Flora of the Venezuelan Guayana, Volume 6: Liliaceae-Myrsinaceae 0915279819, 9780915279814

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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 6 LILIACEAE–MYRSINACEAE

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

MISSOURI BOTANICAL GARDEN PRESS St. Louis

Copyright © 2001 by the Missouri Botanical Garden Press All rights reserved. ISBN 0-915279-81-9 Printed in U.S.A. Published on 21 February 2001 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 Liliaceae (Alan W. Meerow and Robert W. Cruden) . . . . . . . . . . . . . . . . . . . 1 Limnocharitaceae (Robert R. Haynes and Lauritz B. Holm-Nielsen) . . . 16 Lissocarpaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Loasaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Loganiaceae (Paul E. Berry and Alan E. Brant) . . . . . . . . . . . . . . . . . . . . 22 Loranthaceae (Job Kuijt) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Lythraceae (Alicia Lourteig) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 Magnoliaceae (Paul E. Berry and James S. Miller) . . . . . . . . . . . . . . . . . 80 Malpighiaceae (William R. Anderson) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 Malvaceae (Paul A. Fryxell) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186 Marantaceae (Lennart L. Andersson, Helen Kennedy, and Mats Hagberg). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219 Marcgraviaceae (Stefan Dressler) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248 Mayacaceae (Alicia Lourteig) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260 Melastomataceae (Paul E. Berry, Andreas Gröger, Bruce K. Holst, Thomas Morley, Fabián A. Michelangeli, Navina G. Luckana, Frank Almeda, Susanne S. Renner, Alina Freire-Fierro, Orbelia R. Robinson, and Kay Yatskievych) . . . . . . . . . . . . . . . . . . . . . . . . . 263 Meliaceae (Terence D. Pennington and K. S. Edwards) . . . . . . . . . . . . . 528 Mendonciaceae (Dieter C. Wasshausen) . . . . . . . . . . . . . . . . . . . . . . . . . . 550 Menispermaceae (Rupert C. Barneby) . . . . . . . . . . . . . . . . . . . . . . . . . . . 554 Menyanthaceae (Kay Yatskievych) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 578 Mimosaceae (Rupert C. Barneby, James W. Grimes, Paul E. Berry, David Brunner, Enrique Forero, Lourdes Cárdenas, Giovanna De Martino, Helen C. F. Hopkins, and Elêna Maria de Lamare Occhioni) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 580

v

vi C ONTENTS

Molluginaceae (Julian A. Steyermark) . . . . . . . . . . . . . . . . . . . . . . . . . . . Monimiaceae (Paul E. Berry and Ariane Luna Peixoto) . . . . . . . . . . . . . Monotaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moraceae (Cornelis C. Berg) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moringaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Musaceae (Kay Yatskievych) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myricaceae (James S. Miller) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myristicaceae (William A. Rodrigues, Gerardo A. Aymard C., and Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myrsinaceae (John J. Pipoly III and Jon Ricketson) . . . . . . . . . . . . . . .

686 689 691 693 729 731 733 734 748

Appendix: List of new names published in this volume . . . . . . . . . . . . . 776 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 777

Preface and Acknowledgments This volume continues the taxonomic treatments of all native and naturalized vascular plant families known to occur in the Venezuelan Guayana. It includes 28 families of spermatophytes. The remaining families will be published in alphabetical order in three subsequent volumes. The Flora of the Venezuelan Guayana project has been strongly supported since its inception by the Missouri Botanical Garden and especially by its director, Peter H. Raven. The Herbario Nacional de Venezuela, now part of the Fundación Instituto Botánico de Venezuela, has steadily supported this project in many ways; it was there that Julian Steyermark conceived the idea of the flora and did most of his background research. This volume is based upon work supported by the National Science Foundation under Grants Nos. BSR-8717303, BSR-9045532, and BSR-9201044. The Foundation provides awards for research in the sciences and engineering. The awardee is wholly responsible for the conduct of such research and preparation of the results for publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the authors and do not necessarily reflect the views of the National Science Foundation. The Julian A. Steyermark Fund, established by the late Dr. Steyermark at the Missouri Botanical Garden, provided major funding for the flora. The Armand G. Erpf Fund provided financial support for part of the plant illustrations in this volume. Besides the illustrations by Bruno Manara, Karin Douthit did the drawings of Dicella julianii (Fig. 91), Diplopterys cabrerana (Fig. 92), Excentradenia adenophora (Fig. 93), Mezia huberi (Fig. 107), and Spachea elegans (Fig. 109). William R. Anderson did the drawings of Malpighiaceae styles and stigmas (Fig. 71). Review of Menyanthaceae was kindly provided by Garrett Crow. This project has benefited tremendously from the knowledge and assistance of Otto Huber, the main contributor to Volume 1 and an expert on the flora and vegetation of the Venezuelan Guayana. Numerous institutions and individuals in Venezuela have contributed significantly to different aspects of the flora. These include the Corporación Venezolana de Guayana and its affiliate Electrificación del Caroní, C.A., especially through the assistance of Alfredo Lezama, Hermán Roo, Luis Castro, Gabriel Picón, and Antonio Ahogado; the Ministerio del Ambiente y de los Recursos Naturales Renovables; the Instituto Nacional de Parques and the Dirección General de Parques Nacionales; the Servicio Autónomo para el Desarrollo Ambiental del Estado Amazonas; and the Consejo Nacional de Investigaciones Científicas y Tecnológicas. The Fuerzas Aéreas de Venezuela, Fundación Terramar, Charles Brewer-Carías, and the late Parker vii

viii P REFACE AND A CKNOWLEDGMENTS

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

Introduction The Flora of the Venezuelan Guayana is an illustrated, synoptical treatment of the native and naturalized vascular plants that occur in the southern Venezuelan states of Amazonas, Bolívar, and Delta Amacuro (see endpaper map). This area includes spectacular tabletop mountains called tepuis and is known for its high vascular plant endemism. The flora area covers almost 500,000 square kilometers and includes about half the area of the geologically ancient Guayana Shield. The project is centered at the Missouri Botanical Garden and is a collaborative effort with about 200 contributing authors worldwide. It is significant because it is the first effort to produce a comprehensive inventory and identification guide for the plants of such an extensive region of northern South America. Volume 1 of the Flora of the Venezuelan Guayana provides extensive background information for the taxonomic treatments that began in Volume 2. It includes chapters on the physical geography of the region and its human occupation, the history of botanical exploration, the classification and altitudinal zonation of vegetation types in the flora area, floristic and phytogeographical analyses, and the conservation importance of the region. There is also a section of color photographs of landscapes, vegetation types, and plants, as well as a key to the families of seed plants in the flora area. 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 are available separately. 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 contains 20 families from Araliaceae through Cactaceae; Volume 4 contains 35 families from Caesalpiniaceae through Ericaceae; and Volume 5 contains 29 families from Eriocaulaceae through Lentibulariaceae. Family names of flowering plants in the Flora of the Venezuelan Guayana follow those used by A. Cronquist in An Integrated System of Classification of Flowering Plants (Columbia University Press, New York. 1981). Also included are certain families published after Cronquist’s book, such as Euphroniaceae and Monotaceae. A complete listing of the families to be covered in the Flora of the Venezuelan Guayana with the numbers of genera and species currently recorded from the flora area appears in Appendix A of Volume 1. The 28 families that appear in Volume 6 and their ix

x INTRODUCTION

FAMILIES INCLUDED IN THIS VOLUME Genera Species Liliaceae Limnocharitaceae Lissocarpaceae Loasaceae Loganiaceae Loranthaceae Lythraceae Magnoliaceae Malpighiaceae Malvaceae Marantaceae Marcgraviaceae Mayacaceae Melastomataceae

10 2 1 1 4 8 5 1 23 15 9 5 1 49

13 2 2 1 25 37 27 2 153 55 59 16 3 427

Genera Species Meliaceae Mendonciaceae Menispermaceae Menyanthaceae Mimosaceae Molluginaceae Monimiaceae Monotaceae Moraceae Moringaceae Musaceae Myricaceae Myristicaceae Myrsinaceae

5 1 12 1 26 2 1 1 14 1 1 1 4 5 209

30 10 27 1 164 2 3 1 62 1 1 1 27 55 1207

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

INTRODUCTION

xi

Slightly more than half the species in the flora are illustrated by black-andwhite line drawings. Volume 6 includes 640 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 species entries. A key is included if more than one subspecies or variety is present. Forms found in the flora area do not have separate entries, but they are discussed under the next higher rank. Throughout the flora, author abbreviations follow Authors of Plant Names (R. K. Brummitt and C. E. Powell, editors. Royal Botanic Gardens, Kew. 1992). The only exception is the use of the abbreviation “H.B.K.” for species described in Nova Genera et Species Plantarum, by A. von Humboldt, A. Bonpland, and K. S. Kunth (Librairie Grecque-Latine-Allemande, Paris. 1815–1825). This series of seven volumes was published in two versions (quarto and folio), each with different page numbers. Page numbers cited in this flora refer to the quarto version, which is more widely available than the folio version, except for a few instances 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-

xii

INTRODUCTION

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

Contributors Frank Almeda Department of Botany California Academy of Sciences Golden Gate Park San Francisco, California 94118 William R. Anderson Herbarium, University of Michigan North University Building Ann Arbor, Michigan 48109-1057 Lennart L. Andersson University of Göteborg Department of Systematic Botany Carl Skottsbergs Gata 22 B S-413 19 Göteborg SWEDEN

Paul E. Berry Botany Department University of Wisconsin–Madison 132 Birge Hall 430 Lincoln Drive Madison, Wisconsin 53706 Alan E. Brant Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 David Brunner National Fish and Wildlife Foundation 28 Second St., 6th Floor San Francisco, California 94105

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

Lourdes Cárdenas Herbario Victor Manuel Badillo U.C.V., Facultad de Agronomía Apartado 4579 Maracay 2101a, Edo. Aragua VENEZUELA

Rupert C. Barneby (deceased) Herbarium The New York Botanical Garden Bronx, New York 10458-5126

Robert W. Cruden Department of Botany University of Iowa Iowa City, Iowa 52242

Cornelis C. Berg The Norwegian Arboretum University of Bergen N-5067 Store Milde NORWAY

Giovanna De Martino Herbario Victor Manuel Badillo U.C.V., Facultad de Agronomía Apartado 4579 Maracay 2101a, Edo. Aragua VENEZUELA

xiii

xiv

CONTRIBUTORS

Stefan Dressler Forschungsinstitut Senckenberg Senckenberganlage 25 D-60325 Frankfurt/M. GERMANY K. S. Edwards Herbarium Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB ENGLAND, U.K. Enrique Forero Facultad de Ciencias Universidad Nacional Apartado 7495 Santafé de Bogotá COLOMBIA Alina Freire-Fierro Herbario Nacional del Ecuador Apartado 17-15-755C Quito ECUADOR Paul A. Fryxell Section of Integrative Biology School of Biology University of Texas Austin, Texas 78712 James W. Grimes Herbarium Botany School University of Melbourne Parkville, Victoria 3052 AUSTRALIA Andreas Gröger Botanischer Garten Muenchen Menzinger Str. 65 D-80638 Muenchen GERMANY

Mats Hagberg Department of Systematic Botany University of Göteborg P.O. Box 461 SE 405 30 Göteborg SWEDEN Robert R. Haynes Department of Biological Sciences University of Alabama 425 Scientific Collections Building Box 870345 Tuscaloosa, Alabama 35487-0345 Lauritz B. Holm-Nielsen World Bank Education & Social Policies 1818 H Street, NW Washington, DC 20433 Bruce K. Holst Marie Selby Botanical Gardens 811 South Palm Ave. Sarasota, Florida 34236 Helen C. F. Hopkins Department of Biological Sciences Lancaster University Lancaster LA1 4YQ ENGLAND, U.K. Helen Kennedy Herbarium Department of Botany University of British Columbia Vancouver, British Columbia V6T 1Z4 CANADA Job Kuijt Herbarium, Department of Biology University of Victoria Box 1700 Victoria, British Columbia V8W 2Y2 CANADA

CONTRIBUTORS

Alicia Lourteig Herbier, Laboratoire de Phanérogamie Muséum National d’Histoire Naturelle 16 rue Buffon F-75005 Paris FRANCE Navina G. Luckana c/o Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Alan W. Meerow USDA-ARS-SHRS 13601 Old Cutler Road Miami, Florida 33158 Fabián A. Michelangeli Department of Ornithology American Museum of Natural History Central Park West at 79th Street New York, New York 10024 James S. Miller Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Thomas Morley Herbarium Plant Biology Department 220 Biological Sciences Center University of Minnesota St. Paul, Minnesota 55108 Elêna Maria de Lamare Occhioni Herbário Departamento de Botânica Instituto de Biologia, C.C.S. Universdade Federal do Rio de Janeiro Av. Brigadeire Trompowsky s.n. Ilha do Fundão ZC-32, 21941 Rio de Janeiro BRAZIL

XV

Ariane Luna Peixoto UFRRJ - Departamento de Botânica Caixa Postal 74582 23851-970 Seropédica, RJ BRAZIL Terence D. Pennington Herbarium Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB ENGLAND, U.K. John J. Pipoly III Fairchild Tropical Garden 11935 Old Cutler Road Coral Gables, Florida 33156-4299 Susanne S. Renner Biology Department University of Missouri–St. Louis 8001 Natural Bridge Road St. Louis, Missouri 63121-4499 Jon Ricketson Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Orbelia R. Robinson Department of Botany California Academy of Sciences Golden Gate Park San Francisco, California 94118 William A. Rodrigues Universidade Federal do Paraná, Setor de Ciencias Biológicas Herbário de Departamento de Botânica Centro Politécnico - Jardim das Américas CEP 81.531 - 970 Curitiba, Paraná BRAZIL

xvi

CONTRIBUTORS

Julian A. Steyermark (deceased) c/o Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Dieter C. Wasshausen Department of Botany, MSC-166 National Museum of Natural History Smithsonian Institution Washington, DC 20560 Kay Yatskievych Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

LILIACEAE by Alan W. Meerow and Robert W. Cruden Bulbous, cormous, rhizomatous, or tuberous perennials, mostly herbaecous, terrestrial, occasionally epiphytic, caulescent (the stems erect or climbing) or acaulescent. Leaves basal and/or cauline, alternate or whorled, occasionally opposite, sessile and linear or lorate (strap-shaped), or petiolate and lanceolate to widely elliptic; sometimes basally sheathing and forming an aerial pseudostem. Inflorescence spicate, racemose, paniculate, or umbellate (reduced helicoid cymes), leafy or naked, often bracteate; rarely wholly subterranean and appearing obsolete. Flowers 1–many, bisexual, rarely unisexual (the plants then dioecious), sometimes large and showy, sessile or pedicellate, actinomorphic or occasionally zygomorphic. Perianth cratershaped, salverform, funnelform, campanulate, tubular, rotate, or rarely ventricose, consisting of 6 (rarely 3, 4, or > 6) usually undifferentiated, subequal petaloid segments (tepals) in 2 series, connate below into a tube or free ± to the base; outgrowth of the perianth sometimes present, forming a conspicuous corona, or relatively inconspicuous and consisting of a ring of scales or fimbrae at the throat. Stamens 6 (sometimes 3, 4 or > 6), opposite the tepals and sometimes adnate to them, subequal or varying in length, free or variously connate; filaments sometimes variously modified or appendaged; anthers 2-locular, dorsifixed or basifixed, introrse, extrorse, or latrorse, or dehiscing through a terminal pore or slit, rarely connate and forming a tube. Ovary superior or inferior, sometimes adnate to the floral tube and semi-inferior, usually (1)3-locular, with septal nectaries; ovules few to numerous, the placentation usually axile, rarely basal or parietal; style 1(3), united or 3-branched, filiform, occasionally strumose (with cushion-like swellings); stigma capitate, 3-lobed or deeply trifid, sometimes distinctly 3. Mature fruit a loculicidally or septicidally dehiscent (sometimes indehiscent) capsule or a berry. Seeds fleshy or hard, sometimes flattened and winged, sometimes with a black or brown testa or a fleshy sarcotesta, usually with copious endosperm surrounding a straight or curved embryo. Cosmopolitan as broadly delimited here; ca. 300 genera and 6000 species, 10 genera and 13 species in the flora area.

Key to the Genera of Liliaceae by Alan W. Meerow and Robert W. Cruden 1. 1. 2(1).

Ovary superior, or if semi-inferior, the inflorescence a raceme or panicle ................................................................................................................ 2 Ovary inferior, or half inferior, the inflorescence an umbel ..................... 5 Shoot from a corm, some roots enlarged distally as a storage organ; tepals orange-yellow, not persistent; filaments linear, bearing scales at a right angle to the axis of the filament; basal leaves several-ranked, bifacial, bases of previous year’s leaves persistent as a fibrous collar; pedicels 1

2

L ILIACEAE

2.

3(2).

3.

4(3).

4.

5(1). 5. 6(5). 6. 7(6). 7. 8(5).

8. 9(8).

9.

Shoot from a rhizome; tepals blue, white-yellow (rarely green), ± persistent; filaments widened to the base, smooth; basal leaves 2-ranked, unifacial or apparently so, at least below, bases of previous year’s leaves not persistent; pedicels articulated immediately below the flower or unarticulated .......................................................................... 3 Tepals blue, rarely white; pedicels articulated immediately below the flower; capsule 15–25 mm long, not included within the ± persistent tepals; filaments enlarged below the anther; anthers ca. 3 mm long; bract subtending the lowest branch of the inflorescence (2.5–)4–15 cm long ............................................................................................ 5. Eccremis Tepals white-yellow (rarely green); pedicels not articulated; capsule < 12 mm long, usually enclosed by the persistent tepals; filaments without an enlargement below the anther; anthers 0.8–2 mm long; bract subtending the lowest branch of the inflorescence or flower < 1 cm long ............................................................................................. 4 Flowers subtended by 3 bractlets; tepals oblong-obovate, fused near their bases; anthers basifixed; ovary superior; capsules 9-ribbed, septicidal; margin of leaf a ± continuous vein ...................................... 9. Isidrogalvia Flowers ebracteate or subtended by 1 bract; tepals ± lanceolate, free to ovary; anthers dorsifixed near the base; ovary semi-inferior; capsules 6-ribbed, loculicidal; margin of leaf membranous ................. 10. Nietneria Bulbous geophytes; tepals united into a short or long tube ..................... 6 Rootstock a tuberous or corm-like rhizome or absent; true perianth tube absent ..................................................................................................... 8 Scape hollow; flowers zygomorphic ........................................ 7. Hippeastrum Scape solid, at least below middle; flowers actinomorphic ....................... 7 Stamens entirely free ...................................................................... 3. Crinum Stamens fused basally into a conspicuous staminal cup or false corona ............................................................................................ 8. Hymenocallis Rootstock a tuberous or corm-like, densely fibrous rhizome; leaves not resupinate; inflorescence acaulescent; ovary prolonged into a long, terminal, pilose outgrowth resembling a perianth tube .................. 4. Curculigo Rootstock obscure, glabrous; leaves usually resupinate; inflorescence caulescent; ovary not terminally elongated .......................................... 9 Plants never twining vines, producing separate vegetative and flowering stems; perianth zygomorphic; stamens declinate; sarcotesta absent ............................................................................................. 1. Alstroemeria Plants often twining vines, never with separate vegetative and flowering stems; perianth actinomorphic; stamens straight; seeds with a pulpy, often red sarcotesta ....................................................................2. Bomarea

1. ALSTROEMERIA L., Amoen. Acad. 6: 247. 1762. by Alan W. Meerow Herbaceous, erect perennials producing separate vegetative and flowering stems (except in one species). Roots thickened and usually tuberous at the apex. Vegetative stems ± erect, the leaves concentrated distally. Leaves usually resupinate, shortly petiolate to (sub)sessile, linear, lanceolate, oblong, or elliptic, obtuse, thintextured or ± succulent. Flowering stems generally taller than the vegetative ones, with leaves frequently less-developed and sometimes scale-like. Inflorescence um-

Alstroemeria 3

bellate (reduced helicoid cymes), simple or compound, bracteate, bearing 1–30 usually pedicellate flowers. Perianth zygomorphic, variously colored and often spotted; tepals 6, free, in 2 differently sized and shaped whorls, the lowermost tepal of the inner whorl different in size and marking than the other 2. Stamens 6, declinate, inserted at the base of the tepals; filaments filiform, unequal in length; anthers oblong, basifixed. Ovary inferior, turbinate, 3-locular; ovules numerous, globose, the placentation axile; style filiform, declinate; stigma trifid. Fruit a 3-valved, loculicidally dehiscent capsule, with a nipple-shaped projection at the apex. Seeds globose. Neotropics, most diverse in the southwestern Andes (chiefly Chile) and the eastern third of Brazil; 50–60 species, 1 in Venezuela. Alstroemeria amazonica Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 12. 1915. Herbaceous perennial; flowers red, nodding, tubular-campanulate, borne in a simple umbel; leaves of the flowering stems greatly reduced. Savannas, ca. 700 m; Bolívar (35 km west of Kamarata). Brazil (Pará). ◆Fig. 1.

Fig. 1. Alstroemeria amazonica

4

L ILIACEAE

2. BOMAREA Mirb., Hist. Nat. Pl. 9: 71. 1804. Collania Herb., Amaryllidaceae 102. 1837. Sphaerine Herb., Amaryllidaceae 106. 1837 Wichaurea M. Roem., Fam. Nat. Syn. Monogr. 4: 277. 1847. by Alan W. Meerow Herbaceous twining vines or erect to suberect herbs with unbranched, usually glabrous stems. Roots largely fibrous but frequently tuberous at the tip. Leaves alternate, short-petiolate to subsessile, resupinate, ovate, oblong, or lanceolate, acute, many-veined, usually glabrous but rarely with simple, unbranched hairs, slightly succulent; basal leaves sometimes scale-like. Inflorescence umbellate (reduced helicoid cymes), simple or compound, usually subtended by leaf-like bracts, the rays collectively bearing to 100 flowers, rarely 1-flowered. Perianth funnelform or tubular, actinomorphic, frequently red or reddish orange and yellow; tepals 6 in 2 whorls, free; the outer whorl oblong or lanceolate, similar, frequently sepaline, the inner whorl spatula-shaped or clawed, rarely obovate, often thinner and usually longer than the outer whorl, often spotted. Stamens 6, inserted at the base of the tepals; filaments filiform, straight; anthers oblong, basifixed. Ovary half- to completely inferior, 3-locular; ovules numerous, superposed, the placentation axile; style filiform; stigma trifid. Capsule turbinate or subglobose, 3-angled to obscurely 6-ribbed, truncate at the apex, dehiscent, indehiscent, or tardily dehiscent. Seeds subglobose, usually with a pulpy, often red, sarcotesta. Mexico, Guatemala, Costa Rica, Panama, West Indies, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Chile; ca. 100 species, ca. 10 species in Venezuela, 1 of these in the flora area.

Fig. 2. Bomarea edulis

Curculigo 5

Bomarea edulis (Tussac) Herb., Amaryllidaceae 111. 1837. —Alstroemeria edulis Tussac, Fl. Antill. 1: 109, pl. 14. 1808. Glabrous vine with lanceolate leaves; umbels compound, lax, 4–10(–20)-rayed, each ray bearing 1–4 pink and yellow-green, purple-spotted flowers. Lowland forest margins and clearings, 100–200 m; Bolívar (conflu-

ence of Río Caroní and Río Paragua), Amazonas (10 km south of Puerto Ayacucho). Northern Venezuela; Mexico, Central America, West Indies, northern South America. ◆Fig. 2. The edible root tubers are eaten, which may account for the wide range of this lowland species.

3. CRINUM L., Sp. Pl. 291. 1753. by Alan W. Meerow Generally large to massive bulbous geophytes; bulb often large, usually apically articulated into a neck, clumping or solitary. Leaves linear, lorate, lanceolate or ensiform, sessile (rarely subpetiolate), rosulate or distichous, evergreen or deciduous, often keeled at the midrib, frequently finely serrate or scabrous at the margin, sheathing at the base and sometimes forming an aboveground pseudostem, woolly fibers (vascular schlerenchyma) often appearing when torn. Scape emerging laterally, slightly compressed to ancipitous, solid. Flowers 1–many, large, showy, actinomorphic or slightly zygomorphic, salverform, funnelform, or campanulate, (sub)sessile or pedicellate, erect, declinate, or sub-pendulous, often fragrant. Tepals proximally connate into a long perianth tube, straight or curved, (sub)cylindrical, lobes linear to broadly lanceolate, subequal. Stamens 6, free, inserted at the throat of the tube, arcuate, ascending, or declinate. Ovary inferior, 3-locular, ellipsoid-turbinate; ovules 1–many per locule, the placentation axile, but embedded in and somewhat fused to the solid tissue of the ovary; style filiform; stigma capitate. Mature fruit a succulent, subglobose, 3-loculicidal capsule, often rostellate (by the persistence of the perianth tube), frequently bursting irregularly from pressure of the expanding seeds. Seeds subglobose or irregularly shaped, large, fleshy; endosperm primarily with a smooth (rarely papillose) greenish white testa, sometimes surrounded by a thin, gray-brown, water-repellent coat. Tropics and subtropics worldwide (most species in Africa); ca. 100 species, 1 in Venezuela. In South America, there are about 10 species that form a taxonomically difficult complex that may ultimately be reduced to a few variable species. Crinum erubescens Aiton, Hort. Kew 1: 413. 1789, non H.B.K. 1815 [1816]. Robust bulbous perennial, sometimes growing as an emergent aquatic; abaxial surface of the tepals purple-red. Occasional in

wet or swampy forests, near sea level to 300 m; Delta Amacuro (Caño Güiniquina at Morichito, Caño Mánamo at Boca de Tigre), Amazonas (upper Río Putaco). From southern Mexico through Brazil. ◆Fig. 3.

4. CURCULIGO Gaertn., Fruct. Sem. Pl. 1: 63. 1788. Embodium Salisb., Gen. Pl. 43. 1866. by Alan W. Meerow Cespitose plants with short, sometimes tuberous or corm-like rhizomes covered with reticulate fibers. Leaves basal, usually elongate-lanceolate, plicate, small and grass-like or sometimes large. Inflorescence spicate, the spikes sessile among the leaves and few-flowered or the flowers sometimes long-pedunculate, bracteate, the

6

L ILIACEAE

Fig. 3. Crinum erubescens

Fig. 4. Curculigo scorzonerifolia

bracts linear, often densely villous. Flowers small, actinomorphic. Perianth of 6 subequal tepals in 2 series; tepals yellow, free but borne at the apex of a long, slender terminal elongation of the ovary resembling a perianth tube. Stamens 6, inserted at the base of the tepals; filaments short; anthers linear, erect. Ovary inferior, 3-locular, usually rostellate; ovules 2–many per locule; style short, columnar; stigmas 3, oblong, erect. Fruit fleshy, indehiscent. Seeds subglobose, with a black testa. Virtually pantropical; ca. 30 species, 1 in Venezuela. Curculigo can be separated from Hypoxis by its stamen number (3 in Hypoxis) and fruit (a dehiscent capsule in Hypoxis). Curculigo scorzonerifolia (Lam.) Baker, J. Linn. Soc., Bot. 17: 124. 1878. —Hypoxis scorzonerifolia Lam., Encycl. 3: 183. 1789. Small, tufted, cormous herb, generally ≤

30 cm tall. Savannas, 100–500 m; Bolívar (Río Acanán, Río Paragua, base of Serranía de los Pijiguaos), Amazonas (La Esmeralda). Widespread in Mexico, Central America, West Indies, South America. ◆Fig. 4.

Eccremis 7

Fig. 5. Eccremis coarctata

5. ECCREMIS Willd. ex Baker, J. Linn. Soc., Bot. 15: 319. 1876. by Robert W. Cruden Rhizomatous, adventitious roots developing where stem is in contact with the soil. Stems decumbent to erect, 0.5–2 m tall. Basal leaves 2-ranked, sheath semi-ensiform; blade bifacial with a prominent midrib; cauline leaves well-developed. Inflorescence a panicle, each branch subtended by a large spathe-like bract, the lowest usually boat-shaped, (2.5–)4–15 cm long; pedicel articulated immediately below the flower, each subtended by a small bract. Tepals blue, rarely white, broadly ovate, with 5–7 prominent veins and 0–4 minor veins, ± persistent. Filaments yellow, smooth, enlarged below the anther, widened toward the base, epipetalous; anthers yellow, latrorse, ca. 3 mm long. Ovary superior. Capsules broadly oblong, 15–25 × 8–

8

L ILIACEAE

15 mm, septicidal, opening 10–15% of its length, not included within the ± persistent tepals. Seeds ovoid, lustrous black. Widely distributed in the Andes from Colombia to Bolivia and the mountains of Venezuela and adjacent Brazil; 1 species. Eccremis coarctata (Ruiz & Pav.) Baker, J. Linn. Soc., Bot. 15: 320. 1876. —Anthericum coarctatum Ruiz & Pav., Fl. Peruv. 3: 67, t. 299. 1802. Dianella dubia H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 270, t. 675. 1815 [1816]. Robust herb. Wet savannas, swampy

meadows, forests, dwarf forests, open sandy areas, 1800–2500 m; Bolívar (Macizo del Chimantá [Apacará-tepui]), Amazonas (Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Distrito Federal, Mérida, Táchira; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 5.

6. ECHEANDIA Ortega, Nov. Pl. Descr. Dec. 90, t. 18. 1798. by Robert W. Cruden Herbs from corms that are associated with enlarged storage roots. Basal leaves bifacial, several-ranked, linear, narrowly oblong to narrowly elliptic, the bases surrounded by a fibrous collar composed of previous years’ leaf bases. Inflorescence racemose or paniculate, usually with several flowers at a node; pedicels articulated near the middle. Flowers yellow, orange, or white, rarely pale yellow or cream, ± erect to pendent. Perianth segments free to base, narrowly elliptic or elliptic, reflexed to spreading, 3(5)-veined, not persistent. Filaments linear to clavate, smooth or bearing scales at right angles to the axis of the filament; anthers yellow, connate and forming a cone-like structure, or free and either ± versatile and latrorse or nonversatile and dehiscing terminally, dorsifixed near the base. Ovary oblong, superior; ovules ≥ 8 per carpel. Fruit a loculicidal capsule, broadly to narrowly oblong, rarely ± globose, shallowly 3-lobed. Seeds irregularly compressed and folded; seed coat black, colliculose (covered with little round elevations). Southwestern U.S.A., Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Bolivia, northern Argentina; ca. 90 species, 3 in Venezuela, 1 of these in the flora area. Echeandia bolivarensis Cruden, Ann. Missouri Bot. Gard. 76: 350. 1989. Plant several-branched; flowers yellow; cauline leaves 4 or 5, long-attenuate; storage

roots enlarged 3–6 cm from the corm; anthers free. Forests on igneous slopes, 100– 200 m; Bolívar (Los Pijiguaos). Endemic. ◆Fig. 6.

7. HIPPEASTRUM Herb., Appendix 31. 1831. by Alan W. Meerow Bulbs large or small, solitary or stoloniferous, sometimes producing offsets on the surface of the bulb scales. Leaves lorate, ± distichous, appearing after or at the same time as flowers. Scape terete, hollow; spathe bracts 2. Flowers 2–6, pedicellate, ± perpendicular to the scape or declinate, usually large and showy, variously colored, sometimes fragrant. Perianth funnelform, zygomorphic; tepals subequal, the outer series usually longer and narrower than the inner, proximally connate into a usually short, rarely long, tube; paracorona (staminal cup formed by the basal fusion of di-

Echeandia 9

Fig. 6. Echeandia bolivarensis

10

L ILIACEAE

Fig. 7. Hippeastrum elegans

Fig. 8. Hippeastrum puniceum

Hymenocallis 11

lated filaments) of scales, fimbrae, or a callose rim usually present at the throat between the tepals and the staminal filaments. Stamens 6, inserted at the rim of the throat; filaments filiform, declinate, ± biseriate in length, proximally fasciculate; anthers linear or narrowly oblong, versatile. Ovary inferior, 3-locular; ovules numerous, the placentation axile; style long, declinate; stigma trifid, obtusely 3-lobed or capitate. Fruit a globose-trigonous, 3-loculicidal capsule. Seeds usually numerous per locule and flat, discoid or D-shaped, rarely globose and turgid; testa brown or black. Costa Rica, West Indies, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, eastern Brazil, Bolivia, Argentina (most diverse in the Central Andes and eastern Brazil); 40–60 species, 2 in Venezuela, both in the flora area.

