiginosa Stafleu, Fieldiana, Bot. 28: 296. 1952. Tree 9–12 m tall. Montane forests, ca. 1600 m; Bolívar (Gran Sabana at Río Aponguao, ridge between Ptari-tepui and Soropopán-tepui). Endemic. Vochysia saccata Stafleu, Recueil Trav. Bot Néerl. 41: 508. 1948. —Saladillo. Vochysia inundata var. venosa Ducke, Arq. Inst. Biol. Veg. 4: 35. 1938. Tree 6–25 m tall. Riparian forests, 100– 300 m; Amazonas (near Canaripó, bases of Cerros Huachamacari and Duida, Macuruco, Río Yatúa). Brazil (Amazonas: Rio Negro basin).

518

V OCHYSIACEAE

Vochysia spathiphylla Stafleu, Acta Bot. Neerl. 6: 341, fig. 1c,d. 1957. —Godedi. Tree 15–30 m tall. Lower to upper montane forests, 200–1700 m; Amazonas (Cerro Huachamakari, Río Cunucunuma, Sierra de la Neblina). Colombia, Brazil (Amazonas: Rio Negro basin). Vochysia splendens Spruce ex Warm. in Mart., Fl. Bras. 13(2): 101. 1875. —Sanaparo. Vochysia vismiifolia var. robusta Ducke, Arq. Inst. Biol. Veg. 4: 35. 1938, “vismiaefolia.” Tree to 30 m tall. Riparian and evergreen lowland forests, 100–200 m; Amazonas (25 km southwest of San Antonio del Orinoco, San Carlos de Río Negro to Solano, San Fernando de Atabapo). Brazil (Amazonas: Rio Negro basin). ◆Fig. 438. Vochysia steyermarkiana Marc.-Berti, Pittieria 13: 10. 1986. Tree to 24 m tall. Río Negro caatinga, 100–200 m; Amazonas (Yavita to Maroa). Endemic. Vochysia surinamensis Stafleu, Recueil Trav. Bot. Néerl. 41: 439. 1948. Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 varieties, both in the flora area.

Mey., Prim. Fl. Esseq. 12. 1818. —Vochysia tetraphylla (G. Mey.) H. Stone & W. Freeman, Timb. Brit. Guiana 26. 1914, “Vochisia.” —Daujén, Lorjena. Vochysia arcuata Garcke, Linnaea 22: 58. 1849. Tree to 40 m tall. Riparian forests, 200– 500 m; southern Delta Amacuro, widespread in eastern and central Bolívar. Guyana, Suriname, French Guiana, Brazil. ◆Fig. 434. Vochysia tilletii Marc.-Berti, Pittieria 10: 14, fig. 5. 1982. Tree 4–15 m tall. Low, open forests, sandstone slopes with streams, savannas, 500– 1200 m; Amazonas (Caño Yutaje, Cerro Duida, Cerro Huachamacari, Cerro Moriche, Cerro Yaví, Río Coro Coro). Endemic. Vochysia tomentosa (G. Mey.) DC., Prod. 3: 26. 1828. —Cucullaria tomentosa G. Mey., Prim. Fl. Esseq. 13. 1818. Cucullaria excelsa M. Vahl, Enum. Pl. 1: 4. 1804, non Willd. 1797. Tree 10–35 m tall. Evergreen lowland and lower montane forests generally along streams, 100–800 m; Bolívar (Serranía de los Pijiguaos), Amazonas (Río Padamo). Guyana, Suriname, French Guiana, Brazil.

Key to the Varieties of V. surinamensis 1. Spur inflated, 0.3–0.4 mm wide ................ .......................................... var. inflata 1. Spur not or only slightly inflated, 1.9–2.3 mm wide ................ var. surinamensis V.

surinamensis var. inflata Stafleu, Recueil Trav. Bot. Néerl. 41: 440. 1948. Tree to 24 m tall. Montane forests, 1000– 1500 m; Bolívar (Ptari-tepui, Serranía de los Pijiguaos, Sororopán-tepui) Endemic. V. surinamensis var. surinamensis. —Canelito, Daujén, Saladillo. Tree to 40 m tall. Semievergreen forests, 200–400 m; Delta Amacuro (area near Río Toro), northeastern Bolívar, northwestern Amazonas. Guyana, Suriname, French Guiana, Brazil (Pará). ◆Fig. 431. Vochysia tetraphylla (G. Mey.) DC., Prod. 3: 27. 1828. —Cucullaria tetraphylla G.

Fig. 428. Vochysia apopetala

Vochysia 519

Fig. 429. Vochysia catingae

Fig. 430. Vochysia costata

520

V OCHYSIACEAE

Fig. 431. Vochysia surinamensis var. surinamensis

Fig. 432. Vochysia crassifolia

Vochysia 521

Fig. 433. Vochysia glaberrima

Fig. 434. Vochysia tetraphylla

522

V OCHYSIACEAE

Fig. 435. Vochysia ferruginea

Fig. 436. Vochysia venezuelana

Vochysia 523

Fig. 437. Vochysia obscura

Fig. 438. Vochysia splendens

524

V OCHYSIACEAE

Vochysia venezuelana Stafleu, Recueil Trav. Bot. Néerl. 41: 437. 1948. —Sábila, Salado, Salado sabanero, Saldillo. Tree 5–20 m tall. Mainly savannas along rivers and streams, 50–300 m; northwestern Bolívar, Amazonas (Isla Ratón, Puerto Ayacucho to Samariapo, Río Cataniapo, Río Orinoco, Río Ventuari near mouth of Río Parú). Anzoátegui, Apure, Guárico, Portuguesa; Colombia. ◆Fig. 436.

Vochysia vismiifolia Spruce ex Warm. in Mart., Fl. Bras. 13(2): 99. 1875. —Lacre montañero, Salado rojo, Sanaparo. Tree to 30 m tall. Riparian forests, 100– 500 m; Bolívar (Río Canaracuni, Río Tonoro, Serranía de los Pigijuaos), Amazonas (Galipero north of Puerto Ayacucho, Río Cuao, Río Sipapo, San Carlos de Río Negro, Yavita). Colombia, Brazil.

WINTERACEAE by James S. Miller Glabrous, evergreen trees or shrubs. Leaves alternate, simple, entire, pellucidpunctate, often pale on the lower surface, exstipulate. Inflorescences terminal or axillary, fasciculate, umbellate, or cymose, or the flowers solitary and terminal. Flowers hypogynous, actinomorphic or slightly zygomorphic, bisexual or unisexual (Tasmannia), the plants then dioecious or polygamodioecious. Sepals 2–6, valvate, free or slightly connate basally, rarely fully connate; petals (2–)5–many, in (1)2 or more whorls, imbricate, free. Stamens numerous; filaments free, flattened; anthers bithecal, dehiscing longitudinally. Carpels few to many, in a single whorl, free or partly connate, conduplicate and often open; style terminal or the stigma decurrent along the open carpel margin; ovules 1–many per locule, marginal, anatropous. Fruits baccate or a follicle, occasionally the follicles united into a syncarp. Seeds with abundant, oily endosperm. New World tropics (Drimys), Madagascar (Takhtajania), but mostly southwestern Pacific and adjacent Australia; 9 genera and ca. 100 species, 1 species in the flora area. The largest genus, Tasmannia, is treated as a section of Drimys by some authors. 1. DRIMYS J.R. Forst. & G. Forst., Char. Gen. Pl. 83. 1775. Wintera Murray, Syst. Veg. ed. 14, 502. 1784. Trees or shrubs, sometimes growing as epiphytes. Leaves coriaceous. Inflorescences terminal, fasciculate or umbellate, or the flowers solitary, usually subtended by imbricate bracts. Flowers actinomorphic, bisexual. Sepals connate in bud, later splitting into 2 or 3 caducous lobes; petals 4–17(–25), in 1–3 whorls, white. Stamens 15–50(–65), in 2–4(5) whorls; filaments fleshy. Carpels 3–12(–24), free; stigma lateral or subterminal; ovules biseriate on marginal placentas, 6–26 per carpel. Fruits apocarpous, baccate, purple to black. Neotropics (Drimys sensu stricto); 4 species, 2 in Venezuela, 1 of these in the flora area. The other species in Venezuela is Drimys granatensis L. f.

Drimys 525

Drimys roraimensis (A.C. Sm.) Ehrend. & Gottsb., Pl. Syst. Evol. 132: 72. 1979. —Drimys brasiliensis var. roraimensis A.C. Sm., J. Arnold Arbor. 24: 32. 1943. —Hue-tu-yek (Arekuna). Drimys brasiliensis var. ptaritepuiensis Steyerm., Fieldiana, Bot. 28: 235. 1951. Shrub or small tree to 4(–8) m tall; lower surface of leaves pale white or silvery; flowers white, fragrant; fruiting pedicels wine-

red. Montane and cloud forests, dwarf forests, slopes of tepuis, along streams, in swamps of Chimantaea, 1800–2500 m; Bolívar (Auyán-tepui and Macizo del Chimantá east to Roraima-tepui), Amazonas (Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Sierra de la Neblina). Adjacent Brazil. ◆Fig. 439. The peppery-tasting bark of Drimys roraimensis is used for relief of toothache, headache, and fever.

Fig. 439. Drimys roraimensis

526

X YRIDACEAE

XYRIDACEAE by Robert Kral Rosulate or caulescent, fine or coarse, monocotyledonous, terrestrial herbs, mostly of high-hydroperiod acidic soils. Roots mostly slender, diffuse-fibrous, with root hairs. Axis sympodial or monopodial. Leaves alternate, distichous, or spiral, ligulate or eligulate, the bases broad, open-sheathing, frequently equitant and keeled, the blades laterally to dorsiventrally compressed. Inflorescence lateral to terminal, scapose, the scapes 1–few from axils of scape sheaths or inner leaves, naked or with distant pairs of bracteal leaves, sometimes ancipital (flattened and with 2 sharp edges), each scape bearing apically 1 or more imbricate-bracted spikes, heads, or panicles of spikes. Flowers bisexual, 1–many, solitary and subsessile to pedicellate in axils of chaffy, leathery, or scarious bracts. Perianth of 2 differentiated whorls; sepals (2)3, the anterior (inner) one a reduced scale or (usually) membranous and wrapped around the corolla, abcissing as the flower opens, the other 2 subopposite, connivent to basally connate, chaffy, mostly boat-shaped, often keeled, persisting around the ripe capsule; petals 3, equal or unequal, distinct to united and salverform, mostly clawed, the spreading blades broad, yellow to white or blue. Stamens 3, epipetalous; staminodia (1 or 2)3, scale-like, bearded or beardless, or reduced or lacking; anthers 4-sporangiate, usually 2-locular at anthesis, introrsely or laterally dehiscent, dehiscing longitudinally; pollen monosulcate or inaperturate. Gynoecium 3-carpellate, the ovary 1-locular to completely or incompletely 3-locular, the placentation marginal, parietal, basal, free-central, or axile; style terminal, tubular, slender, appendaged or unappendaged, apically 3-branched or variously laminar or fimbriate; stigmas 3. Fruit capsular, mostly loculicidal. Seeds usually numerous, usually with strong longitudinal ridges and finer cross lines, translucent or farinose-opaque; embryo small, situated at base of an abundant mealy endosperm. Pantropics; 5 genera and ca. 300 species, 5 genera and 95 species plus 1 hybrid in the flora area. There are over 200 species in Xyris, the only genus to range into the north temperate zone. For complete descriptive detail of Xyris, including illustrations, please refer to Ann. Missouri Bot. Gard. 75: 511–722. 1988. The remaining genera are confined to South America with Aratitiyopea, Achlyphila, and Orectanthe confined to the Guayana Highlands and Abolboda extending south into central Brazil. For additional details and illustrations of Abolboda, Achlyphila, Aratitiyopea, and Orectanthe, please see Ann. Misouri Bot. Gard. 79: 819–885. 1992. Key to the Genera of Xyridaceae 1.

1.

2(1). 2.

Petals nearly or quite distinct; leaves distichous, mostly equitant; styles unappendaged; pollen lacking spines; corolla actinimorphic; sepals 3 .............................................................................................................. 2 Petals connate; leaves polystichous; styles with appendages or ovary summit appendaged; pollen with spines or productate (spinose, tuberculate, or muricate); corolla actinomorphic or zygomorphic; sepals 2 or 3 .............................................................................................................. 3 Flowers distinctly pedicellate; sepals alike in character; staminodia lacking; style unbranched; stigma capitate, 3-lobed ................... 2. Achlyphila Flowers subsessile; sepals unlike, the lateral ones chaffy and keeled, opposite, exterior, the inner one membranous and enfolding the corolla in

Abolboda 527

3(1).

3.

4(3).

4.

bud; staminodia usually present; style 3-branched; stigmas U-shaped or annular ........................................................................................ 5. Xyris Stylar appendages well above style base; capsule apex thickened; flowers 1–several, sessile or on variously elongated, opposite-bracted scapes in spikes; sepals usually 2 ............................................................ 1. Abolboda Stylar appendages at or near style base; capsule apex not appreciably thickened; flowers many in large, dense, globose or hemispheric heads; sepals 3 ................................................................................................... 4 Corollas excurved (curved outward), yellow (rarely red-purple), irregular; stigma capitate, a lateral-terminal papillate cushion; seeds winged, asymmetrical ......................................................................... 4. Orectanthe Corollas erect, purple, regular; stigma terminal and 3-lobed, the lobes pilose; seeds wingless, ridged and pitted, symmetrical ........ 3. Aratitiyopea

1. ABOLBODA Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 25, 109, pl. 114. 1809. Chloerum Willd. ex Link in Spreng. et al., Jahrb. Gewächsk. 1(3): 74. 1820. Poarchon Mart. ex Seub. in Mart., Fl. Bras. 3(1): 223. 1855. Coarse to low and moss-like, annual or perennial, short-caulescent to acaulescent, rosulate herbs with fine or coarsely fibrous roots, perennating by axillary lateral buds. Axis sympodial. Foliage externally smooth to variously papillose or tuberculate; principal leaves polystichous, the bases typically with sheaths open, broadly clasping, and blades variously linear-triangular or linear, compressed or thickened and angulate, or lingulate, usually narrowly acute at tips, there frequently and excentrically aristate or conic-subulate. Inflorescence a single sessile flower, or 2 or more flowers in spikes and sessile or (more commonly) the flowers in spikes raised and terminal on scapes, the spikes either single or terminal or clustered and terminal or terminal on branches of a branched inflorescence, the scape itself either naked or usually with 1 or more pairs of subopposite, sessile, erect, leafy but short, bracts; spike bracts paired at spike base, upward in subspirals, glumaceous, usually rigid and lanciform-boat-shaped, with usually entire, scarious borders. Sepals 2 or 3, similar to lower bracts but somewhat smaller, the lateral ones subopposite, variously keeled at least apically, exterior, the 3rd sepal, if present, thinner and reduced; petals 3, equal, salverform, the tube variously elongate, the 3 lobes broad, ephemeral, blue, lavender, or white. Stamens 3; filaments short, adnate near base of petal blades and opposite them; anthers mostly narrow and basifixed; staminodia absent or reduced. Ovary 3-locular, the placentation axile; ovules usually numerous; style typically elongate, with 2 or 3, usually pendent, claviform appendages; stigma funnelform and fimbriate. Capsules loculicidal, the 3 valves firm. Seeds usually tumidly and irregularly obovoid, short-apiculate or umbilicate, the broad body dark, spirally longitudinally sharply ridged. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Bolivia; 20 species, 17 plus 1 hybrid in Venezuela, all in the flora area. Key to the Species of Abolboda 1.

1.

Plants without scapes, mostly strongly clumped and moss-like; inflorescences sessile or subsessile, usually hidden among the rosette leaves; leaves usually < 4 cm long ..................................................................... 2 Plants scapose; scapes elevated to at least the level of the leaf blades; leaves various in shape and size ........................................................... 3

528

2(1).

X YRIDACEAE

Leaves of a rosette mostly uniform in outline, smooth, gradate only in size, usually with bases tightly imbricate in tall spirals on elongated stems below the terminal rosette, those of the rosette narrowly triangular and stiff, tapering evenly from base to tip, there with an arista > 1 mm long; flowers several per inflorescence, the corolla blue ....... A. acaulis 2. Leaves of a rosette mostly distinctly dimorphic with many rosettes comprised of short (1 cm or less long) fleshy, angulate, blunt and often granular-papillose leaves, or these mixed with longer, broader-based, longer- and narrower-bladed innovations around flowers and new rosettes, these leaves more narrowly acute or even aristate; flowers 1 or 2 per inflorescence, the corolla blue or white .............................. A. killipii 3(1). Scapes ebracteate or if with bracts, these a single pair (rarely 2 pairs) at or near scape summit or sheathing its very base ................................. 4 3. Scapes with 1 or more pairs of scape bracts, at least 1 of these pairs at or toward middle of scape .......................................................................... 8 4(3). Plants with comparatively coarse foliage, the principal leaves > 15 cm long, their blades 3–5 mm wide; spikes (excluding involucres) 1.5– 2.5 cm long; perennials with short, stout rhizomes ............................. 5 4. Plants with comparatively fine or short leaves, these rarely over 11 cm long, their blades < 3 mm wide; spikes 0.5–1.5 cm long; perennials or annuals ................................................................................................... 6 5(4). Inflorescence overtopped by longer leaves, the leaf blades with strongly thickened, pale, cartilaginous margins; spike bracts and sepals strongly excurved; scape bracts mostly ca. 2 cm long with 5–20 mm of scape between these and the spike base ......................................... A. bella 5. Inflorescence overtopping longer leaves, the leaf blades lacking strongly thickened, pale, cartilaginous margin; spike bracts and sepals not excurved; scape bracts 3–7 cm long, with to 40 mm of scape above, or directly subtending spike and making a leafy involucre ........... A. sprucei 6(4). Annual with soft, filiform leaves and filiform scapes ............... A. americana 6. Perennial with firm leaves and linear scapes .......................................... 7 7(6). Scapes > 2 times longer than principal leaves; scape bracts strongly unequal, forming sheath at scape base ...................................... A. ebracteata 7. Scapes barely if at all overtopping principal leaves; scape bracts lacking ...................................................................................................... A. acaulis 8(3). Spikes glomerate at scape tip ................................................... A. ×glomerata 8. Spikes solitary and terminal at scape tips or tips of scape branches ...... 9 9(8). Flowers per spike 4 or fewer; spikes (except in A. paniculata, A. neblinae) 1.5 cm long or less ................................................................................ 10 9. Flowers (4)5–many per spike; spikes mostly 1.5–8(–10) cm long .......... 16 10(9). Bracts and usually scapes strongly purple-tinged; scape bracts (2)3 pairs or more; bract tips remaining erect, thus fruiting inflorescence narrow; sepals 3, all without keel ..................................................................... 11 10. Bracts and scapes not strongly purple-tinged; scape bracts mostly 1, rarely 2; sepals usually 2 and strongly keeled ................................... 13 11(10). Inflorescence much branched, the spikes terminal on elongate branches; bract pairs many .................................................................... A. paniculata 11. Inflorescence appearing unbranched, the spikes 1 or 2, the second penultimate and sessile; bract pairs 0–4(–7) ..................................... 12

Abolboda 529

12(11). Leaves tapering gradually from base to tip, thus linear-triangular, the blades flattened and with conic-subulate, even aristate, tips .................................................................................................... A. neblinae 12. Leaves narrowed above clasping base, not tapering-bladed, the blades narrowly linear, thickened, their tips with an excentric apiculus .................................................................................................... A. uniflora 13(10). Spike bracts prevalently ovate, the scarious borders abruptly converging to a stubby mucro; lower (outer) spike bracts shorter than the rest of the spike .................................................................................... A. pulchella 13. Spike bracts prevalently lanciform, gradually narrowing to a slender, blunt, mucronate or spinulose tip; lower (outer) spike bracts as long as or longer than the rest of the spike ..................................................... 14 14(13). Leaves and/or scapes smooth or no more than papillate; leaves of a tuft mostly distinctly dimorphic, these and bract apices often spinulosetipped ....................................................................................... A. acicularis 14. Leaves and/or scapes liberally frosted with prominent tubercles, thus tuberculate-scabrid and also often rugose; leaves essentially monomorphic, these and bract apices blunt-tipped ........................................... 15 15(14). Both leaves and scapes tuberculate-rugose; costas of bracts and sepals scabridulous from middle to tip ................................................ A. scabrida 15. Only scapes tuberculate-rugose; costas of bracts and sepals smooth or nearly so .............................................................................. A. dunstervillei 16(9). Spikes narrowly ellipsoid, narrowly lanceoloid, or cylindric; sepals blunttipped and barely projecting beyond subtending bracts .................... 17 16. Spikes broadly ellipsoid, ovoid, globose, or hemispheric; sepal tips acute, the exsertion various ........................................................................... 18 17(16). Blades linear-triangular, gladiate, or linear-lorate, tapering evenly above to acute tips, thin-edged and plane, finely multiveined and comparatively smooth ..................................................................... A. macrostachya 17. Blades narrowly linear, the tips blunt, the edges thick, the surfaces with few coarse and raised veins .................................................. A. linearifolia 18(16). Scapes with 1 pair of bracts; leaves < 10 cm long and < 5 mm wide, the blades toward apex thick, ± 3-angled or with a strong dorsal costa, at blunt tips excentrically spinulose, the margins toward apex ascending spinulose-serrulate; lowest spike bracts length equal to, or slightly longer than, the rest of the spike ................................................. A. ciliata 18. Scapes usually with > 1 pair of bracts; leaves > 10 cm long and usually much broader than 5 mm, the blades flat to very tip, acute and usually not excentrically spinulose, the margins entire; lowest bracts much shorter than the rest of the spike .............................................. A. grandis Abolboda acaulis Maguire, Bull. Torrey Bot. Club 75: 191. 1948. Rosulate, smooth perennial; rosettes dense moss-like cushions; leaves pale green, linear-triangular, 1.5–3(–5) cm long; scapes lacking or short; corolla lobes pale blue. Rocky wet shrub, rapateaceous savannas and tepui meadows, 400–2200 m. Venezuela, Guyana; 2 varieties, both in the flora area.

Key to the Varieties of A. acaulis 1. Spikes sessile ......................... var. acaulis 1. Spikes on scapes ................... var. scaposa A. acaulis var. acaulis Bolívar (Gran Sabana), Amazonas (Cerro Huachamacari, Cerro Sipapo, Cerro Yutajé, Río Coro Coro, Río Manapiare). Guyana.

530

X YRIDACEAE

A. acaulis var. scaposa Kral, Ann. Missouri Bot. Gard. 79: 825. 1992. Habitat and distribution the same as in the type variety, but less frequent, Bolívar (Cerro Guaiquinima, Gran Sabana), Amazonas (Cerro Sipapo, Cerro Yapacana). Guyana (Pakaraima Mountains). Abolboda acicularis Idrobo & L.B. Sm., Caldasia 6: 250. 1954. Low perennial; rosettes tufted; leaves curved, smooth, or papillate-granular, 1–3 (–5) cm long; scapes 1–few, 5–30 cm tall, with 1 evident pair of bracts. Moist white-sand savannas, Amazonian Colombia, Venezuela; 2 varieties, both in the flora area. Key to the Varieties of A. acicularis 1. Leaf blades smooth, most of the principal ones subulate-setose-tipped; spike bracts > 1 cm long, spinulose at tip ................... ..................................... var. acicularis 1. Leaf blades or most of them granular-papillose, also often rugulose, averaging wider, with blunter tips, these apiculate or aristulate; spike bracts ≤ 1 cm long, aristulate or apiculate ..... var. granularis A. acicularis var. acicularis 100–200 m; Bolívar (3 km southwest of Perai-tepui), Amazonas (Caño Caname, Caño Yagua, Cerro Yapacana, Orinoco savannas, Río Guayapo). Amazonian Colombia. A. acicularis var. granularis Maguire, Mem. New York Bot. Gard. 10(1): 17. 1958. 100–200 m; Amazonas (Caño Pimichín, Cerro Venado, Cerro Vinilla, Chapazón, south-southwest of Ocamo, Río Guainía basin, Santa Cruz). Amazonian Colombia. Abolboda americana (Aubl.) Lanj., Recueil Trav. Bot. Néerl. 34: 492. 1937. —Xyris americana Aubl., Hist. Pl. Guiane 140, pl. 14. 1775. Xyris coerulea Lam., Tabl. Encycl. 1: 132. 1791 [1793]. Abolboda imberbis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 256. 1815 [1816]. —Abolboda americana var. imberbis (Kunth) Maguire, Bull. Torrey Bot. Club 75: 193. 1948. Abolboda poeppigii Kunth, Enum. Pl. 4: 27. 1843.

Annual; rosettes forming dense tufts; leaves linear-filiform, 1.5–11 cm long, mostly overtopped by filiform scapes; spikes fewflowered, to ca. 1 cm long, turbinate; corolla pale blue. Wet, sandy savannas or forest transitions, 100–500 m; Bolívar (west of Canaima, Cerro Guaiquinima, Cerro Pitón, Icabarú, Sabana de Triana in Río Chicanán basin, Salto Hacha), Amazonas (Caño Caname, Caño Cumare, Cerro Autana, base of Cerro Duida, Cerro Yapacana, La Esmeralda, Maroa, above Pimichín, San Carlos). Colombia, Trinidad, Guyana, Suriname, French Guiana, Amazonian and northeastern coastal Brazil. Abolboda bella Maguire, Mem. New York Bot. Gard. 10(1): 15, fig. 2k–l. 1958. Low, but rigid, rhizomatous perennial to 15 cm tall; leaves subdistichous with blades 2–3 mm wide, longer than scapes; scapes to 10 cm long with 1 pair of bracts to 2 cm long 5–25 mm below the single spike; spikes fewflowered; corolla dark, bright blue. Whitesand savannas, 100–200 m; Amazonas (base of Cerro Yapacana). Endemic. Abolboda ciliata Maguire & Wurdack, Mem. New York Bot. Gard. 10(1): 17. 1958. Slender perennial; rosettes clumped; leaves 3–7 cm long, blades thickened, angulate, tips blunt and excentrically aristulate, edges scabrid-ciliate; scapes slender, to 20 cm tall, with 1 pair of bracts 3–4 cm long; spikes turbinate, to 1.5 cm long; corolla blue. Tepui meadows, 1900–2300 m; Amazonas (Sierra de la Neblina). Endemic. Abolboda dunstervillei Maguire ex Kral, Ann. Missouri Bot. Gard. 79: 851. 1992. Low perennial; rosettes in dense clumps; leaves stiff, narrowly linear, the blades flat, 30–45 × 0.7–1 mm, blunt with an excentric mucro, the surface of the leaves and scapes variously rugulose-papillate; scapes linear, to 9 cm long with 1 pair of bracts to 15 mm long; spikes turbinate, 10–12 mm long; bracts lanciform; flowers 2 or 3. Tepui summit, ca. 1500 m; Amazonas (Cerro Avispa). Endemic. Abolboda ebracteata Maguire & Wurdack, Mem. New York Bot. Gard. 10(1): 17. 1958. Low, smooth perennial; rosettes tufted; leaves 5–15 cm long, the linear blades 3angled, spinulose-tipped; scapes to 25 cm tall, ebracteate; spikes to 12 mm long; petals

Abolboda 531

purple-blue. Moist white-sand savannas, 100–200 m. Colombia, Venezuela; 2 varieties, both in the flora area. Key to the Varieties of A. ebracteata 1. Lower spike bracts shorter than the rest of spike; longer leaves < 1/2 as long as the scape ............................. var. brevifolia 1. Lower spike bracts exceeding, rarely equaling, rest of spike; longer leaves to 1/2 as long as the scape ........ var. ebracteata A. ebracteata var. brevifolia Maguire, Mem. New York Bot. Gard. 10(5): 8. 1964. Amazonas (Caño Perro de Agua, Caño Yagua, lower Río Orinoco basin, along the Río Ventuari). Adjacent southeastern Colombia. A. ebracteata var. ebracteata Amazonas (Cerro Yapacana). Endemic. Abolboda ×glomerata Maguire, Mem. New York Bot. Gard. 10(1): 14. 1958. Low perennial from stout rhizome; leaves few, to 10 cm long, the blades linear, 2–3 mm wide, firm, flat; scapes to 30 cm tall, flattened; spikes lanceoloid in ellipsoid tight terminal glomerules. White-sand savannas, 100–200 m; Amazonas (Sabana Venado on Caño Pimichín above Puerto Pimichín). Endemic. Not even undeveloped flowers or fruit are present, a mark of a sterile hybrid. Abolboda grandis Griseb., Linnaea 21: 281. 1848. Abolboda grandis var. minor Spruce ex Malme, Bih. Kongl. Svenska Vetensk.Akad. Handl. 26(afd. 3, no. 19): 14. 1901. Low to tall, coarse, rhizomatous perennial; rosettes solitary or clumped; leaves flatbladed, over 10 cm long; scapes elongate with 1–3 pairs of bracts; spikes over 1.5 cm long, many-flowered; corollas large, light blue-violet. Wet savannas, edges of gallery forests, low to high altitudes, Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 2 varieties, both in the flora area. Key to the Varieties of A. grandis 1. Lowest (largest) pair of scape bracts mostly 2–3(–3.5) cm long; spikes broadly ovoid to hemispheric or subglobose, usu-

ally < 2 cm long ............... var. grandis 1. Lowest (largest) pair of scape bracts (3–) 3.5–6 cm long; spikes broadly ovoid to ellipsoid or narrowly ovoid, mostly > 2.5 cm long .............................. var. rigida A. grandis var. grandis Shorter than the following variety; leaves commonly lorate; scapes more slender. 50– 1800 m; Bolívar (Auyán-tepui, south of El Pauji, east of Icabarú, Uaipán-tepui), Amazonas (widespread). Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. A. grandis var. rigida Malme, Bull. Torrey Bot. Club 58: 326. 1931. —Abolboda rigida (Malme) Steyerm., Fieldiana, Bot. 28: 105. 1951, non Gleason 1929. Abolboda pervaginata Malme, Ark. Bot. 25A(12): 16. 1933. Abolboda grandis var. guayanensis Maguire, Mem. New York Bot. Gard. 10(1): 8. 1958. Coarser and taller than the type variety; leaves less often lorate; scapes to 2–4 mm thick. 300–1700 m; Bolívar (southern base of Auyán-tepui, 25 km east-northeast of Canaima, Cerro Guaiquinima, Cerro Guanay, Cerro Jaua, Cerro Pitón, El Paují, Icabarú, Sierra Pakaraima), Amazonas (Cerro Duida, Cerro Guaiquinima, Cerro Guanay, Cerro Parú, Cerro Sipapo, 1 km east of Maroa, Río Siapa just below Raudal Gallineta, Sierra Parima, between Yavita and Pimichín). Colombia, Guyana, Suriname, Amazonian Brazil. There are numerous integrades with var. grandis at lower elevations. Abolboda killipii Lasser, Bol. Soc. Venez. Ci. Nat. 9: 178. 1944. Abolboda psammophila Maguire, Bull. Torrey Bot. Club 75: 193. 1948. Abolboda minima Maguire, Mem. New York Bot. Gard. 10(5): 7. 1964. Delicate, low perennial; rosettes moss-like in dense clumps; leaves dense, dimorphic, one type stubby, 1 cm long or less, 0.2–0.3 mm thick, mostly granular-papillate, the other longer, 1–2 cm long, flatter, mucronate or blunt; inflorescence sessile, 1 or 2(3)-flowered; corolla white or pale blue. Mostly white-sand savannas, 5–500 m; Bolívar (southeast of El Dorado, Río Cusimi, Triana

532

X YRIDACEAE

Sabana on Cerro Pitón), Amazonas (west of Cerro Cuao, Cerro Vinilla, Cerro Yapacana, near Pimichín, Sabana Cucurital, San Antonio, San Carlos de Río Negro). Guyana, Amazonian Brazil. ◆Fig. 440. Abolboda linearifolia Maguire, Mem. New York Bot. Gard. 10(1): 14, fig. 3a–f. 1958. Smooth perennial with stout horizontal rhizome; leaves subdistichous, > 1/2 as long as the scapes, the blades 2–3 mm wide and coarsely veined; scapes to 50 cm tall with a single pair of bracts. White-sand savannas, 100–200 m; Amazonas (Río Atabapo, upper Río Orinoco, Río Pasimoni, Río Siapa). Endemic. ◆Fig. 442. Abolboda macrostachya Spruce ex Malme, Bih. Kongl. Svenska Vetensk.Akad. Handl. 26(afd. 3, no. 19): 15. 1901. Abolboda macrostachya var. angustior Maguire, Mem. New York Bot. Gard. 10(1): 10. 1958. Smooth, tufted or solitary, mostly coarse, stout-rhizomatous perennial (17–)20–120 cm tall; leaf blades linear-triangular, flat or involute; spikes many-flowered, cylindric; corollas pale, bright lavender-blue, 2–4 cm long. Wet, sandy or rocky savannas. Colombia, Venezuela, Guyana, Amazonian Brazil; 2 varieties, both in the flora area. Key to the Varieties of A. macrostachya 1. Scapes with mostly 2 or 3 pairs of bracts, these 1.5–3 cm long; plants mostly < 60 cm tall ................... var. macrostachya 1. Scapes mostly 1(2) pair of bracts, these mostly 4–6 cm long; plants mostly 60– 100 cm tall ................... var. robustior A. macrostachya var. macrostachya 100–1200 m; Bolívar (between Río Horeda and Cerro Gavilán), Amazonas (widespread). Colombia, Amazonian Brazil. A. macrostachya var. robustior Steyerm., Fieldiana, Bot. 28: 104. 1951. Abolboda rigida Gleason, Bull. Torrey Bot. Club 56: 17. 1929. Abolboda excelsa Malme, Ark. Bot 25A(12): 16. 1933. (100–)600–2600 m; Bolívar (widespread), Amazonas (Cerro Sipapo, near Cerro Yapacana). Guyana. ◆Fig. 441.

Abolboda neblinae Maguire, Mem. New York Bot. Gard. 17: 80. 1967. Stout-rhizomatous perennial, strongly tinged with maroon, red, or purple; rosettes in dense domes; leaves linear, to 6 cm long with subulate-spinulose tips; scapes 1 per rosette, to 30 cm tall, with 3 or more pairs of erect bracts to 2 cm long; spikes narrow, 1flowered; corolla lobes large, bright deep blue. Rocky, wet tepui meadows, 1900–2300 m; Amazonas (Sierra de la Neblina). Adjacent Brazil. ◆Fig. 443. Abolboda paniculata Maguire, Mem. New York Bot. Gard. 10(1): 10, fig. 2a–g. 1958. Cespitose, smooth, densely rosulate perennial from a stout rhizome, strongly tinged with maroon, red, or purple; leaves to 10 cm long, 2–3 mm wide; scapes to 1.2 m tall; spikes 1-flowered, narrow, in panicles. High, wet, rocky savannas, edges of gallery forests, 1200–2000 m; Bolívar (Cerro Jaua), Amazonas (Cerro Parú). High-elevation Amazonian Brazil. Abolboda pulchella Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 110. 1813. Xyris vaginata Spreng., Syst. Veg. 1: 183. 1825 [1824]. Abolboda brasiliensis Kunth, Enum. Pl. 4: 26. 1843. Abolboda gracilis Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 323. 1909. Abolboda chapadensis var. pauciflora Hoehne, Com. Lin. Telegr., Bot. 5: 12. 1915. Slender, smooth perennial; rosettes solitary or clumped; leaves 1–14 cm long, mostly linear-triangular, the tips often 3-angled, often narrowing into a spine; scapes to 50 cm tall, with 1 pair of bracts to 2 cm long; spikes ellipsoid to oblanceoloid, few-flowered; corollas blue. Wet, sandy savannas, 100–1300 m; frequent in Bolívar and Amazonas. Amazonian Colombia, Guyana, Suriname, French Guiana, Brazil (Amazonian, Mato Grosso, Minas Gerais). Abolboda scabrida Kral, Ann. Missouri Bot. Gard. 76: 975. 1989. Low, densely cespitose perennial; leaves and scapes scabrid and/or scabrid-rugose; leaves narrow, excurved, to 6 cm long; scapes to 10 cm tall, with 1 pair of bracts, these 2/3

Abolboda 533

Fig. 440. Abolboda killipii

Fig. 442. Abolboda linearifolia

Fig. 441. Abolboda macrostachya var. robustior

Fig. 443. Abolboda neblinae

534

X YRIDACEAE

up scape; spikes 1.1–1.4 cm long, turbinate. Sandy tepui meadows, ca. 1400 m; Amazonas (summit of Cerro Aracamuni). Endemic. Abolboda sprucei Malme, Ark. Bot. 25A(12): 17. 1933. Abolboda schultesii Idrobo & L.B. Sm., Caldasia 6: 246. 1954. Cespitose, rosulate but few-leaved perennial, 20–35 cm tall; leaves shorter or slightly longer than scapes; blades to 8 mm wide, at apex subulate-angulate, entire, the scape bracts 1 pair or 3 pairs, the lowest set to 2 cm below spike or directly under it and forming

a leafy involucre; spikes mostly ovoid, 2 cm long, pale green; corolla lobes large, pale blue-purple. Sandy savannas, 100–200 m; Amazonas (near Maroa). Adjacent Colombia. Abolboda uniflora Maguire, Mem. New York Bot. Gard. 10(1): 12. 1958. Very similar to Abolboda neblinae, but leaves shorter, not over 3 cm long, the blades narrower, 3-angled, tips short-conic, apiculate; scapes often with an additional penultimate sessile spike. Tepui summits, 1700– 2000 m; Amazonas (Cerro Duida). Endemic.

2. ACHLYPHILA Maguire & Wurdack, Mem. New York Bot. Gard. 10(2): 11, fig. 1a–m. 1960. Stiff perennial herbs with diffuse-fibrous roots and shallow, horizontal, frequently forking, elongate rhizomes, the shoots 1–several, stiffly erect, in cross section somewhat elliptic, the edges papillate-costate. Leaves alternate, equitant, distichous, suberect, cloaking stem, the lower ones shortest, scale-like, strongly veined, progressively lengthening up the stem, the principal ones lance-linear, green with tan sheaths about as long as blades and open, broadest at the clasping base and conduplicately ligulate at junction with blade; blades compressed but strongly folded and keeled, tapering gradually from base to narrowly acute, spinulose tip, the margins slightly thickened. Inflorescence solitary and terminal on an ancipitally flattened scape, in outline narrowly obtriangular; bracteal leaves distichous, spathiform, similar to but smaller than foliage leaves, the lowest longest, erect, continuous with scape; inflorescence axis zig-zag, ascending, the internodes clavate to 3-angled, usually 2, the lowest longest, each bearing a secondary spathe. Flowers paired on a short joint in each spathe axil, subtended by a pair of lancesubulate prophylls; pedicels unequal, stiff, 3-angled, erect or excurved, or the uppermost spathe 3-flowered with flowers exceeding or overtopped by the primary spathe. Sepals 3, lanciform, subulate, accrescent, the anterior one more spreading than the laterals; petals 3, distinct, equal, nearly sessile, the blades ovate, obtuse, unequally spreading between the more erect sepals, ca. 10 mm long, yellow. Stamens with filaments 3.5–4 mm long, opposite the petals and shorter, bases dilated; anthers oblong, 2–2.5 mm long, sagittate, dithecal. Ovary columnar, trigonous, the placentation axile; style elongate, erect, stout-based, gradually tapering to a capitate-trilobed stigma. Seeds broadly ovoid or ellipsoid, 1.5–2 mm long, minutely longitudinally striate, translucent. Venezuela, endemic to the flora area; 1 species. Achlyphila disticha Maguire & Wurdack, Mem. New York Bot. Gard. 10(2): 12, fig. 1a–m. 1960. Rhizomes to 3 mm thick, shoots 24–40 cm tall, bases decumbent; principal leaves 6–13 cm long, with sheaths ca. as long as blades, the blade keeled, its tip with a spinulose arista to 4 mm long, margins thickened, yel-

lowish, tuberculate-scabrid, the surfaces papillose-rugulose; scape to 15 cm long, 3–4 mm wide; inflorescence to 8 cm long; sepals 1–1.5 cm long. Tepui near-summit rocky shrub bogs, 1700–2300 m; Amazonas (Sierra de la Neblina). Endemic (probably present on Brazilian side of the border as well). ◆Fig. 444.

Aratitiyopea 535

Fig. 444. Achlyphila disticha

Fig. 445. Aratitiyopea lopezii var. lopezii

536

X YRIDACEAE

3. ARATITIYOPEA Steyerm. & P.E. Berry, Ann. Missouri Bot. Gard. 71: 297, fig. 1A–K. 1984. Stout perennial herbs, the elongate stems from elongate thickened repent bases, subhorizontal, densely leafy, erect only at tips. Leaves in close-set spirals, the ones downstem progressively deciduous; blades flat, elliptic-linear, narrowly acute and slenderly attenuate to inflated, broadly clasping, papery, open sheaths, those toward and beneath the inflorescence shorter, denser, linear-triangular, spreading, ligulate, and lingulate. Inflorescence thus a leafy-involucrate, large, terminal, sessile, hemispheric, multiflorous head. Involucral bracts numerous around head, erect, linear-triangular, papery, and strongly tinged with maroon, red, or purple, the inner somewhat shorter and narrower; receptacular bracts grading smaller, similarly erect. Sepals 3, narrowly triangular-linear, the laterals strongly keeled, the anterior one merely convex, all strongly tinged with maroon, red, or purple, and papery; corolla 3-lobed, exserted beyond sepals, narrowly salverform, purple, the lobes much shorter than the slender tube, ascending, blunt. Stamens 3, epipetalous, opposite the corolla lobes at their base, below level of corolla sinus; filaments slender, slightly longer than the erect, basifixed, dithecal, golden, narrow anthers; staminodia none. Ovary 3-locular, the placentation axile; style exserted, strongly tinged with maroon, red, or purple, filiform; stigma lobes 3, ellipsoid, divaricate, hairy; stylar glands at base of style, 3 suberect on 3 stipes, bulbous with reflexed, plane, rectangular, erose appendages. Capsule valve texture uniform. Seeds numerous, ovoid-ellipsoid, ca. 1 mm long, uniformly low-ribbed longitudinally, each interval with a row of pits. Colombia, Venezuela, Brazil; 1 species. Aratitiyopea lopezii (L.B. Sm.) Steyerm. & P.E. Berry, Ann. Missouri Bot. Gard. 71: 297. 1984. —Navia lopezii L.B. Sm., Bot. Mus. Leafl. 15: 40. 1951. Colombia, Venezuela, Brazil; 2 varieties, 1 in Venezuela. A. lopezii var. lopezii Stem to 2 m long, to 2.5 cm thick; leaves rich green, the principal lower ones 21–23 × 3.5–4 cm, those around heads 10–14 × 2.5–3 cm; outer involucral bracts 3.2–3.3 × 0.6–1.1

cm; sepals 4.5–5.5(–6) cm long; corolla with tube 5–6 cm long and lobes 1.5–2.2 cm long, 3–4 mm wide; stamens with filaments 8–12 mm long; anthers 5–10 mm long; style 8 cm long, with appendages 3–4.5 mm long. Hanging on wet cliffs, in tepui scrub on granite outcrops, meadows on granite, in waterfall spray, 600–1900 m; Amazonas (Cerro Aratitiyope, Cerro Avispa, Cerro Sipapo, Río Siapa at Raudal Gallineta, Serranía Uasadi). Brazil. ◆Fig. 445.

4. ORECTANTHE Maguire, Mem. New York Bot. Gard. 10(1): 2. 1958. Coarse, smooth perennial herbs from a caudiciform rootstock, fleshy-rooted; stems densely leafy, stout, either contracted, erect, and simple, producing an agavelike rosette, to much-elongated and erect or sprawling, branching, the lower parts cloaked by an imbricate stubble of old leaf bases. Leaves polystichous, lax or stiff, their bases broadly clasping, spirally imbricate, many in flat spirals (rosettes) or tall spirals; blades gladiate, lance-linear, or triangular, spreading to ascending, mostly rigid, plane or lingulate, tips sharply acute or pungent; margins firm, thin (later toothed) and sharp. Inflorescence solitary, terminal on 1 or more essentially (except at very base) bractless, terete, fistulose, variously elongated scapes, capitate, large, globose to hemispheric, 5 or more cm thick, of many firm or chaffy bracts and sepals, the involucral bracts ovate-triangular or lance-triangular, in 1 or 2 se-

Orectanthe 537

Fig. 446. Orectanthe ptaritepuiana

Fig. 447. Orectanthe sceptrum

ries of 3. Flowers 3-merous and irregular, each subtended by a short, triangular bract abaxially. Sepals 3, subequal, lanceolate, 3–5 cm long or more, variously longer than the involucral bracts, the lateral ones sharply folded, narrowly acute, and strongly winged-keeled, the anterior sepal plane or slightly convex, without keel, similarly acute; corolla irregular, yellow, rarely strongly tinged with maroon, red, or purple, malodorous, spreading-excurved, gamopetalous, to 8 cm long, the lobes slightly longer than the tube, broadly ovate, the upper lobe largest and somewhat hooded, the lateral lobes more spread, somewhat declinate. Stamens epipetalous, opposite corolla lobes at summit of corolla tube, shorter than the corolla lobes, ascending-spreading; filaments slender, slightly longer than the narrow,

538

X YRIDACEAE

basifixed, 4-locular, introrse anthers, the connective slightly produced beyond the anther tips. Ovary broad, 3-locular, the placentation axile; style slender, projecting outward from the upper corolla lobe but not exsert; stigma broadly and obliquely cupuliform, multifimbriate. Capsule ovoid to obovoid, the valves thick, stiff, somewhat more thickened apically. Seeds many, irregularly curvate-triangular, ca. 2–3 mm long, compressed around the seed cavity to form a broad wing, 1 edge forming a narrow, recurved lobe, the whole thus with a “mitten” outline, the entire surface finely curved-striate and lustrous. Endemic to the Guayana Highlands in Venezuela, Guyana, and Brazil; 2 species, both in the flora area. Key to the Species of Orectanthe 1.

1.

Stems elongate, sprawling, often to 1.5 m or more, densely leafy but the leaves in tall spirals, usually lacking a strong red-brown border or this paler and less distinct; involucral bracts usually in 2 series of 3 ........................................................................................... O. ptaritepuiana Stems mostly erect or ascending, the leaves usually in a dense cluster at or toward the plant base; scapes usually solitary, the leaves with a narrow but strong red-brown border; involucral bracts usually a single series of 3 bracts ........................................................................... O. sceptrum

Orectanthe ptaritepuiana (Steyerm.) Maguire, Mem. New York Bot. Gard. 10(1): 5. 1958. —Abolboda ptaritepuiana Steyerm., Fieldiana, Bot. 28: 104. 1951. Stem elongate, often to 1.5 m or more, densely leafy, but the leaves in tall spirals, the growth often sprawling, erect only at tips. Forests on tepui slopes, tepui summits, 1500–2500 m; Bolívar (Cerro Guaiquinima, Macizo del Chimantá, Ptari-tepui). Adjacent high-altitude Guyana and Brazil. ◆Fig. 446. Orectanthe sceptrum (Oliv.) Maguire, Mem. New York Bot. Gard. 10(1): 3. 1958. —Abolboda sceptrum Oliv., Trans. Linn. Soc. London, Bot. 2: 286. 1887. Orectanthe sceptrum subsp. occidentalis

Maguire, Mem. New York Bot. Gard. 10(1): 5. 1958. Stem stouter than in the preceding, more erect, more caudiciform; rosettes more compact, of flatter spirals of stiffer, more-spreading blades; principal leaves (10–)15–30 × 1.5–4.5 cm; scapes in larger individuals to 1.2 m tall or more, stouter. Boggy tepui summits, high slopes, rapateaceous or shrub bogs, 500–2800 m; frequent in Bolívar and Amazonas. Adjacent western Guyana and northern Brazil. ◆Fig. 447. Leaves of populations in the eastern range tend to be glaucous, and heads there tend to have longer involucral bracts than is true for populations in the central and western parts of the total range, but such characters vary independently.

5. XYRIS L., Sp. Pl. 41. 1753; Gen. Pl. ed. 5, 25. 1754. Kotsjiletti Adans., Fam. Pl. 2: 60. 544. 1763. Xuris Adans., Fam. Pl. 2: 20. 1763. Jupica Raf., Fl. Tellur. 2: 15. 1836 [1837]. Ramotha Raf., Fl. Tellur. 2: 15. 1836 [1837]. Schismaxon Steud., Bot. Zeitung (Berlin) 14: 391. 1856. Annual or perennial, shortly to prominently caulescent, usually rosulate, scapose herbs with diffuse-fibrous roots, usually perennating by lateral offshoot buds. Axis sympodial. Foliage externally smooth to variously papillose, roughened, or pilose; outer leaves often scale-like; principal leaves mainly linear, equitant, disti-

Xyris 539

chous, the sheathing bases open, usually strongly clasping and dilated distally, narrowing gradually or abruptly to blade, where ligulate or eligulate, the sheath edges convergent to form one side, the midrib area the other, of a laterally flattened to terete, angulate, or sulcate, usually linear, entire to ciliate or scabrid blade. Scapes single or few to a shoot, linear, terete to variously compressed, costate or ancipital, mostly overtopping leaves, each subtended by a sheath leaf, this closed and involute-tubular proximally, open and variously bladed to bladeless distally. Inflorescence a terminal, cone-like head or spike of tightly to loosely spirally or distichously imbricate, chaffy bracts, the lowest usually sterile, smaller than the fertile ones to much exceeding them, even foliaceous, the fertile ones each with a sessile to subsessile axillary flower, sometimes the uppermost again sterile. Sepals unequal, the 2 lateral opposing, connivent around the floral base, distinct or variously connate, chaffy, boat-shaped, variously keeled, equilateral or not, the anterior (inner) sepal scarious and enclosing the corolla in bud, fugacious; petals free or rarely connate at base, subequal, long-clawed, the blades usually broad, yellow to orange-yellow or white, spreading. Stamens equal; filaments short, fleshy, adnate just above the petal claw; anthers basifixed, 2-locular, the pollen monosulcate; staminodia 3 (rarely lacking), subequal, mostly 2-armed apically, the branches penicillate with moniliform hairs, less often smooth. Ovary 1-locular to imperfectly or perfectly 3locular; ovules (1–)many, the placentae marginal, parietal, axile, basal, or free-central; style terminal, slender, tubular, strongly 3-branched above the middle, each branch terminating in a truncate, U-shaped or funnelform stigma fringed with glandular hairs. Capsule usually thin-walled, 3-valved, dehiscing along 3 lines alternate with placentae. Seeds mostly small, translucent or farinose, with copious mealy, starchy endosperm; embryo small, basal-lateral; seed coat mostly longitudinally ribbed and cross-lined. Mostly pantropics, some species extending into temperate North America and Asia; ca. 200 species, 79 in Venezuela, 75 of these in the flora area. Key to the Species of Xyris 1. 1.

Placentation evidently marginal or parietal, the capsule valves on dehiscence retaining placentae and seeds .............................................. Group I Placentation free-central, basal, or apparently axile, the capsule valves on dehiscence with or without septa, but not retaining seeds .... Group II Group I of Xyris

1.

1.

2(1).

2.

Seeds fully 1 mm long; scape distally terete with (usually) 3 or more strong costas, the bracts with conspicuous midrib on dorsal area, usually accompanied by several less conspicuous lateral veins ........ X. fallax Seeds 0.4–0.8(–0.9) mm long; scape rarely with more than 2 strong costas, the bracts with midrib on dorsal area not as evident and lateral veins lacking .......................................................................................... 2 Plant base strongly tinged with red or purple; scapes ecostate distally or with costas smooth; seeds mostly opaque and farinous, dark-ribbed, 0.7–0.9 mm long; robust perennial ........................................... X. laxifolia Plant base not strongly tinged with red or purple; scapes either with 1 or 2 strong costas distally (rarely with a few lower ones), these usually

540

3(2).

3.

X YRIDACEAE

papillose or scaberulous, or the plants low, delicate, rhizomatous and with scapes terete, striate; seeds amber, translucent, 0.4–0.7 mm long; smaller annuals or delicate rhizomatous perennials ........................... 3 Plants low, delicate, mat-forming perennials, some stems forming slender, zig-zag, repent rhizomes; bases of leaf sheaths long-ciliate ................................................................................................ X. prolificans Plants erect, not delicate or mat-forming, annual, erhizomatous; bases of leaf sheaths not long-ciliate ........................................................ X. jupicai Group II of Xyris

1.

Margins of leaf sheaths at some or all levels ciliate with various sorts of hairs ....................................................................................................... 2 1. Margins of leaf sheaths glabrous, entire ................................................ 34 2(1). Dorsal area not evident on some or all spike bracts ................................ 3 2. Dorsal area discernible on some or all spike bracts ............................... 14 3(2). Leaves definitely thickened at junction of sheath and blade, not 2-edged ................................................................................................................ 4 3. Leaves distinctly compressed at junction of sheath and blade, with the ventral edges about as narrow as the dorsal edge ............................... 5 4(3). Tips of leaf blades blunt; scapes rugose-tuberculate or rugulose-papillate ................................................................................................. X. delicatula 4. Tips of leaf blades subulate-spinulose; scapes totally smooth, or smooth and puncticulate ......................................................................... X. setigera 5(3). Scapes terete distally, ecostate, smooth and also punctate; leaf apex usually excentrically spinulose-tipped ....................................................... 6 5. Scapes terete or somewhat flattened distally, there costate, usually tuberculate-scabrid, ciliate, papillose and/or rugulose at least on the costas; leaf apex not excentrically spinulose-subulate ........................ 7 6(5). Leaf blades 2–3 mm wide; lateral sepals 6.5–7 mm long; seeds 0.9–1 mm long ............................................................................................... X. riparia 6. Leaf blades 0.5–3 mm wide; lateral sepals 5–6 mm long; seeds 0.5– 0.6 mm long ................................................................................. X. setigera 7(5). Bracts very dark, usually deep olive-brown, the borders entire and not much contrasting in color ...................................................................... 8 7. Bracts with scarious and lacerate borders differing abruptly in texture and color from main body of bract ...................................................... 10 8(7). Surfaces of leaves and scapes smooth or at most papillate only toward leaf sheath base, but edges of leaf blades and of ancipital scape white-ciliolate; lateral sepals free; capsule valves with septa ............................................................................................ X. kukenaniana 8. Surfaces of leaves and scapes completely rugulose-papillose-tuberculate; lateral sepals free or connate; capsule valves with or without septa ................................................................................................................ 9 9(8). Lateral sepals connate; capsule valves lacking septa ................ X. consolida 9. Lateral sepals all free; capsule valves with strong septa ......... X. schneeana 10(7). Borders of bracts red or red-brown ......................................................... 11

Xyris 541

10. Borders of bracts pale .............................................................................. 12 11(10). Spikes ellipsoid to broadly cylindric; foliage above dilated base yellowgreen, scabrid-papillose-rugulose; scapes ancipital distally ...... X. roraimae 11. Spikes broadly obovoid or turbinate or subglobose; foliage except for leaf edges and costas not yellow-green, smooth; scapes not ancipital ............................................................................................. X. thysanolepis 12(10). Apices of bracts, particularly basal and inner ones, acute, folded; spikes ovoid .......................................................................................... X. concinna 12. Apices of bracts mostly obtuse, not folded; spikes broadly ellipsoid, obovoid, or subglobose ............................................................................... 13 13(12). Lowest bracts > 1/2 as long as spikes; seeds 0.6–0.7 mm long; keel of lateral sepals of nearly uniform width from base to apex ..... X. hymenachne 13. Lowest bracts < 1/2 as long as spikes; seeds ca. 1 mm long; keel of lateral sepals distinctly broadened distally ................................. X. gongylospica 14(2). Leaves at junction of sheath and blade definitely not flattened, at most elliptic in cross section, mostly terete or oval, often with a ventral sulcus ......................................................................................................... 15 14. Leaves at junction of sheath and blade definitely flattened .................. 23 15(14). Spikes few-flowered and narrow, narrowly oblong, or linear-lanceoloid .............................................................................................................. 16 15. Spikes several- to many-flowered and of a broader outline ................... 17 16(15). One pair of lower (sterile) bracts longer than the rest, the tips connivent or nearly so over spike top .................................................... X. cryptantha 16. Lower bracts not as above, the lowest pair slightly to considerably shorter than the spike ................................................................ X. oxylepis 17(15). Bract tips narrowly or bluntly acute ....................................................... 18 17. Bract tips broadly angled to rounded ..................................................... 19 18(17). Outer bracts mostly smooth not hirsute-tomentose or hirsute-ciliate; bract tips becoming excurved, bract matrix brown .................. X. frequens 18. Outer bracts densely hirsute-tomentose or hirsute-ciliate with white hairs; bract tips straight, erect or ascending, bract matrix dark castaneous ............................................................................... X. wurdackii 19(17). Dorsal area triangular, often broadly so, comprising most of the bract above the middle; lower bracts of spike with some stiff hairs at tips; leaf blades strongly ribbed and with a strong ventral sulcus ........... 20 19. Dorsal area narrower; lower bracts of spike lacking stiff hairs at tips; leaf blades not strongly ribbed ................................................................... 21 20(19). Leaf blades with broad, pale, villous-tomentose borders; scapes densely white-tomentose ................................................................. X. aracamunae 20. Leaf blades smooth except for trichomiferous tips; scapes not hairy .............................................................................................. X. subglabrata 21(19). Scapes papillate or tuberculate, usually rugulose or rugose, at least above the middle ........................................................................ X. carinata 21. Scapes smooth, usually lustrous, at most punctate ............................... 22 22(21). Bracts pale brown with darker dorsal areas ............................. X. lithophila 22. Bracts dark brown or dark red-brown, dorsal areas paler but inconspicuous ............................................................................................... X. setigera 23(14). Scapes either strongly flattened distally or with 2 costas making 2 strong

542

X YRIDACEAE

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(30).