Key to the Species of Hippeastrum 1. 1.

Perianth tube ≥ 10 cm long ............................................................. H. elegans Perianth tube ≤ 3 cm long ........................................................... H. puniceum

Hippeastrum elegans (Spreng.) H. Moore, Baileya 11: 16. 1963. —Amaryllis elegans Spreng., Pl. Min. Cogn. Pug. 2: 59. 1815. Amaryllis solandriflora Lindl., Coll. Bot. pl. 11. 1821. —Hippeastrum solandriflorum (Lindl.) Herb., Appendix 31. 1821. Amaryllis longiflora Steud., Nomencl. Bot. ed. 2, 1: 70. 1841 [l840]. Scapes with 2–4 very long-tubed, greenish white flowers. Savannas, 100–500 m; Bolívar (Uruyen), Amazonas (upper Río Parucito). Costa Rica, Colombia, Guyana, Suriname, eastern Brazil. ◆Fig. 7. Hippeastrum puniceum (Lam.) Kuntze, Revis. Gen. Pl. 2: 703. 1891, published

in error as Hippeastrum purpureum. —Amaryllis punicea Lam., Encycl. 1: 122. 1783. —Hippeastrum puniceum (Lam.) Voss, Vilm. Blumengärtn ed. 2, 1: 1003. 1896 [1895]. —Hippeastrum puniceum (Lam.) Urb., Symb. Antill. 4: 151. 1903. Amaryllis eguestris Aiton, Hort. Kew. 1: 417. 1789. —Hippeastrum eguestre (Aiton) Herb., Appendix 31. 1821. Flowers orange (rarely pink or white) prominently marked greenish white in the perianth throat. Occasional in moist forests, often associated with human habitation, 100–200 m; Amazonas (near Boca Mavaca). Widespread throughout Central America, the West Indies, and the northern half of South America. ◆Fig. 8.

8. HYMENOCALLIS Salisb., Trans. Hort. Soc. London 1: 338. 1812. Chloretis Herb., Amaryllidaceae 219. 1837. by Alan W. Meerow Bulb globose, ellipsoid, or pear-shaped, tunicate, sometimes articulated apically into a neck or pseudostem comprised of clasping leaf bases that usually remain subterranean. Leaves 2–many, persistent or deciduous, sessile, rarely petiolate, linear-lorate, widely strap-shaped, or oblanceolate, rarely ovate to elliptic. Inflorescence scapose, umbellate; bracts 2–4. Flowers 1–many, usually sessile, erect or suberect, large and fragrant. Perianth crater-shaped, actinomorphic, the tube long, straight or slightly cernuous distally, (sub)cylindrical, green; tepals linear or lanceolate, subequal, spreading widely from the throat and often distally recurved or cernuous, white (rarely pale greenish yellow). Stamens connate below into a conspicuous funnelform or rotate, pure white staminal cup (false corona), often dentate

12

L ILIACEAE

or laciniate between each free filament; free filament inserted at the rim of the cup, filiform, straight, as long or longer than the cup, sometimes erect, often assurgent, frequently green; pollen sticky-granular, orange. Ovary inferior, 3-locular, globose, oblong-ellipsoid, or pear-shaped; ovules 2–10 per locule, the placentation axile and superposed (appearing basal and collateral when 2); style filiform, exserted beyond the stamens and often deflexed laterally, green; stigma capitate. Mature fruit a large, succulent, green, subglobose 3-loculicidal capsule which sometimes bursts irregularly from pressure of the expanding seed. Seeds 1–3 per locule, large, fleshy, with a spongy green testa; radicle sometimes emerging viviparously. Southeastern U.S.A, Mexico, Central America, West Indies, northern South America; ca. 50 species, 2 in Venezuela, 1 of these in the flora area. In the past, Hymenocallis has been considered synonymous with the paleotropical Pancratium L., but it differs markedly in seed morphology and chromosome number. Hymenocallis tubiflora Salisb., Trans. Hort. Soc. London 1: 341. 1812. —Pancratium tubiflorum (Salisb.) Schult. in Roem. & Schult., Syst. Veg. 7: 926. 1830. Pancratium undulatum H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 280. 1815 [1816]. —Hymenocallis undulata (H.B.K.) Herb., Appendix 44. 1821. —Hymenocallis guianensis var. undulata (H.B.K.) Herb., Amaryllidaceae 210. 1837. Pancratium guianense Ker Gawl., Bot. Reg. 1: t. 265. 1818. —Hymenocallis guianensis (Ker Gawl.) Herb., Appendix 44. 1821.

Pancratium petiolatum Willd. ex Schult. in Roem. & Schult., Syst. Veg. 7: 912. 1830. —Hymenocallis petiolata (Willd. ex Schult.) M. Roem., Fam. Nat. Syn. Monogr. 4: 168. 1847. Hymenocallis moritziana Kunth, Enum. Pl. 5: 668. 1850. Bulbous geophyte; leaves broadly elliptic, petiolate. Rainforest understory, 50–500 m, Delta Amacuro (Boca de Meregina near Guarucara, Caño Atoiba north of Caño Araguao, Caño Simoina west of Isla Cocuina), Bolívar (36 km northwest of El Palmar, south of Río Parguaza, Tumeremo). Trinidad, Guyana, Suriname, Brazil. ◆Fig. 9.

Fig. 9. Hymenocallis tubiflora

Isidrogalvia 13

9. ISIDROGALVIA Ruiz & Pav., Fl. Peruv. 3: 69, pl. 302. 1802. by Robert W. Cruden Perennial herbs from slender rhizomes. Leaves narrowly to broadly linear, mostly basal, 2-ranked, unifacial, attenuate, straight to strongly falcate; margin edged by a vein, entire or villous, the unribbed sheath entire, pink to reddish. Scape solitary, erect, with (1)2–17 usually small bracts, glabrous or with hairs lacking glands in the inflorescence. Inflorescence a raceme, each flower-bearing node subtended by a bract and each flower subtended by 3 bractlets; pedicels not articulated. Tepals white to cream (greenish in one species) at anthesis, becoming yellowish or reddish after anthesis, oblong-obovate, fused just above the base, persistent, 5–9 prominent veins. Stamens 6, latrorse, 0.5–0.8 times the length of the tepals; anthers basifixed, 2-locular; filaments linear to subulate, epitepalous and joined to the tepals at the top of the cup formed by the fused tepal bases. Styles united, or divided at apex; stigmas capitate or ± lateral. Capsules 3–9-ribbed, 4.5–10 mm long, septicidal, many-seeded. Seeds 2–3 mm long, fusiform, dark red, with an appendage on the chalazal end which is decurrent along the side to or near the other end and sometimes extends as an appendage, occasionally with no appendage. Widely distributed in the montane areas of Colombia, Venezuela, Guyana, Ecuador, Peru, and Brazil; 6 species, 4 in Venezuela, 2 of these in the flora area. Key to the Species of Isidrogalvia 1.

1.

Leaf margins entire; tepals greenish, 6–9 mm long and lowest fruitbearing pedicel 7–20 mm long; filaments 3–3.5 mm long; anthers 0.9–1.1 mm long ............................................................................ I. duidae Leaf margins densely hairy, rarely entire; tepals, at least the inner ones, white-cream, becoming yellow or green with age, 9–14 mm long or lowest fruit-bearing pedicel < 7 mm long; filaments 3.5–5.5 mm long; anthers mostly 1.2–1.8 mm long .................................. I. schomburgkiana

Isidrogalvia duidae (Steyerm.) Cruden, Syst. Bot. 16: 278. 1991. —Tofieldia duidae Steyerm., Field Mus. Nat. Hist., Bot. Ser. 28: 156. 1951. Plant slender, narrow-leafed; pedicels long, slender; flowers greenish. Margins and banks of streams and rivers, 1400–2000 m; Amazonas (summit of Cerro Duida). Endemic. ◆Fig. 10. Isidrogalvia schomburgkiana (Oliv.) Cruden, Syst. Bot. 16: 276. 1991. —Tofieldia schomburgkiana Oliv. in Thurn, Timehri 5: 206. 1886. Isidrogalvia guianensis Klotzsch in M.R.

Schomb., Reis. Br.-Guiana 3: 1065. 1848 [1849], “Isidrogalvis,” nom. nud. Plant generally robust; flowers white or cream-colored changing color with age, sometimes the inner and outer tepals different-colored, the outer olive-green and the inner whitish. Open dry slopes, sandstone outcrops, rocky and/or open savannas, wet savannas, moist river banks, humid scrub, pygmy forests, swamps, marshy areas, woods dominated by Clusia, Podocarpus, and Stenopadus, (900–)1700–2800 m; widespread in Bolívar and Amazonas. Adjacent Guyana and Brazil. ◆Fig. 11.

14

L ILIACEAE

Fig. 11. Isidrogalvia schomburgkiana Fig. 10. Isidrogalvia duidae

Fig. 12. Nietneria corymbosa

Fig. 13. Nietneria paniculata

Nietneria 15

10. NIETNERIA Klotzsch & M.R. Schomb. ex Benth. in Benth. & Hook. f., Gen. Pl. 3: 825. 1883. by Robert W. Cruden Perennial herbs from rhizomes. Basal leaves 2-ranked, unifacial, strongly ribbed, sometimes falcate; margin entire, membranous. Inflorescence a raceme or panicle; each branch subtended by a bract < 1 cm long, as well as the terminal flower of each branch; scape with 0–5(6) cauline leaves, sometimes reduced to bracts; pedicels not articulated. Flowers yellow to yellow-green at anthesis, most ebracteate. Tepals lanceolate to narrowly lanceolate, mostly 5–7-veined, the inner with a scarious margin, arising from the sides of the ovary, persistent. Filaments smooth, tapered from base to tip, adnate to the base of the tepal; anthers dorsifixed near the base, latrorse. Ovary semi-inferior; styles united nearly to apex; stigmas separate, terminal, ± crest-shaped. Capsule 4–6 mm long, loculicidal, lower portion 6-ribbed. Seeds tannish, darker at the ends, 1 mm long, 0.1 mm diameter, striate. Endemic to the Guayana Shield in Venezuela, Guyana, and Brazil; 2 species, both in the flora area. The specific nature of the these taxa merits additional study. Key to the Species of Nietneria 1.

1.

Leaves usually 4–10 mm wide with 15–28 ribs and/or cauline leaves 3–6, the lowest well-developed and usually overlapping the second; inflorescence a panicle, usually flat-topped; flowers 25 or more; scape > 50 cm tall or < 2 times the length of the basal leaves ....... N. corymbosa Leaves 2–3.5(–4) mm wide with 8–12(–14) ribs and cauline leaves 0–2, rarely 3 and/or overlapping; inflorescence usually a raceme, sometimes with 2-flowered branches below; flowers 8–18(–22); scape 34–51 cm tall and 2.3–3.5 times the length of the basal leaves .................. N. paniculata

Nietneria corymbosa Klotzsch & M.R. Schomb. ex B.D. Jackson, Index Kewensis 2: 314. 1895. —Nietneria corymbosa Klotzsch & M.R. Schomb. in M.R. Schomb., Reis. Br.-Guiana 3: 1066. 1848 [1849], nom. nud. Plant robust; leaves relatively long and broad. Wet savannas and elfin forests on tepuis, 1800–2800 m; widespread in Bolívar and Amazonas. Endemic. ◆Fig. 12. Collections from Sierra de la Neblina have one or two cauline leaves but otherwise are similar to collections from other localities. Small plants may have but two cauline leaves, narrow basal leaves, and < 25 flowers. The inflorescences of such plants tend to be quite short in contrast to the relatively

long and slender inflorescences of Neitneria paniculata. Nietneria paniculata Steyerm., Field Mus. Nat. Hist., Bot. Ser. 28: 153. 1951. Plant slender. Wet savannas, 1000– 1500(–2500) m; eastern Bolívar (Auyántepui, Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá [Apacará-tepui, Murey-tepui], Roraima-tepui). Adjacent Guyana, Brazil (Amazonas: Serra Aracá). ◆Fig. 13. The narrow leaves with few ribs and fewflowered inflorescences are characteristic. Plants from the higher elevations have leaves with more ribs than is typical for the species.

16

L IMNOCHARITACEAE

LIMNOCHARITACEAE by Robert R. Haynes and Lauritz B. Holm-Nielsen Perennial, glabrous herbs with milky juice, growing emersed, submersed, or floating-leaved, in fresh waters. Roots fibrous, few to many, nonseptate, from a stout rhizome or stolon. Stems fleshy, erect, unbranched, the internodes without spinulose teeth, the tips without turions (hardened dormant stem tips that develop into new plants in the subsequent growing season) or tubers. Leaves basal or alternate, petiolate; petioles terete to triangular, mostly 3 or more times the length of the blade, sheathing at the base, the infravaginal scales absent; blades orbicular to lanceolate, without pellucid markings, the margins entire, the apex obtuse to round-acute, the base cordate to attenuate, the venation reticulate, with parallel primary veins from base of blade to apex and reticulate secondary veins. Inflorescences scapose, terminal, umbelliform, erect to floating, an involucrate umbel, lacking a spathe, subtended by an involucre of few to several bracts, the bracts ovate, acuminate, membranous. Flowers hypogynous, bisexual, pedicellate; perianth actinomorphic, of 6 separate segments in 2 series, the outer 3 sepal-like, mostly erect, persistent, enclosing the fruit, the inner 3 petal-like, usually delicate, caducous deciduous. Androecium of 6–many stamens, separate; anthers 2-locular, basifixed, dehiscing by longitudinal slits; pollen 3–7-aperturate, globose, separate. Gynoecium of 3–many, separate or basally coherent carpels, the carpels 1-locular, each with numerous anatropous or campylotropous ovules, the placentation laminar; styles short or absent; stigma linear. Fruit a follicle, dehiscing adaxially. Seeds numerous, glandular-pubescent or costate, U-shaped; endosperm helobial in development, absent in mature seed. Mexico, Central America, Greater Antilles, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa, Asia, northern Australia, some species naturalized in southern U.S.A. and southeast Asia; 3 genera and 9 species, 2 genera and 2 species in the flora area. Key to the Genera of Limnocharitaceae 1. 1.

Plants submersed, with floating leaves; carpels 3–8, linear-lanceolate; blades about as broad as long; styles present ....................... 1. Hydrocleys Plants emersed, the leaves not floating; carpels numerous, semicircular; blades much longer than broad; styles absent ................... 2. Limnocharis

1. HYDROCLEYS Rich., Mém. Mus. Hist. Nat. 1: 368. 1815. Plants submersed, with floating leaves. Stems short, ascending; stolons often present, terete. Leaves basal, floating or submersed, the submersed sessile, phyllodial, the floating leaves long-petiolate, the petioles terete, septate, with a sheathing base; blades orbicular to oblong-lanceolate, the apex mucronate to obtuse, the base rounded to cordate. Inflorescence of few to many flowers, terminating a long septate scape, sometimes proliferating with leaves and stolons, the scapes few to many, terete, septate; bracts elliptic to lanceolate, delicate, separate, shorter than pedicel subtended. Flowers long-pedicellate, the pedicels cylindric, terete. Sepals green, coriaceous, erect, lanceolate, persistent, with or without a midvein, the apex cucullate; petals showy, yellow to white, delicate, oblong-obovate to orbicular, fuga-

Hydrocleys 17

cious, erect to spreading, longer than or shorter than sepals. Stamens 6–many, in 1– several series, the outer often sterile; filaments linear or lanceolate, flattened; anthers linear. Carpels 3–8, terete, linear-lanceolate, basally scarcely cohering, attenuate into the style; style curved inward, papillose at apex. Fruits ± terete, linear-lanceolate, membranous, without dorsal furrows, dehiscing along the inner margins. Seeds numerous, sparsely to densely glandular-pubescent. Southern Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, Ecuador, Brazil, Bolivia, Paraguay, Argentina, naturalized in southern U.S.A.; 5 species, 2 in Venezuela, 1 of these in the flora area. Hydrocleys nymphoides (Willd.) Buchenau, Abh. Naturwiss. Vereine Bremen 2: 2. 1871 [1869]. —Stratiotes nymphoides Willd., Sp. Pl. 4: 821. 1806. Submersed herb to 50 cm tall; leaves floating. Shallow water of lakes and temporary pools, near sea level to 200 m; Delta Amacuro (Clavellina, road between Tucupita and Cocuina), Bolívar (Tumeremo). Widespread elsewhere in Venezuela; southern U.S.A., Guatemala, Puerto Rico, Curaçao, Colombia, Trinidad, Guyana, Suriname, Ecuador, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 14.

Fig. 14. Hydrocleys nymphoides

18

L IMNOCHARITACEAE

2. LIMNOCHARIS Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 116. 1808 [1807]. Plants emersed. Stems short, rhizomatous; stolons occasional, erect. Leaves basal, emersed, long-petiolate; petiole triangular; blade lanceolate to oval, the apex acute to round, the base acute to cordate. Inflorescence of 1–many flowers, terminating an elongate scape, occasionally proliferating, the scapes to 10 in number, shorter than to equal the length of the petioles; bracts separate, delicate throughout, shorter than pedicel subtended. Flowers long-pedicellate, the pedicels somewhat dilated, often winged or inflated, erect to spreading in flower, recurved in fruit, trigonous. Sepals green, broadly ovate, obtuse, appressed; petals yellow, fugacious, ovate to suborbicular, longer than the sepals, erect to slightly spreading above the sepals. Stamens many, the outer ones often sterile; filaments linear, flattened; anthers linear. Carpels 15–20, laterally compressed, verticillate, scarcely coherent at the base; style absent, stigma extrorse. Fruits laterally compressed, semicircular, scarcely coherent, membranous, dorsally furrowed, dehiscent internally. Seeds numerous, transversely multicostate. Mexico, Central America, Greater Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Argentina, naturalized in southeast Asia; 2 species, both in Venezuela, 1 of these in the flora area. Limnocharis laforestii Duchass. ex Griseb., Bonplandia (Hanover) 6: 11. 1858. Limnocharis flava var. minor Micheli in A. DC., Monogr. Phan. 3: 90. 1881. Emergent aquatic herb; flowers yellow. Swampy areas near granitic outcrops along river, 50–100 m; Bolívar (along Río Orinoco at El Burro). Apure, Aragua, Guárico, Lara, Portuguesa, Zulia; Mexico, Central America, Dominican Republic, Colombia, Ecuador, Peru, Brazil, Bolivia, Argentina. ◆Fig. 15.

Fig. 15. Limnocharis laforestii

L ISSOCARPACEAE 19

LISSOCARPACEAE by Paul E. Berry Small, glabrous trees. Leaves alternate, simple, entire, pinnately veined; stipules lacking. Flowers solitary and axillary or in short axillary racemes, 2-bracteolate, actinomorphic, bisexual, sessile or shortly pedicellate. Calyx campanulate, with 4 imbricate lobes; corolla tubular, 4-lobed, the lobes convolute in bud, the throat bearing an 8-lobed corona. Stamens 8, in one series; filaments shortly connate, attached to the corolla tube below the middle; anthers linear, 4-sporangiate, 2-thecal, with an apiculate-prolonged connective, opening longitudinally. Ovary inferior, 4-carpellate; placentas axile, ovules 2 per locule, pendulous; style terminal, clavate; stigma shallowly 4-lobed. Fruit drupaceous, 1- or 2-seeded. Seeds with abundant endosperm and a large straight embryo. Colombia, Venezuela, Guyana, Peru, Brazil, Bolivia; 1 genus and 5 species, 2 species in the flora area. 1. LISSOCARPA Benth. in Benth. & Hook. f., Gen. Pl. 2: 671. 1876. Description and distribution as in family; 5 species, 2 of these in Venezuela, both in the flora area. Key to the Species of Lissocarpa 1.

1.

Mature leaves mostly > 10 cm long, lower order venation usually inconspicuous on lower surface; flowers several, grouped in axillary fasicles; seasonally flooded banks of lowland black-water rivers ....... L. benthamii Mature leaves mostly < 10 cm long; lower order venation prominent on lower surface; flowers solitary or few in a short raceme; montane tepuislope forests at 1000–1800 m elevation ................................ L. stenocarpa

Lissocarpa benthamii Gürke in Engl. & Prantl, Nat. Pflanzenfam. 4(1): 180. 1891. —Palo de carbón, Palo de lameya, Simure. Shrub or small tree 4–15 m tall; fruit orange-red when ripe. Seasonally flooded banks of black-water rivers, 50–200 m; Amazonas (Caño Caname, Caño San Miguel, Caño Yagua, Río Baría, Río Casiquiare, Río Pasimoni, Río Sipapo, Río Temi, San Carlos de Río Negro, upstream from San Fernando de Atabapo). Amazon basin in Colombia and Brazil. ◆Fig. 16. Two collections from cloud forests on ridges and slopes at 1300 m on Sierra de la Neblina (Nee 31144, MO; and Nee 31189, MO) appear more similar to Lissocarpa benthamii than to L. stenocarpa, but are out

of the normal altitudinal and habitat range for L. benthamii. Lissocarpa stenocarpa Steyerm., Ann. Missouri Bot. Gard. 74: 104. 1987. Shrub or small tree 2–10 m tall; flowers yellowish cream, fruit reddish orange. Montane forests, 1000–1800 m; Amazonas (Cerro Duida, slopes of Cerro Huachamacari, slopes of Cerro Marahuaka, saddle between Cerro Duida and Cerro Marahuaka, Cerro Parú). Endemic? (See below). R. Vásquez (Novon 3: 211, fig. 1. 1993) cited 17 specimens from lowland Peru under this species, but a number of them have considerably larger and wider leaves and narrower fruits; they may belong to a yet undescribed species.

20

L ISSOCARPACEAE

Fig. 16. Lissocarpa benthamii

Klaprothia 21

Fig. 17. Klaprothia mentzelioides

22

L OASACEAE

LOASACEAE by James S. Miller Herbs, usually perennial, sometimes climbing, occasionally shrubs or small trees, with hairs of various sorts, these usually silicified and sometimes calcified, sometimes stinging or glandular. Leaves alternate or opposite, simple or lobed, exstipulate. Inflorescences usually cymose or the flowers solitary. Flowers actinomorphic, epigynous. Sepals (4)5(–7), foliaceous, imbricate or convolute, persistent, basally adnate to the ovary; petals (4)5(–7), induplicate-valvate, free or connate in a tube. Stamens (5)10–many; filaments free, basally connate or connate in bundles opposite the petals; anthers bithecal, dehiscing longitudinally; pollen tricolporate. Ovary inferior, 3–5(–7)-carpellate or rarely 1-carpellate, unilocular with the placentation axile; style simple, terminal; stigma capitate or obscurely lobed; ovules numerous or rarely only 1, anatropous. Fruits dry, a loculicidal or septicidal capsule, or rarely indehiscent. U.S.A., Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia, Chile, Argentina, Ethiopia, Somalia, South Africa, Arabia; 14 or 15 genera and ca. 250 species, 1 species in the flora area. 1. KLAPROTHIA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 121. 1823. Annual herb, the stems inconspicuously pubescent, the hairs not stinging. Leaves opposite, serrate, petiolate, exstipulate. Inflorescence terminal, loosely cymose. Flowers actinomorphic, epigynous. Sepals 4, connate for the lower half in a subglobose tube; petals 4, free. Stamens opposite the petals in groups of 4 or 5, with 4 or 5 staminodia. Ovary inferior, 4-carpellate, unilocular, the placentation parietal; ovules few. Fruit a 4-valved, septicidal capsule. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; 2 species, both in Venezuela, 1 of these in the flora area. Klaprothia mentzelioides H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 123. 1823. Herb or vine to 1 m tall; corollas white. Montane forests, 600–1800 m; Amazonas (Si-

erra de la Neblina). Aragua, Distrito Federal, Mérida, Sucre, Táchira; Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 17.

LOGANIACEAE by Paul E. Berry and Alan E. Brant Trees, shrubs, lianas, or herbs. Leaves simple, opposite, rarely whorled, stipulate (stipules sometimes reduced to a fine line). Inflorescence terminal and/or axillary, cymose, generally a branched cyme, sometimes racemiform or spicate, rarely flowers solitary. Flowers actinomorphic to slightly irregular; nectary disk generally lacking. Sepals 4 or 5, free or connate, with imbricate teeth or lobes; corolla sympetalous, with 4 or 5 imbricate, convolute, or valvate lobes. Stamens epipetalous, as many as

Antonia 23

and alternate to the corolla lobes, or 1 in Usteria; anthers 2(4)-locular, opening longitudinally. Ovary superior, rarely semi-inferior, carpels and locules 2(3 or 5), sometimes 1-locular; ovules numerous or seldom few or solitary, the placentation axile or parietal; style terminal; stigma capitate or shortly lobed. Fruit a capsule, berry, or seldom a drupe. Seed sometimes winged; endosperm present; embryo straight. Widespread in the tropics and subtropics, and a few species in temperate areas; ca. 20 genera and ca. 500 species, 4 genera and 25 species in the flora area. We are treating the family here in the more traditional, broad sense, except that Potalia has been treated in the Flora under Gentianaceae. For a more atomized view of the Gentianales family circumscriptions based on cladistic analyses, see Struwe, L., V. A. Albert, and B. Bremer. 1994 [1995]. Cladistics and family level classification of the Gentianales. Cladistics 10: 175–206. Key to the Genera of Loganiaceae by Paul E. Berry 1. 1. 2(1). 2. 3(2). 3.

Herbs to small subshrubs ............................................................... 4. Spigelia Shrubs, trees, or woody lianas ................................................................... 2 Fruit a berry; leaves palmately 3- or 5-veined; lianas, shrubs, or small trees, often spiny or with tendrils rolled up in a spiral .......... 5. Strychnos Fruit a capsule; leaves pinnately veined; trees without spines or tendrils ................................................................................................................ 3 Flowers with an involucre of 8–15 closely spaced scales; corolla white or greenish at anthesis; seeds solitary in each locule .................... 1. Antonia Flowers without a scaley involucre; corolla purple at anthesis; seeds numerous in each locule .......................................................... 2. Bonyunia

1. ANTONIA Pohl, Pl. Bras. Icon. Descr. 2: 13. 1829. by Paul E. Berry Shrub or trees 1–30 m tall. Leaves subcoriaceous, oval-elliptic, obtuse to acute at the apex, rounded to attenuate at the base, 3–8 × 2–5 cm, glabrous to puberulent; adjacent petioles joined by an interpetiolar line. Inflorescence a terminal, subumbellate cyme, many-flowered and trichotomously branched. Flowers 5merous, with an involucre of 8–15 closely imbricate scales. Sepals free, scale-like; corolla white or greenish, firm, the tube pilose internally, the lobes linear, acuminate, spreading-reflexed, valvate in bud. Stamens 5, exserted; filaments awlshaped, with long, thin hairs at base. Ovary obovate, setose on upper part; style filiform, exserted, deciduous; stigma 2-lobed. Capsule oblong, coriaceous, 2(3)-locular and -valvate. Seeds solitary in each locule, rarely 2, oblong, compressed, winged at base and apex. Venezuela, Guyana, Suriname, Brazil, Bolivia; 1 species. Antonia ovata Pohl, Pl. Bras. Icon. Descr. 2: 13, t. 109. 1829. Antonia pilosa Hook., Icon. Pl. 1: t. 64. 1837. —Antonia ovata var. pilosa (Hook.) Progel in Mart., Fl. Bras. 6(1): 254. 1868.

Antonia pubescens Bong., Mém. Acad. Imp. Sci. St-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 3: 2, t. 1. 1835. Antonia ovata var. excelsa Paula, Acta Amazon. 6: 41. 1976.

24

L OGANIACEAE

Shrub or tree 4–10 m tall. In thin, low forests on white sand, open savannas and scrub, sandstone outcrops, 100–1200 m; Bolívar (southwestern talus slopes of Roraimatepui), Amazonas (Caño Caname at confluence with Río Atabapo, Río Manaviche at confluence with Río Orinoco). Guyana, Suriname, Brazil, Bolivia. ◆Fig. 18. José Elias de Paula (Acta Amazon. 6: 41– 42. 1976) recognized three varieties of Antonia ovata: var. ovata for (sub)glabrous shrubs 4–10 m; var. pilosa (Hook.) Progel for densely pilose shrubs; and var. excelsa Paula for large trees to 30 m tall in forests. However, many intermediate forms exist, and until a more thorough study can be done of the variation in the genus, we refrain from recognizing any infraspecific taxa.

Fig. 18. Antonia ovata

2. BONYUNIA M.R. Schomb. ex Progel in Mart., Fl. Bras. 6(1): 267. 1868. by Paul E. Berry Shrubs or small trees 1–10 m tall. Branches dark brown to black or grayish; bark finely fissured. Leaves coriaceous, shortly petiolate. Inflorescence terminal, lax, thyrsoid cymes, few-flowered, 2 or 3 times branched, mostly wider than long. Flowers 5-merous, actinomorphic except for the calyx lobes, subsessile. Calyx tubular-campanulate, persistent, lobes unequally dentate, mostly triangular; corolla at least 2 times longer than calyx, slender, ± cylindrical, often slightly contracted above the base and gradually widened toward the throat, creamy to violet in bud, turning purple at anthesis, the lobes narrowly oblong, valvate in bud, initially erect, then recurving. Stamens 5, included or barely exserted; filaments very short, about half as long as the anthers; anthers basifixed. Ovary superior (pedicel thickened below the calyx making the ovary appear semi-inferior), ± globose, 2-locular; ovules numerous; style pubescent, deciduous; stigma generally 2-lobed. Capsule oblong, 2–3 times as long as wide, pubescent outside, apex acute, 2-valved, septicidal. Seeds oblong, winged all around. Colombia, Venezuela, Guyana, Brazil; 4 species, 3 in Venezuela, all in the flora area. The fourth species in the genus, Bonyunia antoniiflora Progel, occurs in Brazil in Amazonas and Minas Gerais states. Key to the Species of Bonyunia 1. 1.

Leaves obovate or elliptic, rounded or retuse at the apex, glabrous ................................................................................................... B. aquatica Leaves ovate to elliptic, if elliptic then rounded at the apex and pubescent ................................................................................................................ 2

Bonyunia 25

2(1).

2.

Leaves mostly > 5 cm long, ± obtusely acuminate or acute at the apex, cuneate to rounded at the base; calyx lobes often narrowly oblong ..................................................................................................... B. superba Leaves mostly < 5 cm long, obtusely acute or rounded at the apex, subcordate to rounded at the base; calyx lobes triangular ..................... B. minor

Bonyunia aquatica Ducke, Arq. Inst. Biol. Veg. 1: 211. 1935. Shrub to tree 3–15 m tall. Savannas, black-water river banks, edges of granitic

outcrops, 100–300 m; Amazonas (Río Atabapo, Río Guasacavi, Río Guayapo, Río Guianía, tributaries of Río Orinoco, Río Pasimoni, lower Río Ventuari). Colombia (Caquetá, Guainía), Brazil (Amazonas). ◆Fig. 19.

Fig. 19. Bonyunia aquatica

Bonyunia minor N.E. Br., Trans. Linn. Soc. London, Bot. 6: 49, pl. 9, fig. 1–5. 1901. Bonyunia cinchonoides Gleason & Standl., Bull. Torrey Bot. Club 58: 448. 1931. Shrub or small tree 1–10 m tall; leaves orbicular-ovate, lustrous when dry, subsessile. Savannas, shrublands, and edges of gallery forests, (500–)800–1400 m; Bolívar (Ca-

Fig. 20. Bonyunia minor

Fig. 21. Bonyunia superba

26

L OGANIACEAE

naima, Cerro Guiaquinima, Gran Sabana and adjacent areas), Amazonas (Cerro Duida, Cerro Moriche, Cerro Yapacana). Guyana, Brazil (Amazonas, Roraima). ◆Fig. 20. Bonyunia superba M.R. Schomb. ex Progel

in Mart., Fl. Bras. 6(1): 267. 1868. Small tree (2–)7–10 m tall. Gallery forests, savannas, 500–1300 m; Bolívar (Arabopó, Cerro Arepuchi near the Río Caroní, near Kavanayén, Río Acanán near Amaruay-tepui, upper Río Karaurín). Guyana (base of Mount Roraima). ◆Fig. 21.