32.

33(32).

33. 34(1).

ciliate or hairy edges ........................................................................... 24 Scapes in cross section distally terete or at least thickened, 0–many-costate but costas in bicostate types not ciliate or scabrociliate, not making 2 strong edges ................................................................................ 26 Leaf blade and scape edges scabrociliate with shorter, stiffer hairs ................................................................................................... X. bicostata Leaf blade edges and/or scape edges usually with long, straight cilia, rarely smooth, if smooth then thickened ............................................ 25 Spikes narrow; dorsal area of bract narrower; scape narrow, < 2 mm wide ................................................................................................... X. pratensis Spikes broad; dorsal area broadly triangular, occupying most of upper part of bract; scapes mostly ≥ 2 mm wide ......................... X. surinamensis Scapes with strong costas ........................................................................ 27 Scapes smooth, without strong costas or merely fluted ......................... 28 Scapes with 3 or more, sharp, smooth or low-papillate costas; plant base sub-bulbous, the plants short-lived perennials with ascending leaves; spike broadly ovoid or ellipsoid ................................ X. malmeana Scapes with costas fewer, if multicostate the costas papillate-scabrid; plant base not bulbous, the plants annual with flabellately spreading leaves; spike ellipsoid to lanceoloid ............................................. X. tenella Flowers usually ≤ 4; spikes mostly lanceoloid or narrowly ellipsoid, the bracts lacking dorsal areas or with narrow, lanceolate, or streak-like dorsal areas .......................................................................................... 29 Flowers usually > 4; spikes usually ovoid to subglobose, the bracts with dorsal areas (except in X. malmeana) of broad outline ..................... 30 Sheath cilia cowbeb-like; seeds ellipsoid or cylindric, ca. 1 mm long; leaf blades filiform ............................................................................ X. byssacea Sheath cilia long and spreading but firm; seeds ovoid, ca. 0.5 mm long; leaf blades linear .......................................................................... X. tenella Bracts, or some of them, with long hairs on margins or backs or both .............................................................................................................. 31 Bracts without pubescent margins or backs ........................................... 32 Bracts acute, some bract backs as well as edges with white hairs ............................................................................................. X. arachnoidea Bracts rounded, edges villous-ciliate ............................................. X. globosa Matrix of bracts dark brown, the spikes thus dark; scapes lustrous and often reddish or olivaceous; leaf tip excentrically spinose or spinulose ..................................................................................................... X. setigera Matrix of bracts (at level of dorsal areas) pale brown or pale red-brown, thin, lacerate or entire; scapes greenish, dull; leaf tips not spinose or spinulose .............................................................................................. 33 Leaf sheath bases abruptly dilated, the plant base thus bulbous; spike broadly ovoid or broadly ellipsoid, not proliferating; leaves erect or ascending ................................................................................... X. lacerata Leaf sheath bases not abruptly dilated, the plant base not bulbous; spike oblate, often proliferous; leaves flabellately spreading .............. X. oblata Plant aquatic, short-lived perennial, or low and slender-scaped annual with small or narrow spikes; leaf sheaths mostly softer, paler, their

Xyris 543

34.

35(34).

35. 36(35).

36.

37(35). 37. 38(37). 38. 39(38). 39. 40(38). 40. 41(40).

41.

42(41). 42. 43(42). 43. 44(43).

44. 45(37).

bases not sharply contrasting with blades in color or texture .......... 35 Plant terrestrial, mostly densely cespitose and perennial; leaf sheaths hard, lustrous, mainly castaneous to reddish brown and usually strongly contrasting with the blades both in color and texture ........ 52 Scapes ancipital, edges at least in combination broader than the scape body; dorsal area and midrib of at least one of the lowest sterile bracts excurrent as a strong cusp or blade .................................................... 36 Scapes flattened or terete, but not winged as above; dorsal area and midrib of sterile bracts not excurrent or scarsely so ................................ 37 Basal bracts foliaceous, slightly to much longer than the fertile ones, sharply winged-keeled, spreading or ascending and often forming an involucre; sepals acute ........................................................... X. spruceana Basal bracts tending to be incurved, cucullate, the lowest slightly to much longer than the fertile ones, thus spikes not as noticeably involucrate and spreading-foliaceous; sepals obtuse ............................ X. uleana Florets ≤ 4 ................................................................................................. 38 Florets 4–many ........................................................................................ 45 Lateral sepals connate; fertile florets 1 per spike .................................. 39 Laterals sepals free; fertile florets > 1 per spike .................................... 40 Leaf blades with edges cartilaginous-thickened; scapes flattened distally with costas making edges ..................................................... X. esmeraldae Leaf blades filiform, the edges not noticeably cartilaginous thickened; scapes filiform-terete, ecostate ............................................. X. subuniflora Leaf blades strongly flattened, with narrow, pale, thickened borders ................................................................................................ X. guianensis Leaf blades terete or somewhat flattened, without pale thickened borders .............................................................................................................. 41 Leaf sheath apically with a broad ligule, this narrowing abruptly to a terete blade; spikes frequently proliferous; aquatic or emergent plants with foliage often maroon and soft ........................................ X. spathacea Leaf sheath borders narrowing gradually to a more flattened, often strongly ribbed blade, eligulate or with a narrow ligule; spikes not proliferous; plants not aquatic ............................................................ 42 Mature scapes equalled or exceeded by larger leaves; bracts at maturity strongly excurved, thus spikes broadly turbinate ................. X. cyperoides Mature scapes definitely longer than the leaves; bracts and spikes not as above .................................................................................................... 43 Seeds ca. 1 mm long, including an apical, pale coma of small scales or bristles; scapes and leaf blades rugose or rugulose ..................... X. mima Seeds 0.4–0.7 mm long, lacking apical small scales; scapes and leaf blades smooth or nearly so .................................................................. 44 Lateral sepals broad, ciliate or ciliolate, the apex more broadly acute; bract tips more broadly acute, not apiculate, dorsal area triangular, not short-excurrent ................................................................ X. guianensis Lateral sepals narrow, acute, entire; bract tips narrowly acute or apiculate, dorsal areas linear, often short-excurrent ..................... X. toronoana Leaf blades filiform and flaccid; sheath bases broadly scarious-bordered, the scarious borders distally producing a strong ligule much broader

544

X YRIDACEAE

45. 46(45). 46. 47(46). 47. 48(46). 48. 49(48). 49. 50(48).

50.

51(50). 51. 52(34).

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

55(53). 55. 56(55). 56. 57(52). 57. 58(57). 58. 59(58).

than the base of leaf blade; rhizomatous, soft aquatic ........... X. aquatica Leaf blades wider and/or firmer; sheath at apex not as above; plants without rhizomes, of drier sites .................................................................. 46 Spikes elongate, > 4 times as long as wide ............................................. 47 Spikes of a broader, shorter outline, < 4 times as long as wide ............. 48 Bracts polystichous, spikes terete ............................................... X. paraensis Bracts distichous, spikes flattened ........................................ X. stenostachya Lateral sepals subequilateral or slightly inequilateral ......................... 49 Lateral sepals either connate or very inequilateral ............................... 50 Leaf blades lacking strong red border; spikes broadest at or toward base; seeds with an apical coma of scales .............................................. X. mima Leaf blades with a strong red border; spikes broadest at or above middle; seeds lacking an apical coma of scales .............................. X. rubrolimbata Staminodia lacking beard; foliage prevalently (one var. excepted) papillose or rugulose, the leaf blades strongly veined; dorsal areas of fertile bracts mostly narrowly elliptic or linear; seed tips truncate, with a central low apiculus .................................................................... X. savanensis Staminodia bearded; foliage prevalently smooth, the leaf blades either less strongly veined or smooth; dorsal areas of fertile bracts broader; seed tips not truncate .......................................................................... 51 Backs and edges of bracts smooth and entire ............................. X. paraensis Backs of bracts strongly papillose-tuberculate; bract edges at least apically villous or pilose-ciliate .................................................... X. uleana Leaf blades for most of their length either terete or very thickened and deeply ribbed and sulcate, not flattened or 2-edged; ligules prominent, abruptly narrowed to blades ............................................................... 53 Leaf blades flattened, for most of their length 2-edged, the edges either thickened or sharp; ligules evident or not .......................................... 57 Dorsal area of bract imperceptible .......................................................... 54 Dorsal area of bract distinguishable ....................................................... 55 Leaf blades not deeply ribbed or sulcate; sheath bases (usually tips also) nearly black ................................................................................. X. atriceps Leaf blades and upper sheaths coarsely ribbed and sulcate; sheath bases brown (bases of blades and apices of sheaths mostly rose-purple) ............................................................................................... X. sulcatifolia Plant bases nearly black ................................................................ X. neblinae Plant bases deep brown, pale brown, or reddish brown ......................... 56 Mature spikes mostly > 1.5 cm long, fusiform or distinctly broadest above middle; leaf blades ≥ 1 mm thick ............................................ X. juncifolia Mature spikes mostly < 1.5 cm long, ovoid to cylindric or subglobose; leaf blades < 1 mm thick ......................................................... X. lanulobractea Bracts with dorsal area evident .............................................................. 58 Bracts with dorsal area not evident ........................................................ 67 Scapes strongly flattened, at least 2–2.5 mm wide distally, ancipital .............................................................................................................. 59 Scapes distally round or if strongly flattened distally < 2 mm wide there .............................................................................................................. 61 Several lower bracts leaf-like, forming an involucre .............. X. involucrata

Xyris 545

59. 60(59). 60. 61(58). 61. 62(61).

62. 63(61). 63. 64(63).

64. 65(64).

65. 66(64). 66. 67(57). 67. 68(67). 68. 69(68).

69. 70(69). 70. 71(70).

71.

No bracts leaf-like .................................................................................... 60 Scapes 2-spicate; seeds < 2 mm long ........................................... X. bicephala Scapes 1-spicate; seeds > 2 mm long ....................................... X. teinosperma Lowermost sterile bracts distinctly shorter than the fertile ones and grading gradually into them ............................................................... 62 Lowermost sterile bracts as long as or longer than the fertile ones ...... 63 Leaf blade edge not cartilaginous-bordered, smooth; bract borders not pale villosulous, sometimes sparsely short-brown-cililoate; uplands, Sierra de la Neblina ................................................................ X. contracta Leaf blade edge cartilaginous-thickened, entire or cililoate; bract borders pale-villosulous; low-elevation savanna plants ................ X. lomatophylla Lateral sepals connate, keel apex red-villous; plants glaucous; dorsal area of lowest 1 or 2 bracts often excurrent as a green blade .......... X. seubertii Lateral sepals free, not crested as above; plants not glaucous; dorsal area of bracts bladeless ................................................................................ 64 Dorsal area of bracts large, triangular, punctate, occupying most of bract above middle; low-elevation savannas or granite outcrops, Amazonas .............................................................................................................. 65 Dorsal area of bracts narrower, less conspicuous, not evidently punctate; high savannas of tepuis ....................................................................... 66 Bract borders not reddish, eciliate; keel of lateral sepals sparsely scabrid and leaf blade edges scabrociliate only toward base; seeds ca. 0.5 mm long ......................................................................................... X. graniticola Bract borders reddish, long-ciliate; keel of lateral sepals and leaf blade edges ciliate; seeds 1.3–1.4 mm long ........................................... X. huberi Scape edges thick and entire; leaf blades evenly ciliate to tip ... X. frondosa Scape edges thin, comprised of 2 flat, broad, and distinctly pilose-ciliate costas ............................................................................................ X. liesneri Longest leaves rarely with sheaths over 1/2 as long as blades, mostly 1/3 as long or shorter ...................................................................................... 68 Longest leaves commonly with sheaths over 1/2 as long as blades, frequently 2/3 as long or longer ................................................................. 75 Leaf blades < 3 mm wide ......................................................................... 69 Leaf blades mostly > 3 mm wide ............................................................. 72 Keel of lateral sepals red-villous-ciliate toward apex; edges of young bracts red-villose toward apex; Bolívar state (Macizo del Chimantá) ................................................................................................. X. chimantae Keel of lateral sepals ciliate or lacerate but not as above; bract tips entire or brown-ciliolate; Amazonas state ..................................................... 70 Tips of leaf blades acute; leaf blades not coarsely ribbed; seeds ca. 2 mm long .................................................................................. X. valdeapiculata Tips of leaf blades strongly callused, thick, blunt; leaf blades coarsely few-ribbed; seeds < 2 mm long ............................................................ 71 Foliage harsh, particularly leaf blades toward base; leaf sheath apex and blade bases strongly tinged with red or rose; sheath edges gradually narrowing to an eligulate tip, merging with blade base; keel of lateral sepals rusty-ciliolate ............................................................. X. sulcatifolia Foliage smooth; leaf sheath apex tan to straw-colored, lustrous; sheath

546

X YRIDACEAE

72(68). 72. 73(72).

73.

74(72). 74. 75(67).

75.

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

77.

78(75).

78. 79(78). 79. 80(79). 80. 81(78). 81. 82(81). 82.

edge forming a strong, broad ligule apically; keel of lateral sepals narrow, entire or subentire .................................................. X. stenophylloides Scapes ancipital distally, mostly ≥ 2.5 mm wide, variously scabrid, ciliate, or smooth .............................................................................................. 73 Scapes not sharp-edged distally, mostly < 2.5 mm wide, smooth or punctate ....................................................................................................... 74 Spikes ovoid-cylindric, distinctly longer than wide; scape edges smooth to pale-ascending-ciliate; bracts dull, dark greenish black; Bolívar state (Macizo del Chimantá) ................................................. X. melanovaginata Spikes obovoid or hemispheric, ca. as wide as long or wider; scape edges scabrid; bracts dark, lustrous brown; Amazonas state (Cerro Duida) .......................................................................................................... X. tatei Leaf blades with a rusty border, ciliate (often with reddish vein intervals) .............................................................................................. X. culmenicola Leaf blades without a rusty border, eciliate ................................. X. lugubris Bracts toward apex with a scarious, lacerate, pale or reddish border; bases of leaf sheaths abruptly orbicular-dilated; leaf-bearing portion of stem mostly contracted, ≤ 3 cm long ................................................... 76 Bracts not bordered as above; bases of leaf sheaths not abruptly oribicular-dilated; leaf-bearing portion of stem mostly elongated, the leaves thus often forming conspicuous, frond-like plates .................. 78 Scarious bract borders pale to deep red or red-brown; seed ca. 1 mm long ............................................................................................. X. thysanolepis Scarious bract borders almost always pale, off-white; seed 0.5–0.7 mm long ....................................................................................................... 77 Apices of most bracts acute and strongly keeled; stem bases cloaked with very numerous, closely overlapping leaf sheath bases, these a lustrous brown; summit elevations ........................................................ X. concinna Apices of most bracts more rounded, often broadly so, not distinctly folded; stem bases shorter, paler, duller, less closely overlapping; widely distributed from medium to high elevations ......... X. hymenachne Scapes ancipital or not, but never ciliate, at most scabridulous along edges; stems forming frond-like plates of leaves (like giant Fissidens) .............................................................................................................. 79 Scapes ancipital and strongly ciliate distally ......................................... 81 Spikes mostly 2–2.5 cm long, ellipsoid-cylindric or narrowly obovoid; lateral sepals > 1 cm long, narrowly acute ............................. X. xiphophylla Spikes < 1.5 cm long, ovoid or broadly obovoid; lateral sepals < 1 cm long, strongly curved, blunt ......................................................................... 80 Ligular apex often excurved; scapes strongly flattened distally, mostly ≥ 2.5 mm wide; leaf blades commonly > 4 mm wide ............... X. ptariana Ligular apex commonly erect or ascending; scapes narrower, usually ≤ 2 mm wide distally; leaf blades usually < 4 mm wide .... X. witsenioides Scapes 2-spicate ........................................................................... X. bicephala Scapes 1-spicate ....................................................................................... 82 Hairs of leaf and scape edges rusty, forming a dense border; apices of leaves erect; seeds ca. 1.5 mm long ..................................................... 83 Hairs of leaf and scape pale, usually white or blonde; apices of leaves various; seeds various .......................................................................... 84

Xyris 547

83(82). Main stems not rebranching; tips of leaves rounded; spikes hemispheric or subglobose ............................................................................ X. decussata 83. Main stems rebranching, the branches elongate, forming frond-like plates of leaves; tips of leaves narrowly acute; spikes ellipsoid or obovate ............................................................................................ X. ptariana 84(82). Seeds > 2 mm long; spikes hemispheric or subglobose; bracts very lustrous with subcucullate tips; tips of most leaves incurved-blunt .................................................................................................. X. albescens 84. Seeds ca. 1.5 mm long; spikes obovoid or oblong; bracts dull, sooty brown, with flat tips; tips of most leaves erect and acute ................... X. fuliginea Xyris albescens Steyerm., Fieldiana, Bot. 28: 105, fig. 16a, b. 1951. Cespitose perennial to 1 m tall; leaves elongate-linear with red-brown or castaneous bases, the blades densely white-ciliate; spikes roundish, dark, to 1.5 cm long; lateral sepals apically ciliate, free; seeds 2–3 mm long. Moist to wet rocky tepui meadows, mostly at or near summits of tepuis, 1200– 2500 m; Bolívar (Cerro Jaua, east-northeast of Cerro Venado, Kukenán-tepui, Macizo del Chimantá, Ptari-tepui, Roraima-tepui, Sierra de Lema, Uarama-tepui in northeastern Gran Sabana, Yuruaní-tepui), Amazonas (Cerro Marahuaka). Guyana, northern Brazil. ◆Fig. 448. Xyris aquatica Idrobo & L.B. Sm., Caldasia 6: 206, fig. 9. 1954. Lax, smooth, aquatic, cespitose rhizomatous perennial to 0.5 m long, submersed except for scape tips; leaves with broad, thin ligulate sheaths and filiform blades; spikes to 1.5 cm long, the bracts with dorsal areas; the sepals free; seeds ca. 0.5 mm long. Shoalwaters of streams and rivers, 100–300 m; local in Amazonas. Southeastern Colombia, Amazonian Brazil. Xyris aracamunae Kral, Ann. Missouri Bot. Gard. 76: 975, fig. 21. 1989. Cespitose, coarse, dark-based, pale-tomentose perennial; leaves elongate, 2–2.5 mm wide, the edges tomentose; scapes ribbed, to 0.7 m long, tomentose; spikes obovoid, dark, lower bracts retrorsely hirsute; sepals densely fimbriate-ciliate, free; seeds ca. 1 mm long. Tepui meadows, ca. 1500 m; Amazonas (summit of Cerro Aracamuni). Endemic. ◆Fig. 451. Xyris arachnoidea Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 28, fig. 12A–E. 1964.

Cespitose, fibrillose-based, sub-bulbous perennial to 0.4 m tall; leaves with narrow, flattened but thick blades; spikes dark, ovoid, 7–8 mm long; bracts with large dorsal areas, edges with long white hairs; sepals free; seeds ca. 1 mm long. Savannas, 100–200 m; Amazonas (along upper Río Orinoco). Endemic. ◆Fig. 455. Xyris atriceps Malme, Bull. Torrey Bot. Club 58: 325. 1931. Densely cespitose, castaneous- or nearly black-based perennial; leaf blades terete, ventrally sulcate; spikes nearly black, thinedged; bracts without dorsal areas; sepals free. Wet, rocky tepui meadows, 1200–2800 m. Venezuela, probably adjacent Brazil; 4 subspecies, all in the flora area. Key to the Subspecies of X. atriceps 1. Spikes narrowly obovoid; leaf blades lacking sulcus; bracts subentire, with a low costa distally, the bract margins ciliolate with yellowish hairs ................................ ............................... subsp. neblinensis 1. Spikes narrowly to broadly obovoid; leaf blades sulcate at least ventrally; bracts becoming lacerate, ecostate distally, the bract margins various ........................... 2 2. Spikes 1–1.5 cm long; leaf blades ca. 1 mm thick; staminodia absent ......................... ............................ subsp. marahuacae 2. Spikes < 1 cm long; leaf blades < 1 mm thick; staminodia present, bearded ...... 3 3. Spikes broadly obovoid, base strongly attenuate; bract edges distally white-ciliate and erect .............. subsp. atriceps 3. Spikes narrowly obovoid or ellipsoid or ovoid, base short-attenuate if at all so; bract edges sordidly villosulous, usually becoming very lacerate, subsquarrose ........................... subsp. chimantensis

548

X YRIDACEAE

X. atriceps subsp. atriceps 1700–2000 m; Amazonas (southern Cerro Duida). Endemic.

1. Leaf blades < 1 mm wide, at apex more rounded, thicker; fertile bracts slightly keeled, slightly spreading at maturity .......................................... var. tillettii

X. atriceps subsp. chimantensis Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 19. 1964. 1900–2500 m; Bolívar (summits of Macizo del Chimantá). Endemic. ◆Fig. 453.

X. bicostata var. bicostata Ca. 1800 m; Amazonas (Cerro Huachamacari). Endemic.

X. atriceps subsp. marahuacae Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75: 557. 1988. 2500–2800 m; Amazonas (summit of Cerro Marahuaka). Endemic.

X. bicostata var. tillettii (L.B. Sm.) Kral, Ann. Missouri Bot. Gard. 75: 675. 1988. —Xyris tillettii L.B. Sm., Ernstia 9: 3, fig. 1A–E. 1982. 2200–2700 m; Amazonas (Cerro Marahuaka, Sierra de la Neblina). Endemic. ◆Fig. 457.

X. atriceps subsp. neblinensis Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 19. 1964. 1200–2300 m; Amazonas (summit of Sierra de la Neblina). Endemic. A single collection from Macizo del Chimantá in Bolívar is intermediate to subsp. chimantensis (Huber et al. 10151, MYF, VDB, VEN). Xyris bicephala Gleason, Brittonia 3: 155. 1939. Cespitose, coarse perennial with dark bases; leaf blades flat, densely pale-ciliolate; scapes with ovoid, deep brown, paired spikes. Moist to wet savannas, seeps along streams, rapateaceous bogs, 800–2400 m; widespread in Bolívar. Guyana, Suriname, adjacent highaltitude Brazil. ◆Fig. 456. Xyris bicostata Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 30, fig 14F. 1964. Cespitose, low, stiff perennials to 0.3 m tall; leaf sheaths red-brown or dark, ciliate; blades to 1 mm wide, the margins ciliolate; scapes strongly bicostate, costas pale-ciliate or scabrid; spikes 9–10 mm long, red-brown, the bracts with linear dorsal areas; lateral sepals free, entire; seeds ca. 1 mm long. Tepui meadows, Venezuela; 2 varieties, both endemic to Flora area. Key to the Varieties of X. bicostata 1. Leaf blades ca. 1 mm wide, at apex conicacute; fertile bracts strongly keeled, spreading at maturity ..... var. bicostata

Xyris byssacea Kral, Ann. Missouri Bot. Gard. 75: 690. 1988. Cespitose, dark-based perennial to 0.3 m tall; leaves with sheaths elongate, cobwebbyciliate, the blades filiform, smooth, compressed, to 0.4 mm wide; scapes terete; spikes pale brown, ca. 6 mm long; dorsal areas of bracts linear; lateral sepals free, entire; seeds ca. 1 mm long. Rocky tepui meadows, 1500–2200 m; Bolívar (Auyán-tepui, Cerro Guaiquinima), Amazonas (Cerro Yaví, Sierra de la Neblina). Endemic. Xyris carinata Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 36, fig. 21A– E. 1964. Similar to Xyris byssacea, but the leaf sheaths pilose-ciliate, papillose-rugulose, the blades verrucose-rugose; scapes papilloserugose; spikes 7–8 mm long; bract backs papillose; lateral sepals free, ciliate; petals entire; seeds ca. 0.7 mm long. Sandy, rocky savannas and tepui meadows, 1000–2200 m; Bolívar (Auyán-tepui), Amazonas (Cerro Autana, Cerro Guanay, Sierra Parima). Adjacent high-altitude Brazil. ◆Fig. 454. Xyris chimantae Kral & L.B. Sm., Phytologia 53: 432, fig. 1A–H. 1983. Cespitose perennial to 0.6 m tall, the stems elongate; leaf bases castaneous or yellow-brown; scapes nearly ecostate; spikes obovoid, ca. 1 cm long; bracts apically redciliate; lateral sepals free, red-fimbriociliate. Moist tepui meadows, 1900–2500 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 458.

Xyris 549

Xyris concinna N.E. Br., Trans. Linn. Soc. London, Bot. 6: 68. 1901. Similar to Xyris hymenachne, but shorter and harder-based; leaf sheath base orbicular, sparsely ciliolate; bracts more folded-tipped, more acute; petals entire. Wet, rocky tepui meadows, 2500–3000 m; Bolívar (Ilú-tepui, Kukenán-tepui, Roraima-tepui, Uei-tepui). Endemic. ◆Fig. 462. Xyris consolida Kral & L.B. Sm., Phytologia 53: 434, figs. 3 & 4. 1983. Solitary or tufted perennial to 0.6 m tall; leaves flabellately spreading; sheaths ciliate, these and blades rugulose-scabid; scapes multicostate, scabrid; spikes dark, bracts without dorsal areas; sepals connate, ciliolate; seeds ca. 1 mm long. Tepui meadows, 1900–2400 m; Bolívar (Ptari-tepui, Uaipántepui). Endemic. ◆Fig. 461. Xyris contracta Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 33. 1964. Cespitose, smooth perennial to 0.6 m tall; leaves with blade base contracted at junction with ligulate sheath; spikes ellipsoid, ca. 1.5 cm long; lowest, sterile bracts subdecussate, fertile bracts ranked or spiro-distichous; seeds to 2 mm long. Wet shrub tepui meadows, 1200–1900 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Xyris cryptantha Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 16, fig. 2A–E. 1964. Low, slender, cespitose, fibrillose-based perennial; leaves with floccose-ciliate sheaths and filiform, terete blades; scapes round, punctate; spikes to 9 mm long, bracts with large dorsal areas, the lowest pair of bracts overtopping and connivent-tipped over spike top. White-sand savannas, 100–200 m; Amazonas (Caño Pimichín, Caño Yagua, El Venado, near Guarinuma, Río Casiqiare, Río Pasimoni, Río Sipapo, west of San Antonio de Orinoco). Southeastern Colombia, Brazil (Amazonas: Serra Aracá). ◆Fig. 465. Xyris culmenicola Steyerm., Fieldiana, Bot. 28: 108. 1951. Robust cespitose perennial to 0.7 m tall, the bases hard, brown to castaneous; leaves

with blades pilosulous-ciliate, green banded with red-brown; spikes dark, to 2 cm long, blunt, dull; sepals free, rusty-ciliate; seeds ca. 2 mm long. Moist tepui meadows, 1900– 2600 m; Amazonas (Cerro Duida, Cerro Marahuaka). Endemic. ◆Fig. 459. Xyris cyperoides Gleason, Bull. Torrey Bot. Club 56: 17. 1929. Xyris epicarae Kral & L.B. Sm., Phytologia 53: 436, fig. 6. 1983. Tufted, smooth, slender, low perennial, the base pale red-brown; leaves with blades filiform, overtopping scapes; spikes 2-flowered, ca. 4 mm long; bracts with large dorsal areas; lateral sepals free, entire; seeds ca. 0.5 mm long. Sandy savannas, rock ledges around falls, forest transitions, 400–2100 m; Bolívar (Auyán-tepui, Cerro Pitón, Salto Szczerbanari on Cerro Guaiquinima). Guyana, Suriname, Amazonian Brazil. Xyris decussata Gleason, Bull. Torrey Bot. Club 56: 392. 1929. Stiff, low, tufted perennial resembling Xyris bicephala, but 1-spicate, the shorter scapes and leaf edges densely rusty-ciliate, the blade tips broadly rounded; petals entire; seed 1.5–1.7 mm long. Tepui meadows, 1900– 2700 m; rare in Bolívar (Ilú-tepui, Macizo del Chimantá, Roraima-tepui). Adjacent Guyana and Brazil. Xyris delicatula Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 23, fig. 7A–F. 1964. Much like Xyris carinata, but spike bracts lacking dorsal areas, the lateral sepal keels lacerate-ciliate from middle to tip; seeds ca. 0.5 mm long. Tepui summit meadows, 1500– 2300 m; Bolívar (Auyán-tepui, Cerro Guanay), Amazonas (Cerro Sipapo). Endemic. ◆Fig. 452. Xyris esmeraldae Steyerm., Fieldiana, Bot. 28: 109, fig. 16c. 1951. Xyris rugulosa Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 29, fig. 13A– E. 1964. Cespitose, delicate annual 5–30 cm tall; leaf blades reduced or flat, twisted; scapes much-twisted, flattened; spikes 1-flowered, to 5 mm long; bracts decussate; sepals con-

550

X YRIDACEAE

nate. Moist, sandy savannas, 100–500(–800) m; Amazonas (Caño Yagua, Cerro Duida, Cerro Sipapo, Maroa, Piedra Cocuy, near Pimichín, near headwaters of Río Mavaca, Río Mawarinuma, San Carlos de Río Negro). Southeastern Colombia. ◆Fig. 466. Xyris fallax Malme, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 22(afd. 3, no. 2): 12, pl. 1, fig. 5. 1896. Xyris dolichosperma Lanj., Recueil Trav. Bot. Néerl. 34: 488, fig. 5. 1937. Xyris erythema Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 12, fig. 1A–F. 1964. Soft-based, short to tall, tufted perennial; scapes with 3 or more costae; leaves with brown or red-brown, sometimes ciliate sheaths; blades flat and deep green; spikes ovoid; bracts many, with evident dorsal areas; seeds 1–1.5 mm long, placentation parietal. Wet lowland to upland savannas, along tepui streams, 100–1100(–1500 m); common in Bolívar and Amazonas. Guárico; widely distributed in Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil, West Africa. ◆Fig. 468. Xyris frequens Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 32, fig. 16A– E. 1964. Cespitose perennial to 0.4 m tall; leaf sheaths intermittently villous-ciliate; blades terete or subterete, the tips conic, often tipped with pale hairs, the surfaces rugulose proximally; spikes broadly ellipsoid or turbinate, to 1.4 cm long; bracts loosely ranked with large dorsal areas, aging excurved; lateral sepals free, apically ciliolate or fimbriolate; seeds 1.5–1.6 mm long. Moist, white-sand savannas, 100–200 m; Amazonas (along Caño Pimichín, base of Cerro Yapacana). Endemic. Xyris frondosa Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 33, fig. 18A– E. 1964. Low, densely cespitose perennial, the stems elongate; leaves many, arranged in frond-like plates; blades with thickened, ciliate margins and rusty-borders; spikes ellipsoid, to 7 mm long, bracts with red-brown dorsal areas. Tepui meadows, (700–)1500– 2600 m; Bolívar (Cerro Jaua, Macizo del Chimantá). Endemic. ◆Fig. 467.

Xyris fuliginea Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75: 612. 1988. Rigid, cespitose perennial ca. 0.3–0.6 m tall, the plant base dark; leaf sheath castaneous, entire; blades compressed, ciliate; scapes bicostate, ancipital, white-ciliate; spikes dark, sooty-brown, to 1.5 cm long; bracts without dorsal areas; sepals ciliate; seeds ca. 1.5 mm long. Wet, rocky tepui meadows, 2000–2600 m; Amazonas (Cerro Marahuaka). Endemic. ◆Fig. 471. Xyris globosa Nilsson, Kongl. Svenska Vetenskapsakad. Handl. n.s. 24(14): 57. 1892. Bulbous-based perennial to 0.7 m tall; leaf sheaths orbicular-dilated, ciliate; blades narrow, thick-edged, smooth; scapes twisted and terete; spikes ovoid to subglobose, dull brown, fuzzy, the bracts with narrow dorsal areas; sepals free, the winged keel pale-villous-fimbriate; seeds ca. 0.5–0.6 mm long. Sandy savannas, 100–900 m; Bolívar (uncommon), Amazonas (common). Amazonian Colombia. Xyris gongylospica Kral, Novon 8: 393. 1998. Cespitose perennial 0.5–0.7 m tall; leaf sheaths carinate, long-ciliate, pale red or tan; blades narrowly linear-gladiate, scabridulous and rugulose; scapes terete, coarsely multicostate; spikes ca. 1 cm long, subglobose, attenuate; sepals ca. 5 mm long, the lateral ones strongly inequilateral and blunt; seeds ca. 1 mm long. Seeps along streams in riparian scrub, 1000–1100 m; Bolívar (southeastern headwaters of Río Carrao). Endemic. Xyris graniticola Kral, Ann. Missouri Bot. Gard. 75: 573. 1988. Slender, cespitose perennial to 0.3 m tall; leaves narrowly linear, twisted, scabrociliate toward blade base; scapes slender, ecostate; spikes 4–5 mm long, similar to Xyris guianensis. Granite outcrops, ca. 100 m; Amazonas (Río Autana). Endemic. Xyris guianensis Steud., Syn. Pl. Glumac. 2: 285. 1855. Xyris gardneri Malme, Bih. Svenksa Vetensk.-Akad. Handl. 26(afd. 3, no. 19): 8, pl. 1, fig. 1. 1901. Xyris filiscapa Malme, Repert. Spec. Nov. Regni Veg. 3: 112. 1906.

Xyris 551

Low, densely cespitose annual to 0.3 m tall; leaves flabellately spreading, often maroon or red-brown; blades with a narrow pale cartilaginous margin; scapes filiform; spikes ellipsoid, to 7 mm long, the bracts with large dorsal areas; sepals free, papillate or ciliolate; seeds 0.4–0.5 mm long. Sandy, moist savannas, 50–1300(–2000) m; Bolívar (common in Gran Sabana), Amazonas (common). Southeastern Colombia, Guyana, Suriname, Amazonian Brazil (south to Goias). ◆Fig. 469. Xyris huberi Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75: 571, fig. 20. 1988. —Xyris foveolata Kral & L.B. Sm., Phytologia 53: 435. 1983, non Irmsch. Short-lived, tufted perennial from a short subvertical rhizome; leaves spreading; blades flat; margins thickened, yellowish, ciliate, the borders rusty, the surface punctate; scapes punctate; spike bracts with rusty, ciliate borders and pale brown punctate dorsal areas. Riparian white-sand savannas, 100–200 m; southern Amazonas (base of Cerro Yapacana). Endemic. ◆Fig. 460. Xyris hymenachne Mart., Flora 24(2): 55. 1841. Xyris arescens Kunth, Enum. Pl. 4: 3. 1843. Xyris submontana Gleason, Bull. Torrey Bot. Club 56: 393. 1929. Small to medium-sized, pale green, smoothish perennial to 0.6 m tall; leaves flabellately spreading; blades flattened; scapes slender; spikes obovoid to subglobose, to 8 mm long; bracts without dorsal areas and with broad pale scarious borders; seeds 0.6–0.7 mm long. Moist, sandy savannas, streambanks and bars, 100–1800 m; Bolívar (common in Gran Sabana), Amazonas (Cerro Marahuaka, Río Mawarinuma, Sierra de la Neblina). Widespread in Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 470. Xyris involucrata Nees, J. Bot. (Hooker) 2: 397. 1840. Xyris asterocephala Seub. in Mart., Fl. Bras. 3(1): 219. 1855. Tufted, short-lived perennial, the bases dark; leaves flabellately spreading; blades flat, pale green with reddish borders; spikes hemispheric; lower bracts leafy, forming a

spreading, leafy involucre. Lowland to upland savannas, granitic outcrops, 100–1200 m; widespread in Bolívar and Amazonas. Colombia, Guyana, Suriname, Amazonian Brazil. ◆Fig. 474. Xyris juncifolia Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 26, fig. 10A– E. 1964. Tall, rush-like, cespitose brittle perennial; leaves with sheaths long, dull brown or redbrown; blades terete; spikes fusiform-elliptic, bracts with dorsal areas; seeds large. Rocky, wet, shrub savannas, tepui summits, 900– 2000 m; Bolívar (Cerro Guaiquinima, Cerro Jaua), Amazonas (Cerro Duida, Cerro Parú). Endemic. ◆Fig. 472. Xyris jupicai Rich., Actes Soc. Hist. Nat. Paris 1: 100. 1792. Xyris jupicae Michx., Fl. Bor.-Amer. 1: 23. 1803. Xyris anceps Pers., Syn. Pl. 1: 56. 1805, non Lam. 1791. Xyris macrocephala f. minor (Mart.) M. Kuhlm. & E. Kuhn, Fl. Dist. Ibiti 34. 1947. —Xyris laxifolia var. minor Mart., Flora 24(2) Beibl.: 54. 1841. Xyris ambigua Beyr. ex Kunth, Enum. Pl. 4: 13. 1843, pro parte. Xyris communis Kunth, Enum. Pl. 4: 12. 1843. Xyris arenicola Miq., Linnaea 18: 75. 1844. Xyris acuminata Miq. ex Steud., Syn. Pl. Glumac. 2: 284. 1855. Xyris gymnoptera Griseb., Cat. Pl. Cub. 223. 1866, pro parte. Xyris jupicai var. brachylepis Malme, Svensk. Bot. Tidskr. 21: 394. 1927. Short to tall, soft-based, weedy annual, often associated with Xyris laxifolia, the plant base usually brown or straw-colored; scape ridges papillose; placentation parietal; seeds ca. 0.5–0.6 mm long, translucent. Wetlands, ditches, wet savannas, open swamps, 50– 1300 m; locally abundant in Delta Amacuro, Bolívar, and Amazonas. Venezuelan Andes, Apure, Guárico, Monagas, locally elsewhere in Venezuela; widely distributed in southeastern U.S.A., Mexico, Central America, South America except Ecuador and Chile. ◆Fig. 475. Xyris jupicai is the weediest species of Xyris in America.

552

X YRIDACEAE

Xyris kukenaniana Kral, Ann. Missouri Bot. Gard 75: 719. 1988. Low, cespitose perennial to 0.2 m; leaf sheaths pale red-brown; blades flat, whiteciliate or scabrociliate; spikes dark redbrown or sooty brown, ovoid, 7–8 mm long; lower bracts with white ciliate tips; lateral sepals free, sparsely pilose-ciliate. Rocky, shrubby tepui meadows, ca. 2500 m; Bolívar (Kukenán-tepui). Endemic. Xyris lacerata Pohl ex Seub. in Mart., Fl. Bras. 3(1): 26, pl. 26. 1855. Slender, bulbous-based perennial to 0.6 m tall; leaf base dilated, castaneous; blade narrow, to 3 mm wide; spikes ovoid to globose, to 1 cm long, dull brown with scarious-bordered, lacerate bracts, the bracts with distinct dorsal areas; sepals free, ciliolate; seeds 0.5–0.6 mm long. Moist to wet, sandy savannas, also on and around granitic outcrops, 100–400 m; Bolívar (southwest of Caicara del Orinoco, south of Ciudad Piar, Panare, Puerto Nuevo, between Río Horeda and Cerro Gavilán, east of Río Parguaza), Amazonas (Cerro Morrocoy, Estero La Yaguita near Caño Yagua, northeast of Galipero, La Esmeralda, Puerto Ayacucho, Río Manapiare, Río Ventuari, Santa Bárbara). Apure, Guárico; Colombia, Peru, Brazil, Bolivia, Argentina. Xyris lanulobractea Steyerm., Fieldiana, Bot. 28: 109, fig. 16D–H. 1951. Slender, cespitose, wand-scaped perennial; leaf sheath elongate, lustrous redbrown; blades filiform, terete; spikes small, ovoid, fuzzy with white hairs on bract borders. Wet, rocky savannas, 300–1100 m; Bolívar (Gran Sabana). Guyana, Amazonian Brazil. Xyris laxifolia Mart., Flora 24(2): 53. 1841. Coastal southeastern U.S.A., Mexico, Central America, South America except Chile and Uruguay; 2 varieties, 1 in Venezuela. Variety iridifolia (Chapm.) Kral replaces the typical variety in Mexico and the Coastal Plain of the southeastern U.S.A. X. laxifolia var. laxifolia ?Xyris macrocephala Vahl, Enum. Pl. 2: 204. 1805, nom. confus.

Xyris laxifolia var. major Mart., Flora 24(2): 58. 1841. —Xyris macrocephala var. major (Mart.) Nilsson, Kongl. Svenska Vetenskapsakad. Handl. n.s. 24(14): 30. 1892. —Xyris caroliniana var. major (Mart.) Idrobo & L.B. Sm., Caldasia 6: 199, fig. 4. 1954. —Xyris jupicai var. major (Mart.) L.B. Sm. & Downs in Reitz, Fl. Ilustr. Catarin. pt. 1, fasc. 12, 9: 9. 1965. Robust, red- or purple-based, with stiff, flat, deep green leaf blades; scape surface and edges smooth; spikes large, dark, ovoid to cylindric, acute. Wet, alluvial woodland borders, wet savannas, streambanks, shallow acid pools, near sea level to 1300 m; locally abundant and sometimes weedy in Delta Amacuro, Bolívar, and Amazonas. Venezuelan Andes, Apure, Guárico, Monagas, locally elsewhere in Venezuela; widely distributed in Central America, South America except Chile and Uruguay. Xyris liesneri Kral, Ann. Missouri Bot. Gard. 76: 977. 1989. Similar to Xyris frondosa but slightly shorter, the stems less elongate; leaves entire except at tips; margins wiry-thickened; scapes subancipital, bicostate, the costas pilose-ciliate. Moist tepui meadows, ca. 1400 m; Amazonas (Cerro Aracamuni). Endemic. Xyris lithophila Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75: 704. 1988. Slender, cespitose, smooth perennial to 0.4 m tall; leaf sheaths pale red-brown, longciliate; blades filiform, to 0.8 mm wide, the shining edges smooth; scapes terete; spikes few-flowered, to 9 mm long, bracts with dorsal areas reddish, triangular; lateral sepals free, above middle fimbriate-ciliate; seeds ca. 0.7 mm long. Rocky savannas, ca. 1000 m; Bolívar (Salto La Milagrosa south of San Francisco de Yuruaní). Endemic. Xyris lomatophylla Mart., Flora 24(2): 57. 1841. Cespitose perennial to 0.7 m tall; leaves dark-based, flabellately spreading; blades flat with a cartilaginous, sometimes ciliate border; spikes dull, ovoid-cylindric, to 3 cm long; bract edges woolly, pale; sepals free; seeds to 2.5 mm long. Moist, white-sand savannas, 100–400 m; Amazonas (Caño Cotúa

Xyris 553

near base of Cerro Yapacana, Caño Perro de Agua west of San Antonio, Caño Yagua, Carmelitas, El Venado, La Esmeralda, Piedra Cacaguati near Piedra Merey, south of the confluence of Río Manapiare and Río Ventuari). Adjacent Colombia, Amazonian Brazil. Xyris lugubris Malme, Bull. Torrey Bot. Club 58: 324, pl. 24, fig. 1A. 1931. Similar to Xyris culmenicola but leaves eciliate; spikes glossy; sepals more finely and evenly ciliate. Wet, rocky tepui meadows at or near summits, 1500–1800 m; Amazonas (Cerro Duida, Cerro Sipapo, Sierra de la Neblina). Endemic. Xyris malmeana L.B. Sm., Bol. Inspet. Fed. Obras Contra Secas, Río 10: 126. 1939. Xyris glabrata of Griseb. et auct., Fl. Brit. W. Ind. 525. 1864, not as to basionym Xyris savanensis var. glabrata Seub. Resembling Xyris lacerata, but shorter, more slender, the bases sub-bulbous; leaves flat with smooth margins; scapes with 3 or more low but sharp and distinct costas; spikes to 8 mm long, bracts with distinct dorsal areas, bracts with narrower scarious borders; sepals free, ciliolate to entire; seeds ca. 0.5 mm long. White-sand savannas, 100– 1200 m; frequent in Bolívar, rare in Amazonas. Trinidad, Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 476. Xyris melanovaginata Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75: 593. 1988. Robust, dark-based, smooth, cespitose perennial to 1 m tall, producing frond-like plates of stiff, flat, flabellately spreading, smooth or ciliate-edged, thick-margined leaves; spikes ellipsoid to cylindric, to 3 cm long, dark; seeds and sepals similar to those of Xyris albescens but shorter. Rocky, wet tepui meadows, 1800–2600 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 479. Xyris mima L.B. Sm. & Downs, Proc. Biol. Soc. Wash. 75: 250, fig. 4. 1960. Xyris trisperma Kral & L.B. Sm., Phytologia 53: 433, fig. 1a–i. 1983. Very similar in habit and papillosity to Xyris savanensis, but scapes strongly bicostate; staminodia present and bearded; seeds ca. 1 mm long, the tips with a cone of

pale bristles. Sandy savannas, sandy forest transitions, ca. 100–800 m; Bolívar (near Salto Szczerbanari on Cerro Guaiquinima). Amazonian Brazil (Amazonas, Mato Grosso, Pará). Xyris neblinae Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 26, fig. 11A– E. 1964. Very similar to Xyris juncifolia, but bases dull-castaneous, nearly black; staminodia Tshaped, beardless; seeds > 2 mm long, the ribs muricate. Rocky, wet shrub meadows on tepui summits, 1700–2300 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 463. Xyris oblata Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75: 686. 1988. Similar in aspect to Xyris savanensis and similarly rugulose-papillose, but leaf sheaths long-ciliate; spikes distinctly oblate; lateral sepals with fimbriate-lacerate keels; staminodia bearded; seeds ca. 0.8 mm long; spikes appear often to proliferate. Moist white-sand savannas, 100–200 m; Amazonas (along Caño Temi near Yavita). Endemic. Xyris oxylepis Idrobo & L.B. Sm., Caldasia 6: 204, fig. 7a–c. 1954. Xyris fusiformis Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 24, fig. 8A–E. 1964. Wiry, low, cespitose perennial with bases fibrillose-chaffy, brown; leaves with villousciliate sheaths and subterete ribbed blades; spikes few-flowered, narrowly ellipsoid, bracts with strong dorsal areas; lateral sepals free, the keels entire or apically ciliate; seeds 0.7–0.8 mm long. White-sand savannas, 100–300 m; Amazonas (Caño Yagua, Cerro Yapacana, northeast of Galipero, Puerto Ayachucho, Río Atabapo, Santa Cruz). Southeastern Colombia. ◆Fig. 496. Xyris paraensis Poepp. ex Kunth, Enum. Pl. 4: 9. 1843. Mostly low, cespitose, smooth annuals to 0.4 m tall, similar to Xyris savanensis; leaves with eciliate, often maroon-brown sheaths and slender smooth entire blades; spike bracts with broader dorsal areas; lateral sepals free, curved, ciliate; staminodia bearded. Mostly in sandy savannas, often weedy, 100–2200 m. Belize, Colombia, Ven-

554

X YRIDACEAE

ezuela, Trinidad, Guyana, Suriname, French Guiana, Brazil; 3 varieties, 2 in Venezuela, both of these in the flora area.

centation marginal. Shallows and splashrock along streams, ca. 600 m, Amazonas (slopes of Cerro Aracamuni). Endemic.

Key to the Varieties of X. paraensis

Xyris ptariana Steyerm., Fieldiana, Bot. 28: 110, fig. 16I–J. 1951. Densely cespitose perennial, forming domes of ascending stems; leaves darkbased, forming frond-like plates; sheaths with excurved ligules; spikes ellipsoid or obovoid, dark, 1–2 cm long, bracts without dorsal areas; sepals entire to ciliolate, free; seeds 1–1.5 mm long. Boggy or seepy, rocky tepui summit areas, 1300–2700 m; infrequent in Bolívar and Amazonas. Endemic. ◆Fig. 481.

1. Lateral sepals, as viewed from the side, lanceolate and acute; spikes ellipsoid to cylindric, at maturity 1–2 cm long ...................................... var. longiceps 1. Lateral sepals, as viewed from the side, broadly oblong and blunt; spikes predominantly ovoid to nearly round, mostly 5 mm long or shorter ..... var. paraensis X. paraensis var. longiceps (Malme) L.B. Sm. & Downs, Arq. Bot. Estad. São Paulo 4: 28. 1966. —Xyris longiceps Malme, Recueil Trav. Bot. Néerl. 9: 131. 1912. Xyris leptostachya Malme, Recueil Trav. Bot. Néerl. 9: 132. 1912. Rare in Bolívar (Puente Kumerepa north of Río Yuruaní ferry, Río Caruai, Río Hacha), Amazonas (Cerro Parú, Maroa, Río Asisa, upper Río Ventuari). Apure; Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 482. X. paraensis var. paraensis Bolívar (expected along the border with Brazil). Guárico; Belize, Trinidad, Guyana, Suriname, French Guiana, northeastern Brazil. ◆Fig. 483. Xyris pratensis Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 34, fig. 19A– E. 1964. Cespitose, slender, hard-based perennial to 0.6 m tall; leaves with sheaths sparsely pilose-ciliate; blades flat, 2–3 mm wide, the tips often white-bristly-ciliate; spikes becoming obovoid, 8–10 mm long; bracts decussate with pale dorsal areas; lateral sepals shortlacerate and ciliate; petals broadly acute, entire; seeds 3–3.5 mm long. Tepui meadows, 1500–2700 m; Amazonas (Cerro Marahuaka, Cerro Sipapo). Endemic. ◆Fig. 464. Xyris prolificans Kral, Ann. Missouri Bot. Gard. 76: 980. 1989. Delicate, low, mat-forming perennial, perennating by slender zig-zag creeping rhizomes from axils of lower rosette leaves; pla-

Xyris riparia Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 20, fig. 5A–E. 1964. Similar to larger specimens of Xyris setigera as to leaf and general indument but leaves averaging wider (2–3 mm); lateral sepals longer (0.9–1 versus 0.5–0.6 mm). Tepui meadows, 1800–2200 m; Bolívar (Cerro Guaiquinima, Macizo del Chimantá). Endemic. Xyris roraimae Malme in Pilger, Notizbl. Königl. Bot. Gart. Berlin 6: 117. 1914. Perennial, solitary or in small clumps, to 1 m tall or more; leaf sheaths dark, dilated, ciliate; blades pale yellow-green, ciliate, glazed with papillae and tubercles; scapes bicostate, likewise papillate-tuberculate, the costas ciliate; spikes ovoid-cylindric, pale red-brown; bracts without dorsal areas, redscarious-bordered, lacerate; seeds ca. 1 mm long. Moist savannas, 700–1500 m; Bolívar (Gran Sabana). Adjacent Guyana, Brazil (Goiás, Minas Gerais, planalto). ◆Fig. 485. Xyris rubrolimbata Heimerl, Ann. K.K. Naturhist. Hofmus. 21: 70, pl. 4, figs. 4– 6. 1906. Stiff annual to 10 cm tall; leaves broadly flabellately spreading; sheaths entire, ligulate; blades flat, sharp; margins thickened and red-bordered; spikes obovoid, ca. 5 mm long, blunt; bracts with ciliolate margins and papillate dorsal areas; sepals free, curved, ciliolatescabrid; seeds 0.6–0.7 mm long. Wet, sandy savannas, caatingas, 100–300 m; Amazonas (Maroa, Piedra Cocuy, between Río Guainía and Caño Pimichín, upper Río Negro). Amazonian Colombia and Brazil. ◆Fig. 490.

Xyris 555

Xyris savanensis Miq., Linnaea 18: 605. 1844. Cespitose, soft, and usually red-based annual, the foliage and scapes usually maroontinted and rugulose-papillose; leaves with entire sheaths and flat blades with slightly thickened margins; scape terete; spikes tan or brown, bracts with darker narrow dorsal areas; sepals free, curved, ciliate; flowers without staminodes; seeds 0.4–0.5 mm long, the tips truncate, apiculate. Savannas, 100– 1000 m; common in Bolívar and Amazonas. Anzoátegui, Apure, Guárico, Portuguesa; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 478. Xyris schneeana L.B. Sm. & Steyerm., Acta Bot. Venez. 1(7): 149. 1965. Cespitose perennial to 0.3 m tall, the foliage pale dull green and throughout papilloserugose or rugulose; leaves flabellately spreading, with sheaths straw-colored, white-ciliate; spikes dark, ovoid-ellipsoid, ca. 5 mm long, few-flowered; lateral sepals free, dark, serrulate-ciliolate above middle; seeds ca. 1 mm long. Wet, rocky tepui meadows, 2000–2500 m; Bolívar (Macizo del Chimantá). Endemic. Xyris setigera Oliv. ex Thrun, Timehri 5: 207. 1886; Trans. Linn. Soc. London, Bot. 2: 285, pl. 50A, figs. 1–8. 1887. Xyris setigera var. elongata Steyerm., Fieldiana, Bot. 28: 112. 1951. Densely cespitose, stiffish and brittle perennial 0.1–0.6 m tall, the foliage and scapes often very reddened; leaf sheaths mostly deep red-brown, scabrid-rugose, long-ciliate, blades linear or filiform, harshly brown-ciliate; scapes terete, smooth and punctate; spikes very dark, mostly lacerate; sepals free, lacerate-serrulate to entire; seeds ca. 0.5–0.6 mm long. Savannas and tepui meadows, 800–2400 m; Bolívar (Gran Sabana and adjacent tepuis). Adjacent Guyana and Brazil. ◆Fig. 450. Xyris setigera is the most common Xyris found on the Gran Sabana. Xyris seubertii Nilsson, Kongl. Svenska Vetenskapsakad. Handl. n.s. 24(14): 51, pl. 4. 1892. Xyris calcarata Heimerl, Ann. K.K. Naturhist. Hofmus. 21: 68, pl. 4, figs. 1– 3. 1906.