3. SPIGELIA L., Sp. Pl. 149. 1753. by Paul E. Berry Annual or perennial herbs or subshrubs. Leaves often in a whorl or pseudowhorl at stem apex, connected by interpetiolar lines, sheaths, or stipules; blades generally membranous. Inflorescences terminal, spike-like in cincinni. Flowers oriented in the same direction, sessile or nearly so, 2-bracteate. Calyx lobes 5, often slender, free or connate at the base, persistent; corolla usually funnelform, delicate, 5-lobed, valvate in bud, lobes shorter than the tube. Stamens 5; anthers included or rarely inserted. Ovary 2-locular; ovules several per locule; style slender, unbranched; stigma elongate-filiform, articulated in the middle, with distal portion deciduous. Fruit capsular, opening first by 4 valves, then circumscissily. Seeds subglobose or ovoid, sometimes subangulate. Principally South America, extending north into temperate North America; ca. 50 species, 7 in Venezuela, 5 of these in the flora area. Key to the Species of Spigelia 1. 1. 2(1). 2. 3(2). 3. 4(1).

4.

Leaves linear to narrowly lanceolate, < 1 cm wide ................................... 2 Leaves lanceolate to elliptic, > 1 cm wide ................................................. 4 Plants tiny, < 15 cm tall; flowers (including scars of fallen flowers) many, 20–40 per spike ..................................................................... S. polystachya Plants larger; flowers few, 1–7 per spike .................................................. 3 Leaves inconspicuous, linear, 10–13 mm long, 1–2 mm wide ........ S. gracilis Leaves conspicuous, narrowly lanceolate, > 20 mm long, generally 5–10 mm wide ...................................................................................... S. humilis Uppermost leaves sessile, the blades subdeltate, widest near base, much more abruptly narrowed at base than at apex; weak, generally unbranched herbs; corolla 6–9 mm long; style shorter than the calyx ................................................................................................. S. anthelmia Uppermost leaves petiolate, the blades generally elliptic, not markedly wider near base, mostly not abruptly more narrowed at base than at apex; more robust, often branched herbs; corolla 10–15 mm long; style longer than the calyx ............................................................. S. multispica

Spigelia anthelmia L., Sp. Pl. 149. 1753. —Lombricera. Spigelia nervosa Steud., Flora 26: 764. 1843. Annual herb 20–80 cm tall; flowers white to pink. Savannas, morichales, 50–500 m; Delta (Amacuro, Imataca, Tucupita), Bolívar

(near El Manteco). Widespread in Venezuela; U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Trinidad-Tobago, Ecuador, Peru, Brazil, naturalized in Africa and Indonesia. ◆Fig. 22. Spigelia anthelmia is used as a vermifuge against intestinal parasites.

Spigelia 27

Fig. 22. Spigelia anthelmia

Fig. 23. Spigelia gracilis

Fig. 24. Spigelia humilis

28

L OGANIACEAE

Spigelia gracilis A. DC., Prodr. 9: 6. 1845. Slender herb 20–30 cm tall, often in clumps; corolla lilac to purple. White-sand savannas, granitic outcrops, ca. 100–200 m; Amazonas (near Cerro Yapacana, Río Guianía, Río Sipapo, lower Río Ventuari). Colombia (Vichada), Brazil (Bahia). ◆Fig. 23.

Spigelia humilis Benth., J. Bot. (Hooker) 3: 240. 1841. Annual herb 10–30 cm tall; leaves sessile; corolla white to lilac. Rock outcrops, streamsides, disturbed areas, near towns, ca. 100– 500 m; Bolívar (Río Cuyuní), scattered throughout most of Amazonas. Colombia, Guyana, Amazonian Brazil. ◆Fig. 24.

Spigelia multispica Steud., Flora 26: 764. 1843. Spigelia filipes Rusby, Descr. S. Amer. Pl. 81. 1920. Herb 40–110 cm tall; corolla white to lilac. Semiopen sites, roadsides, streamsides in forests, igneous outcrops; 100–1400 m; Bolívar (Gran Sabana, Serranía de Imataca, Sierra de Lema). Widespread in northern Venezuela; Costa Rica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Spigelia polystachya Klotzsch ex Progel in Mart., Fl. Bras. 6(1): 258. 1868. Tiny herb 5–15 cm tall. Rocks along rivers in evergreen lowland forests, 100–300 m; Bolívar (Río Cuyuní near Anacoco). Guyana.

4. STRYCHNOS L., Sp. Pl. 189. 1753. Rouhamon Aubl., Hist. Pl. Guiane 93. 1775. Lasiostoma Schreb., Gen. Pl. 1: 75. 1789. Geniostoma Spreng., Syst. Veg. 1: 588. 1825 [1824], non J.R. Forst. & G. Forst. 1775. Narda Vell., Fl. Flumin. Icon. 3: pl. 24. 1827 [1831]. by Alan E. Brant and Paul E. Berry Lianas, less often shrubs or small trees, usually with tendrils, sometimes with spines, or both. Branches opposite, sometimes with conspicuous elliptic or orbicular lenticels; branchlets terete or flattened; spines simple, terete, curved or straight, to 3 cm long; tendrils generally emerging from the axils of cataphylls, simple, becoming coiled and increasing in diameter toward the tapering apex, to ca. 10 cm long; stipules interpetiolar, usually reduced to fine lines of slightly raised tissue, the nodes somewhat knobby in appearance. Leaves opposite, simple, entire; petioles 0–2 cm long, somewhat channeled on upper side, often narrowly winged on the margin; blades cordate or rounded to acute at the base, rounded to long-acuminate at the apex, membranaceous to coriaceous, 3–7-pliveined (rarely pinnately veined), the lowermost pair opposite or alternate and diverging at 0–3 cm from the base of the blade; indumentum often present, with simple hairs, sometimes barbate (with tufts of hairs) in the inner axils of the leaf veins on the lower surface. Inflorescences terminal or axillary or both, corymbose or globose to elongate-thyrsoid, sometimes racemose, densely to loosely flowered, the ultimate segments usually in 3-flowered cymes with the terminal flowers (sub)sessile and lateral flowers pedicellate, bracts and bracteoles present. Flowers 5- or sometimes 4-merous. Sepals united at the base, the lobes at maturity ± imbricate near base, free apically; corolla with a tubular portion 1–20 mm long, rotate to salverform and usually ± fleshy, the lobes valvate in bud, erect to reflexed at anthesis, shorter to longer than the corolla tube, white to yellowish or greenish white. Stamens alternate with and as many as the corolla lobes; filaments 0–2 mm long and attached at throat of corolla tube; anthers basally or medially attached to the filament, introrse. Style ≤ 20 mm long; stigma ±

Strychnos

29

capitate and shallowly lobed; ovary ovate to spherical, glabrous to pilose, 2-locular or occasionally 1-locular; ovules axile and numerous. Fruit a 1- or 2-locular berry, mostly ellipsoid to spherical or globose or variously flattened, 1–14 cm diameter, the outer layer 0.5–10 mm thick, glabrous, smooth, or variously tuberculate or warty, bluish green, brownish, yellow, or orange, dull or shining, the pulp soft, yellowish or whitish. Seeds 1–many, generally disk-shaped to spherical, sometimes narrowly winged, 5–15 mm long, coriaceous to bony. Pantropics; ca. 190 species, 18 species in Venezuela, 16 of these in the flora area. Many of the lianoid species of Strychnos in the flora area were once (and may still be) used as a major ingredient in the preparation of curare, which is extracted by scraping the bark of the stems and roots and boiling the scrapings in water (usually mixed with other ingredients) until a thick tar or syrup is obtained. The main active compound obtained from Strychnos is strychnine. Curare is locally used as an arrow poison by Amerindian groups and is a widely used common name for the genus (some other names are Bejuco de mavacure, Cruceta, and Hoja de mono). The pulp of the fruit is edible in most species, but the seeds are often poisonous. Key to the Species of Strychnos 1. 1. 2(1). 2. 3(2).

3.

4(1).

4.

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

6.

Leaf nodes, interpetiolar lines, and axillary buds glabrous ..................... 2 Leaf nodes, interpetiolar lines, and axillary buds pubescent, sometimes sparsely so .............................................................................................. 4 Lower surface of leaf blades glabrous, inflorescences axillary and terminal; fruit shell thick, smooth, and dark ................................. S. jobertiana Lower surface of leaf blades pubescent, pubescence visible only under magnification; inflorescences axillary; fruits various .......................... 3 Lower surface of leaf blades with minute, erect pubescence often visible only in the inner axils of the principal veins; fruits ca. 1.5 cm diameter, shell < 1 mm thick, shiny, smooth ......................................... S. panurensis Lower surface of leaf blades with minute, appressed pubescence throughout, most visible in the inner axils of the principal veins; fruits ca. 7 cm diameter, shell ca. 5 mm thick, dull, rough .................................. S. peckii Petioles ± 1 mm long; the raised portion of the stem at the base of the petiole bearing a membranous flap apically; inflorescences terminal; small trees (never lianas) ..................................................................... S. fendleri Petioles > 2 mm long; the raised portion of the stem at the base of the petiole not bearing a membranous flap apically; inflorescences various; lianas (may be shrubby when young) ...................................................... 5 Pubescence of leaf nodes, interpetiolar lines, and axillary buds erect or ascending, some trichomes > 0.5 mm long ............................................ 6 Pubescence of leaf nodes, interpetiolar lines, and axillary buds various, trichomes < 0.5 mm long ..................................................................... 13 Longest trichomes 2–3 mm long; upper and lower leaf blade surfaces ± densely ferruginous-pilose, petioles and twigs bearing a double indumentum with short curved hairs in addition to the longer ones; inflorescences terminal .................................................................. S. toxifera Longest trichomes < 2 mm long ................................................................. 7

30

L OGANIACEAE

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

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

12 13(5). 13. 14(13). 14. 15(14).

15.

16(14).

16.

17(16).

17.

Upper surface of leaf blades sparsely to obviously pubescent over entire surface with erect-ascending or curved trichomes ............................... 8 Upper surface of leaf blades pubescent only on the principal veins, sometimes with a few additional trichomes very near the blade margin .... 9 Lower surface of leaf blades densely ferruginous-barbate in the inner axils of the principal veins; inflorescences axillary .............. S. guianensis Lower surface of leaf blades not barbate in the axils of the principal veins; inflorescences terminal ................................................................ S. diaboli Lower surface of leaf blades with short membranous pockets in the inner axils of the principal veins (at least so on some of the leaves), often appearing barbate in age; inflorescences terminal ................. S. brachiata Lower surface of leaf blades without membranous pockets in the inner axils of the principal veins ................................................................... 10 Leaf blades ciliate, lacking additional trichomes near the margins on upper surface; inflorescences terminal and axillary .......... S. panamensis Upper surface of leaf blades bearing a few scattered long hairs near the margins in addition to being ciliate (sometimes only sparsely so) .... 11 Inflorescences terminal; fruits to 12 cm diameter ........................ S. davidsei Inflorescences axillary; fruits ca. 1.5 cm diameter ................................. 12 Leaf blades generally bicolorous, darker on the upper surface; principal veins on lower surface of leaf often reddish, usually ferruginousbarbate in the inner axils; leaves usually membranous ...... S. guianensis Leaf blades not bicolorous; not appearing barbate in the axils below; leaves coriaceous ........................................................................... S. cogens Trichomes of twigs, leaf nodes, interpetiolar lines, and/or axillary buds erect or ascending, minute .................................................................. 14 Trichomes of twigs, leaf nodes, interpetiolar lines, and/or axillary buds appressed, minute ................................................................................ 18 Lower surface of leaf blades barbate in the inner axils of principal veins; inflorescences terminal or axillary ...................................................... 15 Lower surface of leaf blades not barbate; inflorescences axillary .......... 16 Leaf barbation pale, fulvous, often sparse; inner axils of principal veins also with membranous pockets; plants bearing curved spines in addition to having tendrils; inflorescences terminal ........... S. mattogrossensis Leaf barbation ferruginous; inner axils of principal veins without membranous pockets; plants without spines; inflorescences axillary .............................................................................................. S. bredemeyeri Lower surface of leaf blades universally and minutely appressed-pubescent; fruits large, the shell thick, with roughened pustular eruptions over entire surface even when immature, yellowish brown in color ........................................................................................................ S. peckii Lower surface of leaf blades with a minute, erect or ascending pubescence on the principal veins, rarely glabrous; fruits small, the shell thin, smooth, and shiny ................................................................................ 17 Pubescence strictly erect; corolla lobes ca. 4 mm long, longer than the corolla tube; axillary panicles usually greater than 3 cm long ................................................................................................ S. panurensis Pubescence appressed to ascending; corolla lobes ca. 1.5 mm long, much

Strychnos 31

shorter than the corolla tube; axillary panicles usually not more than 3 cm long .............................................................................. S. bredemeyeri 18(13). Petioles very sparsely appressed-pubescent or glabrous; lower surface of leaves usually barbate in the axils of main veins; fruits ca. 4 cm diameter, dull, rough-textured ................................................. S. mitscherlichii 18. Petioles densely appressed pubescent; lower surface of leaves not barbate; fruits ca. 3 cm diameter, shiny, smooth ............................................... 19 19(18). Inflorescences terminal; corolla lobes densely pilose-bearded at the base within, papillose over the rest of the face within ............ S. rondeletioides 19. Inflorescences axillary; corolla lobes densely pilose bearded at the base within, pilose over the rest of the face within ........................ S. erichsonii Strychnos brachiata Ruiz & Pav., Fl. Peruv. 2: 30. 1799. Strychnos ruizii Sprague & Sandwith, Bull. Misc. Inform. Kew 1927: 130. 1927. Strychnos blackii Ducke, Bol. Técn. Inst. Agron. N. 19: 22. 1950. Liana; inflorescence terminal; corolla tube ca. 10 mm long, densely soft-pubescent outside; fruits large, to 14 cm diameter, shell ca. 2 mm thick, pulp edible. Semideciduous forests, ca. 400 m; Bolívar (Panare village of Corozal ca. 6 km from Maniapure). Apure, Barinas; Colombia, Peru, Brazil, Bolivia. Strychnos bredemeyeri (Schult. & Schult. f.) Sprague & Sandwith, Bull. Misc. Inform. Kew 1927: 128. 1927. —Lasiostoma bredemeyeri Schult. & Schult. f., Mantissa 3: 64. 1827. —Rouhamon bredemeyeri (Schult. & Schult. f.) A. DC., Prodr. 9: 18. 1845. —Bejuco de mono, Cruceta, Hoja de mono. Rouhamon pedunculatum A. DC., Prodr. 9: 561. 1845. Strychnos pedunculatum (A. DC.) Benth., J. Proc. Linn. Soc., Bot. 1: 105. 1856. Strychnos schomburgkiana Klotzsch ex M.R. Schomb., Reis. Br.-Guiana 3: 1144. 1848, nom. nud. Strychnos darienensis Seem., Bot. Voy. Herald 166. 1854. Strychnos trinitensis Griseb., Fl. Brit. W. I. 407. 1861. Liana; inflorescence axillary, corolla tube 3–5 mm long, lobes 1.5–2 mm long; ripe fruits yellow, 1.5–4 cm diameter, shell ca. 0.2 mm thick. Semideciduous to evergreen lowland forests, riparian forests, 50–1100 m; Delta Amacuro (Caño Jota-Sabuca between Caño Mariusa and Laguna del Consejo,

Serranía de Imataca), Bolívar (La Escalera at km 119 south of El Dorado, islands in Lago Guri, middle Río Caura, Río Tírica on slopes of Torono-tepui), Amazonas (Río Casiquiare, upper Río Cuao, Río Orinoco near mouth of Río Ventuari, San Carlos de Río Negro, San Simón de Cocuy, Síquita on east bank of Río Orinoco). Apure, Distrito Federal, Sucre; Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 26. Strychnos cogens Benth., J. Bot. (Hooker) 3: 241. 1841. —Bejuco de cruz, Bejuco de estribo, Curare. Liana; inflorescence axillary, corolla tube ca. 2 mm long, the lobes ca. 4 mm long; mature fruits to 2.5 cm diameter, shell thin (ca. 0.75 mm thick), yellow. Evergreen lowland forests, 50–200 m; Bolívar (south of El Tigre near Rio Cuchivero, Guaniamo, 20 km east of La Paragua), Amazonas (San Carlos de Río Negro, between Tamatama and La Esmeralda). Colombia, Guyana, French Guiana, Peru, Brazil, Bolivia. Strychnos davidsei Krukoff & Barneby, Phytologia 46: 67. 1980. Liana; corolla tube ca. 7 mm long, densely pilose outside, lobes ca. 2 mm long; mature fruits large, to 12.5 cm diameter, shell ca. 1.5 mm thick. Deciduous forests and surrounding gallery forests, 200–300 m; Bolívar (Represa Guri 55 km northeast of Cuidad Piar). Apure. Strychnos diaboli Sandwith, Kew Bull. Misc. Inform., Addit. Ser. 1931: 486. 1931.

32

L OGANIACEAE

Liana; corolla tube ca. 8 mm long, lobes ca. 3 mm long; mature fruits 4–4.5 cm diameter, shell ca. 6 mm thick. Evergreen lowland and semideciduous forests, 100–200 m; Bolívar (20 km east of La Paragua, lower Río Caura 25 km southwest of Maripa, mouth of Río Paragua), Amazonas (middle Río Cataniapo). Colombia (Amazonas), Guyana, Brazil (Amazonas, Roraima). Strychnos erichsonii M.R. Schomb. ex Progel in Mart., Fl. Bras. 6(1): 274. 1868. —Curare, Uruichá. Strychnos urbanii Barb. Rodr., Vellosia ed. 2, 1: 38, pl. 4, fig. A. 1885–1888 [1891]. Strychnos bovetiana Pires, Bol. Técn. Inst. Agron. N. 38: 40. 1960. Liana; corolla tube to 8 mm long, papillose on the outside; fruit to 3.5 cm diameter, shell ca. 2 mm thick. Evergreen lowland forests, 50–300 m; Bolívar (Cerro Camarón southwest of Cerro Guaiquinima, upper Río Paragua), Amazonas (Caño Coramoni off the Río Casiquiare, middle Río Baria, Río Casiquiare, San Carlos de Río Negro, southwestern base of Serranía de la Neblina). Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Strychnos fendleri Sprague & Sandwith, Bull. Misc. Inform. Kew 1927: 129. 1927. —Cruceta real, Palo de loro. Small tree to 10 m tall, with sharp spines, no tendrils; fruit orange, ca. 2.5 cm diameter. Deciduous to semideciduous forests, 50–700 m; widespread in dry parts of northern Bolívar. Anzoátegui, Aragua, Distrito Federal, Falcón, Guárico, Lara, Miranda, Nueva Esparta, Sucre, Zulia; northeastern Colombia, Brazil (Roraima). ◆Fig. 25. Strychnos guianensis (Aubl.) Mart., Syst. Mat. Med. Bras. 121. 1843. —Rouhamon guianensis Aubl., Hist. Pl. Guiane 93. 1775. Lasiostoma curare H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 210. 1825. —Strychnos curare (H.B.K.) Benth., J. Proc. Linn. Soc., Bot. 1: 108. 1856. Strychnos lanceolata Spruce ex Benth., J. Proc. Linn. Soc., Bot. 1: 105. 1856.

Liana; corolla tube ca. 1.5 mm long, lobes ca. 6 mm long; fruit yellow to orange, ca. 1.5 cm diameter. Riparian forests, evergreen lowland forests, edges of Mauritia palm swamps, 50–1000 m; Bolívar (El Carmen on Río Parguaza, La Escalera, Río Caura at mouth of Río Nichare, Río Parguaza, Río Parhueña, upper Río Venamo, Serranía de Imataca), Amazonas (base of Cerro Yapacana, slopes of Cerro Aracamuni, Cucurital near middle Caño Yagua, La Esmeralda, Río Atabapo, Río Baría, Río Cataniapo, Río Guianía, Río Mawarinuma, Río Pasimoni, mouth of Río Samariapo, Río Sipapo, Río Yatúa, San Carlos de Río Negro). Apure, Barinas, Sucre, Táchira; Colombia, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil, Bolivia. ◆Fig. 28. This is one of the most common species of Strychnos in South America. It is widely used in the preparation of curare. Strychnos jobertiana Baill., Adansonia 12: 367. 1879. Strychnos trichostyla Ducke, Bull. Mus. Hist. Nat. (Paris) sér. 2, 4: 746. 1932. Liana; corolla tube ca. 10 mm long; fruit to 4.5 cm diameter, shell ca. 6 mm thick. Evergreen lowlands forests, lower montane gallery forests, 100–500 m; Bolívar (Río Acanán), Amazonas (southeastern base of Cerro Duida, Río Cataniapo, Río Mawarinuma, San Carlos de Río Negro). Táchira; Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. Strychnos mattogrossensis S. Moore, Trans. Linn. Soc. London, Bot. 4: 392. 1895. Liana; leaves membranous; corolla tube ca. 0.2 mm long, lobes ca. 1 mm long; fruits globose, ca. 1.5 cm diameter, the testa fibrous. Riparian forests, near sea level to 300 m; Delta Amacuro (Sacupana), Bolívar (Salto Pará), Amazonas (upper Río Orinoco at Raudal de los Guaharibos on upper Río Amazonas). Apure, Cojedes, Portuguesa, Táchira, Yaracuy, Zulia; Colombia, Amazonian Peru, Brazil, Bolivia. Strychnos mitscherlichii M.R. Schomb., Reis. Br.-Guiana 2: 451. 1848 [1849].

Strychnos 33

—Strychnos smilacina Benth., J. Proc. Linn. Soc., Bot. 1: 105. 1857. High-climbing liana; corolla tube ca. 8 mm long; mature fruits ca. 4 cm diameter, shell to 2 mm thick, dull, rough-textured, smoother when immature. Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 varieties, 1 in Venezuela. S.

mitscherlichii var. mitscherlichii. —Curari-yek (Arekuna). Semideciduous to evergreen lowland forests, riparian and montane forests, 100– 1100 m; Bolívar (20–25 km southwest of El Manteco, La Escalera, Macizo del Chimantá [base of Chimantá-tepui], Río Paragua), Amazonas (Rio Baría, Río Casiquiare, Río Yureba). Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas, Mato Grosso), Bolivia. ◆Fig. 29. Strychnos panamensis Seem., Bot. Voy. Herald 166. 1854. —Curare, Mancalina (Panare). Liana; corolla tube ca. 1.5 cm long; mature fruit to 9 cm diameter, shell ca. 1.5 mm thick. Riparian forests, 100–200 m; Bolívar (San Vicente south of El Tigre), Amazonas (32 km south of San Carlos de Río Negro). Widespread elsewhere in northern Venezuela; Mexico, Central America, Colombia, Guyana, French Guiana, Ecuador, Amazonian Peru, Brazil (Roraima), Bolivia. Strychnos panurensis Sprague & Sandwith, Bull. Misc. Inform. Kew 1927: 132. 1927. —Kumaduwa (Yekwana). Liana (shrub-like when young); corolla tube 1–2.5 mm long, lobes ca. 4 mm long; fruits ± 1.5 cm diameter, yellow. Edges of riparian forests, 50–300 m; Bolívar (Puerto Ordaz, lower and middle Río Caura, middle Río Paragua), Amazonas (Caño Yureba, Río Cataniapo, Río Manapiare, Río Ventuari). Apure, Guárico; Panama, Colombia (Valle), French Guiana, Ecuador, Amazonian Peru, Brazil. ◆Fig. 27. Strychnos peckii B.L. Rob., Proc. Amer. Acad. Arts 49: 504. 1913. —Cachete de vieja.

High-climbing liana, often shrub- or treelike when young; corolla tube ca. 9 mm long; fruits ca. 7 cm diameter, shell brown and ca. 5 mm thick. Riparian and evergreen lowland to lower montane forests, 100–400 m; Bolívar (Río Orinoco near Ciudad Bolívar, middle Río Paragua, Río Suapure, Río Torono, Salto Pará), Amazonas (Río Baría, Rio Casiquiare above Chapazón, Río Cunucunuma, upper Río Orinoco, lower Río Ventuari, San Carlos de Río Negro, San Pedro de Cataniapo, between Tamatama and La Esmeralda). Apure, Aragua, Miranda, Yaracuy; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 30. Strychnos rondeletioides Spruce ex Benth., J. Proc. Linn. Soc., Bot. 1: 104. 1856. —Cumarawa-yek (Arekuna), Curare de pava, Hoja de mono, Manewa (Piaroa). Strychnos smithiana Krukoff, Brittonia 4: 287. 1942. Liana; corolla tube to 8 mm long; fruits to 3.5 cm diameter, shell to 2 mm thick. Riparian forests (especially along black-water rivers), evergreen lowland forests, montane forests, 50–1200 m; Bolívar (between Kavanayén and Río Pacairao), Amazonas (Cariche, Macuruco, Ocamo, Río Casiquiare, Río Emoni in lower Río Siapa basin, Río Mawarinuma, Río Negro, Río Siapa). Colombia, French Guiana, Peru, Brazil, Bolivia. Strychnos rondeletioides is used to prepare curare in both Bolívar and Amazonas states. Strychnos toxifera R.H. Schomb. ex Benth., J. Bot. (Hooker) 3: 240. 1841. —Cruceta, Mankowa (Panare). Strychnos syntoxica Sprague & Sandwith, Bull. Misc. Inform. Kew 1927: 129. 1927. Liana; corolla tube ca. 15 mm long; fruit ca. 7 cm diameter, shell ca. 2.5 mm thick. Semideciduous forests, evergreen lowland forests, riparian forests, 50–400 m; Delta Amacuro (Serranía de Imataca), Bolívar (Panare village of Corozal 6 km from Maniapure, Amazonas (Río Casiquiare, Río Cataniapo, upper Río Cuao, San Carlos de Río Negro). Aragua; Panama, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia.

34

L OGANIACEAE

Fig. 25. Strychnos fendleri

Fig. 26. Strychnos bredemeyeri

Strychnos 35

Fig. 27. Strychnos panurensis

Fig. 28. Strychnos guianensis

36

L OGANIACEAE

Fig. 29. Strychnos mitscherlichii var. mitscherlichii

Fig. 30. Strychnos peckii

L ORANTHACEAE 37

LORANTHACEAE by Job Kuijt Leafy plants, parasitic on branches and trunks of trees and shrubs, in one case (Gaiadendron) parasitic on roots of terrestrial or epiphytic plants; with or without roots at the base and/or on the stem, usually glabrous. Leaves various, simple, mostly decussate; margins entire. Inflorescence variable, determinate or indeterminate, the flowers in monads, dyads, or triads, commonly subtended by bracts which may be fused with the axillary stalk. Flowers bisexual or, by reduction, plants dioecious. Calyx reduced to an inconspicuous rim (calyculus) crowning the inferior ovary; petals 4–7, free or connate, mostly brightly colored when large. Stamens epipetalous; anthers dorsi- or basifixed, commonly dimorphic. Style usually as long as petals or nearly so. Fruit a 1-seeded berry. Seed surrounded by viscin tissue; endosperm white except in Aetanthus and Psittacanthus, where endosperm absent and embryo massive, sometimes polycotylar; elsewhere, embryo dicotylous, green. Largely in the tropics and subtropics around the world; ca. 60 genera and 1000 species, 8 genera and 37 species in the flora area. In the past, the family has usually included Viscaceae (as subfamily Viscoidae) and Eremolepidaceae (often as a tribe of subfamily Viscoideae). All genera are limited to either the Old World or the New World. Key to the Genera of Loranthaceae 1. 1. 2(1). 2. 3(2).

3.

4(2). 4.

5(4).

5.

Inflorescences determinate, i.e., terminated by a single flower; at least the upper lateral units mostly ebracteolate.......................... 1. Cladocolea Inflorescence indeterminate, i.e., not terminated by a single flower; lateral units bracteolate throughout ......................................................... 2 Inflorescence monadic, i.e., lateral units single flowers ........................... 3 Inflorescence dyadic or triadic ................................................................... 4 Flowers mostly sunken in depression in inflorescence axis, flanked by delicate, ribbon-like bracteoles scarcely exceeding the subtending bract; each pollen face with 3 large, circular depressions; leaves with stellate fiber bundles ........................................................... 3. Oryctanthus Flowers not usually in a cavity, the bracteoles large and ± keeled, extending well above the bract or quite separate from it; pollen lacking circular depressions, tricolpate or nearly so; leaves without stellate fiber bundles ....................................................................................... 4. Oryctina Flowers yellow to whitish; both bracts and bracteoles foliaceous; terrestrial, rarely epiphytic .......................................................... 2. Gaiadendron Flowers not yellow, often (partly) red; bracteoles never foliaceous, bracts only very rarely so; not terrestrial, except occasionally seemingly so in Tripodanthus .......................................................................................... 5 Flowers large (petals > 3 cm long) and colorful, usually with red and yellow; endosperm lacking; stem roots absent (but see P. julianus) ............................................................................................ 6. Psittacanthus Flowers small (petals < 2 cm long) and whitish or drab; endosperm present; basal and/or stem roots present .............................................. 6

38

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

7.

L ORANTHACEAE

Median flower of triad with pedicel as long as that of lateral flowers; petals > 10 mm long ........................................................... 8. Tripodanthus Median flower sessile, or pedicel much shorter than that of lateral flowers; petals < 5 mm long .......................................................................... 7 Plants dioecious; anthers mostly versatile, 4-locular, the free filaments not laterally excavated; inflorescence unbranched, axillary only ............................................................................................ 7. Struthanthus Plants dioecious or flowers bisexual; anthers basifixed, 2- or 4-locular, the free filaments conspicuously excavated laterally (sometimes anthers ± sessile); inflorescence branched or unbranched, axillary but often also terminal .................................................................................... 5. Phthirusa

1. CLADOCOLEA Tiegh., Bull. Soc. Bot. France 42: 166. 1895. Loxania Tiegh., Bull. Soc. Bot. France 42: 386. 1895. Glabrous or short-pubescent parasitic shrubs, some with epicortical roots from stems and/or base. Leaves decussate, alternate, or irregular, simple, rarely reduced to scales. Flowers bisexual or plants dioecious. Inflorescence a mostly determinate spike (rarely, a raceme) of ebracteolate monads at least in upper portion; rarely, lateral flowers bracteolate, or even triads below instead of monads. Flowers 4–6merous. Stamens di- or monomorphic, epipetalous; anthers mostly lacking extended connective, sometimes sessile. Style straight or contorted. Fruit 1-seeded. Seed enclosed in viscin; endosperm present, whitish; embryo green, dicotylous. Tropics and subtropics of the Americas, especially well developed in central Mexico; ca. 25 species, 4 in Venezuela, 3 of these in the flora area. Key to the Species of Cladocolea 1. 1. 2(1). 2.

Lateral inflorescence units bracteolate monads, or with triads below ................................................................................................ C. micrantha Lateral inflorescence units ebracteolate monads ..................................... 2 Leaves alternate, orbicular or nearly so, apiculate; inflorescence lacking basal scale leaves ................................................................... C. intermedia Leaves decussate, elliptic to ovate, not apiculate; inflorescence with basal scale leaves ........................................................................... C. roraimensis

Cladocolea intermedia (Rizzini) Kuijt, comb. nov. —Phthirusa intermedia Rizzini, Ernstia 24: 18. 1984. Cladocolea apiculata Kuijt, Brittonia 30: 447. 1987. Sparsely branched, young parts densely furfuraceous; leaves alternate, ovate to orbicular, apiculate, the tooth smooth. Secondary forests, 50–200 m; Bolívar (confluence of Río Paragua and Río Caroní). Endemic. Cladocolea micrantha (Eichler) Kuijt, Syst. Bot. 16: 288. 1991. —Phthirusa micrantha Eichler in Mart., Fl. Bras. 5(2): 65. 1868. —Passovia micrantha (Eichler) Tiegh., Bull. Soc. Bot. France

42: 172. 1895, “Passowia.” —Struthanthus micranthus (Eichler) Baehni & Macbr., Candollea 7: 290. 1937. Phthirusa micrantha var. bolivariensis Rizzini, Rodriguésia 41: 12. 1976. Phthirusa bernardiana Rizzini, Rodriguésia 41: 12. 1976. An extremely inconspicuous plant, to ca. 1 m in diameter; inflorescences exceedingly small, variable in sometimes being strictly monadic, but mostly triadic at the base and monadic above, the monads are mostly but not always bracteolate. On trees, 400–900 m; Bolívar (near Río Icabarú and Río Hacha). Guyana, Suriname, French Guiana, Brazil (Amazonas). ◆Fig. 31.

Gaiadendron 39

Cladocolea roraimensis (Steyerm.) Kuijt, J. Arnold Arbor. 56: 326. 1975. —Phthirusa roraimense Steyerm., Fieldiana, Bot. 28: 224. 1951. Rigid plant with somewhat succulent

stems; leaves decussate, very coriaceous, elliptical to ovate, apex rounded to slightly notched at maturity. Base of sandstone bluffs, 2000–2300 m; Bolívar (southwestern slopes of Roraima-tepui). Endemic.

2. GAIADENDRON G. Don, Gen. Hist. 3: 428. 1834. Taguaria Raf., Sylva Tellur. 125. 1838. Phrygilanthus Eichler in Mart., Fl. Bras. 5(2): 48. 1868, pro parte. Shrubs or small trees, terrestrial or sometimes on tree branches, parasitic on roots of other vascular plants. Leaves dark green and shiny, decussate. Inflorescence a raceme of triads with bracts and bracteoles extended into green foliaceous organs.

Fig. 31. Cladocolea micrantha

Fig. 32. Gaiadendron punctatum

40

L ORANTHACEAE

Flowers bisexual, yellow (whitish). Anthers versatile, dorsifixed. Fruit a rather large berry, pulpy, dull orange. Seed not viscid; endosperm deeply grooved, white. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia; 1 species. Gaiadendron punctatum (Ruiz & Pav.) G. Don, Gen. Hist. 3: 431. 1834. —Loranthus punctatus Ruiz & Pav., Fl. Peruv. 3: 46. 1802. —Phrygilanthus punctatus (Ruiz & Pav.) Eichler in Mart., Fl. Bras. 5(2): 48. 1868. Loranthus lanceolatum Ruiz & Pav., Fl. Peruv. 3: 47, t. 278b, 1802, non P. Beauv. 1808. —Notanthera lanceolata (Ruiz & Pav.) G. Don, Gen. Hist. 3: 428. 1834. —Phrygilanthus lanceolatum (Ruiz & Pav.) Eichler in Mart., Fl. Bras. 5(2): 48. 1868. —Gaiadendron lanceolatum (Ruiz & Pav.) Baehni, Field Mus. Nat. Hist., Bot. Ser. 13: 395. 1937. Loranthus puracensis H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 437. 1818 [1820]. —Gaiadendron punctatum var. puracense (H.B.K.) Steyerm., Fieldiana, Bot. 28: 221. 1951. —Gaiadendron puracensis (H.B.K.) G. Don, Gen. Hist. 3: 432. 1834. —Taguaria puracense (H.B.K.) Raf., Sylva Tellur. 125. 1838. Loranthus tagua H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 436. 1818 [1820]. —Gaiadendron tagua (H.B.K.) G. Don, Gen. Hist. 3: 431. 1834. Gaiadendron poasense Donn. Sm., Bot. Gaz. (Crawfordsville) 56: 61. 1913.

Gaiadendron revolutum Rizzini, Revista Fac. Agron. (Maracay) 8(3): 85. 1975. —Gaiadendron tagua var. revolutum (Rizzini) Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 93. 1982. Gaiadendron tagua var. reductum Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 91. 1982. Shrub or tree to 5 m tall which may grow epiphytically or on the ground, always parasitic on other plants; leaves dark, shiny green, often somewhat revolute at margins; racemes in leaf axils and/or at tips of branches. Upper montane zone, forest edges, 1300–2800 m; widespread in Bolívar and Amazonas. Aragua, Distrito Federal, Lara, Mérida, Miranda, Táchira, Trujillo, Zulia; Nicaragua, Costa Rica, Colombia, Guyana (Mount Roraima), Ecuador, Peru, Bolivia. ◆Fig. 32. Gaiadendron punctatum is a variable species in size of leaves, flowers, bracts, and bracteoles. The flower color is usually bright yellow, but pale yellow or even pure white occurs in some areas (outside the flora area). Numerous species and infraspecific categories have been recognized in the past, but the genus is here regarded as monotypic, and the status of lower categories as uncertain.

3. ORYCTANTHUS Eichler in Mart., Fl. Bras. 5(2): 87. 1868. Foliaceous parasitic shrubs, often with furfuraceous surface on young twigs but otherwise glabrous; epicortical roots from base of plant only. Leaves paired; venation palmate to pinnate, with stellate fiber bundles. Inflorescences solitary in leaf axils or clustered at older nodes, in some in terminal, squamate compound inflorescence, the unit being an indeterminate spike. Flowers sessile, decussate, one per axil of the scale leaves, each with 2 minute, awl-shaped bracteoles, bisexual, (4)6merous. Anthers nearly sessile, basifixed, dimorphic; pollen with 3 conspicuous, circular depressions on both faces. Seed viscid; endosperm whitish; embryo green, dicotylous. Widespread from southern Mexico through Central America, Jamaica, tropical South America to Amazonian Bolivia; ca. 12 species, 6 in Venezuela, 4 of these in the flora area. The mostly Mesoamerican Oryctanthus cordifolius (C. Presl) Urb. might occur in Venezuela, having been reported in the Rupununi River basin, Guyana. It is distinguished from other Venezuelan species by its large, cordate to clasping leaves and keeled stem.

Oryctanthus 41

The genus Oryctina has elsewhere been included in Oryctanthus but is here accorded separate and expanded status (see below). Key to the Species of Oryctanthus 1. 1. 2(1).

2.

3(2). 3.

Stems angled or keeled at least when young; leaves broadly lanceolate ................................................................................................ O. florulentus Stems terete; leaves mostly ovate, obovate or cordate, rarely orbicular ................................................................................................................ 2 Leaves < 3 cm long, obovate to orbicular, spike to 1 cm long; connectival horn absent even from shorter stamens; spikes axillary only ........................................................................................... O. phthirusoides Leaves generally > 4 cm long, cordate or ovate to lanceolate; spikes > 1 cm long, axillary and/or in terminal, compound raceme; connectival horn present at least on shorter stamens ...................................................... 3 Fruits, flowers, and buds inclined forward on spike; terminal, compound inflorescence often formed ...................................................... O. alveolatus Fruits, flowers, and buds perpendicular to the inflorescence axis; inflorescences axillary only, simple ................................................. O. occidentalis

Oryctanthus alveolatus (H.B.K.) Kuijt, Bot. Jahrb. Syst. 95: 504. 1976. —Loranthus alveolatus H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 444. 1818 [1820]. Loranthus amplexicaulis H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 445. 1818 [1820]. —Oryctanthus amplexicaulis (H.B.K.) Eichler in Mart., Fl. Bras. 5(2): 88. 1868. —Oryctanthus alveolatus var. amplexicaulis (H.B.K.) Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 136. 1982. Oryctanthus botryostachys Eichler in Mart., Fl. Bras. 5(2): 89. 1868. Oryctanthus laceratus Kuijt, Bot. Jahrb. Syst. 95: 519. 1976. Oryctanthus alveolatus var. kuijtii Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 136. 1982. Leaves extremely variable, mostly ovate; spikes often in compound, terminal inflorescences, mostly pedunculate; fruits pointing obliquely forward. On a great variety of trees, both native and cultivated, near sea level to 1200 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Lara, Mérida, Miranda, Sucre, Zulia; Mexico?, Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil, Amazonian Bolivia. The great variability of this species has resulted in many additional infraspecific

names, including varieties kuijtii and amplexicaulis. An adequate case can not be made at this time for any subdivisions, and therefore anything more than the species is not recognized. Oryctanthus florulentus (Rich.) Tiegh., Bull. Mus. Nat. Hist. Nat. Paris 2: 339. 1896. —Loranthus florulentus Rich., Act. Soc. Hist. Nat. Paris 1: 107. 1792. Loranthus ruficaulis Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 61, t. 185. 1838. —Oryctanthus ruficaulis (Poepp. & Endl.) Eichler in Mart., Fl. Bras. 5(2): 89. 1868. Stems quadrangular-compressed, furfurascent-lined when young, spikes sessile or nearly so; fruit rather blunt-tipped. Parasitic on trees in forests, forest edges, savannas, city streets, near sea level to 900 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, Bolivia. ◆Fig. 33. Oryctanthus occidentalis (L.) Eichler in Mart., Fl. Bras. 5(2): 89. 1868. —Loranthus occidentalis L., Amoen. Acad. 5: 396. 1760. Stems glabrous; compound inflorescences absent; spike pedunculate, the fruit perpendicular to its axis. On native and cultivated woody plants, 100–200 m; Amazonas (near

42

L ORANTHACEAE

Laja Catipán, Río Pasimoni, Río Yatúa). Costa Rica, Panama, Jamaica, Colombia, Guyana, Suriname, Ecuador, Amazonian Peru, Brazil (Amazonas). Oryctanthus phthirusoides Rizzini, Rodriguésia 41: 27. 1976. Phthirusa maculata Rizzini, Rodriguésia 41: 12. 1976. A very distinctive, small-leaved species. Riparian forests, 100–600 m; Bolívar (La Paragua). Brazil (Amazonas). The species produces new shoots from epicortical roots, an extremely rare phenomenon in Loranthaceae of the Americas.

Fig. 33. Oryctanthus florulentus

4. ORYCTINA Tiegh., Bull. Soc. Bot. France 42: 168. 1895. Maracanthus Kuijt, Brittonia 28: 231. 1976. Rather small mistletoes, glabrous or somewhat furfuraceous; leaves decussate. Dioecious or flowers bisexual (in the flora area). Inflorescence a determinate, pedunculate or sessile spike of bracteolate monads, bracts persistent or deciduous, bracteoles in our species rather large and ± keeled, distinct from subtending bracts. Flowers not sunken in inflorescence axis, 6-merous, petals and stamens somewhat dimorphic. Anthers sessile or on short filaments, pollen sacs 2(4). Style straight and rather stout. Fruit a small berry. Embryo dicotylous. Venezuela, Guyana, Suriname, French Guiana, Brazil; 6 species, 1 in Venezuela.

Phthirusa 43

Oryctina myrsinites (Eichler) Kuijt, Syst. Bot. 16: 290. 1991. —Phthirusa myrsinites Eichler in Mart., Fl. Bras. 5(2): 66. 1868. —Passovia myrsinites (Eichler) Tiegh., Bull. Soc. Bot. France 42: 172. 1895, “Passowia.” Phthirusa savannarum Maguire, Bull. Torrey Bot. Club 75: 301. 1948. —Phthirusa myrsinites var. savannarum (Maguire) Rizzini, Mem. New York Bot. Gard. 29: 25. 1978. A small species characterized by coriaceous leaves and extremely short inflorescences. Forests, near sea level to 500 m; Amazonas (Río Casiquiare, Río Pasimoni, San Carlos de Río Negro). Guyana, Suriname, French Guiana, Brazil (south to Planalto). ◆Fig. 34.

Fig. 34. Oryctina myrsinites

5. PHTHIRUSA Mart., Flora 13: 110. 1830. Passovia H. Karst., Bot. Zeitung (Berlin) 4: 107. 1846. Furarium Rizzini, Rodriguésia 28–29: 154. 1956. Rather small parasitic shrubs, often with furfuraceous young branches but otherwise glabrous, epicortical roots at the base and sometimes also from the branches. Leaves decussate, petiolate. Inflorescence basically an indeterminate spike or raceme of ± paired triads, these units aggregated in squamate, terminal, compound inflorescences in a few species; triads bracteolate, the bracts and bracteoles distinct or showing various degrees of fusion. Flowers bisexual (in a few, unisexual by reduction, the plants then dioecious), mostly 6-merous. Stamens dimorphic; anthers basifixed, either sessile or filaments very short and stout, often conspicuously excavated laterally. Seed viscid; endosperm whitish; embryo green, dicotylous. Mexico (Chiapas), Central America, Jamaica, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Amazonian Bolivia; ca. 40 species, ca. 16 in Venezuela, 8 of these in the flora area. A low-elevation genus of uncertain size, often confused with some species of the related genera Struthanthus, Cladocolea, and Oryctina. Key to the Species of Phthirusa 1. 1. 2(1). 2. 3(2).

Flowers unisexual, plants dioecious .......................................................... 2 Flowers bisexual ......................................................................................... 5 Inflorescence peduncle strongly quadrangular, even winged .... P. podoptera Inflorescence axis not strongly quadrangular, never winged ................... 3 Leaves rather narrowly lanceolate, the lateral veins not evident; inflorescences probably axillary only, always unbranched ............... P. stenophylla

44

3.

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

L ORANTHACEAE

Leaves ovate, lateral veins evident; inflorescences lateral and simple or also terminal, the latter branched (sometimes then the axillary ones also) ........................................................................................................ 4 Leaves narrowly ovate, with strongly attentuate apex; inflorescence probably only axillary and simple ......................................................... P. nitens Leaves broadly ovate, apex not usually attenuate; inflorescence branched when terminal and sometimes also when axillary ......................... P. stelis Triads short-pedunculate, bracts attached to triad peduncle; leaves thin, apex mostly acute ...................................................................... P. pyrifolia Triads completely sessile, often deciduous with bracts, flowers, or fruits; leaves somewhat leathery, apex usually obtuse ................................... 6 Flowers 6-merous; inflorescences to 15 cm long ......................... P. bisexualis Flowers 4-merous; inflorescences rarely longer than 5 cm ...................... 7 Triad bract and bracteoles fused, falling together; leaves mostly thin, narrowly obovate, often emarginate; fruit apex truncate .......... P. guyanensis Triad bracts and bracteoles distinct; leaves coriaceous, ovate, apex rounded; fruit apex rounded ................................................. P. disjectifolia

Phthirusa bisexualis Rizzini, Ernstia 24: 14. 1984. Leaves coriaceous, lanceolate to ovate; inflorescences extremely long, terminal as well as axillary, bearing sessile triads, the acute bracts and bracteoles falling with the fruit. On trees near sandy savannas, 100–200 m; Amazonas (Canaripó, Caño Perro de Agua west of San Antonio). Brazil (Amazonas). Phthirusa disjectifolia (Rizzini) Kuijt, Proc. Kon. Ned. Akad. Wetensch. 93: 114. 1990. —Furarium disjectifolium Rizzini, Rodriguésia 28–29: 155. 1956. Stout plants with stiff, coriaceous leaves usually becoming gray when dry, the petiole and part of lower midrib furfuraceous; inflorescences stout, often terminal, compound, the crowded triads sessile, with prominent bracts and bracteoles. River shores, savanna margins, 100–500 m; Bolívar (Amaruaytepui), Amazonas (Río Parú, San Carlos de Río Negro, San Fernando de Atabapo). Guyana, Brazil (Amazonas, Pará). ◆Fig. 37. Phthirusa guyanensis Eichler in Mart., Fl. Bras. 5(2): 64. 1868. —Passovia guyanensis (Eichler) Tiegh., Bull. Soc. Bot. France 42: 172. 1895, “Passowia.” Phthirusa perforata Rizzini, Revista Fac. Agron. (Maracay) 8(3): 91. 1975. Phthirusa calloso-albida Rizzini, Ernstia 24: 15. 1984. Phthirusa castillana Rizzini, Revista Brasil. Biol. 47: 455. 1987.

Stems ± terete, gray-furfurascent when young; leaves often thin; inflorescences with sessile triads often irregularly distributed. Rocky exposures, 100–200 m; Amazonas (35 km southeast of Puerto Ayacucho). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 39. Phthirusa nitens (Mart.) Eichler in Mart., Fl. Bras. 5(2): 59. 1868. —Loranthus nitens Mart. in Schult. & Schult. f., Syst. Veg. 7: 150. 1829. —Struthanthus nitens (Mart.) Mart., Flora 13: 105. 1830. ?Struthanthus pseudolepidotus Rizzini, Ernstia 32: 14. 1985. Leaves narrowly ovate, with acuminate apex; racemes with rather few triads, these short-pedunculate; dioecious. Habitat not known, 1200–1600 m; Bolívar (Gran Sabana), Amazonas (below Salto Los Monos on tributary of headwaters of Caño Iguapo). Guyana, Brazil (Amazonas). ◆Fig. 38. Phthirusa podoptera (Cham. & Schltdl.) Kuijt, Taxon 43: 198. 1944. —Loranthus podopterus Cham. & Schltdl., Linnaea 3: 218. 1828. Loranthus pterygopus Mart. in Schult. & Schult. f., Syst. Veg. 7: 163. 1829. —Struthanthus pterygopus (Mart.) Mart., Flora 13: 105. 1830. Leaves broadly ovate, tending to be truncate at base and acuminate at the apex; spikes simple or branched, triads sessile; upper anthers 2-locular, lower ones 4-locular; plant dioecious. Habitat not known, 100–300 m; southern Amazonas. Brazil.

Phthirusa

Phthirusa pyrifolia (H.B.K.) Eichler in Mart., Fl. Bras. 5(2): 63. 1868. —Loranthus pyrifolius H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 441. 1818 [1820], non Willd. ex Schult. f. 1830. —Passovia pyrifolia (H.B.K.) Tiegh., Bull. Soc. Bot. France 42: 172. 1895, “Passowia.” Epicortical roots from stem base only; stems compressed-angular, with brownfurfuraceous lines when young; leaves lanceolate, apex often attenuate; inflorescences with numerous, paired, ± sessile triads. In a great variety of habitats, including on many cultivated dicotyledonous trees, near sea level to 1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Common throughout southern Mexico, Central America, Jamaica, tropical South America. Phthirusa stelis (L.) Kuijt, Taxon 43: 193. 1994. —Loranthus stelis L. Sp. Pl. ed. 2, 231. 1762, non G. Forst. 1786. Loranthus retroflexus Ruiz & Pav., Fl. Peruv. 3: 49. 1802. —Phthirusa retroflexa (Ruiz & Pav.) Kuijt, Brittonia 32: 521. 1980. Loranthus aduncus G. Mey., Prim. Fl. Esseq. 149. 1818. —Phthirusa adunca (G. Mey.) Maguire, Bull. Torrey Bot. Club 75: 301. 1948. Loranthus conduplicatus H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 441. 1818 [1820]. Loranthus paniculatus H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 442. 1818 [1820]. —Phthirusa paniculata (H.B.K.) Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 17. 1931. Loranthus orinocensis Spreng. in Roem. & Schult., Syst. Veg. 2: 129. 1825. —Phthirusa orinocensis (Spreng.) Eichler in Mart., Fl. Bras. 5(2): 60. 1868. —Passovia orinocensis (Spreng.) Tiegh., Bull. Soc. Bot. France 42: 172. 1895, “Passowia.” —Phthirusa adunca var. orinocensis (Spreng.) Steyerm., Fieldiana, Bot. 28: 224. 1951. Loranthus magdalenae Cham. & Schltdl., Linnaea 3: 219. 1828. —Phthirusa adunca var. magdalenae (Cham. & Schltdl.) Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 81. 1982 (including formas magdalenae and magnifolia). Passovia suaveolens Karst., Bot. Zeitung (Berlin) 4: 107. 1846.

45

Phthirusa maritima Rizzini, Revista Fac. Agron. Maracay 8: 92. 1975. Phthirusa adunca var. rigidifolia Rizzini in Steyerm. & Brewer-Carías, Bol. Soc. Venez. Ci. Nat. 32: 326. 1976. Phthirusa anastyla Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 69. 1982. Phthirusa pyramidalis Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 54. 1982. Phthirusa cothurnata Rizzini, Ernstia 24: 16-17. 1984. Phthirusa perdivergens Rizzini, Ernstia 24: 19. 1984. ?Phthirusa rubromicans Rizzini, Ernstia 32: 10. 1985. Inflorescences often branching, especially at the tips of shoots; epicortical roots from stems and base; plant dioecious. In a great variety of humid habitats, including many cultivated dicotyledonous trees and shrubs, near sea level to 2200 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Throughout tropical America from Costa Rica to Amazonian Bolivia. ◆Fig. 36. The taxonomy and nomenclature of this species are extremely complex, and the group of species to which it belongs is in urgent need of detailed study. That group is characterized by dioecy, branched (compound) inflorescences which are placed mostly in a terminal position, and the production of epicortical roots from leafy shoots. The tentative view taken here is that nearly all this complex is best treated as a single species. While this approach complicates (or, at least, concentrates) the synonymy even further it would seem to be a reasonable one until a comprehensive study of the group can be made. Phthirusa stenophylla Eichler in Mart., Fl. Bras. 5(2): 60. 1868. Phrygilanthus megathermicus Rizzini, Rodriguésia 41: 13. 1976. —Phthirusa megathermica (Rizzini) Rizzini, Ernstia 24: 17. 1984. Phthirusa huberi Rizzini, Ernstia 24: 17. 1984. Phthirusa percrassa Rizzini, Ernstia 24: 18. 1984. Leaves lanceolate, rather coriaceous, only the midvein evident; inflorescence with few triads, these and lateral flowers stalked. Savanna areas, 100–200 m; Amazonas (Departamento Atabapo). Brazil (Amazonas). ◆Fig. 35.

46

L ORANTHACEAE

Fig. 35. Phthirusa stenophylla

Fig. 36. Phthirusa stelis

Fig. 37. Phthirusa disjectifolia

Phthirusa 47

Fig. 38. Phthirusa nitens

Fig. 39. Phthirusa guyanensis

48

L ORANTHACEAE

6. PSITTACANTHUS Mart., Flora 13: 106. 1830. Apodina Tiegh., Bull. Soc. Bot. France 42: 353. 1895. Psathyranthus Ule, Verh. Bot. Vereins. Prov. Brandenburg 48: 156. 1906 [1907]. Large parasitic shrubs, glabrous, branching percurrent or rarely dichotomous. Stems terete or quadrangular. Epicortical roots lacking (present in P. julianus), the primary attachment often becoming very large. Leaves paired or whorled, mostly coriaceous and petiolate. Inflorescence basically an umbel or raceme of triads or, by reduction, dyads, bracts and bracteoles fused with triad peduncles and pedicels, respectively, and often forming a cupule where investing the base of a flower. Flowers large, 6-merous, bisexual, mostly pedicellate and brightly colored with red and/or yellow. Petals long and slender, the basal (or greater) part often fused into a tube. Stamens epipetalous, mostly dimorphic and filaments long and slender, dorsifixed to the elongated anthers which are mostly at 2 different heights. Ovary with well-defined calyculus; style as long as the petals or nearly so; stigma capitate, placed beyond the anthers. Fruit a large berry. Seed surrounded by gummy viscin; endosperm absent; embryo massive, bright green; cotyledons 2 and flat, or 3 or more and prismatic. Mexico, Central America, Jamaica, Lesser Antilles, tropical South America south to Bolivia and northern Argentina; ca. 80 species, ca. 18 in Venezuela, 14 of these in the flora area. A large-flowered species of Olacaceae, Chaunochiton loranthoides Benth., might be mistaken for Psittacanthus or other large-flowered Loranthaceae even though it is a terrestrial shrub. It has ebracteolate racemes which tend to cluster at the end of shoots, and exceedingly small anthers at the tips of very long filaments, these anthers opening like miniscule flowers. Chaunochiton loranthoides is known from central and southwestern Amazonas state. Key to the Species of Psittacanthus 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(4). 5. 6(5).

6.

Leaf apex attenuate-acuminate to nearly cuspidate ................................ 2 Leaf rounded at the tip or somewhat tapering in upper half, but not acuminate or cuspidate .......................................................................... 3 Tips of petals free even in the closed bud, spreading like small horns; flowers triadic, the mature buds < 4 cm long ................... P. peronopetalus Tips of petals not free in bud, not horn-like; flowers dyadic, the mature buds ≥ 5 cm long ............................................................. P. montis-neblinae Inflorescence a raceme, variously placed, of 4 or more dyads or triads ... 4 Inflorescence an umbel, variously placed, of dyads or triads ................... 8 All flowers sessile, each invested by a cupule completely hiding the ovary; primary bract foliaceous .......................................................... P. cucullaris Flowers pedicellate, part of the ovary always visible above cupule; primary bract not foliaceous ...................................................................... 5 Flower buds straight, 8–9.5 cm long ......................................... P. clusiifolius Flower buds straight or with curved tip, 5 cm long or less ...................... 6 Young stems quadrangular; flower bud apex acute, beak-like, the base of the bud not dilated, the cupule not investing basal portion of ovary ............................................................................................ P. rhynchanthus Young stems terete; flower bud apex rounded, the base dilated or not; cupule covering at least half of the ovary ................................................. 7

Psittacanthus 49

7(6).

Flower buds hairy, curved in upper portion, strongly dilated at base ....................................................................................................... P. cinctus 7. Flower buds glabrous, straight, not dilated at base .......... P. corynocephalus 8(3). Flower buds strongly sigmoid ................................................. P. collum-cygni 8. Flower buds straight or only very slightly curved .................................... 9 9(8). Flowers in dyads ...................................................................................... 10 9. Flowers in triads ...................................................................................... 11 10(9). Leaves 8–13 cm long; petals 5 cm or longer; filaments 1–2 mm long, inserted ca. 5 mm below petal tip ....................................... P. brachynema 10. Leaves 4.5–9 cm long; petals < 4 cm long; filaments 8–10 mm long, inserted 12–15 mm below petal tip ............................................... P. julianus 11(9). Inflorescence with 2 triads ...................................................................... 12 11. Inflorescence with (3)4 triads .................................................................. 13 12(11). Flower buds glabrous ................................................................... P. biternatus 12. Flower buds densely short-hairy ................................................ P. lasianthus 13(11). Flower buds 9–10.5 cm long and 2.5 mm thick, filaments 5–6 cm long; cupule covering < 1/2 of ovary, calyculus ± smooth .................... P. robustus 13. Flower buds 4.5 cm long, to 5 mm thick, filaments to 1.5 cm long; cupule covering 1/2 or more of ovary, calyculus coarsely dentate ........ P. acinarius Psittacanthus acinarius (Mart.) Mart., Flora 13: 108. 1830. —Loranthus acinarius Mart. in Schult. & Schult. f., Syst. Veg. 7: 130. 1829. Psittacanthus warmingii Eichler in Mart., Fl. Bras. 5(2): 36. 1868. Coarse plants with terete stems; leaves to 16 × 6 cm, leathery, but with evident pinnate venation, tending to be falcate; inflorescence an axillary umbel of triads. Forests, near sea level to 400 m; Delta Amacuro (Río Orocoima), Bolívar (Agua Amena along Caicara to Puerto Ayacucho road, upper Río Suapure), Amazonas (Cerro Yapacana, Río Orinoco). Apure; Brazil (Piaui). ◆Fig. 41. Psittacanthus biternatus (Hoffmanns.) Blume in Schult. & Schult. f., Syst. Veg. 7: 1730. 1830. —Loranthus biternatus Hoffmanns. in Schult. & Schult. f., Syst. Veg. 7: 124. 1829. Stems rather stout, terete; leaves to 9 × 6 cm, of which the stout petiole is 1 cm; blade elliptical or obovate to nearly orbicular, extremely leathery and flat, silvery green when fresh; inflorescence axillary, being a pair of triads on inflorescence peduncle of 4–8 mm. Lowland forests, along rivers, 100–200 m; scattered in Amazonas. Brazil. ◆Fig. 42. Psittacanthus brachynema Eichler in Mart., Fl. Bras. 5(2): 29. 1868. Stout plant with massive, terete stems; leaves apparently irregular, whorled in 3s,

paired, or somewhat irregular; leaf blade to 16 × 7 cm, oblanceolate, very leathery, apex rounded, base nearly acute, petiole to 2.5 cm long and 5 mm thick; inflorescence axillary, but apparently also directly from the internode between leaves, being an umbel of 3 dyads. Forests, near sea level to 200 m; Amazonas (Río Yatúa). Brazil (Amazonas). ◆Fig. 44. Psittacanthus cinctus (Mart.) Mart., Fl. Bras. 5(2): 40. 1868. —Loranthus cinctus Mart. in Schult. & Schult. f., Syst. Veg. 7: 134. 1829. Psittacanthus cinctus var. guainiae Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 114. 1982. Inflorescences axillary, being racemes of dyads; buds short-hairy, erect but curved above; style with corrugated thickening 1 cm above base. Forests, 100–200 m; Amazonas (Caño San Miguel, Maroa, San Carlos de Río Negro). Brazil (Amazonas). ◆Fig. 40. Psittacanthus clusiifolius Eichler in Mart., Fl. Bras. 5(2): 30. 1868, “clusiaefolius.” Psittacanthus clusiifolius var. pseudojulianus Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 111. 1982, “clusiaefolius.” Stout plants with terete stems, internodes to 11 cm; leaves decussate, to 15 × 7 cm, grayish green when dry, elliptical to ovate, base obtuse or nearly so; petiole to 15 × 4 mm,

50

L ORANTHACEAE

flat; inflorescence a raceme of 2–4 pairs of dyads. Riparian forests, 100–200 m; Amazonas (30 km east of mouth of Río Guainía). Brazil (Amazonas). ◆Fig. 46. Psittacanthus collum-cygni Eichler in Mart., Fl. Bras. 5(2): 35. 1868. Medium-sized species, internodes terete, to 5 cm long, twigs apparently (at least sometimes) aborting at the tip, or forming an inflorescence there; leaves to 9 × 5 cm, ovate to elliptic, apex rounded, base obtuse; petiole flat, 3–5 mm, grading into base; buds sigmoid. Savannas, roadsides, 100–200 m; Bolívar (Santa Bárbara), Amazonas (between Chapazón & Guirape, near Limón de Parhueña, Río Casiquiare). Apure, Guárico; Brazil (Amazonas, Pará). ◆Fig. 48. Psittacanthus corynocephalus Eichler in Mart., Fl. Bras. 5(2): 36. 1868. Very stout plant with thick, coriaceous leaves; inflorescence terminal and made up of dyads and/or triads; buds and petals massive. Forests, 100–200 m; Amazonas (San Carlos de Río Negro). Colombia, Ecuador, Peru, Brazil (Bahia, Piaui). A rather rare species of wide distribution. Psittacanthus cucullaris (Lam.) Blume in Schult. & Schult. f., Syst. Veg. 7: 1730. 1830. —Loranthus cucullaris Lam., J. Hist. Nat. Paris 1: 444. 1792. —Apodina cucullaris (Lam.) Tiegh., Bull. Soc. Bot. France 42: 353. 1895. Loranthus cupulifer H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 438. 1818 [1820]. —Psittacanthus cupulifer (H.B.K.) Klotzsch in M.R. Schomb., Reis. Br.-Guiana 3: 1162. 1848 [1849]. Loranthus falcifrons Mart. in Schult. & Schult. f., Syst. Veg. 7: 129. 1829. —Psittacanthus falcifrons (Mart.) Mart., Flora 13: 107. 1830. Leaves parallel-veined; triads with sessile flowers and (often) large primary bracts, each ovary completely hidden in elongated cupule which appears to be the ovary. Forests, roadsides, low elevations; no records from the flora area, but to be expected. Apure; Costa Rica (Osa Peninsula), Colombia, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, western Brazil, Amazonian Bolivia. ◆Fig. 43.