Slender perennial to 0.7 m tall; leaves flabellately spreading, with brown or dark entire sheaths and pale green glaucous flat blades; scapes slender and glaucous; spikes ovoid to ellipsoid; lowest bract with a strong cusp on the dorsal area; seeds to 0.4 mm long. Rather dry savannas, 900–2000 m; frequent in Bolívar (Gran Sabana). Adjacent Guyana, Brazil. Xyris spathacea Lanj., Recueil Trav. Bot. Néerl. 34: 484, fig. 4. 1937. Xyris exserta Idrobo & L.B. Sm., Caldasia 6: 229, fig. 21. 1954. Xyris yapobodensis Idrobo & L.B. Sm., Caldasia 6: 232, fig. 22. 1954. Mostly low, lax, soft-based, green-maroon ephemeral 10–30 cm tall; leaves of wet phases filiform, strongly ligulate, the blades terete to 0.5 mm thick, emergent leaf phases shorter, more rigid; spikes elliptic to obovoid, often proliferating, few-flowered, 3–5 mm long; bract thin, with dorsal areas; lateral sepals free, the keel papillose or short-ciliate. Riparian and intermittently flooded whitesand savannas, 100–200 m; Amazonas (Caño Yagua, Cerro Yapacana, Río Sipapo). Southeastern Colombia, Guyana, Suriname, Amazonian Brazil (Amazonas, Mato Grosso, Pará). Xyris spruceana Malme, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26(afd. 3, no. 19): 12, pl. 1, fig. 2, 1901. —Yaguaso. Xyris applanata Idrobo & L.B. Sm., Caldasia 6: 238, fig. 27. 1954. Cespitose annual to 0.3 m tall; leaves soft, flabellately spreading, pale green, with blades 1.5–3 mm wide, papillose to minutely ciliate-edged; scapes broadly thin-winged, the heads turbinate to hemispheric with the lowest bracts forming a leafy involucre. Moist white-sand savannas, forest transitions, 100–600 m; Amazonas (Río Guainía, near Pimichín, San Carlos de Río Negro to Solano). Southeastern Colombia, Brazil (Amazonas: Rio Negro basin). ◆Fig. 489. Xyris stenophylloides Malme, Bull. Torrey Bot. Club 58: 323. 1931. Similar in aspect to Xyris sulcatifolia but smooth; leaves with entire, lustrous brown sheaths and filiform flattened blades; spikes turbinate, 7–8 mm long, red-brown, bracts

556

X YRIDACEAE

without dorsal areas; sepals free, ciliate. Tepui summits, canyon faces, low elevation granitic outcrops, 100–1600 m; Bolívar (Serranía de Parguaza), Amazonas (Cerro Duida, along Río Orinoco, Río Parhueña basin, Río Sipapo 5 km above mouth). Endemic. ◆Fig. 449. Xyris stenostachya Steyerm., Fieldiana, Bot. 28: 113, fig. 16K–L. 1951. Slender annuals to 0.4 m tall; leaves spreading flabellately; sheaths entire; blades flat, to 3 mm wide, the edges thickened; spikes linear, flattened, 1.5–5 cm long; bracts boat-shaped with strong dorsal areas; lateral sepals free, ciliolate; seeds 1–1.5 mm long. Wet, sandy savannas, temporary pools on and around granite, 100–300 m; Bolívar (along Río Orinoco), Amazonas (along Río Orinoco). Reported from southeastern Colombia along the Orinoco. ◆Fig. 484. Xyris subglabrata Malme, Bull. Torrey Bot. Club 48: 322. 1931. Xyris garcia-barrigae Idrobo & L.B. Sm., Caldasia 6: 204, fig. 8. 1954. Tufted, stiff perennial with hard, brown, fibrillose bases; leaves with sheath edges cobweb-like or woolly to pilose-ciliate; blades terete-fluted, the tips with bristly hairs; scapes similar to leaf blades; spikes dark, mostly ovoid; lower bracts tipped with white hairs, the bracts with large dorsal areas; sepals free, villosulous or ciliate-keeled; seeds 0.6–0.7 mm long. Savannas, 100–1200 m; Bolívar (middle Río Ambutuir south of Auyán-tepui), Amazonas (Caño Pimichín, Cerro Duida, La Esmeralda, Río Autana basin, Río Orinoco, middle Río Puraname, Río Ventuari, San Carlos de Río Negro to Solano, Santa Cruz, Yavita). Southeastern Colombia, Brazil (Serra Aracá). Xyris subuniflora Malme, Recueil Trav. Bot. Néerl. 9: 129. 1912. Xyris capillaris Steyerm., Fieldiana, Bot. 28(1): 107. 1951. non Malme 1932. Delicate annual to 10 cm tall; leaves with entire sheaths and filiform but flat blades; scapes filiform; spikes ellipsoid, 1-flowered; sepals connate. Mostly sandy savannas, 100– 1200 m; Bolívar (between Kavanayén and base of Ptari-tepui), Amazonas (Caño Caname, Cerro Vinilla, Piedra Arauicaua, lower Río Pasimoni, between San Carlos de Río Ne-

gro and Solano, between Yavita and Maroa). Southeastern Colombia, Guyana, Suriname, northeastern Brazil (Pará). ◆Fig. 493. Xyris sulcatifolia Kral, nom. nov. pro Xyris scabridula Steyerm., Fieldiana, Bot. 28: 111. 1951, nom. illeg., non Rendle, Cat. Welw. Afr. Pl. 2: 72. 1901. Cespitose perennial to 0.4 m tall, the base roseate or purplish; leaf blades slender, blunt-tipped, these and scapes flexuous and strongly ribbed, papillate-rugulose or scabridulous; spikes small, ovoid, dark redbrown, with papillate-tuberculate bracts. Wet, rocky meadows along tepui summits, 1700–2500 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Duida). Endemic. Xyris surinamensis Spreng., Tent. Suppl. 2. 1828. Xyris eriophylla Rchb., Pl. Excic. Weigelt 4: 4. 1827, nom. nud. —Xyris eriophylla Rchb. ex Kunth, Enum. Pl. 4: 4. 1843. Cespitose, stiff, hard-based perennial; leaves with sheaths castaneous to purplebrown or rose, ciliate; blades twisted, flattened, variously ciliate; scapes strongly flattened and twisted, the costas often pale-pilose-ciliate; spikes dark and broad, to 10 mm long; bracts with large dorsal areas; sepals curved, free, lacerociliate to villous on keel; seeds ca. 1 mm long. Sandy or rocky, moist savannas, 100–1800 m; Bolívar (Amaruaytepui, near Cerro Abismo, El Pauji, Gran Sabana, Icabarú, Kavanayén, Macizo del Chimana [Churí-tepui], lower Río Caruay, Río Yuruaní), Amazonas (Cerro Guanay, Cerro Parú, La Esmeralda, lower Río Pasimoni, Sabana Hechimoní on Río Siapa). Widely distributed in southeastern Colombia, Guyana, Suriname, French Guiana, and Amazonian Brazil. ◆Fig. 486. Xyris surinamensis is extremely variable in degree of hairiness. Xyris tatei Malme, Bull. Torrey Bot. Club 58: 324, pl. 24, fig. 1A. 1931. Robust perennial similar to Xyris albescens but leaves and scape edges eciliate. Wet, rocky savannas, 1100–2000 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 492. Xyris teinosperma Idrobo & L.B. Sm., Caldasia 6: 224. 1954.

Xyris 557

Coarse, cespitose perennial; leaves dark brown at base; sheaths entire; blades broadly linear, deep green, flat, flabellately spreading; scapes to 0.8 m tall, stiff; spike ovoid, 2–3 cm long; petals largest of the genus with blades to 1.2 cm long; seeds the longest of the genus, linear, to 3 mm long with an apical pappus-like coma. Wet savannas, 100–1200 m; Amazonas (Canaripó, Cerro Yapacana, north of Maroa, near Pimichín, Río Atabapo, Río Guainía). Southeastern Colombia, adjacent Amazonian Brazil. ◆Fig. 491. Xyris tenella Kunth, Enum. Pl. 4: 9. 1843. Venezuela, Guyana, Suriname, French Guiana, Brazil, Paraguay; 2 varieties, 1 in Venezuela. X. tenella var. tenella Xyris steyermarkii Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 30, fig. 15A–E. 1964. Xyris yutajensis Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 36, fig. 19A–F. 1964. Cespitose, smooth annual to 0.2 m tall, much like Xyris guianensis, but leaf sheaths long-ciliate proximally; leaf blades not thickmargined; spike bracts with narrow, mostly linear dorsal areas; lateral sepals narrower, sharper, ciliolate to subentire. Sandy savannas, 1000–2300 m; Bolívar (Auyán-tepui, Cerro Guanay, Cerro Jaua, Cerro Marutaní, Cerro Venado, La Escalera, between Luepa and Cerro Venamo, Macizo del Chimantá, Salto Yuruaní), Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Marahuaka, Cerro Yaví, Cerro Yutajé, Río Iguapo, northwest of Yutajé). Guyana, Suriname, French Guiana, Brazil, Paraguay. ◆Fig. 477. Xyris thysanolepis Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 17, fig. 3A–F. 1964. Cespitose, slender perennial to 1 m tall; leaves flabellately spreading, the sheaths castaneous or tan, shading upward to pink or straw-colored, entire or sparsely ciliate; scapes mostly 1-costate, the costa ciliolate; spikes pale red-brown, ellipsoid; bract tips with broad, lacerate, usually red, scarious tips; lateral sepals free or to 1/3 connate; seeds ca. 1 mm long. Wet, rocky savannas, tepui summits, Venezuela; 2 varieties, both in the flora area.

Key to the Varieties of X. thysanolepis 1. Leaf sheath bases lustrous deep brown or pale brown, mostly evidently ciliolate; apices of fertile bracts narrowed, somewhat keeled, the scarious borders pale or pale red-brown; edges of leaf blades densely ciliolate with fine white hairs ......................................... var. sipapoa 1. Leaf sheath bases bright to dull brown, eciliate to sparsely ciliate; apices of fertile bracts broadly rounded, the laceratescarious borders distinctly red; edges of leaf blades entire or inconspicuously ciliolate .................... var. thysanolepis X. thysanolepis var. sipapoa (Maguire & L.B. Sm.) Kral & L.B. Sm., Ann. Missouri Bot. Gard. 75; 604. 1988. —Xyris sipapoa Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 18, fig. 4A–F. 1964. 1200–1600 m; Bolívar (Gran Sabana), Amazonas (Cerro Sipapo). Endemic. ◆Fig. 495. X. thysanolepis var. thysanolepis Xyris jauana L.B. Sm. & Steyerm., Bol. Soc. Venez. Ci. Nat. 32: 277, fig. 2a–g. 1976. 1200–2800 m; Bolívar (Cerro Jaua, Gran Sabana), Amazonas (Cerro Yaví). Endemic. ◆Fig. 494. Xyris toronoana Kral, Ann. Missouri Bot. Gard. 75: 643. 1988. Low, smooth, cespitose perennial to 15 cm tall, similar to Xyris tenella but bases of leaf sheaths entire; scape costas scabrid, the spike bracts with dorsal areas; sepals free and entire. Tepui meadows, ca. 2100 m; Bolívar (Macizo del Chimantá [central Chimantá-tepui]). Endemic. Xyris uleana Malme, Repert. Spec. Nov. Regni Veg. 3: 113. 1906. Cespitose, soft-based annual to 0.15 (–0.22) m tall; leaves flabellately spreading, to 10 cm long, 3.5 mm wide; spikes ovoid, mostly 3–5 mm long; lowest bracts keeled, incurved, about equaling spike, all bracts with large papillose or hirtellous dorsal areas. Wet savannas, lajas, forest transitions. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará, Mato Grosso); 2 varieties, both in the flora area.

558

X YRIDACEAE

Fig. 448. Xyris albescens

Fig. 449. Xyris stenophylloides

Fig. 450. Xyris setigera

Xyris 559

Fig. 451. Xyris aracamunae

Fig. 452. Xyris delicatula

Fig. 453. Xyris atriceps subsp. chimantensis

560

X YRIDACEAE

Fig. 454. Xyris carinata

Fig. 455. Xyris arachnoidea

Fig. 456. Xyris bicephala

Xyris 561

Fig. 457. Xyris bicostata var. tillettii

Fig. 458. Xyris chimantae

Fig. 459. Xyris culmenicola

562

X YRIDACEAE

Fig. 460. Xyris huberi

Fig. 461. Xyris consolida

Fig. 462. Xyris concinna

Xyris 563

Fig. 463. Xyris neblinae

Fig. 464. Xyris pratensis

Fig. 465. Xyris cryptantha

564

X YRIDACEAE

Fig. 466. Xyris esmeraldae

Fig. 467. Xyris frondosa

Fig. 468. Xyris fallax

Xyris 565

Fig. 469. Xyris guianensis

Fig. 470. Xyris hymenachne

Fig. 471. Xyris fuliginea

566

X YRIDACEAE

Fig. 472. Xyris juncifolia

Fig. 473. Xyris uleana var. uleana

Fig. 474. Xyris involucrata

Xyris 567

Fig. 475. Xyris jupicai

Fig. 476. Xyris malmeana

Fig. 477. Xyris tenella var. tenella

568

X YRIDACEAE

Fig. 478. Xyris savanensis

Fig. 479. Xyris melanovaginata

Fig. 480. Xyris witsenioides

Xyris 569

Fig. 481. Xyris ptariana

Fig. 482. Xyris paraensis var. longiceps

Fig. 483. Xyris paraensis var. paraensis

570

X YRIDACEAE

Fig. 484. Xyris stenostachya

Fig. 485. Xyris roraimae

Fig. 486. Xyris surinamensis

Xyris 571

Fig. 487. Xyris xiphophylla

Fig. 488. Xyris wurdackii subsp. wurdackii

Fig. 489. Xyris spruceana

572

X YRIDACEAE

Fig. 490. Xyris rubrolimbata

Fig. 491. Xyris teinosperma

Fig. 492. Xyris tatei

Xyris 573

Fig. 493. Xyris subuniflora

Fig. 494. Xyris thysanolepis var. thysanolepis

Fig. 495. Xyris thysanolepis var. sipapoa

574

X YRIDACEAE

Xyris valdeapiculata Kral, Ann. Missouri Bot. Gard. 75: 588. 1988. Slender smooth perennial to 0.4 m tall; leaf blades flat, 1–2.5 mm wide; scapes prominently bicostate-ciliate; spikes fewflowered, turbinate, to 1 cm long; fertile bracts with narrow keel, lacking dorsal area. Wet, rocky tepui meadows, ca. 2200 m; Amazonas (crests of Sierra de la Neblina). Endemic. Xyris witsenioides Oliv. ex Thurn, Timehri 5: 207. 1886. Similar to Xyris ptariana but somewhat shorter; leaves and scapes narrower; ligule similar but not excurved; spikes to 1.4 cm long; seeds ca. 1–1.3 mm long. Seepage, wet ledges and cliffs, wet tepui meadows, 1200– 2700 m; common in Bolívar and Amazonas. Adjacent Guyana and Brazil. ◆Fig. 480. Fig. 496. Xyris oxylepis

Key to the Varieties of X. uleana 1. Scapes at most low-costate, terete, rarely with 1–few low costas; leaves ≤ 1 mm wide ......................... var. angustifolia 1. Scapes distally ancipital, the costas produced at 2 flat wings comprising the edges; leaf blades > 1 mm wide ............... .......................................... var. uleana X. uleana var. angustifolia Lanj., Bull. Misc. Inform. Kew 1939: 562. 1939. Xyris connectens Malme, Ark. Bot. 19(13): 2. 1925. 100–1400 m; Bolívar (Gran Sabana, Puente Kumerepa north of Río Yuruaní ferry), Amazonas (Chacorro de Rincones near Galipero, Río Autana, northwest of Yutajé). Guyana, Suriname, eastern Amazonian Brazil. X. uleana var. uleana Xyris duidensis Malme, Bull. Torrey Bot. Club 58: 325. 1931. Xyris vaupesana Idrobo & L.B. Sm., Caldasia 6: 237, fig. 26a–f. 1954. 100–1900 m; Bolívar (Gran Sabana), Amazonas (widespread). Colombia, Guyana, Suriname, Brazil. ◆Fig. 473.

Xyris wurdackii Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 24, fig. 9A–E. 1964. Low, stiff, cespitose perennial, the bases sub-bulbous, fibrillose, dark; leaves erect, few, 0.1–0.3 m long, longer or shorter than the scapes; sheaths long-ciliate with villous or shaggy white hairs, and terete, ribbed blades; scapes similar to leaf blades; spikes obovoid, to 10 mm long, with dark sharp bracts largely concealed by shaggy, white, villous tomentum. Colombia, Venezuela; 2 subspecies, 1 in Venezuela. X. wurdackii subsp. wurdackii Savannas, forest transitions, 100–200 m; Amazonas (Caño Caname, Río Guainía, Río Guasacavi, Río Temi). Endemic. ◆Fig. 488. Xyris xiphophylla Maguire & L.B. Sm., Mem. New York Bot. Gard. 10(5): 20, fig. 6A–E. 1964. Robust, stiff, cespitose, smooth, darkbased perennial; stems often elongate; leaves imbricate, dark-based, forming plates; scapes slender but stiff, to 0.8 m tall, distally flattened, entire; spikes cylindric-obovoid, to ca. 2 cm long, dark; lateral sepals to 12 mm long, keel entire or papillate; seeds to 2 mm long. Wet tepui meadows, 1800–2300 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 487.

Zamia 575

ZAMIACEAE by Dennis Wm. Stevenson Plants dioecious, stems subterranean to arborescent, when arborescent often clothed in persistent leaf bases. Crown of few (1–6) to many (100+) pinnately compound leaves; pinnae 10–60 in subopposite to opposite pairs, densely pubescent when young, glabrous when mature, venation dichotomous. Strobili usually emerging from center of crown, 1–several; sporophylls flat to peltate; microsporangiate (pollen) strobili composed of numerous microsporophylls with sporangia located on the abaxial surface; microsporangia clustered in groups of 2–5 and dehiscing by longitudinal slits. Megasporangiate (ovulate) strobili composed of numerous ovulebearing, stalked megasporophylls, each with 2 ovules. Seed coat when mature consisting of an outer fleshy layer which is usually red or orange-red but may be green, white, or yellow, and an inner stony layer. Pantropical to warm temperate regions of North America, Central America, West Indies, South America, Africa, and Australia; ca. 8 genera and 100 species, 1 genus and 2 species in the flora area. 1. ZAMIA L., Sp. Pl. ed. 2: 1659. 1763, nom. cons. Stems subterranean and tuberous, when arborescent trunks smooth. Leaves generally 2–10; petioles with stipules, smooth or armed with scattered prickles. Pinnae ovate to linear, entire or more commonly serrate especially at the tips. Strobili pedunculate, tomentose, tan to dark brown or rarely green; sporophylls arranged in definite vertical orthostichies, the exposed tips thick, hexagonal, truncate, broader than high, and often with a central depression. Seeds red to orangered, rarely yellow or cream. U.S.A. (southern Florida), Mexico, Central America, West Indies, south in South America to northern Bolivia; 60 species, 3 in Venezuela, 2 of those in the flora area. Key to the Species of Zamia 1. 1.

Petiole and rachis densely prickled throughout, prickles often branched; leaflets papyraceous .............................................................. Z. amazonum Petiole sparsely prickled in lower 1/2, rachis smooth, prickles unbranched; leaflets coriaceous ...................................................................... Z. lecointei

Zamia amazonum D.W. Stev., Flora de Colombia 21: 33, fig. 3. 2001. Stem subterranean; leaves 2–6, to 2.5 m long, oval to elliptic; petiole often ferruginous, armed with small to stout branched prickles; rachis terete, 0.5–1 m long, with 10–50 pairs of leaflets, armed with prickles in the lower third; leaflets chartaceous to papyraceous, oblong-lanceolate to lanceolate, acuminate at apex, often denticulate in upper half. Open to fairly closed forest understory with sandy soils subject to sporadic

flooding, 50–500 m; Amazonas (Río Mawarinuma, Río Siapa, Yavita). Widespread in Amazonian Colombia, Peru, and Brazil, with disjunct populations in the Chocó area of Colombia. Zamia amazonum is most often found in sandy soils that experience sporadic flooding for short periods. Zamia lecointei Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 9, t. 1. 1915. —Zamia ulei subsp. lecointei (Ducke) Ducke, Arch.

576

Z AMIACEAE

Inst. Biol. Veget. Rio de Janeiro 2: 27. 1935. Zamia obidensis Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 20. 1922. Zamia jirijirmensis R. E. Schult., Mutisia 15: 2, fig. p. 5, 1953. Stem subterranean; leaves 2–4, erect or slightly curved, oblong, petiole glabrous, to 75 cm long, sparsely armed with stout prickles; rachis terete, unarmed, to 1 m long, 30– 40 pairs of leaflets; leaflets subcoriaceous to coriaceous, linear-lanceolate, subfalcate, long acuminate at apex, margins entire or

rarely with 2 or 3 indistinct teeth near the apex of lower margin. Open and/or frequently burned areas on well-drained, sandy to pure sand soils or igneous rock formations, 50–200 m; Bolívar (Río Parguaza), Amazonas (Puerto Ayacucho, Rincones de Chacorro, Río Pacimoni-Yatúa). Colombia, Peru (one collection), Brazil (north of the Rio Amazonas). ◆Fig. 497. Zamia lecointei is often found growing in white sands within the flora area. The drier the habitat the more coriaceous and narrow are the leaflets.

Fig. 497. Zamia lecointei

Renealmia 577

ZINGIBERACEAE by Hiltje Maas and Paul J. M. Maas Perennial, aromatic herbs. Rhizome often fleshy. Stems terete, straight or spirally contorted. Leaves with an open leaf sheath, ligulate, distichous. Inflorescence terminal on a leafy shoot, or on a separate, leafless shoot. Flowers bisexual, zygomorphic, solitary or arranged in cincinni in the axil of bracts and bracteoles. Calyx tubular, shortly 3-lobed; corolla tubular at the base, the dorsal lobe somewhat larger than the 2 lateral ones; labellum staminodial, almost always the most conspicuous feature of the flower. Lateral staminodes 2; stamen 1, sometimes petaloid; anther with 2 thecae, the connective sometimes appendaged. Style filiform, placed in a groove between the thecae; stigma cup-shaped or bilamellate; nectarial glands at the base of the flower tube, or sunken in the apex of the ovary (septal glands). Ovary inferior, 3-locular (rarely 2-locular, seldom 1-locular), placentation axile or parietal; ovules many, anatropous. Fruit an indehiscent or loculicidally dehiscent capsule or fleshy berry, often crowned by the calyx. Seeds 1–many, arillate. Tropics and subtropics, particularly in Asia; 47 genera and 1000 species, 1 genus and 8 species in the flora area. 1. RENEALMIA L. f., Suppl. Pl. 7. 1781 [1782], nom. cons.. Tall to low, aromatic herbs. Leaves distichous, sheaths open, smooth, striate or reticulate; petiole small or absent; ligule small, truncate; blade ± narrowly elliptic, mostly superficially glabrous (but nearly always covered with microscopic trichomes). Inflorescence a thyrse or raceme or very rarely a spike, terminating a separate, leafless shoot or rarely terminating a leafy shoot; indument composed of simple, furcate, or plurifurcate trichomes; bracts green, pink, red, or yellow, herbaceous to membranous mostly deciduous, subtending 1–10-flowered cincinni; bracteoles tubular and closed before anthesis, or cup-shaped and open even before anthesis; pedicels short or relatively long. Calyx tubular or turbinate, shortly 3lobed; corolla yellow to white, tubular at the base, 3-lobed; labellum composed of a basal claw and an erect and tubular or horizontally spreading limb. Stamen subsessile. Ovary 3-locular; nectarial glands implanted at the top of the ovary, partly or completely enclosing the style base; stigma cup-shaped. Capsule loculicidally dehiscent from base to apex. Seeds 1–200, glossy brown, irregularly ellipsoid, with a large, membranous, red, orange, yellow, or white aril. Neotropics, tropical Africa; ca. 75 species, 11 in Venezuela, 8 of these in the flora area. Key to the Species of Renealmia 1. 1. 2(1). 2.

Inflorescence a raceme (the branches 1-flowered); fruit many-seeded (> 50), thick-walled (2–8 mm in living material) ................................. 2 Inflorescence a thyrse (the branches 2–many-flowered); fruit few-seeded (1–25), thin-walled (1–1.5 mm in living material) ............................... 3 Bracteole tubular and closed before anthesis; pedicels 2–8 mm long; calyx circumscissile; plants usually from low altitudes ...................... R. alpinia Bracteole cup-shaped and open even before anthesis; pedicels 1–2 mm long; calyx not circumscissile; plants usually from high altitudes ................................................................................................ R. thyrsoidea

578

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

Z INGIBERACEAE

Leaves very narrow (1.5–3.5 cm wide); scape of inflorescence 3–10 cm long; leafy stems 20–50 cm high ........................................ R. monosperma Leaves much wider; scape and leafy stems much longer ......................... 4 Leaf sheaths distinctly reticulate, chocolate-brown in herbarium material .......................................................................................................... 5 Leaf sheaths smooth, striate, or only very slightly reticulate ................. 6 Capsule red; calyx deciduous during fruiting; cincinni 1–3(4)-flowered ................................................................................................ R. guianensis Capsule green, maturing black; calyx persistent during fruiting; cincinni 2–6(–10)-flowered .................................................................. R. floribunda Scape of inflorescence prostrate; leaves 40–100 × 6–20 cm ..... R. orinocensis Scape of inflorescence erect; leaves 10–65 × 5–15 cm .............................. 7 Capsule red, not prominently 3-valvate; calyx turbinate, 4–7 mm long ................................................................................................. R. aromatica Capsule black, prominently 3-valvate, crowned by a small (1–3 mm long), tubular calyx remnant ........................................................... R. microcalyx

Renealmia alpinia (Rottb.) Maas, Acta Bot. Neerl. 24: 474. 1975 [1976]. —Amomum alpinia Rottb., Soc. Med. Havn. Collect. 2: 245, 248, t. 1. 1775. Renealmia exaltata L. f., Suppl. Pl. 79. 1781 [1782]. Renealmia raja Petersen in Mart., Fl. Bras. 3(3): 44. 1890. Herb. Secondary forests, flooded forests, swamps, or on stream banks, usually at low altitudes, 50–900 m; Delta Amacuro (Catalina, Tucupita), Bolívar (widespread), Amazonas (widespread). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Falcón, Lara, Mérida, Monagas, Portuguesa, Táchira, Zulia; Mexico, Central America, Lesser Antilles, widespread in tropical South America. Renealmia aromatica (Aubl.) Griseb., Abh. Königl. Ges. Wiss. Göttingen 7: 275. 1857. —Alpinia aromatica Aubl., Hist. Pl. Guiane 3. 1775. Renealmia occidentalis var. ß cylindrica Petersen in Mart., Fl. Bras. 3(3): 45. 1890. Renealmia lutea J.R. Johnst., Contr. Gray Herb. 37: 199, t. 26, fig. 1. 1909. Renealmia occidentalis var. macrocarpa Loes., Notizbl. Bot. Gart. Berlin-Dahlem 10: 63. 1927. Renealmia occidentalis var. occidentalis sensu Maas, Fl. Neotrop. Monogr. 18: 103, fig. 39a-e. 1977; Fl. Venezuela 9(2): 221. 1982. Herb. Forests, sea level to 1500 m; Delta

Amacuro, Bolívar (El Dorado, Río Paragua), Amazonas (upper Río Orinoco). Apure, Aragua, Barinas, Carabobo, Nueva Esparta, Portuguesa, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 498. Renealmia floribunda K. Schum. in Engl., Pflanzenr. IV. 46(Heft 20): 300. 1904. Renealmia pedicellaris Gleason, Bull. Torrey Club 52: 192. 1925. Herb. Rain forests, savanna forests, forest margins, often on white sand, 100–400 (–1400) m; Bolívar (Macizo del Chimantá, Río Cuyuní basin, upper Río Paragua), Amazonas (Cerro Yapacana base, Maroa, Piedra Arauicaua, Río Casiquiare, Río Cunucununma, San Carlos de Río Negro, San Pedro, Yavita). Trinidad, Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. Renealmia guianensis Maas, Acta Bot. Neerl. 24: 477. 1975 [1976]. Herb. Forests, near sea level to 900 m; Delta Amacuro (near El Palmar, Río Cuyubini), Bolívar (Altiplanicie de Nuria). Guyana, Suriname, French Guiana, Brazil (Amapá, Bahia, Pernambuco). Renealmia microcalyx Maas & H. Maas, Notes Roy. Bot. Gard. Edinburgh 46: 315. 1990. Herb. Primary forests, secondary forests along roadsides, savannas, 100–200 m; Amazonas (Río Negro basin). Brazil (Amazonas: Rio Negro).

Renealmia 579

Fig. 498. Renealmia aromatica

Fig. 499. Renealmia orinocensis

Renealmia monosperma Miq., Linnaea 22: 80. 1849. Herb. Rain forests, 50–1000 m; Delta Amacuro (Río Amacuro), Bolívar (Gran Sabana, Río Abacapá, Río Caroní, Río Paramichi). Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amazonas, Roraima), Bolivia. Renealmia orinocensis Rusby, Descr. S. Amer. Pl. 8. 1920. Herb. Rain forests, near sea level to 1000 m; Delta Amacuro (lower Río Orinoco near Santa Catalina), Bolívar (Río Chicanán, Si-

580

Z INGIBERACEAE

erra de Lema). Guyana, Suriname, French Guiana. ◆Fig. 499. Renealmia thyrsoidea (Ruiz & Pav.) Poepp. in Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 25, t. 134. 1838. —Amomum thyrsoideum Ruiz & Pav., Fl. Peruv. 1: 2, t. 2. 1798. Herb. Montane forests. Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, 1 in Venezuela. Renealmia thyrsoidea subsp. chrysantha

Maas is known only from Guyana and Suriname. It is characterized by golden yellow bracts. R. thyrsoidea subsp. thyrsoidea Bracts red to orange. 1700–2100 m; Amazonas (Cerro Yutajé, Sierra de la Neblina). Anzoátegui, Aragua, Barinas, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Subandean regions of Costa Rica, Panama, Trinidad, Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia.

ZYGOPHYLLACEAE by Gerardo A. Aymard C. and Nidia L. Cuello A. Shrubs, small to medium-sized trees, or annual or perennnial herbs. Branches opposite, often swollen or jointed at the nodes, sometimes strongly scented and with sticky resin. Leaves opposite, rarely alternate, mostly even-pinnately compound, occasionally simple, 2-foliolate, or 3–7-parted, petiolate; leaflets sessile or nearly so, opposite, entire, generally inequilateral, flat or fleshy and sometimes terete; stipules 2, persistent or rarely deciduous, interpetiolar, leathery or spinescent. Inflorescences axillary or terminal, racemose or fasciculate (rarely cymose), or flowers solitary on axillary peduncles. Flowers bisexual, actinomorphic, rarely zygomorphic; peduncle ebracteate. Sepals (4)5(6), free or rarely basally connate, persistent or deciduous. Petals (4)5(6), free, clawed, twisted in bud or sometimes lacking. Stamens (5)6–10(–12), in 1 or 2 whorls, often unequal, opposite the petals, hypogynous; filaments broadened at the base, free, naked or with basal scales; anthers introrse, 2-thecate, opening longitudinally. Ovary with nectary disk usually well developed, usually intrastaminal, or modified into a gynophore or base of the ovary, superior, 4-, 5-, or 10–12-locular, placentation axile; style 1, sometimes very short; stigmas 1–several, capitate or lobed; ovules 1–several in each locule, pendulous. Fruit a 2–6lobed loculicidal or septicidal berry, drupe, capsule, or a schizocarp breaking into few to several mericarps or nutlets. Seeds with endosperm hard and oily or lacking endosperm; embryo straight or curved; cotyledons linear or oblong. Pantropics; 25–30 genera and 200–250 species, 3 genera and 4 species in the flora area. Bulnesia arborea (Jacq.) Engl., commonly called “Vera,” is a tree of deciduous forests in the northern part of the Río Orinoco area. It is well known for its extremely hard wood, which is very durable and resistant to decay, and is sometimes planted in parks and gardens.