Psittacanthus julianus Rizzini, Revista Fac. Agron. (Maracay) 8(3): 95. 1975. Stout but rather small plant, sometimes with 1 or 2 stout epicortical roots from the haustorium, internodes terete, 1–2 cm long; leaves paired, obovate, leathery, often grayish green when dry, apex rounded, base cuneate; venation palmate; inflorescence axillary, being an umbel of 2 dyads. Savannas, 100–200 m; southwestern Amazonas. Endemic. ◆Fig. 47. Psittacanthus lasianthus Sandwith, Bull. Misc. Inf. 1939: 18. 1939. Leaves coriaceous, epidermis (including guard cells), heavily cutinized, the epidermal cells each with conical upper surface; inflorescence an umbel of 2 triads; flower buds densely covered with short hairs; lower half of petals with interlocking, marginal lobes. Lowland forests, 100–200 m; Amazonas (Río Emoni in lower Río Siapa basin, Río Pasiba). Guyana. Psittacanthus montis-neblinae Rizzini, Mem. New York Bot. Gard. 29: 29. 1978. Rather small-leaved plant; leaves generally narrowly lanceolate, stems terete; flowers straight, lacking neck, apparently opening only at the tip, dyadic, pendent, orangered with yellow tip. Upper montane gallery forests, 2000–2400 m; Amazonas (Sierra de la Neblina). Adjacent Brazil. Psittacanthus peronopetalus Eichler in Mart., Fl. Bras. 5(2): 31. 1868. Rather large plant, stems terete, internodes to 7 cm long; leaves to 11 × 4 cm, ovate, sometimes attenuate at the tip; inflorescences axillary but often at the very tip of the branch, which seems to abort, with 2 or 3 pairs of triads; petal tips recurved in bud. Common along river banks, 100–200 m; Amazonas (between mouth of Río Pasimoni and its junction with Río Baría and Río Yatúa). French Guiana, Brazil (Amazonas). ◆Fig. 45. Psittacanthus rhynchanthus (Benth.) Kuijt, Ann. Missouri Bot. Gard. 74: 529. 1987 (including subsp. rhynchanthus). —Loranthus rhynchanthus Benth., Bot. Voy. Sulphur 102. 1845. Psittacanthus chrismarii Urb., Bot. Jahrb. Syst. 24: 331. 1897.

Psittacanthus 51

Fig. 40. Psittacanthus cinctus

Fig. 41. Psittacanthus acinarius

Fig. 42. Psittacanthus biternatus

52

L ORANTHACEAE

Fig. 43. Psittacanthus cucullaris

Fig. 44. Psittacanthus brachynema

Psittacanthus 53

Fig. 45. Psittacanthus peronopetalus

Fig. 46. Psittacanthus clusiifolius

54

L ORANTHACEAE

Fig. 47. Psittacanthus julianus

Fig. 48. Psittacanthus collum-cygni

Fig. 49. Psittacanthus robustus

Struthanthus 55

Psittacanthus semiarticulatus Rizzini, Rodriguésia 18/19: 142. 1956. Psittacanthus calyculatus var. wurdackii Rizzini, Rodriguésia 41: 15. 1976. —Psittacanthus rhynchanthus subsp. wurdackii (Rizzini) Kuijt, Ann. Missouri Bot. Gard. 74: 529. 1987. Stems obscurely quadrangular; leaves to 10 × 1.2 cm, with a few (usually 3) striking parallel veins, narrowly lanceolate; inflorescence terminal; bud sharply curved in upper 1/3, with slender, acute tip. Rock outcrops, savanna and river edges, 100–200 m; Bolívar (between Río Horeda near Ciudad Bolivar and Cerro Gavilán, ca. 10 km above mouth of Río Parhueña). Anzoátegui, Apure, Cojedes, Guárico, Monagas, Portuguesa, Zulia; southern Mexico, Central America, north Colombia.

Psittacanthus robustus (Mart.) Mart., Flora 13: 108. 1830. —Loranthus robustus Mart. in Schult. & Schult. f., Syst. Veg. 7: 125. 1828. Loranthus formosus Cham. & Schltdl., Linnaea 3: 212. 1828. Loranthus speciosus Pohl in DC., Prodr. 4: 309. 1830. Psittacanthus intermedius Rizzini, Revista Fac. Agron. (Maracay) 8(3): 94. 1975. Inflorescence seemingly terminal and subterminal, each an umbel of 4 triads; petals and filaments unusually long and slender, calyculus smooth-margined; petals recurving strongly when dry, twisted into a double or triple curl when dry. River forests, forest edges, 100–400 m; Amazonas (25 km north of Samariapo on road to Puerto Ayacucho). Central Brazil. ◆Fig. 49.

7. STRUTHANTHUS Mart., Flora 13: 102. 1830. Scandent, leafy, mostly glabrous parasitic shrubs, stems terete or quadrangular, epicortical roots commonly developing both from the stems and base. Leaves decussate. Inflorescence an indeterminate spike or raceme of pairs of triads, in a few species determinate; solitary or clustered at the older nodes, very rarely branched and terminal; lateral flowers of triads often pedicellate, central one sessile. Flowers unisexual (plants dioecious), 6-merous, rather small, mostly greenish white, with prominent aborted organs of the opposite sex in both male and female flowers. Filaments dorsifixed to versatile; anthers dimorphic. Style straight or (in some species outside the flora area) contorted, nearly the full length of the petals. Seeds with whitish endosperm and copious viscin; embryo green, dicotylous. Northwestern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil, Amazonian Bolivia, Paraguay, northern Argentina, Uruguay; ca. 60 species, ca. 10 in Venezuela, 5 of these in the flora area. Key to the Species of Struthanthus 1.

1.

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

Leaves usually oblanceolate and mucronate, the petiole ca. 1/3 as long as the blade; triads sessile or nearly so, at least 5 pairs per inflorescence ...................................................................................................... S. gracilis Leaves ovate to lanceolate, not mucronate, petiole 1/4 as long as blade or less; triads with peduncles at least 2 mm long, no more than 4 pairs per inflorescence ........................................................................................... 2 Median flower of triads with pedicel ca. 2 mm long, equaling those of lateral flowers ............................................................................ S. vulgaris Median flower of triads sessile .................................................................. 3 Leaf blades broadly ovate, base truncate to very obtuse; lateral flowers of triads pedunculate; fruit to 20 mm long ............................ S. syringifolius Leaf blades lanceolate, base acute; lateral flowers of triads sessile; fruit < 10 mm long ............................................................................................. 4

56

4(3). 4.

L ORANTHACEAE

Inflorescence peduncle no more than 5 mm long ................ S. dichotrianthus Inflorescence peduncle 15–25 mm long ................................... S. eichlerianus

Struthanthus dichotrianthus Eichler in Mart., Fl. Bras. 5(2): 75. 1868. Struthanthus terniflorus Eichler in Mart., Fl. Bras. 5(2): 75. 1868. Struthanthus dichotrianthus var. dichotrianthus Rizzini, Revista Brasil. Biol. 10: 408. 1950. Struthanthus dichotrianthus var. lasserianus Rizzini, Revista Brasil. Biol. 10: 408. 1950. Struthanthus granulatus Rizzini, Revista Fac. Agron. (Maracay) 8(3): 96. 1975. Struthanthus dissimilis Rizzini, Rodriguésia 41: 14. 1976. ?Struthanthus yavitensis Rizzini, Rodriguésia 41: 28. 1976. Struthanthus translucens Rizzini in Luces & Steyerm., Fl. Venez. 4(2): 30. 1982. Rather delicate species, stems terete to slightly angular, nodes callussed in axils around inflorescence bases even when young, obviously so when older; leaves commonly ca. 4 × 1.5 cm, lanceolate, apex and base acute, blade thin; inflorescences often clustered in leaf axils, delicate, mostly with 2(4) triads. Forests, open spaces, 50–1100 m; Delta Amacuro (Río Orocoima on Brazo Imataca of Río Orinoco), Bolívar (Gran Sabana), Amazonas (Yavita). Anzoátegui, Apure, Aragua, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 50. Struthanthus dichotrianthus is extremely variable as to stoutness, leaf shape and size, and number of triads. It apparently intergrades with the more delicate species Struthanthus phyllyraeoides (H.B.K.) Blume, from which it seems difficult to distinguish. The 2 varieties described by Rizzini require more detailed study and are not at present recognized. Struthanthus eichlerianus Rizzini, Rodriguésia 41: 29. 1976. Forests, ca. 100 m; Bolívar (Serranía Baraguán). Aragua. Struthanthus eichlerianus is perhaps no more than a long-pedunculate form of Struthanthus dichotrianthus. If a valid species, it is known only from 2 collections.

Struthanthus gracilis (Gleason) Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 443. 1967. —Phthirusa gracilis Gleason, Bull. Torrey Bot. Club 58: 359, fig. 4, 1931. Struthanthus mucronatus Steyerm., Bol. Soc. Ven. Ci. Nat. 26: 418. 1966. —Struthanthus gracilis var. mucronatus (Steyerm.) Rizzini in Luces & Steyerm., Fl. Ven. 4(2): 17. 1982. Struthanthus chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 443. 1967. Struthanthus cupulifer Rizzini, Bol. Soc. Venez. Ci. Nat. 32: 327. 1976. Sparsely branched, the young twigs much elongated (nodes to 10 cm), often with recurved leaves on prehensile petioles; mature leaves to 6 × 3.5 cm, mostly obovate and the apex mucronate; inflorescence mostly single in leaf axil. Rocky plateaus, forest slopes, 1000–2000 m; Bolívar (Macizo del Chimantá, Ptari-tepui), Amazonas (Cerro Aracamuni). Adjacent Guyana. Struthanthus gracilis is very reminiscent of a small S. orbicularis (H.B.K.) Blume, to which it is undoubtedly closely related. Struthanthus syringifolius (Mart.) Eichler in Mart., Fl. Bras. 5(2): 78. 1868. —Loranthus syringifolius Mart. in Schult. & Schult. f., Syst. Veg. 7: 141. 1829. Phthirusa ptariana Steyerm., Fieldiana, Bot. 28: 224. 1951. Phthirusa hippocrateoides Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 442. 1967. Stout plant; leaves to 10 × 5 cm, often red when fresh and young, broadly ovate, with acute, slightly attenuate apex and truncate to very obtuse base; fruit to 2 cm long. Forests, 800–2200 m; Bolívar (Gran Sabana, Macizo del Chimantá, Ptari-tepui). Aragua, Lara, Mérida, Miranda, Yaracuy, Zulia; Guyana, Suriname, Brazil. ◆Fig. 51. Struthanthus vulgaris Eichler in Mart., Fl. Bras. 5(2): 85. 1868. Rather large plant, stems often strongly but finely lenticellate; leaves ovate, apex

Struthanthus 57

somewhat attenuate; inflorescences several per axil, each with 2 pedunculate triads; all flowers pedicellate, pedicels ca. 2 mm long.

Forests, 500–1900 m; Bolívar (Kamarkawarai-tepui, middle Río Paragua basin). Aragua; Brazil.

Fig. 50. Struthanthus dichotrianthus

Fig. 51. Struthanthus syringifolius

58

L ORANTHACEAE

8. TRIPODANTHUS (Eichler) Tiegh., Bull. Soc. Bot. France 42: 178. 1895. —Phrygilanthus subgen. Tripodanthus Eichler in Mart., Fl. Bras. 5(2): 45. 1868. Large to very large, glabrous parasitic shrubs, producing often profuse masses of epicortical roots from stems, with some roots becoming established on the host’s subterranean organs. Leaves decussate or somewhat alternate. Inflorescence a determinate raceme of paired triads, all flowers pedicellate, bisexual, white, 6-merous. Stamens dimorphic; anthers versatile, dorsifixed, epipetalous. Seeds with whitish endosperm; embryo rather small, greenish; cotyledons 2. South America; 2 species, 1 in Venezuela.

Fig. 52. Tripodanthus acutifolius

L YTHRACEAE 59

Tripodanthus acutifolius (Ruiz & Pav.) Tiegh., Bull. Soc. Bot. France 42: 179. 1895. —Loranthus acutifolius Ruiz & Pav., Fl. Peruv. 3: 48. 1802. —Phrygilanthus acutifolius (Ruiz & Pav.) Eichler in Mart., Fl. Bras. 5(2): 50. 1868. Loranthus eugenioides H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 435. 1818 [1820]. —Tripodanthus eugenioides (H.B.K.) Tiegh., Bull. Soc. Bot. France 42: 179. 1895. Phrygilanthus acutifolius coriaceus Rizzini, Rodriguésia 28–29: 96. 1956. Phrygilanthus acutifolius eugenioides

lateovatus Rizzini, Rodriguésia 28–29: 96. 1956. Phrygilanthus acutifolius eugenioides vulgaris Rizzini, Rodriguésia 28–29: 96. 1956. Plant sometimes very large, often with profuse root development on host stems; leaves rather thin, acuminate; flowers white and sweetly scented, triadic, even the median ones pedicellate. River shores, margins of open spaces in forests, 1200–2200 m; Bolívar (Auyán-tepui, Cerro Jaua, Chimantá-tepui, Roraima-tepui). Ecuador, Peru, Brazil, Bolivia, northern Paraguay, northern Argentina, northern Uruguay. ◆Fig. 52.

LYTHRACEAE by Alicia Lourteig Herbs, shrubs, or trees. Stems prostrate, erect, or woody. Leaves simple, entire, opposite, decussate, or rarely partly alternate or verticillate (in 3s or 4s); stipules absent or represented by several setae. Flowers generally solitary or in cymes, racemes, spikes, or rarely in panicles. Bracts small, often caducous; bractlets small, in pairs. Flowers bisexual, actinomorphic or zygomorphic, 4–6-merous. Calyx persistent, campanulate, turbinate, or tubular, the lobes acute, intersepalic appendages commonly present; petals free, sometimes caducous, reduced, or absent, inserted on the rim of the calyx tube alternate with the lobes. Stamens isomerous, 2–4 (Rotala), 11 (most Cuphea), or (6)8–15, inserted on the calyx at different heights or on a ring; filaments of unequal length; anthers dorsifixed or basifixed (Crenea), longitudinally dehiscent, connective usually conspicuous. Disk glandular, roundish, dorsal, asymmetric (Cuphea) or inconspicuous. Ovary superior, 2-carpellate, usually 1-, 2-, or 4(5)-locular; gynophore present or absent; placenta central, axile; ovules 2–many, anatropous, ascendent; style filiform, included to long-exserted, rarely reduced; stigma 2-lobed, capitate, discoid or inconspicuous, papillose. Fruit a capsule ± included in the calyx, dehiscent or indehiscent. Seeds mostly suborbicular, polyhedric, or flat, sometimes winged, without endosperm; embryo straight. Tropical and temperate regions of both hemispheres; ca. 25 genera and 500 species, 5 genera and 27 species in the flora area. Key to the Genera of Lythraceae 1. 1. 2(1). 2.

Flowers zygomorphic; calyx tubular, 6-merous ............................... 2. Cuphea Flowers actinomorphic; calyx globose or turbinate-campanulate, 4–6-merous ............................................................................................. 2 Herbs; stamens 2–4, included; ovary incompletely 2-locular .......... 5. Rotala Herbs or shrubs to small trees; stamens (4)8–15, ± exserted; ovary completely 2–many-locular .......................................................................... 3

60

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

L YTHRACEAE

Petals violet; leaves and young branches violet-glandular ............. 4. Pehria Petals white; plants eglandular ................................................................. 4 Anthers basifixed; flowers solitary or in 2- or 3-flowered cymes; branches never spinescent ........................................................................... 1. Crenea Anthers dorsifixed; flowers fragrant, in few-flowered corymbs or panicles; branches often spinescent ....................................................... 3. Lawsonia

1. CRENEA Aubl., Hist. Pl. Guiane 523. 1775; Lourteig, Caldasia 15: 121, map. 1986. Dodecas L. f., Suppl. Pl. 36. 1781 [1782]. Glabrous herbs or shrubs to 4 m tall. Roots fibrous, much-branched. Stems often woody, partially rhizomatous, reddish, covered by a gray film, branched, tetragonous, 4-winged. Leaves opposite, decussate, fleshy or not, obovatesubspatulate to elliptic-lanceolate or elliptic-oblong, obtuse, often mucronulate. Inflorescences of solitary flowers or in 2-, 3-, or 5-flowered cymes; bracts 2 mm long, caducous; bractlets generally caducous; peduncle to 15 mm long. Flowers 4-merous; pedicel bracteolate near the apex. Calyx green, violet, or green with red margin, turbinate-campanulate, 4-lobed, intersepalic appendages absent; petals 4, suborbiculate. Stamens 8(?) or 12–15, inserted ± at the middle of the calyx tube, as long as the calyx or slightly exserted; anthers yellow, basifixed, oblong, erect, connective very narrow, mucro red. Ovary sessile, globose, 4(5)-locular; ovules numerous, anatropous; styles 2 or 3 times the length of the ovary; stigmas subcapitate. Capsule globose, membranous, 4(5)-locular, indehiscent, partially covered by the calyx, many-seeded. Seeds in compact masses, subfalcate, trigonous, ± curved, rugose, apiculate, convex face reddish, the 2 concave (interior) faces white; cotyledons narrowly elliptic; radicles cylindric, long. Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Brazil; 2 species, 1 in Venezuela. Crenea patentinervis (Koehne) Standl. grows only on the Pacific coast of Colombia and northern Ecuador. Crenea maritima Aubl., Hist. Pl. Guiane 523, pl. 209. 1775. —Dodecas maritimus (Aubl.) Griseb., Fl. Brit. W. I. 270. 1860. Dodecas surinamensis L. f., Suppl. 245. 1781. —Crenea surinamensis (L. f.) Koehne, Bot. Jahrb. Syst. 3: 320. 1882. Herb or shrub to 1 m tall; roots partially adventitious; internodes commonly shorter than the leaves; leaves 15–90 × 5–25 mm, obovate-subspatulate or lanceolate, fleshy, obtuse, decurrent on the petiole, midvein prominent on the lower surface; secondary veins 4–9 pairs, very fine, sometimes inconspicuous; flowers solitary or in cymes of 2 or

3; bractlets 1–1.75 mm, acuminate at the apex; pedicel 2–3 mm long; calyx 3–6 mm long, membranous, green or violet, 8-veined; calyx lobes triangular, about as long as the tube; petals 5–6 mm long, white; stamens 12–15; anthers 1.75–2.5 mm; styles white; stigmas red; capsule 5–8 mm diameter, green-violet. Mainly in mangroves and estuaries, near sea level to 50 m; Delta Amacuro (Caño Capure, Caño Güiniquina, Caño Mánamo south of Tucupita, Caño Pedernales, Isla Burojoida at mouth of Caño Araguao). Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 53.

2. CUPHEA P. Browne, Civ. Nat. Hist. Jamaica 216. 1756. Herbs or small shrubs, rarely glabrous, usually with pubescence of pluricellular, glandular or malpighiaceous hairs, sometimes cystolithic-pubescent (with crystalline inclusions of the epidermis that have the gross appearance of hairs). Leaves sessile or petiolate, opposite and decussate, or 3(4)-verticillate, rarely

Cuphea 61

Fig. 53. Crenea maritima

partly alternate; stipules represented by several reddish setae at the insertion of the petiole. Inflorescences of solitary, axillary flowers, or extra-axillary with interpetiolar and infrapetiolar pedicels and decussate to the bracts, or in terminal, axillary cymes, or simple or compound racemes. Flowers zygomorphic; pedicels 2bracteolate, the bracteoles generally at a constant height for each species. Calyx tubular, 12-veined, dorsiventrally flattened, obtuse or spurred at the base, often narrowed at the throat, then widened to the mouth, pubescent inside, very rarely glabrous or with 2 rows of hairs on dorsal veins; calyx lobes 6, usually acute; intersepalic appendages 6, narrower than the lobes, linear or subulate; petals 2, 4, or 6, rarely absent, pinkish to violet, very rarely white or yellow, the dorsal pair usually different in size or color from the 4 ventral ones. Stamens 11, rarely 6 or 9, very rarely 4, inserted on the calyx tube, the 2 dorsal ones lower and much shorter than the others, the 5 episepalic ones longest, of which the 3 ventral ones are glabrous or visibly less pilose, the other 4 epipetalic, generally pubescent; anthers dorsifixed, introrse, longitudinally dehiscent. Glandular disk dorsal (its shape is an important sectional or species-specific character). Ovary sessile, asymmetrically ovoid or fusiform, tapering gradually into an included to long-exserted style, 2-carpellate, incom-

62

L YTHRACEAE

pletely 2-locular; placenta thick, central, cylindric, free; ovules few to numerous; stigma inconspicuous. Capsule membranous, ovoid or oblong, asymmetric, enclosed in the calyx, (1–)many-seeded; when ripe, the placenta inverts, the ovary and calyx open longitudinally, and the seeds are violently ejected. Seeds small, suborbiculate to obovate, minutely foveolate, acute, the margin obtuse, acute or narrowly winged; embryo straight; cotyledons reniform; radicle very short. U.S.A., Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia, Paraguay, Argentina, Old World tropics and subtropics; ca. 250 species, 28 in Venezuela, 22 of these in the flora area. Species found in the flora area belong to section Brachyandra (5 species: Cuphaea carthagenensis, C. elliptica, C. micrantha, C. repens, C. tenuissima); section Melvilla (3 species: C. annulata, C. galeato-calcarata, C. melvilla); section Trispermum (3 species: C. anisoclada, C. antisyphilitica, C. gracilis); and section Amazonianae (11 species, mostly endemic to the Venezuelan Guayana region, which appears to be a center of speciation for this well-delimited group: C. bolivariensis, C. cardonae, C. cataractarum, C. curiosa, C. dactylophora, C. distichophylla, C. maigualidensis, C. odonellii, C. pleiantha, C. rhodocalyx, C. rigidula). Key to the Species of Cuphea 1. 1. 2(1).

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

Calyx 12–40 mm long, thick, tubular, greenish or bright-colored ............ 2 Calyx to 12 mm long, slender, narrowed at the throat, then widened to the mouth, green or ± reddish-veined ......................................................... 4 Leaves 1–5 mm wide, linear; calyx greenish, spurred and abruptly enlarged dorsally near the base into a helmet-shaped protrusion ..................................................................................... C. galeato-calcarata Leaves 8–55 mm wide, elliptic or lanceolate; calyx orange-yellow, red, or violet, spurred but without a dorsal protrusion ................................... 3 Calyx bright orange-yellow with a red border; petals reddish purple ................................................................................................... C. annulata Calyx red to violet; petals usually absent ..................................... C. melvilla Creeping herbs; leaves mostly 3–5-verticillate; bractlets ca. 1/3 the length of the calyx .................................................................................... C. repens Erect or prostrate herbs; leaves mostly opposite, decussate; bractlets smaller .................................................................................................... 5 Petals yellow; calyx pubescence dense, hirsute-glandular and malpighiaceous .................................................................................... C. anisoclada Petals whitish or pinkish to violet; calyx pubescence different from above ................................................................................................................ 6 Plants glandular, setose, cystolithic-pubescent (not malpighiaceous) ..... 7 Plants malpighiaceous-pubescent or with other types of hairs intermixed .............................................................................................................. 12 Inflorescences borne on the upper half of the branches, with bracts; cespitose subshrubs .................................................................... C. gracilis Inflorescences borne among the leaves, without bracts; erect herbs or subshrubs ..................................................................................................... 8 Leaves narrowly linear (to 6 cm long), hirsute ......................... C. tenuissima Leaves elliptic to ovate, rarely linear and then not as long as above ...... 9

Cuphea 63

9(8). Glandular disk cylindric, erect; seeds elliptic ............................... C. elliptica 9. Glandular disk ovoid, horizontal or deflexed; seeds suborbiculate ........ 10 10(9). Plants with whitish, appressed-cystolithic pubescence or occasionally with fine setae intermixed ............................................... C. antisyphilitica 10. Plants with setose-glandular pubescence ............................................... 11 11(10). Leaves nearly glabrous; calyx inflated in fruit and almost closed at the mouth .............................................................................. C. carthagenensis 11. Leaves setose and pubescent; calyx enlarged in fruit but open at the mouth ..................................................................................... C. micrantha 12(6). Calyx scarcely pubescent to almost glabrous .......................................... 13 12. Calyx ± densely malpighiaceous-pubescent and with some setae ......... 15 13(12). Corolla 2-colored, the ventral petals whitish or pinkish, the 2 dorsal ones purplish with hairs on the midveins dorsally near the base .................................................................................................. C. pleiantha 13. Corolla pinkish, the petals glabrous ........................................................ 14 14(13). Stem malpighiaceous-pubescent and violet-setose ...................... C. cardonae 14. Stem scattered malpighiaceous-pubescent ............................... C. rhodocalyx 15(12). Inflorescence terminal, long and branched; plant with long (to 2.5 mm) setae ........................................................................................... C. rigidula 15. Inflorescence mostly solitary flowers within the foliage; plant without setae ......................................................................................................... 16 16(15). Leaves rigid, sessile, much-crowded ................................... C. distichophylla 16. Leaves membranous, distant along the branches ................................... 17 17(16). Semiprostrate herb; stems long (to 1 m), much-branched; savannas ................................................................................................... C. odonellii 17. Erect or decumbent herbs or shrubs 0.25–0.6 cm tall ............................ 18 18(17). Corolla whitish; flowers alternate on the upper half of the branches ........................................................................................... C. cataractarum 18. Corolla ± violet or 2-colored; flowers otherwise disposed ....................... 19 19(18). Corolla 2-colored, the 2 dorsal petals violet with the midveins dorsally hairy near the base, the ventral ones pinkish ............................ C. curiosa 19. Corolla pinkish ......................................................................................... 20 20(19). Stems malpighiaceous-pubescent and setose, the setae violet ................... ........................................................................................ C. maigualidensis 20. Stems malpighiaceous-pubescent only .................................................... 21 21(20). Glandular disk ovoid, horizontal ............................................ C. bolivariensis 21. Glandular disk cylindric, erect .............................................. C. dactylophora Cuphea anisoclada Lourteig, Mem. New York Bot. Gard. 9: 358, fig. 69. 1957. Subshrub 50–60 cm tall, pubescent, glabrescent; several stems from the base erect or ± decumbent, branched to the apex, internodes shorter than the subsessile leaves, pubescence malpighiaceous, white, hairs with branches of unequal length, with some simple hairs intermixed; blade coriaceous, 15–23 × 4–7 mm, elliptic or oblong, acute, mucronate, base slightly cordate, margin ±

recurved, seldom ciliate, both surfaces malpighiaceous-pubescent, more so on the veins on the lower surface, glabrescent; pedicel 3–3.5 mm, pubescent and setose; bractlets ovate to suborbiculate, 0.75–1 mm long, the apex pubescent and ciliate; calyx 11–20 mm, not much enlarged at the throat, with a roundish, incurved spur, pubescence dense, hirsute-glandular and malpighiaceous; intersepalic appendages shorter than the calyx lobes; calyx inside with pubes-

64

L YTHRACEAE

cence on the veins, increasing to the base, woolly behind the stamens; petals yellow, oblong-to obovate-subspatulate, 5–6 × 2–2.5 mm, the ventral ones ca. 7 × 3 mm; stamens included or reaching the rim of the calyx; glandular disk thick, deflexed; ovary glabrous; style included, glabrous; ovules 3; seeds ca. 3 mm, suborbiculate or subellipsoid. Headwaters of rivers, waterfalls, ca. 400–1700 m; Bolívar (base of Ilú-tepui, Río Caroní near Urimán, Salto Aponguao, Salto Hacha), Amazonas (Cerro Ualipano). Endemic. ◆Fig. 58. Cuphea annulata Koehne in Mart., Fl. Bras. 13(2): 304, pl. 56, fig. 5. 1877. Scandent shrub to 2 m tall; branches somewhat compressed and trigonous at apex, pubescent, glabrescent, the internodes shorter than the leaves, to 22 mm long; leaves sessile; blade subcoriaceous, bright green, 20–55 × 8–20 mm, elliptic or lanceolate, base cuneate, obtuse or subcordate, the upper surface glabrous or some hairs on the midvein, the lower surface with some setae, margin setose-glandular, 10–12 pairs secondary veins; flowers solitary among the leaves; pedicel 4–5 mm long; bractlets ovate, ca. 1 mm long, setaceous-acuminate in the lower half or at the middle; calyx to 25 mm long, tubular, bright orange-yellow; calyx lobes short, acute, red; intersepalic appendages thick, wide, the apex setiform with fine hairs and sparse glandular hairs; spur obtuse or incurved; calyx inside finely pubescent on the veins, woolly behind the stamens, with a transversal fringe of short, scarious hairs near the bottom; petals 6, small, purplish red, 1–1.5 mm long; stamens purplish red, exserted; glandular disk, thick, deflected, subcordate, obtuse; ovary glabrous; ovules 5; style long-exserted; seeds 2 or 3, dark brown, suborbicular-flattened, ca. 3 mm long, slightly marginate, the border somewhat concave at the place of the caruncle, apex retuse. Sandstone outcrops, gallery forests, shrublands, 100–1600 m; Bolívar (Cerro Marutaní), Amazonas (widespread). Amazonian Colombia, Brazil, Bolivia. ◆Fig. 55. Cuphea antisyphilitica H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 202. 1823 [1824]. Cuphea antisyphilitica f. subhirsuta

Koehne, Bot. Jahrb. Syst. 29: 158. 1900. Cuphea gracilis auct. non H.B.K. 1824, nec Seem. 1854. Herb to 70 cm tall, ± decumbent; pubescence whitish, appressed-cystolithic and hirsute, often with setose-glandular hairs intermixed; stems slender, branched; leaves decussate, subsessile, 3–25 × 2–9 mm, oblong to elliptic or lanceolate, base subcordate or acute; flowers solitary on the upper part of the branches; pedicel 0.5–2 mm; bractlets in the distal 1/2 of the pedicel, 0.5–1 mm long, ovate, acute, hirsute; calyx purplish, 4–4.5 mm long, pubescent inside; calyx lobes wide, acute; intersepalic appendages shorter than the lobes; petals 1.5–2 mm long, violet (sometimes faded); stamens included or reaching the rim of the calyx; glandular disk thick, deflexed; ovary glabrous or pilose on the apex; style included; ovules 3 (very rarely more); seeds brownish, ca. 2 mm, suborbiculate-flattened, thick. Apure, Táchira; Colombia, Venezuela, Guyana, Brazil; 2 varieties, both in the flora area. Key to the Varieties of C. antisyphilitica 1. Leaves ovate to elliptic-linear, base cordate to roundish; plant with appressed cystolithic-pubescence, the hairs uniform and rigid, and very rarely also with some setae .................... var. acutifolia 1. Leaves elliptic, rarely linear, base obtuse, rarely subcuneate; plant with fine sparse setae and uniform cystolithicpubescence .......... var. antisyphilitica C. antisyphilitica var. acutifolia Benth., J. Bot. (Hooker) 2: 118. 1840. Cuphea antisyphilitica f. gracillima Koehne in Mart., Fl. Bras. 13(2): 285, pl. 51, fig. 4 (sub antisyphilitica). 1877. Cuphea gracilis var. major Koehne in Mart., Fl. Bras. 13(2): 284, pl. 51, fig. 3 (sub gracilis). 1877. Mauritia palm swamps, streamsides, white-sand savanna-shrub mosaics, bauxitic-lateritic outcrops, 50–800 m; Bolívar (between Ciudad Piar and base of Cerro Bolívar, 45 km east of Los Pijiguaos, Represa Guri), Amazonas (15 km northwest of La Esmeralda, near Puerto Ayacucho, Río Manapiare, Río Parucito). Eastern Colombia, common in Brazil. ◆Fig. 56.

Cuphea 65

C. antisyphilitica var. antisyphilitica Wet areas, savannas, Mauritia palm swamps, 100–1400 m; Bolívar (widespread), Amazonas (Caño Asisa, Río Manapiare). Apure; widespread in eastern Colombia, Guyana, Suriname, French Guiana, Amazonian Bolivia. ◆Fig. 57. Cuphea bolivariensis Lourteig, Phytologia 60: 30. 1986. Subshrub to 30 cm tall; stem woody at base, decumbent, 0.5 cm diameter, densely appressed-malpighiaceous-pubescent, the hairs fine, with arms long and unequal; leaves densely appressed-pubescent, 6.5–12 × 1–2 mm, linear, obtuse, 1-veined, the vein red, visible on lower surface; flowers decussate; pedicel thick, elongated in fruit to 2.5 mm long; bractlets subulate, 0.2–0.3 mm long, glabrous; calyx ca. 4 mm long, malpighiaceous-pubescent, enlarged to the throat (± inflated in fruit); calyx lobes wider than the intersepalic appendages; calyx inside pubescent on the veins, woolly behind the stamens (hairs white and purplish); petals oblong, ca. 2.5 × 1.5 mm; stamens inserted near the middle of the calyx tube; glandular disk thick, ovoid, obtuse, subhorizontal-descendent; ovary slightly hirsute-pubescent; ovules 3; style pilose at the base, exserted after anthesis; seed brownish, ca. 1 mm long when immature. Streamsides, wet savannas, ca. 500 m; Bolívar (Urimán). Endemic. Cuphea cardonae Lourteig, Phytologia 60: 42, fig. 4A’. 1986. Cuphea rhodocalyx var. setosa Lourteig, Sellowia 16: 136, fig. 2 1964. Subshrub to 60 cm tall, profusely branched; branches woody, setose, the setae violet, thickened at base, appressed-malpighiaceous-pubescent, the hairs white, fine, with branches of unequal lengths; young shoots, tetragonous, subwinged; foliage dense; internodes 2–5 mm long; petiole ca. 2 mm long, dorsally pilose; blade 4–12 × 1–2 mm, oblong-elliptic, obtuse, with 2 or 3 pairs of alternate, ascendent secondary veins, the lower surface with sparse malpighiaceous hairs, the upper surface glabrous, marginate, the base cuneate, decurrent; inflorescence short, with bracts; flowers toward the apex, alternate; pedicel pilose; bractlets

suborbiculate, obtuse or acute ca. 0.25 mm in the upper half; calyx straight, slightly widened toward the mouth, glabrous or with a few appressed, fine, malpighiaceous hairs; calyx lobes acute, the 2 dorsal ones wider; intersepalic appendages thick, greenish-purplish, somewhat shorter than the lobes; calyx inside pilose, woolly behind the stamens; glandular disk violet, terete, erect; ovary dorsally hirsute-pilose; ovules 5; style with a few hairs, exserted after anthesis; seeds brownish, suborbiculate, flattened, ca. 1 mm long. Rocky soil along rivers, ca. 400 m; Bolívar (Río Pauo tributary of upper Río Caura). Endemic. Cuphea carthagenensis (Jacq.) J.F. Macbr., Publ. Field Columbian Mus., Bot. Ser. 8: 124. 1930. —Lythrum carthagenense Jacq., Select. Stirp. Amer. Hist. 148. 1763. Cuphea balsamona Cham. & Schltdl., Linnaea 2: 363. 1827. Herb or subshrub, erect to 60 cm tall, glandular-pubescent and setose; stem reddish, sometimes woody at base, to 1 cm diameter; leaves elliptic to lanceolate, acute, ± pubescent, the veins on the lower surface glandular and setose; petioles 1–5 mm, slightly pubescent, with a few setae; pedicel interpetiolate, 1–3 mm long; bractlets ovate, acuminate 0.5–1 mm at the apex; calyx 4.5–6 mm, slightly pubescent, with a few setae, pilose inside behind the stamens and on the dorsal veins; petals 6, pinkish to purplish, obovate or oblong, subspatulate, ca. 1.5 mm; stamens inserted at the middle of the calyx tube, pilose; glandular disk small, lobed; ovules 4–8; fruiting calyx inflated, almost closed at mouth; seeds 3–5, reddish brown, obovate, ca. 1.5 mm long, narrowly winged. Moist areas, Mauritia palm swamps, 100– 400 m; Bolívar (Altiplanicie de Nuria, east of Miamo, Upata). Southeastern U.S.A., Central America, Antilles, Colombia, Guyana, Ecuador, Peru, Brazil, Paraguay, Argentina, New Caledonia, Taiwan. Cuphea cataractarum Spruce ex Koehne in Mart., Fl. Bras. 13(2): 226 (in key). 1877. Subshrub to 60 cm tall, woody at base; stem profusely branched from the base; branches covered by the foliage; internodes

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10–12 mm, shorter than the overlapping sessile leaves; pubescence malpighiaceous, finely appressed, also with fine caducous setae; blade 6–22 × 2–7 mm, elliptic to linear, very finely malpighiaceous-pubescent on both surfaces, 1-veined, the larger ones with an extra pair of basal veins and 1 or 2 pairs of secondary veins, base obtuse, apex acute or obtuse; flowers alternate in the upper half of the branches; pedicel 1.5–3 mm; bractlets ovate to elliptic, pubescent, acute at the apex; calyx 5.5–7 mm, pubescent and setose on the veins in the lower part; calyx lobes wide; intersepalic appendages short, pubescent; calyx inside conspicuously pubescent, woolly behind the stamens; petals whitish, spatulate, narrow, ca. 3 × 0.75–1.25 mm; stamens included; glandular disk suboblong to obovoid, subscandent; ovary glabrous or with some hairs at the apex; ovules 6 or 7; style glabrous or with a few hairs; seeds brownish, suborbicular or subovoid, flattened, ca. 1.25 mm long. Rocks in forests, edges of waterfalls, river banks, 100–400 m; Amazonas (near Culebra, Río Cunucunuma, Salto Yureba). Endemic. ◆Fig. 59. Cuphea curiosa Lourteig, Phytologia 60: 38, fig. 3. 1986. Subshrub to 35 cm tall, very finely appressed-malpighiaceous-pubescent, the hair branches different lengths, and setose; rhizome to 15 cm long, branched, with abundant fibrous roots; stem woody at base, to 0.5 cm diameter, profusely branched and densely foliated, malpighiaceous-pubescent and setose; internodes ca. 7 mm; leaves crowded, more so at the apex; blade 8–30 × 1–5 mm, lanceolate to elliptic, decurrent on petiole, acute, finely malpighiaceous-pubescent, the hairs longitudinally disposed (rarely glabrous above), very rarely some setae; 3-veined, generally 2 pairs (or 1) of secondary veins; margin somewhat recurved; inflorescences to 7 cm in the superior branches, with 1-veined, 3–5 × 1.5–2 mm bracts; flowers alternate; pedicel 1–1.5 mm long; bractlets 0.2–0.5 m, thick, elliptic, glabrous at the apex; calyx violet, 3–3.5 mm long, thin, enlarged at the throat, spurred, veins prominent, uniformly malpighiaceouspubescent, very rarely with some short setae; dorsal calyx lobes wider; intersepalic appendages subulate, pubescent; calyx inside

pilose on the veins, woolly behind the stamens; petals pinkish violet (the 2 dorsal ones darker), oblong-subspatulate, 1–1.75 mm long, the 2 dorsal ones obovate-subspatulate, ca. 2 mm, midvein dorsally white-hairy near the base; stamens included; glandular disk reddish-purplish, thick, terete, erect, ca. 0.5 mm long; ovary ± pilose dorsally; ovules 5 or 6; style pilose, included; seeds brownish, suborbiculate, ca. 1 mm long, plano-convex. Venezuela; 2 varieties, both in the flora area. Endemic. Key to the Varieties of C. curiosa 1. Calyx uniformly malpighiaceous-pubescent; in lowlands ............. var. curiosa 1. Calyx uniformly malpighiaceous-pubescent and also with long (0.5–1 mm) setae along the veins; in mountains ................ .......................................... var. oresbia C. curiosa var. curiosa Along rapids, in gallery forests and savannas on rocky outcrops, 100–400 m; Bolívar (Río Canaracuni, Río Caura, Río Guaña, Río Nichare, Río Parguaza), Amazonas (Río Siapa, Río Yureba). Endemic. ◆Fig. 63. C. curiosa var. oresbia Lourteig, Bradea 8: 28. 1996. Streamsides, on wet rocks and soil, 2000– 2100 m; Bolívar (Sierra de Maigualida), Amazonas (Sierra de Maigualida). Endemic. ◆Fig. 61. Cuphea dactylophora Koehne in Mart., Fl. Bras. 13(2): 266. (in key) 1877. Subshrub, densely malpighiaceouspubescent, the hair branches unequal lengths; rhizomes to 20 × 0.5 cm diameter, fibrous, much-branched; stem procumbent, woody at base, sometimes tortuous, profusely branched, densely malpighiaceous-appressed-pubescent; internodes 5–30 mm, very rarely longer than leaves; petiole 0.5–4 mm, thick; blade linear to lanceolate, rarely elliptic 10–45 × 1–8 mm, base acute, decurrent on petiole, apex acute, rarely obtuse; midvein reddish, noticeable on both surfaces, secondary veins 2–4 pairs, ascendent, visible on the lower surface; flowers alternate, on the summit of the branches (exceptionally

Cuphea 67

with bracts); pedicel 1–4 mm; bractlets suborbicular or oblong, 0.2–1 mm, glabrous or pubescent near the apex; calyx 5–7 mm, narrowed at the throat and then widened to the mouth; calyx lobes triangular, acute; intersepalic appendages smaller than the lobes, densely malpighiaceous-pubescent; calyx inside pilose, woolly behind the stamens; petals pinkish violet, orbicular- or obovatespatulate; glandular disk terete, erect; stamens sometimes somewhat unequal, all pilose; ovary ca. 2 mm, pilose or only so at the apex; ovules 3; style included, pilose; seeds browish, suborbiculate, flattened, ca. 1 mm, slightly marginate. Frequent in highland meadows, streambanks, rocky areas, also montane forests, ca. (200)1000–1500 m; Bolívar (widespread in Gran Sabana, Sierra de Lema), Amazonas (Caño Yutajé). Guyana, Brazil. Cuphea distichophylla Lourteig, Mem. New York Bot. Gard. 9: 357, fig. 68. 1957. Subshrub to 30 cm tall; rhizome woody; stem erect or decumbent, 4 mm diameter at base, branched, malpighiaceous-pubescent and with fine simple hairs and long purplish setae; leaves rigid, very crowded, decussate, subsessile; petiole short, thick, purplish, pubescent; blade discolorous, 10–18 × 1.5–3 mm, lanceolate to linear, acute, midvein wide, conspicuous on the lower surface, both surfaces malpighiaceous-pubescent; pedicel ca. 2 mm, purplish; bractlets at the apex of the peduncle, ovate, malpighiasceous and glandular-pubescent; calyx 4.5–5 mm, purplish, whitish-malpighiaceous-pubescent and with purplish setae; calyx lobes acute; intersepalic appendages smaller than the lobes, generally setose; calyx inside pubescent on the veins and woolly behind the stamens; petals purplish, obovate to oblong, spatulate, 3.5–5 × 2–3 mm, the 2 dorsal ones dorsally hairy on the midvein near the base; stamens included, the episepalic ones nearly reaching the rim of the calyx; glandular disk very thick, subhorizontal or subascendent, ca. 0.6 mm, slightly lobed; ovary small, ca. 1 mm long, pubescent; ovules 2 or 3; style thick, included, 1–1.25 mm long, pubescent; seeds brownish, suborbiculate, 1.5–1.6 mm long, slightly marginate. Mountain slopes, 1400–1900 m; Amazonas (Cerro Sipapo). Endemic.

Cuphea elliptica Koehne, Bot. Jahrb. Syst. 2: 145. 1881. Herb or subshrub to 45 cm tall, base sometimes woody; stem with long (to 2 mm) setae, the young branches with fine yellowish, glandular hairs; leaves subsessile or petiolate, to 3 mm long; blade 8–40 × 3–10 mm, elliptic or ovate-acuminate, midvein noticeable, the base cuneate or decurrent on petiole, the upper surface with sparse, glandular setae, the lower surface with sparse, glandular setae on the veins and cystolithic hairs, the margin ciliate; cymes contracted, axillary; pedicel 1.5–3 mm long, the upper part with subtriangular setaceous bractlets 0.2 mm long; calyx purplish, 5–7 mm long, somewhat narrowed, barely widened to the mouth; calyx lobes wide, acuminate; intersepalic appendages much shorter with long setae; spur incurved with few hairs at base; calyx inside glabrous, woolly behind the stamens; petals purplish ca. 1.5 mm long; stamens inserted shortly above the middle of the tube length; glandular disk erect, oblong, slightly keeled below; ovary glabrous; ovules 4–6; style short; seeds 2 mm long, ellipsoidflattened, narrowly winged. Humid soils, flooded savannas, near sea level to 100 m; Delta Amacuro (Capure), Bolívar (Ciudad Bolívar, Upata to Guasipati), Amazonas (Puerto Ayacucho, Samariapo). Anzoátegui, Apure, Guárico; Panama, Colombia, Brazil. Cuphea galeato-calcarata Lourteig, Bradea 8: 65. 1996. Subshrub to 0.5 m, branched, pubescence of fine, straight, short hairs, with much-appressed, very short, rarely 2-armed, cystolithic hairs and erect glandular setae intermixed; stem subligneous, bark reddish brown and peeling; foliage densely imbricate at the top; leaves subsessile, 5–31 x 1–5 mm, sublinear, acute, base subauriculate, midvein impressed on the upper surface, prominent on the lower, covered by cystolithic hairs; upper surface glabrous or cystolithicpubescent, more so on the lower surface; margin with appressed cystolithic hairs and some caducous setae; inflorescences at the top of branches sometimes hidden, 1-flowered; peduncles 6–8 mm, 2-bracteolate; bractlets 0.5–0.7 mm, suborbiculate, pilose and ciliate; calyx greenish, 14–18 mm, tubu-

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lar, straight or slightly convex dorsally, slightly broadened to the mouth, uniformly cystolithic-pubescent, and with very short fine hairs and glandular setae; calyx base spurred, abruptly enlarged dorsally into a helmet-shaped protrusion near the base; calyx lobes 0.5–0.7 mm, wide-triangular, shortly ciliate or glabrous; intersepalic appendages fleshy, less than half as long as the calyx lobes, glabrous or with 1 or 2 setae; calyx inside woolly-pubescent behind the stamens, shortly and finely villous in the tube, densely so in the basal zone where mixed with hairs or wide, ± flat cavities; petals 6, greenish yellow, triangular-suborbiculate or ovate-oblong, equaling the calyx lobes or somewhat larger, 1-veined, acute, submucronate, caducous; stamens 11, inserted on the distal 1/4 of the calyx tube, the 2 dorsal ones shorter and almost glabrous, the others with woolly filaments with glabrous apex and nearly reaching the rim of the calyx; disc ovoid, thick, deflexed, dorsally concave; ovary semiovoid, glabrous, 3-ovulate; style long, glabrous, subexserted after anthesis; stigma capitate; seeds not seen. Granitic outcrops on margins of river banks, 1500–1700 m; Amazonas (near Río Asita, Sierra Uasadi). Endemic. Cuphea gracilis H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 199. 1823 [1824]. Cuphea gracilis var. media Koehne in Mart., Fl. Bras. 13(2): 284. 1877. Cuphea multicaulis Koehne in Mart., Fl. Bras. 13(2): 228 (in key). 1877. Cuphea pauciflora Koehne in Engl., Pflanzenr. IV. 216(Heft 17): 132. 1903. Cespitose herb to 30 cm tall; rhizome ca. 1 cm diameter, woody; stem decumbent, branched from base, pubescence whitish-appressed with some glandular hairs intermixed, becoming glabrescent; internodes commonly shorter than the leaves; leaves 7– 26 × 2–8 mm, ovate-lanceolate to linear-lanceolate, marginate, midvein noticeable, uniformly cystolithic-pubescent on both surfaces, with some glandular hairs, hirsute on the veins, glabrescent; flowers solitary, on the upper half of the stems, with small bracts; pedicel to 2 mm long, near the apex; bractlets elliptic, 0.5–1 mm, acute, ciliate; calyx purplish, 5.5–6 mm long, thin, narrowed at the throat, and widened to the

mouth; calyx lobes wide, acute; intersepalic appendages small; calyx inside with fine pubescence, some glandular hairs, and long hairs on the veins, woolly behind the stamens; petals purplish, ca. 2.5 mm long; stamens reaching the rim of the calyx; glandular disk thick, subglobose, deflexed, concave below; ovary glabrous or pubescent at apex; ovules 3; style thin, glabrous or pubescent, included; seeds brownish, suborbiculate, flattened, thick, 2 mm diameter. Savanna borders, river edges, 100–200 m; Amazonas (near Puerto Ayacucho, Raudal de Atures, Raudal de Maipures, Río Casiquiare, Río Samariapo). Trujillo; Colombia, Guyana. ◆Fig. 66. Cuphea maigualidensis Lourteig, Bradea 8: 28. 1996. Subshrub to > 30 cm tall; stem branched from base; branches 2- or 3-forked, ascendent, dark, subligneous; pubescence of white-malpighiaceous hairs, the branches mostly unequal lengths, and violet setae thickened at base; internodes 6–20 mm long; petiole thick, pubescent 1–2 mm long; blade 3.5–20 × 1.5–5 mm, elliptic, diminishing to the apex, obtuse or subacute; base obtuse or subcuneate, the lower surface with prominent midvein with 2 pairs of ascendent secondary veins, sparsely pubescent with some uniform setae, glabrescent the upper surface appressed-malpighiaceous pubescent; pedicel ca. 1 mm long, straight; bractlets elliptic, ca. 0.75 mm long, pubescent near the apex; calyx violet, 5.5–6 mm long, tube straight, much-enlarged at the throat, densely appressed-malpighiaceous-pubescent and with setae on the veins its entire length; calyx lobes acute, acuminate, the dorsal one larger, the margin very shortly ciliate inside with some cilia at the apex; intersepalic appendages equaling half the length of the calyx lobes or shorter, thick, subacute, setose; calyx inside long-pilose on the veins, woolly behind the stamens; petals violet, 3.5–4.75 × 2.5–3.5 mm, obovate-oblong or suborbiculate, cuneate; stamens inserted on the distal 1/3 of the calyx tube, included, all pilose; seeds brownish-reddish, suborbiculate, flattened, ca. 1.5 mm long. Streamsides, ca. 2000 m; Bolívar (Sierra de Maigualida), Amazonas (Sierra de Maigualida). Endemic. ◆Fig. 60.

Cuphea 69

Cuphea melvilla Lindl., Edwards’s Bot. Reg. 10: pl. 852. 1824. —Amor seco, Coral. Melvilla speciosa A. Anderson, Trans. Soc. London Encour. Arts 25: 207. 1807, nom. nud. —Cuphea speciosa (A. Anderson) Kuntze, Revis. Gen. Pl. 3(2): 96. 1898. Shrub to 2 m tall, with appressedmalpighiaceous-pubescence and simple and glandular hairs; petioles 3–4(–10) mm long; blades 20–140 × 10–55 mm, elliptic, lanceolate, or rarely ovate, acute, cuneate or obtuse at base, malpighiaceous-pubescent on both sides, hirsute on veins; racemes terminal, ca. 15 cm long, lax-flowered; flowers decussate; pedicel 7–11 mm long; bractlets ovate, ca. 1.5 mm from the base, pubescent; calyx red or violet, green at mouth, straight or somewhat curved, malpighiaceous-pubescent, enlarged toward the base in fruit; spur narrow, incurved; calyx lobes acute, ciliate, sometimes glandular; intersepalic appendages short, ovoid, pilose and glandular, somewhat longer than calyx lobes; calyx inside finely pubescent; stamens inserted on the distal 1/3 of the calyx tube, ± exserted; petals usually absent; glandular disk deflexed, subcordate, thick; ovary pubescent, multiovulate; styles very long, pubescent, exserted; seeds purplish, subcordate, 1.5–2 mm long, concave on both faces, marginate. Moist and shady places in forests, near sea level to 300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Monagas; widely distributed in South America south to Argentina. ◆Fig. 54. Cuphea melvilla is cultivated as an ornamental plant. Cuphea micrantha H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 196. 1823 [1824]. Erect herb to 40 cm tall; stem slender, pubescent and with hirsute-glandular yellowish setae to 2 mm long; internodes commonly longer than the leaves, 1.5–4 mm long; leaves subsessile; blade 15–55 × 5–12 mm, ovate or oblong-lanceolate, acuminate, subcordate, with sparse, long, spreading or ± appressed setae on both surfaces, margin ciliate; cymes terminal, foliate or shortened, axillary; pedicel 1.5–3 mm long; bractlets 0.5–0.3 mm long, ovate, setaceous-ciliate near the apex; calyx purplish, somewhat narrowed at the throat and widened to the

mouth; calyx lobes wide; intersepalic appendages subulate, setaceous; spur small, obtuse, hirsute-glandular on the veins, elsewhere with scanty, fine and short pubescence; calyx inside with hairs on the veins and woolly behind the stamens; petals pinkish violet, ca. 2 mm long; glandular disk, thick, subhorizontal; stamens inserted on the distal 1/3 of the tube length, included; ovules 3; style short; seeds 1.5–2 mm long, suborbiculate, flattened, thick, narrowly marginate. Disturbed savannas, 50–300 m; Bolívar (Caicara, Ciudad Bolívar, Represa Guri). Frequent in northern Venezuela; Central America, Antilles, Guyana, Brazil, Bolivia, Paraguay. Cuphea odonellii Lourteig, Notas Mus. La Plata, Bot. 94: 281, 2 figs. 1959. Herb, malpighiaceous-pubescent, the hair branches unequal lengths, subappressed, white; stem prostrate or decumbent to 1 m long, profusely branched; leaves subsessile or petiolate to 0.5 mm long; blade 8–30 × 1.5–6 mm, linear-lanceolate, acute, upper surface sometimes glabrous, midvein reddish, prominent on the lower surface, secondary veins to 6 pairs fusing to form a marginal vein; flowers alternate, rarely decussate; pedicel ca. 1 mm long; bractlets near the apex of the pedicel, ovate, 0.5–0.7 mm long, acute, pubescent, exceptionally large and leaf-like; calyx 5–7 mm long, malpighiaceous-pubescent, inflated in fruit, widened to the mouth; calyx inside pubescent on the veins, woolly behind stamens; spur obtuse; petals pink to violet, oblong to subspatulate, 1–2.5 mm long, obtuse; stamens included; glandular disk thick, deflexed; ovary ca. 2 mm, glabrous; ovules 3–6; style ca. 2 mm, glabrous; stigma subcapitate; seeds usually 2, brown, 2–2.3 mm, obovoid, flattened. River banks, savannas, sandy humid soils at edges of gallery forests, near sea level to 500 m; Bolívar (Maripa, Río Villacoa, Serranía de los Pijiguaos), Amazonas (Caño Coro Coro, south of Cerro Moriche, base of Cerro Yapacana, Puerto Ayacucho to Samariapo, Raudal de Atures). Guárico, Sucre; Peru, Brazil, Bolivia. ◆Fig. 65. Cuphea pleiantha Lourteig, Mem. New York Bot. Gard. 9: 356, fig. 67. 1957. Subshrub to 50 cm tall, scanty malpigh-

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iaceous-pubescent, the hair branches unequal lengths, appressed, white; rhizome subligneous, ca. 1 cm diameter; stem subligneous to 1 cm diameter, successively branched; branches ascendent, reddish, malpighiaceous-pubescent; internodes 3–6 mm long, shorter than the leaves; leaves sessile, very crowded; blade lanceolate to linear, 12–40 × 1–6 mm, subacute, glabrous on upper surface, slightly malpighiaceous-pubescent on lower surface; midvein red, visible on both surfaces; margin recurved; racemes ca. 7 cm long with alternate flowers; bracts 2–5 × 2 mm, ovate-lanceolate or oblong; pedicel 1–3 mm long, thick, glabrous or nearly so; bractlets near the apex, thick, reddish, oblong or ovate, 0.5–0.6 mm long, acute, glabrous; calyx 5.5–6 mm long, purplish, sparsely malpighiaceous, the hairs very fine, widened to the throat; calyx lobes acute; intersepalic appendages smaller than the lobes; calyx inside pubescent on the veins, woolly behind the stamens; ventral petals whitish or slightly pinkish, obovate- to oblong-spatulate, the 2 dorsal ones dorsally purplish with white hairs on the lower part of the midvein; stamens included, the longer ones almost reaching the rim of the calyx; glandular disk large, to 1.25 mm high , erect, terete; ovary finely pubescent; ovules 7–9; style pubescent; seeds (immature) suborbiculate, flattened, slightly marginate. Among wet boulders, near waterfalls, (100–)1300– 1500 m; Amazonas (Cerro Yutajé, Yutajé). Endemic. ◆Fig. 64.

Cuphea rhodocalyx Lourteig, Mem. New York Bot. Gard. 9: 355, fig. 66. 1957. Herb to 60 cm tall, appressedmalpighiaceous-pubescent to glabrous; rhizome short, branched, with fibrous roots; stem procumbent, woody at base, 5 mm diameter, or erect and branched from base, pubescence of few appressed, malpighiaceous, whitish hairs or glabrous; internodes shorter than the leaves, 2–3 mm long; leaves subsessile, decurrent on stem, giving a 4winged aspect; blade 5–10 × 1–3 mm, linearlanceolate, acute, midvein visible on both surfaces, margin recurved, sparse malpighiaceous hairs on lower surface or glabrous; flowers alternate in the upper part of the branches among the foliage, or in racemes, with bracts; pedicel ca. 0.5 mm long; bractlets near the apex, large thick, oblong to ovate, acute, ca. 0.5 mm long; calyx pinkishviolet, 2.5–4 mm long, widened to the mouth, inflated in fruit; calyx lobes small; intersepalic appendages thick, obtuse, shorter than the lobes, glabrous or with very few malpighiaceous hairs; calyx inside densely pilose, more so behind the stamens; petals violet, obovate- to oblong-spatulate; stamens included; glandular disk, 0.6 mm, thick, subulate, obtuse, erect; ovary 1.5 mm long, pubescent at the apex; ovules 4 or 5; style glabrous, 0.5 mm long; seeds brownish, ± 1.5 mm, subovoid-flattened, base acute, slightly marginate. River banks, gravel bars, forest slopes, 1000–1600 m; Amazonas (Cerro Marahuaka). Endemic.

Cuphea repens Koehne in Mart., Fl. Bras. 13(2): 251, pl. 43, fig. 4. 1877. Small creeping herb, densely malpighiaceous-pubescent; stem filiform, long, branched; leaves 3–5-verticillate, alternate at branch tips, sessile, 2–15 × 0.4–1 mm, linear, acute, margin recurved; flowers extraaxillary; bractlets at the apex of the pedicel, 1.5–2 mm long, linear; calyx 3–4 mm long, narrow, woolly inside behind the stamens, pilose on the veins; petals reddish to violet, ovate to elliptic; stamens inserted at the midpoint of the calyx tube; filaments pilose, included; ovary glabrous or pilose; ovules 9; seeds minute, subglobose. Savannas, lowland marshes; not yet found in the flora area. Apure; Amazon basin (Colombia, Brazil, Bolivia).

Cuphea rigidula Benth., J. Bot. (Hooker) 2: 316. 1840. Herb or subshrub to 30 cm tall; pubescence of fine, appressed, malpighiaceous hairs of equal or unequal branch lengths, and also with abundant, white or violet, very long (to 2.5 mm long) setae; petiole 1–1.5 mm long; blades 15–60 × 8–12 mm, elliptic to lanceolate, decurrent on petiole, acute, the upper surface with scanty malpighiaceous pubescence and abundant setae, the lower surface with malpighiaceous pubescence and setae on the veins; cymes of racemes of alternate flowers; bracts 2–5 × 1–2 mm, oblong to elliptic, acute, little-pubescent to glabrous and setose-ciliate; pedicel 1–1.5 mm long; bractlets in the upper half, oblong-elliptic, acute, 1–1.5 mm long, glabrous; calyx 4–6

Cuphea 71

Fig. 54. Cuphea melvilla

Fig. 55. Cuphea annulata

72

L YTHRACEAE

Fig. 56. Cuphea antisyphilitica var. acutifolia

Fig. 57. Cuphea antisyphilitica var. antisyphilitica

Cuphea 73

Fig. 58. Cuphea anisoclada

Fig. 59. Cuphea cataractarum

74

L YTHRACEAE

5 mm 5 mm

5 mm

1 mm

1 mm

5 mm

5 cm

Fig. 60. Cuphea maigualidensis

Fig. 61. Cuphea curiosa var. oresbia

Cuphea 75

Fig. 62. Cuphea rigidula

Fig. 63. Cuphea curiosa var. curiosa

Fig. 64. Cuphea pleiantha

76

L YTHRACEAE

Fig. 65. Cuphea odonelli

Fig. 66. Cuphea gracilis

Pehria 77

mm long, narrowed at the throat, then widened to the mouth; calyx lobes wide, acute; intersepalic appendages subulate, smaller than the lobes; calyx inside finely pubescent on all veins or only on the dorsal ones, woolly behind the stamens; petals pinkish, obovatesubspatulate, the 2 dorsal ones violet(?); stamens inserted in the distal 1/3 of the calyx tube; glandular disk violet, oblong, horizontal-subascendent; ovary included, dorsally pilose; ovules 9 or 10; style densely pilose; seed brownish, suborbiculate, 1–1.25 mm long, flattened, slightly marginate. Among moist rocks, ca. 700 m; Bolívar (upper Río Caura at Salto Maraveni). Guyana. ◆Fig. 62. Cuphea tenuissima Koehne in Mart., Fl. Bras. 13(2): 254, pl. 43, fig. 7. 1877. Erect herb 20–35 cm tall; stem branched

in the upper half, finely pubescent and loosely hirsute; internodes long; blades about the same length as the internodes or shorter, 20–60 mm, very narrow, linear, almost subulate, finely pubescent and loosely hirsute, margin incurved; inflorescences fewflowered; bracts unequal, 4 x 1 mm; pedicel 1–3 mm long; bractlets near the apex, elliptic, ca. 0.4 mm; calyx 5–5.4 mm long, finely hirsute; calyx inside finely pubescent in the lower part, woolly in the upper part; petals subequal, elliptic-linear, acute; stamens inserted at half length of the calyx tube, included, very short; ovary included; style twice as long as the ovary; glandular disk ovate-cordate, deflected; ovules 3; seeds obovate-roundish, ca. 1.5 mm long. Gravel depressions in savannas, 50–100 m; Bolívar (14 km southeast of Caicara). Brazil.

3. LAWSONIA L., Sp. Pl. 349. 1753; Gen. Pl. ed. 5, 191. 1763. Shrub or small tree to 7 m tall. Branches often spinescent, glabrous, tetragonous when young, the internodes shorter than the leaves. Leaves decussate, 17–55 × 7–20 mm, lanceolate, elliptic or oblong, decurrent on petiole, entire or slightly crenate, pinnately veined, often mucronate; stipules subulate. Flowers fragrant, borne in few-flowered axillary cymes, or corymbs sometimes grouped into terminal panicles to 40 cm long; pedicel bibracteolate; bractlets early caducous. Calyx turbinate, ca. 4 mm long, 4-veined, narrowly winged, wings decurrent on pedicel, lobes 4, ovate; petals 4, white to yellowish, wrinkled, larger than the calyx lobes, 4.5–5.5 mm long, reniform, clawed, entire or finely crenate. Stamens 8, seldom 4, or very rarely 9–12, episepalous, inserted in pairs on a ring slightly below the petals, ca. 5 mm long, exserted; anthers oblong. Ovary subglobose, sessile, 2–4-locular; ovules numerous; style thick, slightly longer than the stamens, 4 mm long; stigma capitate. Capsule subglobose, brown, 4.5–7 mm diameter, indehiscent. Seeds subtetrahedric, brownish, 2–2.5 mm long, spongy-reticulate. Probably native to the Old World, now cultivated and escaped in all tropical regions; 1 species. Lawsonia inermis L., Sp. Pl. 349. 1753. —Resedá. Lawsonia speciosa L., Sp. Pl. 349. 1753. Lawsonia alba Lam., Encycl. 3: 106. 1789. Seasonally flooded riparian areas, near sea level to 100 m; Delta Amacuro (Tucupita), Bolívar (Ciudad Bolívar, Tem-

blador). Apure; other distribution as in genus. The powdered leaves of Lawsonia inermis are the source of the dye henna, the flowers are used to scent oils, and the bark is sometimes used medicinally. The plant is often cultivated for its ornamental value.