Guaiacum 581

Key to the Genera of Zygophyllaceae 1. 1. 2(1). 2.

Trees 5–15 m tall; petals blue, purple, or white; each locule with 8– 10 ovules .................................................................................. 1. Guaiacum Annual or occasionally perennial herbs; petals yellow or white to bright orange; each locule with 1–5 ovules ...................................................... 2 Leaflets in 3 or 4 pairs; each locule with 1 pendulous ovule; petals white to bright orange; fruit separating into 8–12 nutlets ......... 2. Kallstroemia Leaflets in 5–10 pairs; each locule with 2–5 ovules on axillary placentae; petals yellow; fruit separating into 5 nutlets ........................... 3. Tribulus

1. GUAIACUM L., Sp. Pl. 381. 1753. Trees to 15 m tall, wood very hard, heavy, and resinous. Leaves 3–15 cm long, opposite, petiolate, even-pinnately compound; leaflets in 2–6 pairs, glabrous, entire, inequilateral at least at base, broadly elliptic or obovate, 1–4.5 × 1–2.5 cm, stipules minute, caducous. Inflorescence terminal, fasciculate at the distal nodes, peduncles

Fig. 500. Guaiacum officinale

582

Z YGOPHYLLACEAE

1–4 cm long. Sepals 4 or 5, broadly ovate, unequal, 4–5 mm long; petals 4 or 5, obovate, slighty clawed, 12–13 × 7–8 mm, blue, purple, or white. Stamens 8–10 on an inconspicuous disk; filaments filiform; anthers cordate or sagittate at base. Ovary with a short stipe (gynophore), locules 2–5; ovules 8–10 per locule. Fruit a flattened septicidal capsule, with 2–5 prominent longitudinal lobes or rounded ridges or wings. Seeds solitary in each wing, ovoid to ellipsoid, 10–12 × 6 mm, enclosed in a fleshy aril. Mexico, Central America, Antilles, Colombia, Venezuela, Ecuador, Peru, Bolivia, cultivated in other tropical regions; ca. 5 species, 1 in Venezuela. Guaiacum officinale L., Sp. Pl. 381. 1753. —Guayacán. Evergreen tree 5–10 m tall; flowers fragrant; wood extremely hard. Cultivated and possibly naturalized, 50–300 m; northern Bolívar (Ciudad Bolívar, Isla Anacoco, middle Río Orinoco, Santa Rita, Tumeremo). Anzoátegui, Apure, Aragua, Carabobo, Dependencias Federales, Distrito Federal, Falcón, Guárico, Nueva Esparta, Portuguesa, Sucre,

Zulia; Panama, Antilles, Colombia, Guyana, Peru. ◆Fig. 500. Guaiacum officinale is the plant widely known as “lignum vitae” or “palo santo,” an extract of which was widely used in medicine as a stimulant and to increase perspiration. It was introduced into Europe in about the 17th century and was thought to be a cure for more than ten different diseases. It is also planted in parks and gardens as an ornamental for its blue flowers and dark green foliage.

2. KALLSTROEMIA Scop., Intr. Hist. Nat. 212. 1777. Annual or occasionally perennial herbs. Stems herbaceous to suffrutescent, prostrate to decumbent or ascending. Leaves opposite, even-pinnate, 1–8.5 × 1.5–5 cm; leaflets entire, elliptic to broadly oblong or obovate, 3–29 × 1.5–17 mm, in 3 or 4 pairs, one leaflet of each pair alternately smaller or aborted; petiolule ≤ 1 mm long; stipules 2, interpetiolar, foliaceous, narrowly to broadly falcate, acuminate, 3–10 × 1–3 mm. Inflorescences usually represented by solitary, axillary or pseudoaxillary flowers. Flowers pedunculate, 0.7–6 cm diameter, appearing alternate on stem. Sepals 5 or 6, free, foliaceous, lanceolate to broadly ovate, 2–16 × 1–5 mm, hirsute, hispid, or strigose, persistent or rarely caducous; petals 4–6, free, obovate or obcordate, 3–3.5 × 1.5–2 cm, white to bright orange. Stamens 10–12, inner ones smaller and with basal gland, shorter than petals; filaments filiform, subulate, inserted in 2 whorls in the disk, disk fleshy, 10–12-lobed; anthers globose or ovoid to oblong-linear, bilobed, sub-basifixed to versatile. Ovary sessile, glabrous to pubescent, 8–12locular, 1 pendulous ovule in each locule; style simple; stigma capitate to clavate. Fruit 8–12-angled, muricate or tuberculate, breaking into 8–12 indehiscent, 1seeded nutlets, leaving the persistent axis. Seeds oblong to ovoid; endosperm lacking; embryo straight. Neotropics (most diverse in southern Mexico and southeastern U.S.A.), naturalized in West Africa and India; ca. 25 species, 2 in Venezuela, both of these in the flora area. Key to the Species of Kallstroemia 1. 1.

Leaflets oblong to elliptic; sepals ovate, almost covering the fruit; ovary and fruit ovoid, glabrous, occasionally strigose ....................... K. maxima Leaflets elliptic to obovate; sepals linear-lanceolate not covering the fruit; ovary and fruit pyramidal, densely pilose ................... K. pubescens

Tribulus 583

Fig. 501. Kallstroemia maxima

Kallstroemia maxima (L.) Hook. & Arn., Bot. Beechey Voy. 282. 1838. —Tribulus maximus L., Sp. Pl. 386. 1753. Annual herb; stems branched, prostrate to decumbent; flowers white to pale orange. Disturbed areas, 50–100 m; Bolívar (Ciudad Bolívar, Upata). Anzoátegui, Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Trujillo, Zulia; U.S.A. (California, Georgia, Florida, New Mexico, South Carolina, Texas), Mexico, Central America, Antilles, Colombia, Ecuador, Brazil (Bahia). ◆Fig. 501.

Kallstroemia pubescens (G. Don) Dandy, Kew Bull. 10: 138. 1955. —Tribulus pubescens G. Don, Gen. Hist. 1: 769. 1831. Kallstroemia caribaea Rydb., N. Amer. Fl. 25(2): 111. 1910. Annual herb; stems branched, prostrate to decumbent; flowers white to yellow. Disturbed areas, 50–100 m; Bolívar (Ciudad Bolívar). Anzoátegui, Distrito Federal, Falcón, Miranda, Nueva Esparta, Sucre, Táchira, Zulia; U.S.A. (Florida), Mexico, Central America, Antilles, Colombia, Ecuador, Peru, introduced into west Africa and India.

3. TRIBULUS L., Sp. Pl. 386. 1753. Annual or occasionally perennial herbs. Stems herbacous to slightly woody, prostrate, sometimes ascending, pubescent or rarely glabrous. Leaves opposite, even-pinnate; leaflets oblong to ovate-oblong or elliptic, asymmetric at base, similar, in 5–10 pairs, one of each pair alternately small or aborted; lanceolate or subulate, membranous. Inflorescences of solitary axillary flowers borne on long peduncles. Sepals 5, free, lanceolate, pubescent outside, deciduous or rarely persistent; petals 5, free, deciduous, obovate, 2–22 × 1–18 mm, narrowed at the base, bright yellow. Stamens (5)10; inner filaments shorter, without scales; intrastaminal glands 10, free or connate beneath the base of the ovary; anthers oblong-cordate. Ovary sessile, globose to ovoid, covered with erect bristle-like trichomes, 5-locular, with lobed basal disk; ovules 5–10 per locule. Fruit of 5 hard mericarps, tuberculate or spinose, separating at maturity into 5 bony carpels, each divided by oblique or

584

Z YGOPHYLLACEAE

transverse septae into 3–5, 1-seeded locules; seeds oblong-obovate; endosperm absent; embryo straight. Paleotropics, introduced and naturalized in the New World; ca. 25 species, ca. 3 in Venezuela, 1 of these in the flora area. Tribulus zeyheri Sond., Fl. Cap. 1: 353. 1860. Widespread in the Paleotropics, naturalized in the Neotropics; 2 subspecies, 1 in Venezuela. T. zeyheri subsp. macranthus (Hassk.) Hadidi, Fl. Trop. East Afr. 7. 1985. —Tribulus macranthus Hassk., Flora 48: 403. 1865. Prostrate herb; pedicels 15–40 mm long; petals yellow. Weedy areas near towns, 50– 100 m; Bolívar (near Puerto Ordaz). Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Nueva Esparta, Sucre, Zulia; Central America, Antilles, Colombia, Peru. ◆Fig. 502.

All herbarium specimens of the genus Tribulus in Venezuela are referred to T. cistoides L. (Lasser, T. 1971. Zygophyllaceae. In: Flora de Venezuela. Vol. 3). However, we use the name T. zeyheri subsp. macranthus here because the specimens from the flora area match the main morphological characters used to separate T. zeyheri subsp. macranthus from T. cistoides. Tribulus zeyheri subsp. macranthus has the style 0.5– 1 mm long or absent; stigma pyramidal or cylindrical, 2–4 mm long; petals 4–20 × 4–20 mm; pedicel to 2 times as long as the subtending leaf. In contrast, T. cistoides has the style 2–4 mm long; stigma hemispherical; petals 8–10 × 4–5 mm; pedicel shorter than to as long as the subtending leaf.

Fig. 502. Tribulus zeyheri subsp. macranthus

Addendum 585

Addendum When Volume 4 of the flora was published, we did not realize at the time that Ceratophyllum was present in the flora area. Therefore the family is added on here at the end of the final volume.

CERATOPHYLLACEAE by Paul E. Berry Rootless, floating to submerged herbs, sometimes with root-like branches anchoring the plant, glabrous. Stems usually branched (one per node). Leaves 1–4 times forked, rigid and often brittle, in whorls of 3–10, with apical minute teeth and bristles; stomata and stipules lacking. Flowers unisexual (plants monoecious), solitary, extra-axillary, staminate and pistillate flowers usually on alternating nodes. Perianth segments 8–12, linear, basally connate. Androecium with (3–)10–20(–27) stamens, spirally arranged on a flat receptacle, not clearly differentiated into filament and anther, but the connective extended into 2 points, extrorsely dehiscent, pollen smooth and inaperturate. Gynoecium with 1 carpel and a single orthotropous ovule. Fruit an achene with a persistent style and often with a spiny margin. Seed with thin testa, cotyledons 2, linear, from an annular primordium. Cosmopolitan; 1 genus and 6 species, 2 species in Venezuela, 1 of these in the flora area. The second species in Venezuela in the northern part of the country is Ceratophyllum demersum L. When sterile, the genus resembles other aquatics such as Myriophyllum (Haloragidaceae), Najas (Najadaceae), or Chara (a green alga), but it can be readily distinguished by its forked leaves. 1. CERATOPHYLLUM L., Sp. Pl. 992. 1753. Description and distribution as for family. Ceratophyllum muricatum Cham., Linnaea 4: 504. 1826. Aquatic herb, algae-like in appearance, with whorled, linear-bifurcate leaves; fruit spiny-margined. Cosmopolitan; 3 subspecies, 1 in Venezuela. Ceratophyllum muricatum subsp. australe (Griseb.) Les, Syst. Bot. 13: 85. 1988. —Ceratophyllum australe Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 14. 1879. Creeks and rivers, near sea level to 100 m; Delta Amacuro (El Paso at Capure). Widely scattered in lowland northern Venezuela; widely distriubted in the Neotropics and subtropics. ◆Fig. 503.

Fig. 503. Ceratophyllum muricatum subsp. australe

Appendix A List of new names published in this volume

Sterculia kayae P.E. Berry, nom. nov. .................................................................... 266 Xyris sulcatifolia Kral, nom. nov. .......................................................................... 556

586

Index Entries in Roman type = accepted names of taxa and vernacular names Entries in italics = synonyms Page numbers in bold = illustrations

—A— Abari, 182 Abolboda, 527 acaulis, 529 var. acaulis, 529 var. scaposa, 530 acicularis, 530 var. acicularis, 530 var. granularis, 530 americana, 530 var. imberbis, 530 bella, 530 brasiliensis, 532 chapadensis var. pauciflora, 532 ciliata, 530 dunstervillei, 530 ebracteata, 530 var. brevifolia, 531 var. ebracteata, 531 excelsa, 532 ×glomerata, 531 gracilis, 532 grandis, 531 var. grandis, 531 var. guayanensis, 531 var. minor, 531 var. rigida, 531 imberbis, 530 killipii, 531, 533 linearifolia, 532, 533 macrostachya, 532 var. angustior, 532 var. macrostachya, 532

var. robustior, 532, 533 minima, 531 neblinae, 532, 533 paniculata, 532 pervaginata, 531 poeppigii, 530 psammophila, 531 ptaritepuiana, 538 pulchella, 532 rigida, 531, 532 scabrida, 532 sceptrum, 538 schultesii, 534 sprucei, 534 uniflora, 534 Achetaria, 146 guianensis, 146, 147 scutellarioides, 147 Achlyphila, 534 disticha, 534, 535 Achras caimito, 118 venosa, 124 Achrouteria pomifera, 94 Acopanea, 316 ahogadoi, 318 Acuchipuru mira, 118 Adarro-Itipi, 418 Adéketón-yek, 438 Adiscanthus, 5 fusciflorus, 5, 6 Aegiphila, 409 bracteolosa, 411 cowanii, 413

587

elata, 411 var. elata, 411 var. macrophylla, 411 filipes, 411 glandulifera, 411 var. pyramidata, 411, 414 guianensis, 412 hystricina, 412 hoehnei, 412 var. venezuelensis, 412 integrifolia, 412, 416 var. guianensis, 412 intermedia, 412 laevis, 412 laxiflora, 412, 417 lewisiana, 412, 416 macrantha, 412, 415 macrophylla, 411 membranacea, 412 mollis, 413, 415 var. intermedia, 413 var. mollis, 413 var. surfaceana, 413 parviflora, 413 pendula, 413 perplexa, 413 racemosa, 413 roraimensis, 413, 417 spruceana, 413 steyermarkii, 413 var. macrophylla, 413 var. obtusifolia, 413 surfaceana, 413

588

I NDEX

[Aegiphila] venezuelensis, 413 var. serrata, 414, 414 var. venezuelensis, 414 Agalinis hispidula, 149 Agardhia, 504 Aguacatillo blanco, 182 Akaymota-yek, 459 Albahaca, 13 Alectra, 147 aspera, 147 fluminensis, 147 stricta, 147, 148 Alegria, 352 Algodoncillo, 272 Allophylus, 48 amazonicus, 48 occidentalis, 49 parimensis, 48 punctatus, 48 racemosus, 49, 49 Almendrón cunavichero, 175 Alpinia aromatica, 578 Alsodeia, 456 brevipes, 457 camptoneura, 458 falcata, 458 var. grandifolia, 460 flavescens, 458 guianensis, 460 var. lindeniana, 459 lindeniana, 459 macrocarpa, 459 ovalifolia, 459 paniculata, 460 pubiflora, 460 racemosa, 461 sprucei, 463 Amarun-yek, 118 Amasonia, 417 arborea, 418 campestris, 418, 418 lasiocaulos, 418 obovata, 418 spruceana, 418 Amblyorhinum, 403 Amomum alpinia, 578 thyrsoideum, 580 Ampelocera, 386 edentula, 386, 387

Amphilochia, 504 Amphirrhox, 447 longifolia, 447, 448 Anakasha, 453 Anamorpha, 261 Anamu, 122 Anapora-yek, 175 Anepurayek, 175 Angelón, 149 Angelonia, 149 salicariifolia, 148, 149 Angostura, 6 cuspare, 6 pentagyna, 10 ramiflora, 10 trifoliata, 6, 7 ucayalina, 11 ulei, 11 Anisantherina, 149 hispidula, 149, 150 Antichorus, 350 Aparo, 212 Apeiba, 345 albiflora, 346 aspera, 346 burchellii, 346 echinata, 348 var. macropetala, 346 glabra, 346 hypoleuca, 348 macropetala, 346 membranacea, 346, 347 petoumo, 347, 348 schomburgkii, 348 surinamensis, 348 tibourbou, 347, 348 var. krukofii, 346 var. membranacea, 348 var. rugosa, 346 uittienii, 348 Apocaulon, 8 carnosum, 8, 8 Aratitiyopea, 536 lopezii, 536 var. lopezii, 535, 536 Arautamakatadek, 15 Arbol de vaca, 124 Arbol del danto, 182 Archytaea, 314 angustifolia, 315, 315 multiflora, 315 triflora, 315, 315 Aruhuhu, 70

Arujujuj, 70 Arunillo, 459 Arusi-moroconi, 70 Aru-simu, 70 Asterophorum, 348 Astropus, 273 Aubletia, 345 Avicennia, 419 germinans, 419, 419 f. venezuelensis, 420 var. cumanensis, 419 var. guayaquilensis, 420 nitida, 419 schaueriana, 420 f. candicans, 420 tomentosa var. cumanensis, 419 var. guayaquilensis, 419 Awadi, 182 Ayenia, 249 magna, 250, 250 —B— Bacopa, 151 aquatica, 152 callitrichoides, 150, 152 chamaedryoides, 163 ciliata, 153 conferta, 154 domingensis, 154 gratioloides, 152 innominata, 152 laxiflora, 153 monnierioides, 153 punctata, 153 reflexa, 154 repens, 150, 153 reptans, 153 salzmannii, 153 serpyllifolia, 153 sessiliflora, 154 stricta, 154 Balata, 107 Bampara, 124 Bampara amarilla, 123 Barbacenia, 405 alexandrinae, 406 celiae, 405, 405 Bartramia, 358 indica, 360 Bassovia macrophylla, 246

I NDEX

Batata de lagartilla, 212 Bejuco barba de mono, 70 Bejuco barbasco, 169 Bejuco corona e cristo, 186 Bejuco cresta de gallo, 69 Bejuco de arrendajo, 212 Bejuco de caro, 496 Bejuco de corona, 186, 187 Bejuco de fuente, 495 Bejuco de judío, 494 Bejuco de lora, 255 Bejuco de loro, 70 Bejuco de mono, 496 Bejuco de pereza, 412 Bejuco de raya, 494 Bejuco de sapo, 494, 495 Bejuco de seguera, 449, 450 Bejuco marrón, 69 Bejuco mayo, 432 Bejuco moreno, 80 Bejuco negro, 368 Bejuco parcha morada, 494 Bejuco Santa Ana, 70 Bejuco tres filos, 70 Bejuco tres filos rebalsero, 70 Benjaminia, 154 reflexa, 154, 154 utriculariiformis, 154 Berga de toro, 70 Beyrichia scutellarioides, 147 Boboyo winae, 390 Bocsuo amarillo, 37 Bocsúo negro, 36 Boehmeria, 392 cylindrica, 392, 393 hirta, 394 Bojúa, 37 Bonnetia, 316 ahogadoi, 318, 323 bolivarensis, 318 celiae, 318 chimantensis, 318 cordifolia, 318 crassa, 318 dinizii, 319 euryanthera, 318 fasciculata, 319 guaiquinimae, 319 huberiana, 319 jauaensis, 319 kathleenae, 319, 322

lanceifolia, 319, 322 liesneri, 319 maguireorum, 319, 324 martiana, 320 var. venadoensis, 320 multinervia, 319, 324 neblinae, 319, 322 paniculata, 319, 321 parviflora, 319 phelpsii, 320 ptariensis, 320 roraimae, 320, 323 roseiflora, 320 rubicunda, 320 sessilis, 320, 321 steyermarkii, 320, 323 tepuiensis, 321, 323 subsp. minor, 321 toronoensis, 318 tristyla, 321, 323 subsp. nervosa, 321 wurdackii, 321, 324 Bonnetiaceae, 310, 313 Bonplandia, 6 angostura, 6 trifoliata, 6 Bontia germinans, 419 Borrachera, 438 Bosí, 37 Bosnó, 39 Bosú, 36, 37 Bosúa, 37, 39 Bosuo, 36 Bosúo, 37 Bouchea, 420 prismatica, 420 var. longirostra, 420, 421 Boxúo amarillo, 37 Boxúo negro, 39 Boya de mono, 103 Brachistus macrophyllus, 246 Braddleya, 447 Bradleya, 447 Bretónica, 263 Bretónica morada, 265 Brotera, 352 Brotobroma, 259 Brugmansia, 205 Brunfelsia, 196 imatacana, 196, 197

589

uniflora, 197 Bubroma grandiflorum, 270 Buchnera, 154 elongata, 156 grandiflora, 159 lithospermifolia, 156 lobelioides, 156 longifolia, 156 macrocarpa, 156 palustris, 156, 156 pusilla, 156 rosea, 156 spruceana, 156 ternifolia, 156 weberbaueri, 157 Byttneria, 251 aristeguietae, 252 caripensis, 252 catalpifolia, 252 subsp. catalpifolia, 252 divaricata, 253 var. divaricata, 253, 253 genistella, 253, 254 obliqua, 253, 254 piresii, 254, 255 rhamnifolia, 255 scabra, 255 uaupensis, 255 venezuelensis, 253 —C— Cabeza de baba, 122 Cabeza de babocilla, 122 Cabeza de culebra, 357 Cabeza de mono, 346 Cabeza de negro, 346, 348, 350, 352 Cabeza de tamatama, 357 Cacaguillo, 463 Cacao, 269 guianensis, 270 sylvestris, 270 Cacao de monte, 270 Cacao montañero, 261 Cacao montero, 270 Cacao-rana, 270 Cachito, 109, 120 Cacho, 98 Cacho de toro, 70 Cacho de venado, 122, 459 Cacho venado, 118

590

I NDEX

Caconapea, 151 auriculata, 153 conferta, 154 debilis, 153 gratioloides, 152 sessiliflora, 154 stricta, 154 Cadillo, 276, 360 Café de venado, 448 Cafecillo, 452 Caijiro, 186 Caimito, 93 Caimito amarillo, 121 Caimito blanco, 120 Caimito largo, 93 Caimito morado, 93 Caimito negro, 121 Cajón, 109 Cajou, 109 Cakak-orai-yek, 134 Callicarpa integrifolia, 412 Calyptrion, 448 Camaruco, 266 Caná-acu-dajay, 356 Candela, 418 Canelito, 518 Caniba, 516 Canoa, 354 Capraria, 157 biflora, 157, 158 f. hirsuta, 157 crustacea, 161 Capsicum, 197 annuum, 197 var. glabriusculum, 198, 198 frutescens, 198 hispidum var. glabruiusculum, 198 macrophyllum, 246 solanaceum var. macrophylla, 246 Capure, 94, 118, 124 Capurillo, 94, 110 Caramacarte, 120 Carapilla, 178 Carapillo, 60 Carapo blanco, 86 Carbón, 172 Cardiospermum, 49 corindum, 50, 50

grandiflorum, 50 halicacabum, 51 var. halicacabum, 51 var. microcarpum, 51 macrolophum, 57 microcarpum, 51 Cariaquito, 427 Cariaquito blanco, 427 Cariaquito colorado, 427 Cariaquito morado, 428 Caro Montañero, 56 Carpodiptera schomburgkii, 350 Caruache, 41 Casabe colorado, 70 Casabote, 172 Castilleja, 157 arvensis, 157, 158 Cedro blanco, 175 Cedro dulce, 175 Cedrus alternifolia, 352 Celedonia, 374 Celtis, 387 iguanaea, 388, 389 Cenato, 356 Cenicero, 441 Ceratophyllaceae, 585 Ceratophyllum, 585 australe, 585 muricatum, 585 subsp. australe, 585, 585 Cerezillo, 422 Cerezo, 175 Cestrum, 198 acuminatissimum, 201 alternifolium, 200 var. mitanthum, 200 baenitzii, 201 glabrescens, 200, 202 grande, 200 latifolium, 200, 202 var. tenuiflorum, 200 megalophyllum, 201 neblinense, 200 oliganthum, 200 var. latifolium, 200 prieurei, 200 racemosum, 200 var. grande, 200 reflexum, 201, 201 schlechtendahlii, 201

schulzianum, 201 strigilatum, 203 subuniflorum, 203 tenuiflorum, 200 var. glabrescens, 200 tubulosum, 203 Chaetochilus, 216 Chalcas, 18 exotica, 19 paniculata, 19 Chaparrillo, 52 Chaparro de agua, 52 Chaparro manteco, 94 Chapominaji, 348 Chicle, 99, 107, 109 Chicle de capure, 99 Chicle rosado, 120 Chimaco, 502, 506 Chipitudek, 109 Chiquita blanca, 14 Chirin-yo, 117 Chloerum, 527 Christiana, 348 africana, 349, 350 cordifolia, 350 madagascariensis, 350 Chromolucuma, 91 rubriflora, 91, 92 Chrysophyllum, 91 amazonicum, 93 ambelaniifolium, 117 anomalum, 117 aquaticum, 109 argenteum, 93 subsp. auratum, 93 auratum, 93 beardii, 134 cayennensis, 118 cuspidatum, 98 ficoides, 100 humbodltianum, 109 lucentifolium, 94 subsp. pachycarpum, 94 monachinoanum, 134 obtusatum, 109 pomiferum, 94, 95 prieurii, 94, 96 reticulatum, 122 sanguinolentum, 94 subsp. balata, 94, 96 subsp. sanguinolentum, 95 subsp. spurium, 95

I NDEX

schomburgkianum, 136 sparsiflorum, 95, 95 steyermarkii, 95 ucuquirana-branca, 97 ulei, 100 Chudumiro, 122 Chupón, 107, 120, 134, 136, 137 Chuponcillo, 117, 119 Ciponima guianensis, 291 Ciruela, 175 Ciruelillo, 175 Cissus, 493 alata, 494, 497 bonariensis, 495 cervii, 495 cordifolia, 496 descoingsii, 494, 498 erosa, 494, 498 var. salutaris, 495 gibertii, 495 gongylodes, 495 guaranitica, 495 haematantha, 495 hassleriana, 496 lucida, 494 martiniana, 496 ovata, 496 palmata, 495, 497 var. balansaeana, 495 paraguayensis, 495 parkeri, 496 pubescens, 494 quadrialata, 494 rhombifolia, 494 salutaris, 494 sexangularis, 495 sicyoides, 496 smilacina, 496 spinosa, 495 tamoides, 496 trianae, 496 var. martiniana, 496 tricuspis, 495 venezuelensis, 496, 499 verticillata, 496 subsp. laciniata, 496 subsp. verticillata, 496 Citharexylum, 420 amazonicum, 422 fruticosum, 423 var. brittonii, 423

macrophyllum, 422, 422 poeppigii, 422 f. anomalum, 422 spinosum, 422, 423 venezuelense, 423 sp. A., 423 Citrosma bifida, 181 cristata, 182 cuspidata, 182 decipiens, 182 guianensis, 182 reginae, 183 Clavija, 325 chermontiana, 328 imatacae, 326, 328 lancifolia, 328 subsp. chermontiana, 327, 328 Clavito, 200 Clavo de especie amarilla, 379 Clavo de especie de sabana, 379 Clerodendrum, 423 fragrans, 423 var. pleniflora, 423 philippinum, 423, 424 ternifolium, 424 thomsonae, 424 Coco de mono, 357 Cocuna, 231 Cojón de Berraco, 266 Cola de pava, 10, 14, 328 Comida de guacharaca, 49 Conchocarpus, 9 heterophyllus, 9 longifolius, 10, 10 ramiflorus, 10 ucayalinus, 10, 11 Conejemuro, 441 Conejilla, 451 Conejito, 380 Congrillo, 175 Congrio blanco, 507 Congrio morado, 506 Conobea, 158 aquatica, 159, 160 innominata, 152 scoparioides, 159 Conohoria, 456 flavescens, 458 riana, 463

Conók-pichoray, 315 Conserva, 124 Conuleum guianense, 182 Corchorus, 350 acutangulus, 351 aestuans, 351 argutus, 351 var. benthamii, 351 hirtus, 351, 351 var. orinocensis, 351 orinocensis, 351 pilolobus, 351 Corocuro, 119 Corona guaica, 187 Corynostylis, 448 arborea, 449, 449 carthagenensis, 449 hybanthus, 449 pubescens, 450 volubilis, 449, 450 Cotoperí, 82 Cotoperí montañero, 81 Cotoperíz, 82 Cotoperíz montañero, 81 Cotupería, 81 Cuberu, 106 Cucullaria, 512 excelsa, 518 ferruginea, 516 grandis, 517 tetraphylla, 518 tomentosa, 518 Cucuna, 231 Cuehiemuni, 70 Culebra, 182 Cumanagota, 182 Cumanagoto, 182 Cuna baruk, 70 Cupana, 69 Cupania, 51 cinerea, 52, 54 hirsuta, 52, 54 hispida, 52 kavanayena, 60 kukenanica, 52 latifolia, 52, 53 macrophylla, 82 papillosa, 52 roraimae, 52 rubiginosa, 52 scrobiculata, 53, 54 semidentata, 52

591

592

I NDEX

[Cupania] spruceana, 64 tepuiensis, 54 Cupuasú, 270 Cuquirana, 99 Cusparia, 6 angostura, 6 febrifuga, 6 officinalis, 6 pentagyna, 10 ramiflora, 10 trifoliata, 6 ucayalina, 11 Cuweyoja, 390 Cuyuri, 94 Cyphomandra, 203 allophylla, 226 bolivarensis, 203 tobagensis, 203, 204 —D— Dallama, 103 Daphnopsis, 334 americana, 335 subsp. caribaea, 335 caribea, 335 dircoides, 335 guaiquinimae, 336 longipedunculata, 336, 337 nevlingiana, 336 steyermarkii, 337, 338 Datura, 205 inoxia, 205, 206 Daujen, 504 Daujén, 518 Dawicú blanco, 458 Debraea, 500 floribundum, 502 violacea, 502 Decagonocarpus, 11 cornutus, 11 var. congestus, 11 oppositifolius, 11, 12 Dendrophthora, 465 basiandra, 468 crispula, 467 decipiens, 467 densifrons, 467, 470 elliptica, 467, 470 var. stenophylla, 467 fendleriana, 468, 471

hylaeana, 467 inaequidentata, 477 intermedia, 468 jauana, 468 lacryma-jobi, 467 macbridei, 468 microsoma, 468 nitidula, 469, 470 obliqua, 469 oligantha, 469 parvispicata, 468 roraimae, 469, 471 rubicunda, 467 tenuiflora, 469 tepuiana, 469, 471 Deprea, 206 orinocensis, 206, 206 Dewaado, 332 Dewaka, 110 Dilodendron, 55 bipinnatum var. elegans, 56 costaricense, 56 elegans, 55, 56 Diomma, 32 fruticosa, 32 ulei, 32 Diploon, 97 cuspidatum, 97, 98 venezuelana, 98 Dipterodendron costaricense, 56 elegans, 56 venezuelense, 56 Ditmaria, 500 floribunda, 502 violacea, 502 Dodonaea, 56 viscosa, 56, 57 Drimys, 524 brasiliensis var. ptaritepuiensis, 525 var. roraimensis, 525 roraimensis, 525, 525 Duranta, 424 coriacea, 425 erecta, 425, 425 obtusifolia, 425 plumieri var. alba, 425 repens, 425 var. canescens, 425 var. lopez-palacii, 425

sp. A, 426 sp. B, 426 —E— Ecclinusa, 98 atabapoensis, 99, 101 balata, 94 bullata, 99 grisebachii, 136 guianensis, 99, 102 lanceolata, 99, 101 orinocoensis, 99, 100 parviflora, 99 ramiflora, 100 spuria, 95 ucuquirana-branca, 97 ulei, 100 Ectozoma ulei, 207 Ehchipipin, 438 Ehparayo’, 16 Elaeoluma, 102 crispa, 103 glabrescens, 103, 104 nuda, 103, 104 schomburgkiana, 103, 104 Enea, 385 Enourea capreolata, 69 Erinus procumbens, 163 Erisma, 500 blancoa, 502, 503 calcaratum, 502, 502 floribundum, 502 var. floribundum, 502 japura, 502 laurifolium, 503 micranthum, 504 pallidiflorum, 502 parvifolium, 502 var. pallidiflorum, 502 pulverulentum, 504 splendens, 504 uncinatum, 503, 504 violaceum, 502 Erizo, 37, 348 Ero-cuaja, 412 Eroguata, 494 Ero-Kuaja, 496 Eroteum, 300 calophyllum, 301

I NDEX

guianense, 302 Ertela, 12 trifolia, 13, 12 Erythrochiton, 14 brasiliensis, 13, 14 Escoba, 452 Escobedia, 159 grandiflora, 159, 160 parimensis, 159 Escobillo, 440 Esenbeckia, 14 grandiflora, 15 subsp. grandiflora var. grandiflora, 15 obovalifolia, 15 pilocarpoides, 15 subsp. pilocarpoides, 15, 15 venezuelensis, 15 Espina, 252 Eurya calophylla, 301 sect. Freziera, 300 subg. Freziera, 300 guianensis, 302 roraimensis, 302 —F— Fagara, 34 amapaensis, 36 apiculata, 37 caribaea, 37 compacta, 37 ekmanii, 37 mar-tinicensis, 37 ocumarensis, 37 pentandra, 37 pterota, 37 rhoifolia, 39 syncarpa, 39 Farfara roja, 418 Fedejodo y Hede-hede, 448 Fiikcki, 427 Fleurya aestuans, 392 Franchetella atabapoensis, 118 neblinaensis, 124 Franciscea uniflora, 197 Freziera, 300 calophylla, 301, 302

carinata, 302, 302 cuatrecasasii, 301 esmeraldana, 301 guianensis, 302 hieronymi, 301 roraimensis, 302 Fruta de paloma, 49, 52, 200 Fruto de paloma, 60 —G— Galipea, 16 cusparia, 6 davisii, 16, 16 febrifuga, 6 heterophylla, 9 longifolia, 10 officinalis, 6 pentagyna, 10 stelligera, 29 Gaspadillo, 448, 455, 458, 459, 460, 463 Gaspadillo blanco, 448, 460, 463 Gaspadillo grande, 15 Gaspadillo negro, 455 Gateado, 137 Gerardia hispidula, 149 Ghinia, 438 spicata, 438 Globifera umbrosum, 164 Gloeospermum, 450 sphaerocarpum, 450, 451 ulei, 463 Glossostylis aspera, 147 Glycoxylon maguirei, 136 rubrum, 134 Godedi, 518 Goma de guaca, 124 Gomphiluma martiana, 121 Gordonia, 312 fruticosa, 312, 313 semiserrata, 312 Gratiola dubia, 161 repens, 153 Gua-chimacán, 19 Guacimillo, 363

593

Guácimo, 256, 353 Guácimo blanco, 256 Guácimo cimarrón, 353 Guácimo montañero, 256 Guácimo negro, 256 Guácimo rebalsero, 357 Guaiacum, 581 officinale, 581, 582 Guakabakadeupadek, 453 Guámari, 390 Guána, 346 Guanaguanare, 378 Guapeba coriacea, 119 stylosa, 121 Guarai blanco, 506 Guarapo, 506 Guarapo amarillo, 506 Guarataro, 440, 441, 445 Guarataro lechoso, 441 Guaraunera, 269 Guaritoto, 392, 400 Guaritoto montanero, 401 Guayabón, 136 Guayacán, 582 Guayare, 266 Guazuma, 255 ulmifolia, 256 var. tomentella, 256 var. ulmifolia, 255, 256 —H— Haemocharis, 312 caracasana, 313 pubescens, 312 Hala por atrás, 186 Hawkesiophyton, 207 panamense, 207 ulei, 207, 207 Heliamphora, 138 chimantensis, 140 elongata, 140 folliculata, 140 heterodoxa, 140 var. exappendiculata, 140, 142 var. glabra, 141 var. heterodoxa, 141, 142 hispida, 141 ionasii, 141, 142 macdonaldae, 144

594

I NDEX

[Heliamphora] minor, 141, 143 neblinae, 144 var. viridis, 144 nutans, 141, 143 sarracenioides, 141 tatei, 141 var. macdonaldae, 144 var. neblinae, 144 var. parva, 144 var. tatei, 143, 144 tyleri, 144 Helicteres, 256 apetala, 266 baruensis, 257, 258 brachypetala, 259 bracteosa, 259 brevispira, 257 guazumifolia, 257, 258 heptandra, 259 pentandra, 258, 259 pronifolia, 259 Herpestis, 151 auriculata, 153 caprarioides, 163 chamaedryoides, 163 ciliata, 153 domingensis, 154 laxiflora, 153 procumbens, 163 ranaria, 153 reflexa, 154 reptans, 153 salzmanni, 153 serpyllifolia, 153 sessiliflora, 154 stricta, 154 Herrania, 259 camargoana, 261 lemniscata, 260, 261 Hicaquillo, 448 Hierba de pasmo, 182 Hierba San Juan, 265 Hierbabuena de laguna, 434 Hinirji, 124 Hoglashdi, 459 Hoja babaso, 411 Hoja de bachaco, 182 Hortia, 17 coccinea, 17, 18 neblinensis, 17 regia, 17

vandelliana, 17 Houssayanthus, 57 macrolophus, 57, 57 Hubokawa, 390 Huele de noche, 200 Hue-tu-yek, 525 Huevo de gato, 228 Huevo de gato amarillo, 231 Huevo de sapo, 215 Huevo frito, 109 Humarí, 412 Hybanthus, 451 calceolaria, 451, 452 circaeoides, 452 ipecacuanha, 451 oppositifolius, 452 phyllanthoides, 452 Hydranthelium, 151 callitrichoides, 152 —I— Icaco, 136, 458 Ihkodï tawï, 448 Ilex lucumifolia, 121 Ilysanthes, 159 dubia, 161 riparia, 161 Ionidium calceolaria, 451 circaeoides, 452 ipecacuanha, 451 opposiifolium, 452 phyllanthoides, 452 Irana, 99 Ivira pruriens, 268 Ixora alternifolia, 200 —J— Jabón, 455 Jaboncillo, 75 Jabua, 502 Jacquinia, 328 arborea, 328 armillaris, 328, 329 revoluta, 328 Jaguajeshi, 270 Jalapatrás, 37

Jasmín grande, 26 Jazmín de monte, 11 Jazmín del monte, 197 Jiejiema, 362 Jö’nö jodedü, 463 Jobo, 178 Jona joreri, 460 Jonojodedö, 458 Jööna jodedü, 460 Juachetuma, 124 Juan de la calle, 197 Juanasi, 386 Juichamo, 496 Jupica, 538 —K— Kahka, 137 Kahka yo, 137 Kakka-yek, 136 Kallstroemia, 582 caribaea, 583 maxima, 583, 583 pubescens, 583 Kanke, 122 Katunyo’, 348 Kayani, 270 Këmëhkë tuku’ya këmën, 186 Kerepo-yek, 269 Kieseria, 316 sessilis, 320 Killipiodendron, 300 Kotsjiletti, 538 Kun-a-wa-lu-yek, 71 Kú-no-to-wo-ku-deu, 453 Kurunmota-yek, 458 —L— Labatia bilocularis, 118 laevigata, 121 parviflora, 120 torta, 123 tovarensis, 121 Lacre montañero, 524 Lantana, 426 achyranthifolia, 426, 428 aculeata, 427 alba, 430 armata, 427 var. guianensis, 427

I NDEX

camara, 427, 428 var. aculeata, 427 var. mista, 427 var. moritziana, 427 canescens, 427 cujabensis, 427 fucata, 427 geminata, 430 glutinosa, 427 mista, 427 moritziana, 427 radula, 427, 429 ruiz-teranii, 428, 428 trifolia, 428, 429 f. hirsuta, 428 f. oppositifolia, 428 f. rubierensis, 428 var. rigiduscula, 428 Laplacea, 312 camelliaefolia, 312 caracasana, 313 fruticosa, 312 var. chimantae, 313 pubescens, 312 var. camelliaefolia, 312 var. minor, 313 semiserrata, 312 Laportea, 392 aestuans, 392, 393 Lasiadenia, 338 ottohuberi, 338 rupestris, 339, 339 Laurel bobo, 182 Laurel orillero, 412 Laurus americanus, 335 Leonia, 453 cymosa, 453 glycycarpa, 453, 454 Lettsomia, 300 guianensis, 302 Leucococcus occidentalis, 399 Lightia, 259 lemniscata, 261 Lignonia, 454 Limoncillo, 175 Lindernia, 159 barrosorum, 161 brachyphylla, 161 crustacea, 161 diffusa, 161, 162

dubia, 161 microcalyx, 162, 162 rotundifolia, 162 Lindleya, 312 fruticosa, 312 semiserrata, 312 Linostoma sect. Lophostoma, 340 Lippia, 429 alba, 430 betulifolia, 434 geminata, 430 micromera, 430, 430 origanoides, 430, 430 schomburgkiana, 430 Lophanthus, 273 Lophostoma, 340 amoenum, 340, 340 Loranthus piperoides, 481 quadrangulare, 482 Lorjena, 518 Lubaria szczerbanii, 22 Lucuma glabrescens, 103 glomerata, 120 gomphiaefolia var. blepharantha, 121 gomphiefolia, 121 rigida, 122 temare, 118 Luehea, 352 alternifolia, 352 althaeiflora, 354 candida, 353, 353 cymulosa, 353 ferruginea, 352 flavescens, 355 nobilis, 353 rosea, 354 speciosa, 352 tessmannii, 353 Lueheopsis, 354 althaeiflora, 354, 355 duckeana, 354 flavescens, 355 rosea, 354 violacea, 354 Luma, 256 Lycianthes, 208 guianensis, 209 pauciflora, 208, 209

595

—M— Macagua, 269 Macude, 212 Macuderi, 412 Macuillamia, 151 limosa, 153 repens, 153 Ma-eri, 494 Ma-eri hasubi, 496 Mai pihpe, 122 Maiz tostado, 389 Majagua, 354, 356 Majagua hoja fina, 121 Majagua orillera, 357 Majagua rabipelado, 357 Majaguillo, 363 Majaguillo erizado, 346 Makararo, 123 Malachra berterii, 276 Malvavisco amarillo, 380 Mamón, 65 Mamoncilla, 463 Mamoncillo, 65, 82, 453 Manabea, 409 laevis, 412 Manilkara, 103 bidentata, 106 subsp. bidentata, 105, 106 subsp. surinamensis, 107 bolivarensis, 107 huberi, 107 williamsii, 106 Manjar, 69 Manman, 496 Mano de mato, 494 Manzana del diablo, 229 Mapurite, 36, 39 Mapurite blanca, 37 Mapurite blanco, 39 Mapurite morado, 37 Mapurito, 37 Mapurito amarillo, 39 Maraca, 332 Marfíl, 441 Markea, 209 camponoti, 212 formicarum, 210, 212 longiflora, 210, 212 panamensis, 207

596

I NDEX

[Markea] porphyrobaphes, 212 reticulata, 211, 212 sessiliflora, 211, 212 Marua-yek, 32 Masarandu, 107 Masaranduba, 107 Mata de hormiga, 13 Matapalo, 136 Matayba, 58 adenanthera, 59 affinis, 61 arborescens, 59, 62 atropurpurea, 60 camptoneura, 60 chimantensis, 61 discolor var. atropurpurea, 60 elegans, 60, 63 fallax, 61 guianensis, 60 inelegans, 60 jauaensis, 61 kavanayena, 60, 63 longipes, 60 subsp. tepuiensis, 61 macrolepis, 61 macrostylis, 61 oligandra, 61 var. occidentalis, 61 var. ptariana, 61 opaca, 61 var. fallax, 61, 63 var. opaca, 61 peruviana, 61, 64 ptariana, 61 subsp. guaiquinimae, 64 subsp. ptariana, 62, 64 reducta, 60 robusta, 64 sororopaniana, 60 spruceana, 62, 64 tovarensis, 60 venezuelana, 61 yutajensis, 64 Mauirruda, 270 Mawanani, 270 Mawaya-yeha, 494 Mayabo, 91 Mecardonia, 162 procumbens, 163, 163 Medebadi, 182 Melaito, 118

Melananthus, 212 ulei, 212, 213 Melasma melampyroides, 147 spathaceum, 167 Melero, 425 Melicocca bijuga, 65 Melicoccus, 65 bijugatus, 65, 66 oliviformis, 65 Meliosma, 40 gentryi, 41 herbertii, 41 var. herbertii, 41, 41 var. tepuiensis, 41 aff. meridensis, 41 nubicola, 41 tepuiensis, 41 uberrima, 41 vasquezii, 41 Mella laxiflora, 153 Melochia, 261 arenosa, 262, 264 benthamii, 263 carpinifolia, 263 carthaginensis, 263 cinerea, 262 concinna, 262 conglobata, 263 glandulifera, 262 graminifolia, 262 hirsuta, 263 var. tomentosa, 265 llanorum, 262 manducata, 262 melissifolia, 262 nodiflora, 263, 264 obovata, 263 parvifolia, 263 var. parvifolia, 263 scordiifolia, 263 tomentosa, 263 var. frutescens, 263 var. tomentosa, 263 ulmifolia, 263, 264 villosa, 263 var. tomentosa, 265 var. villosa, 265 Melosa, 166 Merecure, 412 Merecurillo, 124, 453

Micranthemum, 163 orbiculatum, 164 pilosum, 164 umbrosum, 164, 164 Micropholis, 107 casiquiarensis, 108 egensis, 108, 112 eugeniifolia, 120 guyanensis, 109 subsp. duckeana, 109 subsp. guyanensis, 109 humboldtiana, 109 maguirei, 109, 111 melinoniana, 109, 111 mensalis, 110 obscura, 110 spectabilis, 110 splendens, 110 steyermarkii, 120 suborbicularis, 110 venamoensis, 110, 112 venulosa, 110 wurdackii, 109 Mimusops balata var. schomburgkii, 106 bidentata, 106 huberi, 107 surinamensis, 107 Mnasium sphaerocephala, 332 Mochochito, 390 Mojo–mera, 494 Molenillo, 458, 459, 463 Molinillo, 459, 460 Mollia, 355 glabrescens, 356 grandiflora, 356 lepidota, 357 subsp. casiquiarensis, 357 var. casiquiarensis, 357 scabra, 357 speciosa, 357, 358 var. alata, 357 stellaris, 357 tomentosa, 357 ulei, 357, 358 sp. A, 357 Moniera, 12 Monocardia, 151 humilis, 153 lilacina, 153

I NDEX

violacea, 153 Moradito, 432 Moreillo, 504 Mougeotia, 261 Moyeja, 95 Mulato, 60 Muntingia, 362 calabura, 363, 363 Muntingiaceae, 342, 362 Mureillo, 504 Murraya, 18 exotica, 19 paniculata, 18, 19 Mutui-yek, 453 Myllanthus, 22 maroana, 22 neblinensis, 22 Myrsine schomburgkiana, 103 —N— Naiadothrix reflexa, 154 Naiyek, 504 Naranjuelo, 37 Navia lopezii, 536 Nazareno, 432 Neblinaria, 316 celiae, 319 rubicunda, 320 Neogleasonia, 316 multinervia, 319 wurdackii, 319 Neoraputia, 19 cowanii, 19 paraënsis, 19, 20 Neoxythece atabapoensis, 99 gabrielensis, 120 maguirei, 122 roraimaensis, 122 tomentosa, 123 Nettoa, 350 Nicolás, 26 Nicotiana, 212 tabacum, 214, 215 Niguo, 363 Nispero montañero, 124 Nomimadi, 453 Nortenia thouarsii, 166

—O— O’mochahcho ipeñ, 351 Oceanopapaver, 350 Ojo, 69 Ojo de Grillo, 262 Oligococe, 403 Once horas, 51 Onotillo, 348 Ophiocaryon, 42 duckei, 42 manausense, 42 sp. A, 42, 43 Orectanthe, 536 ptaritepuiana, 537, 538 sceptrum, 537, 538 subsp. occidentalis, 538 Oregano, 430 Orégano de playa, 262 Oreja del tigre, 272 Orinaré Bareua, 270 Ortega, 400 Oxytandrum, 345 Oxythece caracasana, 134 robusta var. brevipetiolasta, 119 scrobiculata, 123 steyermarkiana, 103 —P— Padamá, 270 Palo azul, 82, 175 Palo bolsa, 362 Palo de araguato, 354 Palo de arco, 441, 445 Palo de arco morado, 441, 445 Palo de babilla, 122 Palo de bachaco, 182, 183 Palo de chamanare, 110 Palo de conoto, 516 Palo de cuchara, 448 Palo de cuyure, 94 Palo de guabina, 507 Palo de hambre, 441 Palo de sol, 110 Palo de troja, 448 Palo de vela, 172 Palo manteco, 94 Palo paripari, 183 Palo piapoco, 510

Palo rayao, 103 Palo triste, 94 Paneco, 37 Panucoa, 61 Paraguita, 494 Paralabatia orinocoensis, 122 Paramo-yek, 118 Para-para, 75 Para-to-re-yek, 123 Parietaria microphylla, 397 Paripari, 182, 183 Parrillo, 175 Pascualito, 95 Pasita, 227 Passaveria lanceolata, 99 Passoura, 456 guianensis, 460 Pata de paují, 460 Patascoya, 300 Pate grulla, 448 Patillo-chicle, 99 Paullinia, 66 alsmithii, 69 anisoptera, 69, 72 bracteosa, 69 caloptera, 69, 73 capreolata, 69, 71 caracasana, 77 cupana, 69 cururu, 70 fuscescens, 70 imberbis, 70 ingaefolia, 70, 74 leiocarpa, 70, 74 micrantha, 70, 73 novemalata, 71 nuriensis, 70 paullinioides, 70 pinnata, 70, 74 rubiginosa, 71 rufescens, 71, 73 rugosa, 71, 72 serjaniifolia, 71 sorbilis, 69 verrucosa, 71, 73 Pavito, 328 Paypayrola, 454 grandiflora, 455 guianensis, 455 longifolia, 455, 455

597

598

I NDEX

[Paypayrola] ventricosa, 455 Payrola, 454 Pedicularis melampyroides, 147 Peine de mono, 346 Pendare, 99, 107 Pendare de sapo, 99 Pendare purue, 107 Pendarito, 119 Penenimpuatsi, 346 Pentamerista, 309 neotropica, 309, 310 Pepino, 257 Perguillo, 122 Perguillo negro, 124 Perguo, 106 Perguo balata, 106 Perguo blanco, 106 Perguo morado, 106 Periclista, 454 Pese, 516 Petrea, 431 arborea, 432 aspera, 432 blanchetiana, 431 bracteata, 431, 433 glandulosa, 432 kohautiana, 432 macrostachya, 432 pubescens, 432 var. albicalyx, 432 volubilis, 432, 432 Phenakospermum, 283 guyannense, 282, 283 Phenax, 394 hirtus, 394 sonneratii, 394, 394 vulgaris, 394 Phoradendron, 472 acinacifolium var. surinamense, 480 apertum, 483 aphyllum, 475, 484 appunii, 483 ayliffei, 484 baileyae, 483 var. exile, 483 berryi, 475, 485 bilineatum, 475 bolivianum, 477 caerulesens, 479 caracasanum, 479

var. parvifolium, 479 carinatum, 475 cayennense, 482 chaguaramasarum, 479 chrysocarpum var. stehlei, 484 chrysocladon, 475, 491 crassifolium, 476, 486 crenato-callosum, 477 crispulum, 467 demerarae, 476 dendrophthora, 467 densifrons, 467 dimidiatum, 481 diminutivum, 476 dipterum, 476 duidanum, 468 var. hymenaeifolium, 469 dunstervillorum, 481 ellipticum, 467 emarginatum, 479 var. truncato-retusum, 479 essequibense, 483 excisum, 480 exiguum, 476 exile, 483 fendlerianum, 468 flavens, 482 var. longispicum, 475 glauco-lutescens, 481 granaticola, 479 var. taeniicaule, 479 guascanum, 469 haughtii, 477 herbert-smithii, 477 herminieri, 484 hexastichum, 477, 487 huricola, 482 ichythystoma, 480 inaequidentatum, 477, 492 insigne, 480 intermedium, 468 jenmanii, 477 johnstonii, 483 kelloggii, 478 knoopii, 475 leptarthrum, 477 longiarticulatum, 476 longipetiolatum, 478 lyonii, 479

macbridei, 468 mairaryense, 478 martinicensis, 476 membranifolium, 478 microstachyum, 478 morsicatum, 479 mucronatum, 479 nervosum, 479, 488 nitidulum, 469 northropiae, 480 obliquum, 469 obtusissimum, 480, 489 ottonis, 479 var. oblongifolium, 479 pachyphyllum, 475 pellucidulum, 480, 487 perrottetii, 480, 488 piperoides, 481, 489 platycaulon, 481, 489 prancei, 477 pseudomucronatum, 480 ptarianum, 478 pteroneuron, 481 pulleanum, 482 quadrangulare, 482, 486 racemosum, 482, 490 roraimae, 469 rubrum var. longipedunculatum, 482 rugulosum, 481 sanctae-martae, 483 scariosum, 481, 482 schultesii, 483 semivenosum, 480 var. agostinii, 480 var. longipes, 480 spectabile, 468 var. altimontanum, 468 steyermarkii, 483 strongyloclados, 483, 485 sulfuratum, 478 surinamense, 483 taeniicaule, 479 tatei, 468 tenuiflorum, 469 tepuianum, 469 theloneuron, 484 tovarense, 476 treleasei, 484 triflorum, 483 trinervium, 483, 490 tubulosum, 479

I NDEX

tunaeforme, 484 undulatum, 484, 491 venezuelense, 482 var. filispicum, 482 verticillatum, 483, 496 williamsii, 480 zuloagae, 483 Phoxanthus, 42 manausensis, 42 Phyla, 434 betulifolia, 434, 434 Physalis, 215 angulata, 215, 216 cordata, 215 lagascae, 216 orinocensis, 206 pubescens, 216 surinamensis, 216 Physocodon, 261 Pi’tzejécute, 41 Piacayure, 328 Pica pica, 227 Picapica, 394 Pica-pica, 266 Picaton, 400 Picatón, 494, 496 Pico de lombriz, 109, 120 Pico de paloma, 70 Picramnia, 177 guianensis, 178 juniniana, 178, 179 latifolia, 178 macrostachys, 178 magnifolia, 178 nuriensis, 178 pentandra, 178 platystachya, 178 sellowii subsp. spruceana, 179 spruceana, 179, 179 tristamina, 178 Picramniaceae, 168, 177 Picrolemma, 169 pseudocoffea, 169 sprucei, 169, 170 Piedrero, 440 Pilea, 395 chrysosplenioides, 397 dauciodora, 395, 396 var. crenata, 395 fendleri, 395 involucrata, 397, 397 leptophylla, 395

microphylla, 397, 397 pubescens, 396, 398 var. involucrata, 397 Pilocarpus, 20 racemosus, 21 subsp. racemosus var. racemosus, 21, 21 Pilón, 175 Pimúyure, 120 Pionandra allophylla, 226 Piqueguaro, 440 Piraquiña, 175 Pirequina, 175 Piripea palustris, 156 Piriqueta, 373 caroliniana, 374 var. integrifolia, 374 cistoides, 374 subsp. caroliniana, 374 subsp. cistoides, 374, 375 var. genuina, 374 foliosa, 374 lanceolata, 374 tomentosa, 374 undulata, 375, 376 villosa, 374 viscosa, 376 subsp. tovarensis, 376 subsp. viscosa, 375, 376 Pleurostima, 405 celiae, 405 Poarchon, 527 Podoluma, 112 pacimoniensis, 124 Porteria, 403 Pouteria, 112 ambelaniifolia, 117 amygdalicarpa, 117, 126 anomala, 117, 126 arcuata, 117 atabapoensis, 118 auyantepuiensis, 103 bangii, 118, 127 bilocularis, 118 caimito, 118, 127 var. strigosa, 121 canaimaensis, 118, 125 cayennensis, 118, 128 cladantha, 118

599

coriacea, 119 crassiflora, 119 cuspidata, 119 subsp. cuspidata, 119, 128 subsp. dura, 119 subsp. robusta, 119 duckeana, 109 dura, 119 egregia, 119, 129 elegans, 120, 129 aff. engleri, 124 eugeniifolia, 120, 129 fimbriata, 120 gabrielensis, 120 glomerata, 120 subsp. glomerata, 120 subsp. stylosa, 121 gomphiifolia, 121, 130 guianensis, 121, 125 hispida, 121, 131 laevigata, 121, 131 lucumifolia, 121 maguirei, 122 mensalis, 110 aff. minima, 124 nuda, 103 orinocoensis, 122, 132 pimichinensis, 122, 132 pittieri, 120 plicata, 122 reticulata, 122 subsp. reticulata, 122, 133 rigida, 122 subsp. rigida, 122 subsp. tomentosa, 123, 130 scrobiculata, 123, 130 sipapoensis, 123 spectabilis, 110 spuria, 95 steyermarkii, 120 stipitata, 123 surinamensis, 137 surumuensis, 123 torta, 123 subsp. glabra, 123, 133 tovarensis, 121 trilocularis, 124 ucuqui, 124 venamoensis, 110