4. PEHRIA Sprague, J. Bot. 61: 232. 1923. Grislea Loefl., Iter. Hispan. 245. 1758, non L. 1753. Shrub to 5 m tall; pubescence short, ± glabrescent, and with uniformly scattered, violet, glandular hairs intermixed. Stem tetragonous, the internodes 2.5–5 cm long, much shorter than the leaves. Leaves opposite, decussate, membranous, 4–12

78

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× 0.8–4 cm, elliptic to lanceolate, pinnately 10–12-veined, secondary veins impressed on the upper surface, prominent on the lower side. Inflorescences 3–14-flowered axillary cymes; bracts and bractlets small; peduncle tetragonous. Flowers 4(5)merous. Calyx purple, urceolate-tubular, 8–10-veined, 6–10 mm long, shortly pubescent, lobes short, intersepalic appendages very short; petals violet, spatulate, 2–5

Fig. 67. Pehria compacta

Rotala 79

mm long, glandular. Stamens 8(–10), red, inserted in the lower part of the calyx tube; filaments thin, 2 or more times as long as the calyx, the episepalic ones somewhat shorter; anthers suborbiculate, dorsifixed. Ovary globose, on a short gynophore, 2-locular, 2-valved; ovules numerous. Capsule globose, 2-valved, 4–5 mm long, enclosed within the calyx, dehiscent loculicidally, the valves bifid at the apex. Seeds numerous, brownish, pyramidal, ca. 1 mm long, thin, finely foveolate. Honduras, Nicaragua, Colombia, Venezuela; 1 species. Pehria is closely related to Adenaria and is often misidentified as that. Pehria compacta (Rusby) Sprague, J. Bot. 61: 232. 1923. —Grislea compacta Rusby, Descr. S. Amer. Pl. 68. 1920. —Bucarito, Coralito, Indiecito. Grislea secunda Loefl., Iter Hispan. 245.

1758, non L. 1753. Semideciduous forests, 100–200 m; Bolívar (El Manteco, Temblador). Common throughout much of northern Venezuela; Honduras, Nicaragua, Colombia. ◆Fig. 67.

5. ROTALA L., Mant. Pl. 143, 175. 1771. Creeping herbs, or taller and erect. Leaves opposite, decussate, or verticillate, entire, sessile to subsessile. Flowers small, solitary, axillary, subsessile, bibracteolate, usually 4-merous. Calyx globose, campanulate, or urceolate, usually lobed, sometimes with intersepalic appendages. Petals fugacious or absent. Stamens 2–4, episepalic; anthers small, roundish. Ovary ± ovoid, incompletely 2–4-locular; ovules ca. 13 or more; style small or inconspicuous; stigma capitate. Capsule cartilaginous, 2–4-valved, valves finely and densely transversally striate, septicidally dehiscent. Seeds small, ovate to orbicular. Tropical and temperate regions of both hemispheres; 44 species, 2 in Venezuela, both in the flora area. Key to the Species of Rotala 1. 1.

Stamens 2; leaves 3-verticillate, at least in part ........................ R. mexicana Stamens 4; leaves decussate ......................................................... R. ramosior

Rotala mexicana Cham. & Schltdl., Linnaea 5: 567. 1830. Rotala apetala F. Muell., Fragm. 3: 105. 1862. —Rotala mexicana var. apetala (F. Muell.) Lourteig, Bol. Soc. Argent. Bot. 5: 145. 1954. Hypobrichia spruceana Benth. in Griseb., Cat. Pl. Cub. 106. 1866. —Rotala mexicana var. spruceana (Benth.) Koehne in Mart., Fl. Bras. 13(2): 195, t. 39, fig. 2b. 1877. Hypobrichia spruceana var. tenuifolia Griseb. Cat. Pl. Cub. 106. 1866. Small creeping herb, sometimes submerged, or the top emersed; leaves usually verticillate, rarely partially decussate, 5–15 × 1–1.5 mm, lanceolate to linear, obtuse, rarely retuse; calyx (3)4- or 5-veined, ca. 1

mm long; calyx lobes about 1/3 of the calyx length, subtriangular, acute; intersepalic appendages absent; petals absent; stamens inserted at different heights on the calyx tube, included; ovary subglobose, 2- or 3-locular; ovules ca. 13; capsule globose, 2- or 3-valved, enclosed within the calyx ± all its length; seeds suborbicular. Savannas, 100–200 m; northwestern Bolívar (Caicara to Puerto Ayacucho road, Serranía Carichana), Amazonas (La Esmeralda, Raudal de Atures, near Samariapo, San Juan de Manapiare). Guárico, Monagas; pantropics. Rotala ramosior (L.) Koehne in Mart., Fl. Bras. 13(2): 194, t. 39, fig. 1. 1877. —Ammannia ramosior L., Sp. Pl. 120. 1753.

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Erect or procumbent herb ca. 30 cm tall; leaves to 24 × 3–7 mm, elliptic, linear to oblanceolate, obtuse; calyx 4-merous, ca. 1 mm long; calyx lobes acute; intersepalic appendages about the same length as the lobes; petals 4, pinkish or whitish, early caducous; stamens 4, inserted at about the middle of the calyx tube, included; ovary globose-ovoid; ovules numerous; style very short; stigma capitate; capsule ovoid or globose, to 3 mm long, enclosed in the calyx, 3- or 4- valved; seeds numerous, flattened-obovoid. Roadside ditches, wet areas, near sea level to 50 m; Delta Amacuro (Caño Acomito near San José de Macareito, Tucupita to La Horqueta), Bolívar (Caicara to Río Cuchivero, Moitaco). Apure, Monagas; temperate U.S.A., pantropics, naturalized in the Philippines and the Mediterranean. ◆Fig. 68.

Fig. 68. Rotala ramosior

MAGNOLIACEAE by Paul E. Berry and James S. Miller Evergreen or deciduous trees or shrubs. Leaves alternate, simple, entire or rarely lobed (Liriodendron), the terminal bud enclosed by a large, sheathing, deciduous stipule that leaves annular scars on the branches; petioles present, sometimes with an adaxial scar. Flowers usually solitary, terminal, pseudolateral, or axillary, large, bisexual or rarely unisexual (Kmeria), hypogynous, actinomorphic. Perianth seldom clearly differentiated into sepals and petals, spirally arranged in several to numerous whorls, imbricate. Stamens numerous, spirally arranged; filaments free, usually flattened; anthers elongate, the thecae often running nearly the entire length of the filaments, dehiscing longitudinally. Carpels usually numerous, spirally arranged, usually free, but often aggregated into a cone-like structure at fruiting, conduplicate and sometimes not fully sealed, usually with a style and decurrent stigma; ovules 2 or more on a marginal placenta, anatropous. Fruits apocarpic or syncarpic, dehiscing by a ventral suture (apocarpic follicles) or circumscissilely (syncarpic), or indehiscent and samaroid (Liriodendron). Seeds 1–few per carpel, arillate, suspended by a thread-like funiculus, with abundant, oily endosperm and a small embryo. Tropical, subtropical, and warm-temperate regions of both hemispheres; 8–13 genera and 200–250 species, 1 genus and 2 species in the flora area. There are differing opinions concerning generic circumscriptions in the family, especially in the Magnolia alliance. This treatment follows Lozano-Contreras (Dugandiodendron y Talauma en el Neotrópico. Acad. Ci. Ex. Fis. Nat. Col. Jorge Alvarez Lleras 3: 1–147. 1994) and recognizes both Talauma and Dugandiodendron,

Dugandiodendron 81

whereas Nooteboom (Blumea 31: 83. 1985) included these genera in a more broadly circumscribed Magnolia. 1. DUGANDIODENDRON Lozano, Caldasia 11: 33. 1975. Evergreen trees and shrubs. Leaves alternate, simple, entire, petiolate; stipules tightly enclosing the buds, but not adnate to the petioles. Flowers solitary, terminal (pseudolateral), bisexual, large. Perianth somewhat fleshy, poorly differentiated into sepals and petals; the outermost sepaloid segments 2–4, reflexed; petaloid segments 5–10, with a narrower and thicker base. Stamens linear, numerous (58–182), in 4–6 spiral series; filaments very short, anthers introrse, with a long, filiform apical connective. Gynoecium of 4–40 spirally arranged, united carpels; ovules 2 per carpel. Fruit a circumscissilely dehiscing syncarp, the carpidia 3–9-ridged, abaxially dehiscing and the external part falling off either singly or in a mass, leaving a cone-like receptacle with the seeds hanging from it. Seeds oblong-reniform, with a red or orange aril. Colombia, Venezuela, Ecuador, probably Guyana; 14 species, 2 in Venezuela, both in the flora area. Fig. 69. Dugandiodendron chimantense

Fig. 70. Dugandiodendron ptaritepuianum

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Dugandiodendron and Talauma can be distinguished from Magnolia by the very differently dehiscing fruiting structure. Talauma is differentiated from Dugandiodendron mainly by its petioles having noticeable adaxial scars where the stipules were adnate in the bud stage. Key to the Species of Dugandiodendron 1. 1.

Leaf blades elliptic, sericeous along the midrib; leaves < 7 cm wide, apex acute .................................................................................... D. chimantense Leaf blades orbicular or nearly so, > 9 cm wide, the apex rounded ........................................................................................ D. ptaritepuianum

Dugandiodendron chimantense (Steyerm. & Maguire) Lozano, Caldasia 12: 9. 1977. —Magnolia chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 443, fig. 2. 1967. Tree to 15 m tall; internodes covered with woolly indument; leaves 11 × 6 cm, woolly on lower surface, 13–15 secondary veins on either side of the midvein; flowers white, fragrant, with 3 sepals and 7 or 8 petals; carpels 5 or 6; fruit ellipsoid. Moist tepui-slope forests, 1800–2300 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 69. Dugandiodendron ptaritepuianum (Steyerm.) Lozano, Caldasia 11: 42.

1975. —Magnolia ptaritepuiana Steyerm., Fieldiana, Bot. 28: 233. 1951. —Aura-tá-na-ca-tá (Arekuna), Cabeza de mono. Magnolia roraimae Steyerm., Fieldiana, Bot. 28: 234. 1951. Tree 4–13 m tall, with pale gray bark; leaves white-woolly on lower surface, ± revolute-margined, 15–17 secondary veins on either side of the midvein; flowers white, fragrant, arched-ascending; sepals 3, petals 6; carpels 13–20. Moist tepui-slope forests on both sandstone and granitic substrates, 2100–2300 m; Bolívar (Ptari-tepui, Roraimatepui, Sierra de Maigualida). Endemic. ◆Fig. 70.

MALPIGHIACEAE by William R. Anderson Trees, shrubs, and vines, always perennial; hairs unicellular, usually medifixed or submedifixed. Leaves usually opposite, often bearing large multicellular glands on the petiole or blade (usually the abaxial side) or both; stipules usually present; blade simple, usually entire, rarely lobed or pseudodentate. Flowers usually bisexual, subtly to strongly bilaterally symmetrical. Sepals 5, eglandular or, most often, the lateral 4 or all 5 bearing (1)2 large multicellular abaxial glands; petals 5, distinct, clawed, alternating with the sepals, imbricate, the innermost (flag) petal posterior and often different from the lateral 4. Stamens mostly 10, fewer by reduction in some genera; anthers mostly dehiscent by longitudinal slits. Gynoecium superior, comprising (2)3 distinct to connate carpels, each fertile locule containing 1 pendent anatropous ovule; styles mostly 1 per carpel and distinct, sometimes connate or reduced in number. Fruits dry or fleshy, dehiscent or indehiscent, samaroid, nut-like, or drupaceous. Seeds without endosperm. Tropics and subtropics (mostly between 30°N and 35°S, mostly New World but also found in the Old World, especially Africa and Madagascar); about 67 genera with over 1250 species, 23 genera and 153 species in the flora area.

M ALPIGHIACEAE 83

Much of this treatment is modified from my paper on the Malpighiaceae of the Guayana Highland (Mem. New York Bot. Gard. 32: 21–305. 1981), which contains full descriptions of many of these species. Successful use of the keys to genera and species requires an understanding of what I mean by certain morphological terms. The ancestral inflorescence of the Malpighiaceae was a raceme of cincinni, but in many genera the cincinni have been reduced to one-flowered units. Each flower is borne on a pedicel, whose base is defined by a joint; below the joint the stalk is called the peduncle, and the peduncle bears two bracteoles, which can be borne anywhere on the peduncle but are most commonly at or near its summit; the peduncle is subtended by a single bract. The peduncle has been lost in several evolutionary lines, in which case the pedicel is described as sessile, subtended then by a cluster of the bract and two bracteoles. The Malpighiaceae are notable for having highly stereotyped flowers (5 sepals mostly with paired abaxial glands on at least the lateral 4 in the New World, 5 clawed petals, mostly 10 stamens, mostly 3 carpels with distinct styles), and as a consequence identification to genus of flowering specimens can be difficult. Understanding the variation in styles and stigmas, and the terminology used to describe that variation, is essential. Most byrsonimoid genera (Blepharandra, Burdachia, Byrsonima, Diacidia, Glandonia, Lophanthera, and Pterandra in the flora area) have subulate styles, i.e., slender styles that taper gradually distally to minute stigmas (◆Fig. 71, A–C). In most cases the stigma is terminal on the style but in some species it is slightly internal, i.e., the stigmatic tissue is on the internal angle of the style apex. In all other genera in the flora area the styles are less tapered distally, such that the stigmas are larger, and their position is correspondingly easier to discern. Some of those genera have the stigmas terminal or nearly so, i.e., the stigmatic tissue is distributed ± evenly over the entire apex of the style. Our genera with consistently terminal stigmas on thick styles are Banisteriopsis, Bunchosia, Diplopterys, and Spachea, but similar styles also occur in some species of Heteropterys, Malpighia, Mascagnia, and Tetrapterys (◆Fig. 71, D–F; see also figures of Diplopterys cabrerana and Spachea elegans below). The rest of our Malpighiaceae, including all or most species of Clonodia, Dicella, Excentradenia, Heteropterys, Hiraea, Jubelina, Lophopterys, Malpighia, Mascagnia, Mezia, Stigmaphyllon, and Tetrapterys, have the styles stigmatic on the internal angle of the apex; in the descriptions below these are called internal stigmas (◆Fig. 71, G–L; see also figures of Dicella julianii, Excentradenia adenophora, Hiraea faginea, and Mezia huberi below). Dorsally the apex of those styles is variable, and that variation has great systematic utility. Such a style tip may be dorsally rounded, truncate, acute, apiculate, or extended into a hook, and in Stigmaphyllon that extension often bears flap-like lateral outgrowths called folioles. In contrast to their relatively uniform flowers, the fruits of Malpighiaceae are exceedingly diverse. Much of that diversity is represented in this flora. The principal dispersal agents for our Malpighiaceae are birds, wind, and water, but a few fruits have no obvious adaptation for dispersal. Three genera of shrubs and trees, representing different clades in the family, have independently evolved fleshy bird-dispersed fruits ranging in size from a pea to a small plum; they are Bunchosia, Byrsonima, and Malpighia. No one has ever actually studied their dispersal, so my statement that they are dispersed by birds represents an assumption, and it is possible that small mammals are also involved. Among the many genera of vines, most of which probably share a common ancestor, the usual fruit is a dry schizocarp, with

84

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A. Burdachia sphaerocarpa

B. Byrsonima coniophylla

G. Heteropterys H. Heteropterys megaptera steyermarkii

C. Lophanthera D. Banisteriopsis longifolia maguirei

I. Jubelina magnifica

J. Mascagnia sinemariensis

E. Bunchosia decussiflora (2 styles connate)

K. Mascagnia schunkei

F. Tetrapterys pusilla

L. Stigmaphyllon sinuatum

Fig. 71. Malpighiaceae styles and stigmas each mericarp being a wind-dispersed samara. Two basic types of samaras have evolved, probably representing two aerodynamically optimal designs. Samaras with a single dorsal wing (the maple type) are found in Banisteriopsis, Heteropterys, and Stigmaphyllon. Samaras with lateral wings, like that of elm, plus many variations, occur in Excentradenia, Hiraea, Jubelina, Lophopterys, Mascagnia, Mezia, and Tetrapterys. There are also two other types of wind-dispersed fruits in (or expected in) our flora. Diacidia is a genus of shrubs and trees in which the fruit proper is a tiny indehiscent nut without any adaptation for dispersal, but in all species except one (which is probably closest to the ancestor of the group) the sepals grow into dry veiny wings as the fruit matures and presumably aid in dispersal. Something similar happens in Dicella, a woody vine with a much larger nut-like fruit subtended at maturity by much-enlarged spreading wings derived from the sepals. The latter two genera are not closely related to each other or the samara-bearing genera; their adaptations for dispersal by wind clearly represent independent innovations. As might be expected in an area with extensive rivers, dispersal by water is common, both as the ancestral method and as a secondary adaptation in plants whose relatives are mostly dispersed by wind. Lophanthera is a genus of riparian trees in which the schizocarpic fruit breaks apart into small dry unwinged cocci half-filled with aerenchyma. Spachea is a related genus, probably sister to Lophanthera, and while the cocci are not as obviously adapted for floating, it seems probable that they are at least partially dispersed by water. Burdachia and Glandonia are sister genera of trees, in both of which the fruit is a relatively large, indehiscent, corky or fibrous nut, beautifully adapted for dispersal by water; all their species grow in lowland forests near water. Then there are the vines and shrubs that are obviously derived from groups with samaras. Clonodia and Diplopterys are small genera in which the large wings of their ancestors have been replaced by crests and ruffles that probably help

M ALPIGHIACEAE 85

the mericarps float by increasing the surface area and trapping air; Diplopterys is clearly derived from Banisteriopsis, while Clonodia probably originated in either Heteropterys or Mascagnia. Almost all the larger genera of samara-producing vines contain one or more species in which the samaras have had the principal wing(s) reduced or lost, often augmented by new supernumerary winglets or various kinds of aerenchyma or air-filled chambers. In our flora, that tendency is well developed in species of Heteropterys, Hiraea, Jubelina, Mezia, Stigmaphyllon, and Tetrapterys. Finally, there are a few groups in which the fruits seem to have no obvious adaptation for dispersal. Pterandra is a genus of trees and shrubs in which the distinct carpels grow into small dry indehiscent cocci without wings or flesh, although the areole may be surrounded by spongy tissue (see C. Anderson’s revision, cited below under Pterandra). They are probably dispersed by water or by wind with other small bits of detritus. Given the nature of its fruits, the genus has achieved an impressive distribution, from Panama to southern Brazil, and an endemic species occurs at 1500 m on Cerro Sipapo in our area; one wonders how it got there! In Blepharandra and Diacidia galphimioides the fruit is a tiny indehiscent nut without wings, flesh, or aerenchyma. These fruits also seem likely to be dispersed by wind with detritus. If the fruit were retained in the old flower the whole unit might aid dispersal by wind, but my observations on B. hypoleuca indicate that the fruits fall out and collect under the plant. The more derived species of Diacidia (Sipapoa sensu Maguire) have the sepals enlarged as an adaptation for dispersal by wind. In addition to the 23 genera treated here, there are two additional genera at 1000 m or higher on the Serra Aracá of Amazonas, Brazil, either or both of which may eventually be found in the Venezuelan Guayana. These are Acmanthera (A. Juss.) Griseb. and Verrucularia A. Juss. Both are shrubs or trees with eglandular leaves and bracteoles, slender subulate styles, and unwinged fruits, so they will come out with Blepharandra, Byrsonima, Diacidia, and Pterandra in the keys to genera. In both Acmanthera and Verrucularia the inflorescence is an unbranched thyrse or pseudoraceme, corymbose or elongated, terminating a main axis or a lateral axis with one pair of leaves. Acmanthera has the outer anther locules winged for their whole length, white or pink petals, sessile pedicels, and deciduous stipules 15– 110 mm long with the four at a node coherent or connate to form a sheath. In Verrucularia the outer anther locules bear a line or cluster of vesicular outgrowths toward the apex, the petals are yellow, the pedicels are pedunculate, and the persistent stipules are up to 2 mm long and distinct or basally connate. There is one species each of Acmanthera and Verrucularia on Serra Aracá. Acmanthera parviflora W.R. Anderson is described in my revision of the genus (Contr. Univ. Michigan Herb. 11: 41–50. 1975), and Verrucularia piresii W.R. Anderson is described in my 1981 paper on the Guayana Highland, cited above. Key to the Genera of Malpighiaceae based on flowering material 1. 1. 2(1).

Styles slender and subulate, tapering to minute stigmas; shrubs or trees ................................................................................................................ 2 Styles slender to stout, of uniform thickness or widened at apex, the stigmas large; vines, shrubs, or trees .......................................................... 8 Leaves bearing large glands on petiole or abaxial surface of blade; some bracteoles often bearing large apical or abaxial glands ....................... 3

86

2.

M ALPIGHIACEAE

Leaves and bracteoles eglandular (except for tiny pellucid dots in blade and gland-tipped marginal teeth or cilia on bracts and bracteoles in some species) .......................................................................................... 5 3(2). Anthers with 2 dark longitudinal wings on outer locules; carpels connate only along a narrow central axis, separating in fruit ...... 15. Lophanthera 3. Anthers unwinged; carpels broadly and persistently connate ................. 4 4(3). Stipules connate intrapetiolarly, persistent; flower buds spheroidal; connective of anthers enlarged, greatly exceeding the apically rounded locules; filaments glabrous ..................................................... 4. Burdachia 4. Stipules connate interpetiolarly, caducous, leaving a large interpetiolar scar; flower buds pyramidal; connective of anthers exceeded by extensions of the apically tapered locules; filaments densely hirsute ............................................................................................... 11. Glandonia 5(2). Inflorescence a tight umbellate fascicle, sessile or subsessile, axillary to leaves or bracts or leaf scars on older stems; carpels distinct; petals abaxially sericeous, at least on claw ..................................... 20. Pterandra 5. Inflorescence an elongated terminal thyrse or pseudoraceme; carpels connate, at least in flower; petals glabrous or rarely bearing a few hairs ................................................................................................................ 6 6(5). Anthers glabrous or bearing medifixed or submedifixed hairs, the apex without specially modified or directed hairs; hairs on leaves (if any) mostly medifixed or submedifixed or branched, rarely basifixed or sub-basifixed .......................................................................... 5. Byrsonima 6. Anthers bearing few to many basifixed awns or hairs, the apical ones ± stiff and directed slightly forward; hairs on leaves (if any) mostly basifixed or sub-basifixed ...................................................................... 7 7(6). Petals all yellow; anthers with 2(–4) stout apical awn-like hairs, these strongly differentiated from other hairs on stamen, if any; stamens 6–10; ovary with only 2 locules developed and containing ovules .................................................................................................... 7. Diacidia 7. Petals white and/or pink, or 4 white and the posterior pale yellow; anthers with apical hairs hardly or not at all different from other hairs on stamen; stamens 10; ovary with all 3 locules fertile, except in some populations of B. fimbriata (a few pink-flowered species of Byrsonima have anthers that mimic those of Blepharandra, from which they differ in having stipules connate to form an intrapetiolar pair, connective of anthers much exceeding fertile part of locules, and ovary sericeous at apex) .................................................................................. 2. Blepharandra 8(1). Petals pink or rose and/or white, or lilac .................................................. 9 8. Petals yellow, or yellow with a red central blotch or red flecks, or yellow turning red, or brownish ..................................................................... 15 9(8). Stipules large, 3–6 mm long, intrapetiolar and completely connate; leaf blades bearing 2–4 impressed glands in adaxial surface near apex ................................................................................................... 21. Spachea 9. Stipules small, up to 2 mm long, interpetiolar or borne on base of petiole, distinct; leaf blades without adaxial glands ....................................... 10 10(9). Inflorescence an unbranched axillary umbel or corymb; shrubs or small trees ....................................................................................... 17. Malpighia

M ALPIGHIACEAE 87

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

14.

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

17. 18(17). 18. 19(18). 19. 20(19). 20. 21(19). 21. 22(21). 22.

23(15).

Inflorescence an elongated pseudoraceme or compound panicle or cyme, or if an axillary corymb the plant a vine; woody vines or shrubs ...... 11 Petals (at least the lateral 4) with a prominent abaxial wing ................ 12 Petals abaxially smooth or at most carinate ........................................... 13 Ultimate units of inflorescence tight corymbs or umbels of 4–10 flowers ............................................................................................ 12. Heteropterys Ultimate units of inflorescence elongated pseudoracemes comprising (15–)20–50 flowers .................................................................... 6. Clonodia Styles with stigmas quite terminal and without any sort of dorsal extension at apex ....................................................................... 1. Banisteriopsis Styles with apex stigmatic on internal angle and dorsally rounded, truncate, acute, or extended into a hook .................................................... 14 Calyx with 1 large central gland on each of the 4 lateral sepals, the anterior sepal eglandular; ultimate units of inflorescence umbels of 4 flowers or corymbs of 6; bracts and bracteoles 4–8 mm long ........ 14. Jubelina Calyx with the 4 lateral sepals biglandular, the anterior eglandular; ultimate units of inflorescence pseudoracemes of (4–)7–50 flowers; bracts and bracteoles up to 3 mm long, mostly smaller ................ 18. Mascagnia Styles with stigmas quite terminal and without any sort of dorsal extension at apex .......................................................................................... 16 Styles with apex stigmatic on internal angle and dorsally rounded, truncate, acute, or extended into a hook or flap-bearing appendage ........ 23 Petals completely concealed by sepals during enlargement of bud, emerging only when flower opens ................................................. 18. Mascagnia Petals (at least the outermost) exposed during enlargement of bud ..... 17 Stipules borne on base of petiole; carpels 2 or 3, completely connate in ovary, developing into an indehiscent fleshy fruit; styles as many as carpels, distinct or connate and then apparently only 1; trees or shrubs ................................................................................................. 3. Bunchosia Stipules interpetiolar or absent; carpels 3, centrally connate in ovary, the fruit dry and schizocarpic; styles 3, distinct; vines or shrubs ............ 18 Calyx bearing 10 glands; bracteoles wider than floriferous bracts and often longer, often bearing marginal or abaxial glands ...... 23. Tetrapterys Calyx bearing 8(9) glands or eglandular; bracteoles as large as bracts or smaller, eglandular .............................................................................. 19 Petals abaxially sparsely to densely sericeous ........................................ 20 Petals glabrous ......................................................................................... 21 Petals abaxially sparsely sericeous, long-fimbriate ................. 9. Diplopterys Petals abaxially densely sericeous, denticulate to lacerate .... 1. Banisteriopsis Pedicels sessile ...................................................................... 1. Banisteriopsis Pedicels raised on peduncles 1–6 mm long ............................................. 22 Flowers borne ultimately in umbels of 4–6; leaf blades abaxially glabrous or very thinly sericeous to glabrate ..................................... 23. Tetrapterys Flowers borne ultimately in pseudoracemes of 6–25, the terminal 4 sometimes crowded into an umbel; leaf blades abaxially sparsely to densely sericeous ............................................................................ 1. Banisteriopsis Bracteoles larger than bracts, globose-cymbiform, borne just below flower, enclosing bud until flower opens; pedicels absent or up to 2(–5)

88

M ALPIGHIACEAE

23.

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

27.

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

30.

31(30). 31. 32(29). 32. 33(32). 33. 34(33). 34. 35(34).

35.

36(34).

mm long in fruit, the peduncles well developed .......................... 19. Mezia Bracteoles mostly similar to bracts or smaller than them, if larger not enclosing bud until flower opens; pedicels well developed relative to peduncles ................................................................................................. 24 Calyx with 1 large central gland on each of the 4 lateral sepals, the anterior sepal eglandular .................................................. 16. Lophopterys Calyx with 2 glands on each of the 4 lateral sepals or on all 5 sepals, or all sepals eglandular ................................................................................. 25 Pedicels sessile ......................................................................................... 26 Pedicels pedunculate ................................................................................ 29 Flowers borne ultimately in pseudoracemes of 2–13 .............. 18. Mascagnia Flowers borne ultimately in umbels of 3–15 or more ............................. 27 Stipules well developed, epipetiolar, often subulate, usually borne at or beyond middle of petiole, if borne near base of petiole the stipules ≥ 2.5 mm long; inflorescences axillary ....................................... 13. Hiraea Stipules none or very small and triangular, borne on or beside base of petiole, if borne on base of petiole the stipules < 1 mm long; inflorescences terminal or axillary and terminal ............................................ 28 Petioles biglandular at base ................................................... 12. Heteropterys Petioles biglandular at apex, or eglandular with glands on abaxial base of blade ............................................................................... 22. Stigmaphyllon Petals abaxially ± densely sericeous or tomentose ................................. 30 Petals glabrous or at most very sparsely sericeous ................................ 32 Full-sized styles 2, posterior, the anterior style often present as a short, slender rudiment; ovary with only the 2 posterior locules developed and fertile; anthers with locules densely hairy ........................... 8. Dicella Styles 3, equal or the anterior slightly shorter; ovary with all 3 locules developed and fertile; anthers glabrous or at most sericeous on connective ........................................................................................................ 31 Bracteoles eglandular or bearing a row of stalked marginal glands .............................................................................................. 18. Mascagnia One of each pair of bracteoles bearing 1 large eccentric abaxial gland .............................................................................................. 23. Tetrapterys Sepals erect or appressed in anthesis ..................................................... 33 Sepals revolute at apex in anthesis ......................................................... 37 Apex of styles (2 or all 3) dorsally extended into a long hook or flapbearing appendage ......................................................... 22. Stigmaphyllon Apex of all styles dorsally rounded, truncate, acute, or short-hooked ... 34 Flowers borne ultimately in elongated to congested pseudoracemes .... 35 Flowers borne ultimately in umbels of 4–6 ............................................. 36 Bracteoles smaller than bracts, borne well below apex of peduncle; lateral 4 sepals biglandular, anterior eglandular; carpels with 1 crest on each side ....................................................................................... 18. Mascagnia Bracteoles mostly larger than bracts, borne at or slightly below apex of peduncle; all 5 sepals biglandular; carpels with 2 (or more) crests on each side ............................................................................... 23. Tetrapterys Stipules none or borne on base of petiole; carpels smooth-sided ............................................................................................ 12. Heteropterys

M ALPIGHIACEAE 89

36. 37(32). 37. 38(37). 38. 39(37). 39. 40(39). 40. 41(40). 41. 42(40). 42. 43(42).

43.

Stipules interpetiolar, distinct or connate; carpels with lateral crests, developing into lateral wings in fruit .................................. 23. Tetrapterys Sepals all abaxially biglandular .............................................................. 38 Sepals all eglandular or the lateral 4 biglandular and the anterior eglandular ............................................................................................ 39 Petals completely concealed by sepals during enlargement of bud, emerging only when flower opens ....................................... 18. Mascagnia Petals (at least the outermost) exposed during enlargement of bud .............................................................................................. 23. Tetrapterys Petioles eglandular ................................................................ 12. Heteropterys Petioles biglandular ................................................................................. 40 Flowers borne ultimately in umbels of 4, sometimes with an additional pair of flowers borne below the terminal umbel ................................. 41 Flowers borne ultimately in elongated to congested pseudoracemes .... 42 Petioles 1–4 mm long ............................................................. 12. Heteropterys Petioles 10–20 mm long ...................................................... 10. Excentradenia Petiole glands borne between middle and apex .................... 12. Heteropterys Petiole glands borne at or slightly above base ........................................ 43 Mature leaf blades abaxially metallic-sericeous, the hairs so dense and appressed as to completely conceal epidermis; inflorescence compound, paniculate, the ultimate pseudoracemes containing 2–16 flowers; bracts eglandular or biglandular; bracteoles eglandular .............. 18. Mascagnia Mature leaf blades abaxially sericeous to glabrescent, the hairs never dense enough to completely conceal epidermis; inflorescence simple, containing 20–60 flowers; bracts eglandular; 1 of each pair of bracteoles bearing 1 large eccentric abaxial gland ........... 12. Heteropterys Key to the Genera of Malpighiaceae based on fruiting material

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

5(2).