600

I NDEX

[Pouteria] venosa, 124 subsp. amazonica, 124, 125 Pouzolzia, 399 occidentalis, 398, 399 Pradosia, 134 beardii, 134, 135 caracasana, 134 grisebachii, 136, 137 schomburgkiana, 136 subsp. schomburgkiana, 135, 136 subsp. sericea, 136 surinamensis, 136, 137 Prieurella maguirei, 122 Primo chipón, 110 Priniga-moza, 400 Priva, 434 lappulacea, 435, 435 f. albiflora, 435 Pseudocladia cassiquiarensis, 121 neblinanensis, 118 orinocoensis, 121 Pulgo negro, 107 Pulguillo, 99, 106, 118, 119 Pulguillo amarillo, 119 Pulvillo mágico, 455 Punuwoyo, 124 Punwoyo, 103, 124 Purgo, 107, 118 Purguillo chicle, 124 Purguillo felix, 110 Purguillo terciopelo, 124 Purguo, 107 —Q— Qualea, 504 acuminata, 506, 507 apodocarpa, 510 calcarata, 502 ser. I callophylloidea, 508 cassiquiarensis, 510 cyanea, 506 decorticans, 506 dinizii, 506, 508 esmeraldae, 510 ferruginea, 510 homosepala, 506 obtusata, 510

paraensis, 506 polychroma, 506 psidiifolia, 506 pulcherrima, 506, 508 retusa, 510 var. coriacea, 510 rigida, 510 rubiginosa, 510 var. angustior, 510 rupicola, 506 schomburgkiana, 507, 507 speciosa, 506 sprucei, 507 suprema, 507 themistoclesii, 507 trichanthera, 512 sect. Trichanthera, 508 tuberculata, 507 verruculosa, 510 wurdackii, 508 Quamoclit rodriguesii, 262 Quassia, 169 amara, 169, 171 Quejido, 253 Quina, 6 Quina amarilla, 6, 34 Quina negra, 34 Quino blanco, 16 —R— Rabo de guaca, 418 Rabo de iguana, 69, 77 Rabo de Maquire, 507 Rabo de zorro, 418 Rabo pelado, 200 Radia tubiflora, 406 Ragala sanguinolenta, 94 Ramotha, 538 Ranaria, 151 monnierioides, 153 Raputia, 22 maroana, 22, 23 neblinensis, 22, 23 paraënsis, 19 szczerbanii, 22, 24 Rastrojero blanco, 413 Rauia, 24 spicata, 26 subtruncata, 25, 26

ulei, 11 Ravenia linearis, 29 paruana, 29 ruellioides, 29 var. ptariana, 29 tatei, 29 Raveniopsis, 26 abyssicola, 28 breweri, 28 capitata, 28 cowaniana, 29 fraterna, 29 jauaensis, 29 liesneri, 29 linearis, 29, 30 nubicola, 29 paruana, 29 peduncularis, 29, 30 ruellioides, 29, 30 var. angusta, 29 sericea, 29, 30 stelligera, 29, 30 var. plicata, 29 steyermarkii, 30 tomentosa, 30, 30 trifoliolata, 31 Rebalsero, 121 Renealmia, 577 alpinia, 578 aromatica, 578, 579 exaltata, 578 floribunda, 578 guianensis, 578 lutea, 578 microcalyx, 578 monosperma, 579 occidentalis var. ß cylindrica, 578 var. macrocarpa, 578 var. occidentalis, 578 orinocensis, 579, 579 pedicellaris, 578 raja, 578 thyrsoidea, 580 subsp. thyrsoidea, 580 Rhamnus iguanaeus, 389 micranthus, 390 Riana, 456 guianensis, 463 Riedlea, 261 dichotoma, 263

I NDEX

heterotricha, 265 littoralis, 262 ramuliflora, 263 urticifolia, 263 Rinorea, 456 brevipes, 457 camptoneura, 458 endotricha, 458 falcata, 458 flavescens, 458, 462 lindeniana, 459 var. lindeniana, 459 macrocarpa, 459, 462 melanodonta, 459 ovalifolia, 459, 461 paniculata, 460, 461 pubiflora, 460 var. grandifolia, 460 var. pubiflora, 460, 462 racemosa, 461 riana, 463 sprucei, 462, 463 Rinoreocarpus, 463 salmoneus, 463 ulei, 463, 464 Robinia rubiginosa, 54 Rosado, 118 Ruizterania, 508 apodocarpa, 510 cassiquiarensis, 510 esmeraldae, 510, 511 ferruginea, 510, 511 obtusata, 510 subsp. multivenosa, 510 subsp. obtusata, 510 retusa, 510, 511 rigida, 510 rubiginosa, 510 var. angustior, 510 trichanthera, 512 verruculosa, 510 Rutaceae, 1 Rutaneblina, 31 pusilla, 31, 31 —S— Sabiaceae, 39 Sábila, 524 Saladillo, 516, 517, 518 Salado, 504, 517, 524 Salado de rebalse, 506

Salado morado, 504 Salado rebalsero, 506 Salado rojo, 524 Salado sabanero, 524 Salao, 517 Saldillo, 524 Salmonia, 512 Samán montañero, 37 Sanaparo, 518, 524 Sanaparo fino, 517 Santalaceae, 44 Sapindaceae, 46 Sapindus, 75 arborescens, 59 saponaria, 75, 75 Sapotaceae, 89 Saracha solanacea, 246 Sardina, 217 Sarraceniaceae, 138 Sarrapia, 86 Sauce, 379 Sauce sabanero, 217 Saunan-yek, 95 Sawara kurimi, 117 Schinus fagara, 37 Schismaxon, 538 Schlechtendalia, 355 Schmidelia amazonica, 48 occidentalis, 49 punctata, 48 Schoenobiblus, 341 amazonicus, 341, 342 Schuechia, 504 Schwenckia, 216 americana, 217, 218 var. angustifolia, 217 var. hirta, 217 elegans, 217 glabrata, 217 grandiflora, 217, 219 guianensis, 217 guineensis, 217 heterantha, 221 hirta, 217 huberi, 220, 221 micrantha, 221 patens, 217 ulei, 217 Sciaphila, 369 albescens, 369, 370

601

brevipes, 371 guianensis, 369 purpurea, 369, 370 rubra, 370 spruceana, 371 Sciuris officinalis, 6 Scoparia, 164 dulcis, 164, 165 Scrophularia fluminensis, 147 Scrophulariaceae, 144 Seje piapoco, 510 Serae-serae, 440 Sereno, 380 Serjania, 76 adusta, 77 caracasana, 77, 78 clematidea, 77, 78 communis, 77 var. communis, 77, 79 grandifolia, 77 ierensis, 77 mariquitensis, 77 membranacea, 77 paucidentata, 77 pyramidata, 79, 80 rhombea, 80 Sesejudi, 441 Shibwichi, 120 Shinnate, 411 Sicklera tetrandra, 246 Sida villosa, 263 Sideroxylon amygdalicarpum, 117 bangii, 118 cuspidatum, 119 egense, 109 elegans, 120 guyanensis, 109 robustum, 119 venulosum, 110 Simaba, 169 cedron, 172, 173 guianensis, 172 subsp. guianensis, 172 subsp. huberi, 174, 175 monophylla, 175 multiflora, 175 obovata, 174, 175 orinocensis, 174, 175

602

I NDEX

]Simaba[ sp. A., 175 Simarouba, 175 amara, 175, 176 monophylla, 175 Simaroubaceae, 168 Siparuna, 180 bifida, 181 cristata, 182 cuspidata, 182 decipiens, 182, 184 ficoides, 182 guianensis, 182, 183 micrantha, 183 obstipa, 182 reginae, 183, 183 sancheziana, 183 surinamensis, 183 Siparunaceae, 180 Sitella, 273 involucrata, 278 Smilacaceae, 184 Smilax, 185 auraimensis, 186 chimantensis, 186 cumanensis, 186, 188 duidae, 187 jauaensis, 193 lappacea, 186, 188 lasseriana, 193 maypurensis, 186 pittieriana, 186, 189 poeppigii, 187, 190 schomburgkiana, 187, 191 siphilitica, 187, 192 staminea, 188, 193 f. obtusata, 193 triplinervia, 193 Sohnreyia, 32 Solanaceae, 194 Solandra, 221 grandiflora, 221, 222 longiflora, 222 Solanum, 222 acerifolium, 226 adhaerens, 226 allophyllum, 226, 232 altissimum, 226, 233 americanum, 226, 232 arboreum, 227, 233 arcuatum, 226 arecunarum, 231 asperrimum, 227

asperum, 227, 234 aturense, 227, 235 badilloi, 227 bicolor, 227, 235 boissieri, 230 bolivarense, 229 brachyacanthum, 228 callicarpifolium, 227 campaniforme, 227, 237 caripense, 227 confusum, 231 coriaceum, 228, 234 crinitum, 228, 238 cyananthum, 228 var. jubatum, 228 cyrrhosum, 230 davidsei, 228 demerarense, 231 var. inerme, 231 dolichostylum, 227 donnellsmithii, 226 echinocarpum, 228 enoplocalyx, 226 flavescens, 228 var. albescens, 228 formosum, 228 fraternorum, 231 geminifolium, 230 globiferum, 229 granelianum, 230 guianense, 209 hastatum, 226 heterophyllum, 231 hirtum, 228, 239 hostmannii, 230 humboldtii, 226 hyporrhodium, 245 ipomoea, 229 jamaicense, 228 javitense, 229 jubatum, 228 juripeba, 231 laetum, 230 lanceifolium, 228, 237 var. brachyacanthum, 228 lanugiflorum, 231 leucocarpon, 228, 239 macranthum, 228 mammosum, 229, 240 mesopliarthrum, 231 miquelii, 230 molle, 229

monachophyllum, 229 nodiflorum, 226 nuricum, 229 obscurum, 228 obtusifrons, 228 oleraceum, 226 oocarpum, 229, 241 oppositifolium, 229, 242 orenocense, 228 orientale, 229 orinocense, 231 sect. Pachystemonum subsect. Lycianthes, 208 paludosum, 229, 241 pauciflorum, 209 pensile, 229, 240 platanifolium, 229 platyphyllum, 231 prehensile, 230 prunifolium, 230 pterocaulon, 226 puberuloba, 229 quinquangulare, 226 radula, 227 rhamphidacanthum, 231 ripivagum, 227 rufistellatum, 230, 236 rugosum, 230, 243 sacupanense, 229 salignum, 230 salviifolium, 227 scabridum, 245 scandens, 229 schlechtendalianum, 230, 236 schomburgkii, 230 seaforthianum, 230, 244 secundum, 227 sempervirens, 230 sessiliflorum, 230, 245 siparunoides, 227 stamineum, 231 stramonifolium, 231, 244 var. inerme, 231 styracifolium, 231 styracoides, 230 subinerme, 231, 243 surinamense, 228 swartzianum, 231 tepuiense, 231 timotense, 228 tobagense, 203 topirito, 228

I NDEX

topiro, 231 toxicarium, 231 trichocarpum, 231 triste var. crassipes, 229 umbratile, 231 umbrosum, 228 undecimangulare, 231 velutinum, 231, 242 vestissimum, 245 Soridium, 371 guianense, 369 spruceanum, 370, 371 Spatellaria, 447 Spathe, 32 Spathelia, 32 chimantaensis, 32 fruticosa, 32 jauaensis, 32 neblinaensis, 32 ulei, 32, 32 Spathularia, 447 longifolia, 448 Speirostyla, 348 tiliifolia, 350 Sphaerotheca scoparioides, 159 Sphenoclea, 248 zeylanica, 247, 248 Sphenocleaceae, 247 Spiciviscum, 472 Spiranthera, 33 guianensis, 33, 34 parviflora, 34 Sponia, 389 integerrima, 390 Stachytarpheta, 436 angustifolia, 436, 437 f. elatior, 436 var. elatior, 436 cayennensis, 437 f. albiflora, 437 dichotoma, 437 f. albiflora, 437 elatior var. jenmanii, 436 jamaicensis, 438 lopez-palacii, 438 mutabilis, 438 pilosiuscula, 438 roraimensis, 438 var. pubescens, 438 sprucei, 437, 438

surinamensis, 436 Stemmatosiphon nitens, 294 Stemodia, 164 foliosa, 165, 166 Sterculia, 265 abbreviata, 266 amazonica, 266 apetala, 266, 267 kayae, 266 parviflora, 266 pruriens, 268, 268 var. parviflora, 266 rugosa, 267, 269 Sterculiaceae, 248 Steudelia, 453 Strelitziaceae, 281 Strigilia glabrata, 286 guianensis, 287 psilophylla, 286 Strukeria, 512 Styracaceae, 283 Styrax, 284 costanus, 286 duidae, 285 subsp. duidae, 285 subsp. neblinae, 287 subsp. paruae, 285, 288 fanshawei, 286 filii, 287 glaber, 286, 289 var. micranthus, 286 glabratus, 286 guaiquinimae, 286 guanayanus, 286, 288 var. yutajensis, 287 guyanensis, 287, 288 var. japurensis, 287 jauaensis, 286 lauraceus, 286 longipedicellatus, 287, 289 micrasterus, 286 neblinae, 287 occidentalis, 286 psilophyllus, 286 roraimae, 286 sipapoanus, 287 squamulosus, 286 subleprosum, 286 tepuiensis, 286 subsp. guaiquinimae, 286

603

subsp. huachamacarii, 287 wurdackiorum, 287, 289 yutajensis, 287 Symplocaceae, 290 Symplocos, 290 acananensis, 291, 292 chimantensis, 291 guianensis, 291 jauaensis, 291, 293 martinicensis, 291 neblinae, 292, 294 nitens, 294 var. claussenii, 294 pycnophylla, 293, 295 roraimensis, 295 schomburgkii, 294, 295 ulei, 293, 295 yapacanensis, 292, 295 Syzygiopsis sericea, 117 —T— Tabaco, 215 Tabacón borrachera, 227 Tabaquero, 412 Tabaquero peluo, 227 Tabaquillo, 227, 411, 412, 413 Tacca, 296 parkeri, 296, 296 var. lanceolata, 296 sprucei, 296 ulei, 296 Taccaceae, 295 Tagua-tagua, 495 Tahitia, 348 Takrum-yek, 494 Taligalea campestris, 418 Talisia, 80 acutifolia, 81 allenii, 82 amaruayana, 82 amaruyana, 82 cararensis, 82 caudata, 81 chartacea, 81 dwyeri, 82 firma, 81 glandulifera, 81 grandifolia, 82

604

I NDEX

[Talisia] hemidasya, 81, 84 heterodoxa, 82 hexaphylla, 82 subsp. hexaphylla, 82 laevigata, 82 macrophylla, 82, 83 medrii, 81 nervosa, 82, 83 obovata, 82 olififormis, 65 panamensis, 82 pentantha, 82 retusa, 82 sancarlosiana, 82 tirirensis, 82 Tambak, 175 Tamonea, 438 spicata, 438, 439 Taonabo dentata, 306 punctata, 305 Tar seeho waina he héhéchë, 69 Tariri guianensis, 178 Taymarito, 119 Tee-u-ben-yek, 123 Temara, 118 Temare, 95, 109, 118, 120, 124 Temare amarillo, 118 Temare de montaña, 99 Temare montañero, 95 Temari, 118 Temarito, 121, 136 Temarito banero, 136 Temure, 118 Tepe camino, 200 Tepuianthaceae, 297 Tepuianthus, 297 auyantepuiensis, 298 sarisariñamensis, 298 subsp. duidensis, 298, 299 subsp. sarisariñamensis, 299 savannensis, 299, 299 yapacanensis, 299 Ternstroemia, 303 brevistyla, 303 browniana, 306 campinicola, 303

candolleana, 303 var. rotundata, 303 crassifolia, 304, 308 delicatula, 306 dentata, 306 discoidea, 304, 308 distyla, 304 duidae, 304, 308 f. latifolia, 304 dura, 304, 307 gleasoniana, 306 grandiosa, 307 guanchezii, 304 laevigata, 304 longipes, 307 maguirei, 305 monosperma, 304 paucifolia, 304 punctata, 305 pungens, 305, 308 retusifolia, 305 revoluta, 305 schomburgkiana, 307 tristyla, 305 verticillata, 305 sp. A., 306, 307 sp. B., 306 Ternstroemiaceae, 300, 310 Tetoncillo de rebalse, 103 Tetrameristaceae, 309 Theaceae, 310, 311 Theobroma, 269 bicolor, 270 cacao, 270 subsp. sphaerocarpum, 270, 271 grandiflorum, 270, 271 guianense, 270 microcarpum, 270 sphaerocarpa, 270 subincanum, 270, 271 sylvestris, 270 Theophrastaceae, 325 Thesium, 44 tepuiense, 44, 45 Thinouia, 84 myriantha, 85, 85 Thurnia, 330 polycephala, 331, 332 sphaerocephala, 331, 332 Thurniaceae, 330 Thymelaeaceae, 332

Tiaebo hiEi hele y Tiebo here, 448 Tiestigo, 82 Tiliaceae, 342, 343 To’oóto, 69 Tonina, 70 Töpishécute, 41 Topotopo, 215 Toqui-voquiris, 427 Torenia, 166 crustacea, 161 thouarsii, 166, 166 var. nivea, 166 Tornillo, 257, 259 Tostadito, 431, 432 Totumillo, 423, 441 Totumillo blanco, 422, 423 Totumillo morado, 440, 441 Totumillo negro, 422 Totumillo sabanero, 423 Toulicia, 85 anomala, 89 guianensis, 86, 86 petiolulata, 86 pulvinata, 86 Traira-mira, 507 Traira-miri, 507 Trema, 389 integerrima, 389, 390 micrantha, 390, 390 Tres filos, 77 Tribroma, 269 Tribulus, 583 macranthus, 584 maximus, 583 pubescens, 583 zeyheri, 584 subsp. macranthus, 584, 584 Trichilia caudata, 61 ptariana, 61 Trigonia, 364 ehrendorferi, 365 kaieteurensis, 365 microcarpa, 365, 365 nivea, 365 var. nivea, 365, 366 reticulata, 365 sericea, 365, 367 spruceana, 367, 368 villosa, 368 var. villosa, 366, 368

I NDEX

Trigoniaceae, 364 Triumfetta, 358 bartramia, 360 bogotensis, 359 bolivariensis, 359 hispida, 359 lappula, 359, 360 rhomboidea, 360 rubricaulis, 360 semitriloba, 360 valenciensis, 359 Triuridaceae, 368 Triuris, 371 hyalina, 371, 371 Trompeta de angel, 222 Tropaeolaceae, 372 Tropaeolum, 372 orinocense, 372, 372 Tú ne yó, 175 Tu-ku-da-mo, 95 Tukuinanwuyek, 506 Tuliwaedi hele, 460 Tu-pata-yén, 187 Tupirito, 231 Tupiro de danto, 226 Túpiru, 230 Tupukuiden, 71 Turnera, 376 acuta, 378, 381 annectens, 379, 382 argentea, 379 aturensis, 376 breviflora, 379 carpinifolia, 378 castilloi, 379 cicatricosa, 379, 383 cistoides, 374 corchorifolia, 379 guianensis, 379, 380 hexandra, 379 huberi, 379 humboldtii, 379 lineata, 379, 381 macrophylla, 379, 382 muricata, 379 odorata, 379, 384 paruana, 380 parviflora, 379 pilosula, 378 pumilea, 380, 384 scabra, 380, 381 schomburgkiana, 380, 383

steyermarkii, 380 tomentosa, 374, 379 ulmifolia f. serissima, 379 var. intermedia, 380 valleana, 279 venosa, 380, 383 waltherioides, 380, 384 Turneraceae, 373 Typha, 385 domingensis, 385, 385 Typhaceae, 385 —U— Uatakurán, 182 Ucuquirana, 95 Ucuquirana de selva baja, 95 Udidi, 412 Uladendron, 272 codesuri, 272, 272 Ulmaceae, 386 Uña de gato, 227 Uña de gavilán, 37 Uña de pereza, 172 Urania guyannensis, 283 Urera, 399 baccifera, 400, 400 caracasana, 401, 401 laciniata, 402 Uri tote hyEi, 453 Urtica aestuans, 392 baccifera, 400 caracasana, 401 cylindrica, 392 involucrata, 397 Urticaceae, 391 Urvillea, 87 macrolopha, 57 ulmacea, 87, 87 Uva, 496 Uvita, 200 —V— Valeriana, 403 alpina, 403 candolleana, 403 phaseoli, 403 scandens, 403

605

var. candolleana, 403, 404 var. dentata, 403 var. subcordata, 403 Valerianaceae, 402 Vandellia, 159 crustacea, 161 diffusa, 161 Vara blanca, 200, 357 Vasivaea, 361 alchorneoides, 361, 362 Vela de muerto, 172 Vellosiella, 167 spathacea, 167, 167 Vellozia, 406 alexandrinae, 406 duidae, 406 tubiflora, 406, 406 Velloziaceae, 404 Verbena, 436, 437, 438 angustifolia, 436 cayennensis, 437 dichotoma, 437 jamaicensis, 438 mutabilis, 438 prismatica, 420 Verbenaceae, 407 Verga de toro, 449 Veru caimito, 93 Viola arborea, 449 calceolaria, 451 hybanthus, 449 ipecacuanha, 451 oppositifolia, 452 Violaceae, 445 Viscaceae, 464 Viscum crassifolium, 476 dichotomum, 481 dimidiatum, 481 ellipticum, 467 flavens, 482 fockeanum, 481 glandulosum, 482 hexastichum, 477 latifolium, 482 martinicensis, 476 mucronatum, 479 obliquum, 469 obtusissimum, 480 perrottetii, 480 piperoides, 481

606

I NDEX

[Viscum] quadrangulare, 482 racemosum, 482 trinervium, 483 tunaeforme, 484 undulatum, 484 verticillatum, 496 Visenia, 261 Vitaceae, 492 Vitex, 439 appunii, 441 calothyrsa, 440, 442 capitata, 440, 443 compressa, 441, 443 klugii, 441, 444 multiflora, 441 orinocensis, 441 var. multiflora, 441, 444 var. orinocensis, 441 pacimonensis, 440 schomburgkiana, 441 spongiocarpa, 441 var. longidentata, 441 sprucei, 441 var. longidentata, 441 var. vaupesensis, 441 stahelii, 441, 445 triflora, 442, 445 f. quinquefoliolata, 445 var. quinquefoliolata, 445 Vitis alata, 494 bakeri, 495 gibertii, 495 gongylodes, 495 palmata, 495 rhombifolia, 494 sicyoides, 496 f. laciniata, 496 var. smilacina, 496 spinosa, 495 tricuspis, 495 Viudita, 230 Vochisia, 512 Vochy, 512 Vochya, 512 ferruginea, 516 Vochysia, 512 angustifolia, 516 apopetala, 516, 518 arcuata, 518

bautistae, 516 calophylla, 516 cassiquiarensis, 516 catingae, 516, 519 complicata, 516 costata, 516, 519 crassifolia, 516, 520 curvata, 516 elegans, 516 expansa, 516 ferruginea, 516, 522 glaberrima, 517, 521 grandis, 517 var. genuina, 517 var. uaupensis, 517 inundata var. venosa, 517 javitensis, 516 jonkeri, 517 julianensis, 517 aff. laxiflora, 517 liscanoi, 517 lucida, 517 obscura, 517, 523 ortegae, 517 pauciflora, 517 pinkusii, 517 punctata, 517 rubiginosa, 517 saccata, 517 spathiphylla, 518 splendens, 518, 523 steyermarkiana, 518 surinamensis, 518 var. inflata, 518 var. surinamensis, 518, 520 tetraphylla, 518, 521 tilletii, 518 tomentosa, 518 venezuelana, 522, 524 vismiifolia, 524 var. robusta, 518 Vochysiaceae, 500 Volkameria fragrans, 423 Vouarana, 88 anomala, 88, 89 —W— Wacadu, 109 Waka-dame, 441

Waltheria, 273 albicans, 276 americana, 278 berteroi, 276, 280 carmensarae, 277 caroliniana, 374 collina, 277 hirsuta, 281 indica, 278, 280 involucrata, 278, 280 machrisiantha, 281 operculata, 279 subcordata, 276 tubiflora, 281 viscosissima, 281 Wana, 348 Waramiku-yek, 336 Waranshikuai, 445 Waremba-dek, 93 Wa-won-yek, 121 Wayarayuyek, 15 Wene iye, 103 Wesemonama, 448 We-won-yek, 121 Wibelia, 454 Wikstroemia, 312 camelliaefolia, 312 fruticosa, 312 Wintera, 524 Winteraceae, 524 Witheringia, 246 macrophylla, 246 solanacea, 246, 246 —X— Xuris, 538 Xyridaceae, 526 Xyris, 538 acuminata, 551 albescens, 547, 558 ambigua, 551 americana, 530 anceps, 551 applanata, 555 aquatica, 547 aracamunae, 547, 559 arachnoidea, 547, 560 arenicola, 551 arescens, 551 asterocephala, 551 atriceps, 547 subsp. atriceps, 548

I NDEX

subsp. chimantensis, 548, 559 subsp. marahuacae, 548 subsp. neblinensis, 548 bicephala, 548, 560 bicostata, 548 var. bicostata, 548 var. tillettii, 548, 561 byssacea, 548 calcarata, 555 capillaris, 556 carinata, 548, 560 caroliniana var. major, 552 chimantae, 548, 561 coerulea, 530 communis, 551 concinna, 549, 562 connectens, 574 consolida, 549, 562 contracta, 549 cryptantha, 549, 563 culmenicola, 549, 561 cyperoides, 549 decussata, 549 delicatula, 549, 559 dolichosperma, 550 duidensis, 574 epicarae, 549 eriophylla, 556 erythema, 550 esmeraldae, 549, 564 exserta, 555 fallax, 550, 564 filiscapa, 550 foveolata, 551 frequens, 550 frondosa, 550, 564 fuliginea, 550, 565 fusiformis, 553 garcia-barrigae, 556 gardneri, 550 glabrata, 553 globosa, 550 gongylospica, 550 graniticola, 550 guianensis, 550, 565 gymnoptera, 551 huberi, 551, 562 hymenachne, 551, 565 involucrata, 551, 566 jauana, 557 juncifolia, 551, 566

jupicae, 551 jupicai, 551, 567 var. brachylepis, 551 var. major, 552 kukenaniana, 552 lacerata, 552 lanulobractea, 552 laxifolia, 552 var. laxifolia, 552 var. major, 552 var. minor, 551 leptostachya, 554 liesneri, 552 lithophila, 552 lomatophylla, 552 longiceps, 554 lugubris, 553 macrocephala, 552 f. minor, 551 var. major, 552 malmeana, 553, 567 melanovaginata, 553, 568 mima, 553 neblinae, 553, 563 oblata, 553 oxylepis, 553, 574 paraensis, 553 var. longiceps, 554, 569 var. paraensis, 554, 569 pratensis, 554, 563 prolificans, 554 ptariana, 554, 569 riparia, 554 roraimae, 554, 570 rubrolimbata, 554, 572 rugulosa, 549 savanensis, 555, 568 scabridula, 556 schneeana, 555 setigera, 555, 558 var. elongata, 555 seubertii, 555 sipapoa, 557 spathacea, 555 spruceana, 555, 571 stenophylloides, 555, 558 stenostachya, 556, 570 steyermarkii, 557 subglabrata, 556 submontana, 551 subuniflora, 556, 573 sulcatifolia, 556 surinamensis, 556, 570

607

tatei, 556, 572 teinosperma, 556, 572 tenella, 557 var. tenella, 557, 567 thysanolepis, 557 var. sipapoa, 557, 573 var. thysanolepis, 557, 573 tillettii, 548 toronoana, 557 trisperma, 553 uleana, 557 var. angustifolia, 574 var. uleana, 566, 574 vaginata, 532 valdeapiculata, 574 vaupesana, 574 witsenioides, 568, 574 wurdackii, 574 subsp. wurdackii, 571, 574 xiphophylla, 571, 574 yapobodensis, 555 yutajensis, 557 —Y— Ya’ra ipun yo, 357 Ya’ra ipun yo’, 353, 354 Yachiyek, 39 Yacu-yek, 175 Yakantayuru, 182 Yaraboba, 390 Yaradek, 315 Yarakaru nishesahri, 453 Yavaraji, 110 Yerba mora, 226 Yoquito, 136 Yoron, 137 Yuco, 124 Yuquito, 110, 117, 121 Yuquito banero, 103 Yuri, 502 Yuú, 124 Yuu-ka-mohi, 328 —Z— Zamia, 575 amazonum, 575 jirijirmensis, 576 lecointei, 575, 576 obidensis, 576

608

I NDEX

[Zamia] ulei subsp. lecointei, 575 Zamiaceae, 575 Zanthoxylum, 34 amapaense, 36 apiculatum, 37 caribaeum, 37, 38

compactum, 37 ekmanii, 37 fagara, 37, 39 huberi, 37 juniperinum, 37 martinicense, 37 ocumarense, 37 pentandrum, 37 pterota, 37

rhoifolium, 39 syncarpum, 38, 39 sp. A., 39 sp. B, 38, 39 Zapatero, 54, 59, 60, 266 Zarza, 496 Zingiberaceae, 577 Zygophyllaceae, 580