Fruits themselves unwinged, the sepals sometimes accrescent or winglike in fruit; fruits schizocarpic or indehiscent ..................................... 2 Fruits winged, the wings reduced in some species to winglets or dissected crests; fruits schizocarpic .................................................................... 12 Fruits schizocarpic, the mericarps dry, or the carpels distinct even in flower ...................................................................................................... 3 Fruits indehiscent, dry or fleshy ............................................................... 5 Leaves and bracteoles eglandular (except for tiny pellucid dots in leaf blades of some species) .......................................................... 20. Pterandra Leaves bearing large glands on petiole or abaxial surface of blade; some bracteoles often bearing large apical or abaxial glands ....................... 4 Styles slender and subulate, the stigmas minute; anthers longitudinally winged; leaf blades without adaxial glands ..................... 15. Lophanthera Styles stout, truncate or subpeltate at apex; anthers unwinged; leaf blades bearing 2–4 impressed glands in adaxial surface near apex ................................................................................................... 21. Spachea Woody vines; sepals greatly enlarged in fruit, forming narrowly elliptic or obovate wings 20–55 mm long, these strongly unequal, the posterior 2 longest, the anterior shortest ....................................................... 8. Dicella

90

M ALPIGHIACEAE

5. 6(5). 6.

7(6).

7. 8(7).

8.

9(6).

9.

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

11. 12(1). 12. 13(12). 13. 14(13). 14. 15(14).

Shrubs or trees; sepals accrescent or not in fruit but not longer than 13 mm, equal or subequal ...................................................................... 6 Leaves and bracteoles eglandular (except for gland-tipped marginal teeth or cilia on bracts and bracteoles in some species) ................................ 7 Leaves bearing large glands on petiole or abaxial surface of blade; bracteoles eglandular or some bracteoles bearing large abaxial glands ................................................................................................................ 9 Fruits 4–15 mm diameter or larger, the stone covered by a fleshy exocarp; hairs on leaves, if any, mostly medifixed or submedifixed or branched, rarely basifixed or sub-basifixed ........................................... 5. Byrsonima Fruits up to 3.5 mm diameter, dry at maturity, the stone covered by a very thin, nonfleshy coat; hairs on leaves (if any) mostly basifixed ............ 8 Fruits with only 2 locules; anthers with 2(–4) stout apical awn-like hairs, these strongly differentiated from other hairs on stamen, if any; sepals slightly to greatly accrescent in fruit ........................................ 7. Diacidia Fruits with up to 3 fertile locules (only 2 in some populations of B. fimbriata); anthers with apical hairs hardly or not at all different from other hairs on stamen; sepals not or hardly accrescent in fruit ........................................................................................... 2. Blepharandra Fruits without a soft fleshy exocarp, dry and corky or fibrous at maturity; inflorescence terminal, with each bract subtending a short cincinnus of 1–6 flowers; styles slender and subulate, distinct, the stigmas minute .............................................................................................................. 10 Fruits with an edible fleshy yellow, orange, or red exocarp at maturity; inflorescence lateral, with each bract subtending 1 flower; styles stout, distinct or connate, with large stigmas ............................................... 11 Stipules connate intrapetiolarly, persistent; filaments glabrous ................................................................................................. 4. Burdachia Stipules connate interpetiolarly, caducous, leaving a large interpetiolar scar; filaments densely hirsute ............................................ 11. Glandonia Flowers borne in pseudoracemes; bracteoles (1 or both) often bearing 1 large abaxial gland, sometimes 2; styles 2 or 3, distinct or connate and then apparently only 1; stipules borne on base of petiole ..... 3. Bunchosia Flowers borne in umbels or tight corymbs; bracteoles all eglandular; styles 3, distinct; stipules borne on stem between petioles .... 17. Malpighia Mericarps with wings very short relative to size of nut, often dissected and irregular or rudimentary .............................................................. 13 Mericarps samaroid, with either dorsal or lateral wing(s) well developed .............................................................................................................. 18 Flowers borne ultimately in a pseudoraceme, sometimes reduced to a single pair ............................................................................................. 14 Flowers borne ultimately in umbels or corymbs of (3)4(–8) ................... 16 Mericarps bearing a dorsal winglet but no lateral winglets, the sides smooth or at most somewhat rugose ................................. 12. Heteropterys Mericarps bearing a dorsal crest or winglet and several winglets or rounded or aculeate outgrowths on each side..................................... 15 Pseudoracemes comprising (15–)20–50 flowers; woody vines, occasionally shrubby; blade of larger leaves (2.5–)4–6(–7) cm wide ............ 6. Clonodia

M ALPIGHIACEAE 91

15. 16(13). 16. 17(16).

17. 18(12). 18. 19(18). 19. 20(19). 20. 21(18).

21.

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

23.

24(22). 24. 25(24).

Pseudoracemes comprising 2–12(–22) flowers; shrubs 0.2–1 m tall; blade of larger leaves 0.9–2.8 cm wide .......................................... 23. Tetrapterys Stipules epipetiolar, usually borne at or beyond middle of petiole, often subulate ....................................................................................... 13. Hiraea Stipules interpetiolar, short, triangular .................................................. 17 Mericarps with a dorsal crest or winglet 1–5 mm wide and essentially 4 roughly parallel ridges or winglets 0.5–10 mm wide on each side, these irregular, dissected, and interconnected with transverse ridges ............................................................................................... 9. Diplopterys Mericarps with a dominant triangular dorsal wing 4–9 mm long and several lateral ribs or crests radiating from the areole ..... 22. Stigmaphyllon Samaras with dorsal wing dominant, the nut bearing on its sides only short winglets or crests or quite smooth ............................................. 19 Samaras with lateral wing(s) dominant, the dorsal wing smaller or reduced to a winglet or crest, occasionally absent ................................. 21 Wing of samara with the abaxial edge thickened, the veins diverging and branching from it toward the thinner adaxial edge ......... 12. Heteropterys Wing of samara with the adaxial edge thickened, the veins diverging and branching from it toward the thinner abaxial edge ........................... 20 Styles with stigmas quite terminal and without any sort of dorsal extension at apex ....................................................................... 1. Banisteriopsis Styles with apex stigmatic on internal angle and a prominent dorsal extension of apex in the form of a hook or flap ............. 22. Stigmaphyllon Bracteoles larger than bracts, globose-cymbiform, borne just below flower, enclosing bud until flower opens; pedicel absent or up to 2(–5) mm long in fruit, the peduncle well developed, 7–25 mm long in fruit ....................................................................................................... 19. Mezia Bracteoles similar to bracts or smaller than them, or if larger not enclosing bud until flower opens; pedicel well developed relative to peduncle .............................................................................................................. 22 Calyx with 1 large central gland on each of the 4 lateral sepals, the anterior eglandular ..................................................................................... 23 Calyx eglandular or with 2 glands on each of the 4 lateral sepals or on all 5 sepals ................................................................................................. 24 Lateral wings of samara directed sideways, semicircular or up to twice as long as wide; bracteoles 4 mm long or longer; flowers borne ultimately in umbels of 4 or corymbs of 6; fertile locule of samara accompanied on each side by a parallel sterile cavity developed in base of lateral wing during maturation ................................................................... 14. Jubelina Lateral wings of samara directed forward, 3 or more times as long as wide; bracteoles 1.5–3 mm long; flowers borne ultimately in elongated pseudoracemes of 4–50; fertile locule of samara without parallel sterile cavities adjacent to it ......................................................... 16. Lophopterys Pedicels sessile ......................................................................................... 25 Pedicels pedunculate ................................................................................ 26 Flowers borne ultimately in pseudoracemes of 2–16; stipules short (≤ 0.5 mm long), triangular, borne on petiole at or somewhat above base .............................................................................................. 18. Mascagnia

92

M ALPIGHIACEAE

25.

Flowers borne ultimately in umbels of 4; stipules mostly subulate, borne on petiole usually at or beyond middle, if borne near base of petiole the stipules ≥ 2.5 mm long ................................................................ 13. Hiraea 26(24). Samaras with 4 discrete lateral wings, 2 on each side (in some species the 4 lateral wings may be accompanied by several long aculeate outgrowths between them) ....................................................... 23. Tetrapterys 26. Samaras with 1 continuous lateral wing or 2, 1 on each side ................ 27 27(26). Flowers mostly borne ultimately in pseudoracemes of (2–)4–50, these occasionally congested into corymbs or umbels; bracteoles eglandular or all bearing several small stalked marginal glands ............. 18. Mascagnia 27. Flowers borne ultimately in umbels of 4; one of each pair of bracteoles bearing 1 large, sessile, eccentric abaxial gland ............ 10. Excentradenia 1. BANISTERIOPSIS C.B. Rob. in Small, N. Amer. Fl. 25: 131. 1910. Banisteria sensu A. Juss., Griseb., and Nied., non L. 1753. Vines, shrubs, or rarely small trees. Leaves bearing glands on petiole or abaxial surface of blade or both; stipules small, distinct, interpetiolar. Floriferous bracts and bracteoles eglandular; pedicels usually sessile, raised on a peduncle in a few species. Petals yellow, pink, or white, usually the lateral 4 spreading or reflexed and the posterior erect. Stamens 10, all fertile; anthers alike or more commonly heteromorphic. Ovary with the 3 carpels adnate to a common torus, all fertile; styles 3, of uniform thickness or thicker distally, the stigmas terminal. Fruit breaking apart into 3 samaras separating from a short pyramidal torus, each samara having its largest wing dorsal, thickened on the adaxial (upper) edge, the veins terminating in the thinner abaxial edge; much shorter winglets, crests, or irregular outgrowths present on sides of nut in some species; nut usually with a functional carpophore. Mexico, Central America, West Indies, and South America (all countries except Chile and Uruguay); 94 species, 19 in Venezuela, 10 of these in the flora area. See Banisteriopsis, Diplopterys (Malpighiaceae) by Bronwen Gates [Fl. Neotrop. Monogr. no. 30. 1982]. Key to the Species of Banisteriopsis 1. 1.

2(1).

2.

3(1).

3.

Leaf blades very densely and persistently metallic-sericeous abaxially ................................................................................................................ 2 Leaf blades glabrous or thinly sericeous to glabrate abaxially, the hairs at maturity never dense enough to completely hide the epidermis ................................................................................................................ 3 Petals pink, paler in age, the posterior yellow in proximal half; pedicels raised on a peduncle 0.5–3(–7) mm long; nut of samara rugose, tuberculate, muricate, or bearing irregular winglets on sides ........ B. muricata Petals all yellow; pedicels mostly sessile, rarely raised on a peduncle up to 1 mm long; samara unknown, but its nut probably smooth-sided, or possibly bearing 1 winglet on each side ........................................ B. lyrata Bracts and bracteoles deciduous before or during anthesis, or immediately afterwards; petals rose-pink turning white in age; nut of samara hairy on inner surface of locule ...................................................... B. caapi Bracts and bracteoles persistent; petals all yellow; nut of samara glabrous within locule ........................................................................................... 4

Banisteriopsis 93

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

5.

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

8.

9(7). 9. 10(9). 10. 11(10).

11.

12(6). 12. 13(12). 13.

Sepals all eglandular ................................................................................. 5 Lateral 4 sepals biglandular, anterior eglandular .................................... 6 Flowers produced mostly on leafless stems, the inflorescences axillary to scars of leaves of previous seasons; styles densely bearded for 1/4–1/2 their length, the long spreading hairs especially extensive on anterior style; nut of samara bearing on each side 3–7 crest-like wings radiating from the areole ............................................................................ B. cristata Flowers produced on currently leafy stems; styles glabrous; nut of samara unappendaged on sides, smooth or with reticulate veins prominent ............................................................................................... B. martiniana Specimens with flowers ............................................................................. 7 Specimens with fruits .............................................................................. 12 Petals, especially the lateral 4, densely sericeous abaxially .................... 8 Petals glabrous ........................................................................................... 9 Petioles of larger leaves usually bearing 2 large glands at or slightly below apex; bracts and bracteoles 1–1.7 mm long, slightly concave to nearly flat, spatulate or subrotund, spreading to reflexed; anthers glabrous; nut of samara bearing several winglets on each side, nearly parallel to the areole ...................................................................... B. krukoffii Petioles eglandular; bracts and bracteoles up to 1 mm long, cymbiform or triangular, erect to appressed; anthers with the locules pubescent; nut of samara with several to many ridges or crests on each side, radiating from the areole .............................................................................. B. lucida Lateral petals subentire to denticulate, especially at base of limb; pedicels raised on a peduncle 0.5–2(–3) mm long ................................ B. wurdackii Lateral petals fimbriate or lacerate or at least distinctly dentate all around margin of limb; pedicels sessile .............................................. 10 Leaf blades deeply cordate at base, the lobes often equaling petiole; inflorescence usually glabrous, rarely sericeous to glabrate .... B. pulcherrima Leaf blades truncate, rounded, or shallowly cordate at base; inflorescence hairy, the hairs usually persistent ...................................................... 11 Inflorescence golden-sericeous; connective of anthers opposite 3 anterior sepals only slightly exceeding locules, by up to 0.5 mm; reduced leaves in inflorescence entire, the margin eglandular or bearing tiny glands ................................................................................................... B. maguirei Inflorescence minutely brown- or white-tomentose; connective of anthers opposite 3 anterior sepals greatly exceeding locules, by 0.6–1 mm; reduced leaves in inflorescence usually bearing well-developed glands or ciliate processes on margin ................................................... B. martiniana Nut of samara unappendaged on sides, the veins sometimes prominent ........................................................................................ go to couplet 10 Nut of samara bearing winglets, crests, or ridges on sides .................... 13 Flowers borne ultimately in short pseudoracemes of 10–25 ..... B. wurdackii Flowers borne ultimately in umbels of 4(–6) ...................... go to couplet 8

Banisteriopsis caapi (Griseb. in Mart.) C.V. Morton, J. Wash. Acad. Sci. 21: 486. 1931. —Banisteria caapi Griseb. in Mart., Fl. Bras. 12(1): 43. 1858. —Ayahuasca, Caapi, Yagé.

Banisteriopsis inebrians C.V. Morton, J. Wash. Acad. Sci. 21: 485. 1931. Woody vine; blade of larger leaves 10–22 × 4–10 cm, abaxially sparsely sericeous to glabrate; inflorescence with flowers borne ul-

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timately in umbels of 4; bracts and bracteoles deciduous before or during anthesis, or rarely immediately afterward; sepals all eglandular or the lateral 4 biglandular; petals rose-pink turning white in age, fimbriate; anthers with locules sparsely pilose to glabrate, some with the connective much enlarged and glandular; samaras 21–47 × 8–22 mm, the nut ribbed or rarely short-aculeate on sides, hairy on inner surface of locule. Areas of evergreen lowland forests, 100–200 m; Amazonas (Caño Iguana, Gavilán, Isla el Silencio in Río Orinoco basin, Río Marueta, San Carlos de Río Negro). Amazonian Colombia, Ecuador, Peru, Brazil, Bolivia, northernmost Argentina near Bolivia. Banisteriopsis caapi is widely cultivated by native peoples of western Amazonia, and used by them as a principal ingredient in hallucinogenic concoctions. Unequivocally natural populations are not known, but the plants are often found naturalized near present or former villages. Banisteriopsis cristata (Griseb.) Cuatrec., Ciencia (Mexico) 23: 141. 1964. —Banisteria cristata Griseb., Linnaea 22: 16. 1849. Banisteria orbicularis Nied., Repert. Spec. Nov. Regni Veg. 33: 69. 1933. Woody vine or shrub with twining branches; blade of larger leaves 3.5–9(–12) × 4–8 cm, soon glabrate on both sides except sericeous margin and midrib; inflorescences axillary to scars of leaves of previous seasons, the flowers borne ultimately in umbels of 4; bracts and bracteoles persistent; calyx eglandular; petals yellow, glabrous; anthers glabrous; styles bearded for up to 1/2 their length; samaras 29–39 × 9–11 mm, the nut bearing on each side 3–7 crest-like wings radiating from areole. Savannas, deciduous forests, 200–400 m; Bolívar (area of Ciudad Bolívar, Lago Guri, San Félix). Anzoátegui, common in northern coastal states of Venezuela; Guyana, Suriname. Banisteriopsis krukoffii B. Gates, Fl. Neotrop. Monogr. 30: 200. 1982. —Bejuco de ratón. Woody vine; petiole usually biglandular at or near apex; blade of larger leaves 10–21 × 5–9 cm, abaxially thinly sericeous with very

short hairs; inflorescence with flowers borne ultimately in umbels of 4; bracts and bracteoles 1–1.7 mm long, slightly concave to nearly flat, spatulate or subrotund, spreading to reflexed, persistent; 4 lateral sepals biglandular, anterior eglandular; petals yellow, abaxially densely sericeous, denticulate or short-fimbriate; anthers glabrous; nut of samara bearing several winglets on each side, nearly parallel to areole. Evergreen lowland and lower montane wet forests, 100–600 m; Bolívar (La Escalera), Amazonas (Río Mawarinuma, Solano). Brazil (Amazonas). Banisteriopsis lucida (Rich.) Small, N. Amer. Fl. 25: 133. 1910. —Banisteria lucida Rich., Actes Soc. Hist. Nat. Paris 1: 109. 1792. Woody vine; petioles eglandular; blade of larger leaves 8–16 × 3.5–7.5 cm, abaxially sparsely sericeous, sometimes glabrescent; inflorescence with flowers borne ultimately in umbels of 4(–6); bracts and bracteoles up to 1 mm long, cymbiform or triangular, erect to appressed, persistent; 4 lateral sepals biglandular, anterior eglandular; petals yellow, abaxially densely sericeous, dentate to lacerate; anther locules pubescent; samaras 41–73 × 14–25 mm, the nut with the sides prominently ridged or rarely cristate, the ridges radiating from areole. Evergreen lowland to montane forests, 100–1300 m; Delta Amacuro (Caño Güiniquina), Bolívar (Caño Majagua tributary of Río Parucito, Gran Sabana, upper Río Caroní, upper Río Caura, upper Río Paragua), southern Amazonas (Cariche, Río Baría/Mawarinuma, Río Casiquiare, Río Padamo, Río Siapa, Sierra de la Neblina, Tamatama). Apure, Aragua, Distrito Federal, Falcón, Lara, Sucre, Táchira, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Banisteriopsis lyrata B. Gates, Brittonia 31: 108. 1979. Woody vine; petioles biglandular at apex; blade of larger leaves 7–13 × 4.5–6.5 cm, abaxially densely and persistently goldenmetallic-sericeous; inflorescence with flowers borne ultimately in short pseudoracemes of 3 or 4 pairs or umbels of 4, usually with 1

Banisteriopsis 95

or 2 more proximal pairs below; pedicels sessile or rarely raised on a peduncle up to 1 mm long; bracts and bracteoles persistent; lateral 4 sepals biglandular, anterior eglandular; petals yellow, glabrous, long-fimbriate; anthers glabrous, some with the connective much enlarged and glandular; samaras unknown, the nut probably smooth-sided, possibly bearing 1 winglet on each side. Lowland forests, 100–200 m; Delta Amacuro (town of Sierra Imataca east-southeast of Los Castillos), Bolívar (Río Caura, Río Paragua). Brazil (Pará, Rondônia). Banisteriopsis maguirei B. Gates, Brittonia 31: 108. 1979. Woody vine; blade of larger leaves 6–12.5 × 3.5–10 cm, rounded or shallowly cordate at base, abaxially sparsely sericeous; inflorescence densely to sparsely golden-sericeous, the flowers usually borne ultimately in pseudoracemes of up to 7 pairs, rarely in umbels of 4 flowers; reduced leaves of inflorescence with margin eglandular or bearing tiny sessile glands; bracts and bracteoles persistent; lateral 4 sepals biglandular, anterior eglandular; petals yellow, glabrous, long-fimbriate; anthers glabrous; samaras 20–28 × 8–10 mm, the nut unappendaged on sides, sometimes with prominent veins. Tepui meadows, probably mostly in wooded ravines along streams, 1000–1900 m; western Bolívar (Cerro Guanay, Cerro Guaiquinima, Cerro Jaua, Sierra de Maigualida), Amazonas (Cerro Aracamuni, Cerro Coro Coro, Cerro Cuao, Cerro Duida, Cerro Guanay, Cerro Huachamacari, Cerro Marahuaka, Cerro Parima, Cerro Sipapo, Cerro Yutajé). Endemic. Banisteriopsis martiniana (A. Juss.) Cuatrec., Webbia 13: 498. 1958. —Banisteria martiniana A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 284. 1840. Banisteria leptocarpa Benth., London J. Bot. 7: 130. 1848. —Banisteriopsis leptocarpa (Benth.) R.O. Williams, Fl. Trinidad 1: 131. 1929. Woody vine, rarely shrubby; leaf blades obtuse to shallowly cordate at base, abaxially sparsely sericeous to soon glabrate; inflorescence minutely brown- or white-tomentose, the flowers borne ultimately in umbels of 4; reduced leaves in inflorescence with ±

well-developed glands or cilia on margin; bracts and bracteoles persistent; sepals all eglandular or lateral 4 biglandular and anterior eglandular; petals yellow, dentate to lacerate, glabrous; anthers glabrous, those opposite 3 anterior sepals with the connective much enlarged; samaras 18–35 × 5–12 mm, the nut unappendaged on sides, sometimes with prominent veins. Both recognized varieties occur in the Venezuelan Guayana. Key to the Varieties of B. martiniana 1. Hairs of inflorescence rusty brown; leaf blades plane, 4.5–12.5 cm long, bearing cupulate marginal glands; Delta Amacuro and Bolívar ...... var. martiniana 1. Hairs of inflorescence white or gray; leaf blades falcate, 7–17 cm long, bearing minute marginal glands; Amazonas .................................... var. subenervia B. martiniana var. martiniana Climbing in forests and sprawling in open places, near sea level to 2000 m; common in Delta Amacuro and eastern Bolívar, also northwestern Bolívar. Aragua, Sucre; Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 72. B. martiniana var. subenervia Cuatrec., Webbia 13: 501. 1958. Banisteriopsis martiniana var. laevis Cuatrec., Webbia 13: 502. 1958. Evergreen lowland forests, 100–300 m; Amazonas (Río Yatua, San Carlos de Río Negro). Amazonian Colombia, Peru, Brazil. Banisteriopsis muricata (Cav.) Cuatrec., Webbia 13: 503. 1958. —Banisteria muricata Cav., Diss. 9: 423, pl. 246. 1790. Heteropterys argentea H.B.K., Nov. Gen. Sp. 5: pl. 450. 1821.—Banisteria argentea (H.B.K.) Spreng., Syst. Veg. 2: 388. 1825. —Banisteriopsis argentea (H.B.K.) C.B. Rob. in Small, N. Amer. Fl. 25: 133. 1910. Banisteria metallicolor A. Juss. in A. St.Hil., Fl. Bras. Merid. 3: 46. 1832 [1833]. —Banisteriopsis metallicolor (A. Juss.) O’Donell & Lourteig, Lilloa 9: 259. 1943. Banisteria schomburgkiana Benth., London J. Bot. 7: 129. 1848. —Banisteriopsis

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Fig. 72. Banisteriopsis martiniana var. martiniana

Blepharandra 97

schomburgkiana (Benth.) C.B. Rob. in Small, N. Amer. Fl. 25: 133. 1910. Woody vine, sometimes shrubby; blade of larger leaves 5.5–14 × 3–8 cm, abaxially (in the flora area) very densely and persistently metallic-sericeous (silver or golden); inflorescence with flowers borne ultimately in umbels of 4; pedicels raised on a peduncle 0.5– 3(–7) mm long; bracts and bracteoles persistent; sepals all eglandular or the lateral 4 biglandular; petals glabrous, pink, paler in age, the posterior yellow proximally; anthers glabrous, some with the connective much enlarged and glandular; samaras (16–)24–40 (–50) × (9–)11–16(–20) mm, the nut rugose, tuberculate, muricate, or bearing irregular winglets on sides. Wet forests to drier, ± disturbed woods, often growing at edge of wooded areas along rivers and ravines, 100– 400 m; common in northern Bolívar. Elsewhere in Venezuela known from most states; Mexico, Central America, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Banisteriopsis pulcherrima (Sandwith) B. Gates, Brittonia 31: 109. 1979. —Banisteriopsis elegans var. pulcherrima Sandwith, J. Arnold Arbor. 24: 223. 1943. Woody vine or shrub; blade of larger leaves 8–16 × 4–12 cm, deeply cordate at base, glabrous, usually with tooth-like marginal glands or ciliate processes; inflores-

cence usually glabrous, rarely sericeous to glabrate, the flowers borne ultimately in umbels of 4; reduced leaves in inflorescence with ciliate processes up to 6 mm long around margin; lateral 4 sepals biglandular, anterior eglandular; petals yellow, fimbriate, glabrous; anthers glabrous, those opposite the 3 anterior sepals with connective much enlarged; samaras 27–39 × 10–13 mm, the nut unappendaged on sides, sometimes with prominent veins. Wooded slopes, roadsides, savannas, 300–2200 m; southeastern Bolívar (Gran Sabana). Western Guyana. Banisteriopsis wurdackii B. Gates, Brittonia 31: 109. 1979. Woody vine; blade of larger leaves 10–19 × 4.5–10.5 cm, abaxially sparsely sericeous; inflorescence with flowers borne ultimately in pseudoracemes of 10–25; pedicels raised on a peduncle 0.5–2(–3) mm long; bracts and bracteoles persistent; lateral 4 sepals biglandular, anterior eglandular; petals yellow, glabrous, subentire to denticulate, especially at base of limb; anthers glabrous; samaras 25–46 × 10–15 mm, the nut bearing on each side a single winglet parallel to areole. Semideciduous to evergreen lowland and lower montane forests, 50–400 m; northwestern Bolívar (El Manteco, Río Parguaza, Río Villacoa). Mérida; Costa Rica, Panama, Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia.

2. BLEPHARANDRA Griseb., Linnaea 22: 7. 1849. Trees or shrubs, the hairs often basifixed or sub-basifixed. Leaves eglandular; stipules intra- and epipetiolar, distinct from each other but often basally connate with opposite stipules to form an interpetiolar sheath; blade with many (12–20 or more) fine parallel lateral veins interconnected by a fine elaborate reticulum. Inflorescence a thyrse or pseudoraceme composed of 1–several-flowered cincinni; bracts and bracteoles without abaxial glands. Sepals all abaxially biglandular; petals white, pink, or red (posterior petal pale yellow in 2 species), glabrous or bearing a few hairs on the claw. Stamens 10; filaments distinct, hirsute with straight basifixed hairs; anthers with locules bearing at least apical tufts of basifixed hairs and often hirsute on sides as well. Ovary of 3 completely connate carpels, all fertile or the anterior empty, glabrous; styles 3, slender and subulate with minute terminal stigmas. Fruit a tiny (2–3.5 × 2.5–3.5 mm), spheroidal or ovoidal, 3-angled, dry, indehiscent, nut-like capsule with a bony and often rugose endocarp. Southern Venezuela, Guyana, Amazonian Brazil; 6 species, 4 in Venezuela, all in the flora area.

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

2.

3(1).

3.

Stipules acute or acuminate at apex, deciduous before leaves; 500–2600 m ................................................................................................................ 2 Stipules rounded at apex, persistent on petiole; 50–400 m ...................... 3 Sepals, most bracteoles, and some bracts bearing gland-tipped marginal processes 0.5–2 mm long; leaves light green abaxially, not or only thinly glaucous; leaves, vegetative internodes, and abaxial surface of stipules glabrous ...................................................................... B. fimbriata Sepals, bracteoles, and bracts entire or denticulate; leaves white- or yellowish-glaucous abaxially; petioles, internodes, and abaxial surface of stipules at least initially densely hairy; leaf blades densely hairy to glabrate abaxially ................................................................... B. hypoleuca Petals all pink, turning white in age; sepals glabrous adaxially or sparsely sericeous near margin, sericeous abaxially; free lobes of stipules 2.5– 4 mm long; pubescence of inflorescence light to dark brown; pedicels 5– 6(–7) mm long; blade of larger leaves 1.5–3 cm wide, the margin thick and nonrevolute ................................................................... B. angustifolia Four lateral petals white, posterior petal pink; sepals densely sericeous on both sides; free lobes of stipules 4–7 mm long; pubescence of inflorescence white; pedicels 8–10(–13) mm long; blade of larger leaves (2.5–)3–7 cm wide, the margin thin and often revolute ..... B. heteropetala

Blepharandra angustifolia (H.B.K.) W.R. Anderson, Mem. New York Bot. Gard. 32: 55. 1981. —Byrsonima angustifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 153, pl. 449. 1821 [1822]. Much-branched shrub or small tree 1–6 m tall; stipules persistent on petiole, the free lobes 2.5–4 mm long, rounded; petioles 7– 11(–18) mm long; blade of larger leaves 4.5– 7.5(–8.5) × 1.5–3 cm, elliptic or rectangular or narrowly obovate, thick and nonrevolute at margin; petals pink turning white in age. Savannas, usually on white sand, 100–400 m; Amazonas (base of Cerro Yapacana, Río Atabapo, Río Casiquiare, Río Guainía, Río Orinoco, Río Pasimoni). Brazil (Amazonas: upper Rio Negro). ◆Fig. 74. Blepharandra fimbriata MacBryde, Canad. J. Bot. 52: 2437. 1974. Weak shrub, prostrate to erect, to 2 m tall; vegetative internodes and leaves glabrous, except for tufts of hairs at nodes; stipules 11– 30 × 4–7 mm, acute or acuminate at apex, deciduous before leaves; blade of larger leaves 3.5–8 × 1.7–3.8 cm, elliptic, not or only thinly glaucous abaxially, usually revolute at margin; some bracts, most bracteoles, and sepals

glandular-fimbriate; 4 lateral petals white, posterior petal pale yellow. Open scrub and rocky places along rivers, 600–1800 m; Bolívar (near Canaima, Cerro Guaiquinima, Uei-tepui). Guyana (Ayanganna Plateau). ◆Fig. 75. Blepharandra heteropetala W.R. Anderson, Mem. New York Bot. Gard. 32: 57. 1981. Shrub or tree 2–7 m tall; stipules persistent on petiole, the free lobes 4–7 mm long, rounded; petioles (6–)8–12 mm long; blade of larger leaves 6–10 × (2.5–)3–7 cm, elliptic or ovate, often revolute at margin; 4 lateral petals white, posterior petal pink. Shrubby vegetation or scrub forests on white sand, 50– 400 m; Amazonas (Cerro Cariche, Cerro Morocoto below San Fernando del Atabapo, Río Atabapo, between upper Río Orinoco and Cerro Autana, Río Sipapo). Brazil (Amazonas: campinas north and east of Manaus). Blepharandra hypoleuca (Benth.) Griseb., Linnaea 22: 7. 1849. —Coleostachys hypoleuca Benth., London J. Bot. 7: 125. 1848.

Blepharandra 99

Fig. 73. Blepharandra hypoleuca

Fig. 74. Blepharandra angustifolia

Fig. 75. Blepharandra fimbriata

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Byrsonima cretacea Gleason, Bull. Torrey Bot. Club 58: 378. 1931. —Blepharandra cretacea (Gleason) Steyerm., Fieldiana, Bot. 28: 280. 1952. Blepharandra ptariana Steyerm., Fieldiana, Bot. 28: 282. 1952. Shrub or small tree 1–8 m tall; vegetative internodes sericeous to glabrate; stipules 5–17(–22) × 3–9 mm, acuminate at apex, deciduous before leaves; blade of larger leaves 4.5–15.5 × 3–9.5 cm, elliptic or ovate, abaxially glaucous and densely hairy to glabrate; bracts, bracteoles, and sepals entire or denticulate; 4 lateral pet-

als white, posterior petal pale yellow. Sandy upland savannas and shrubby associations, often along rocky streams, 500– 2600 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Jaua, Cerro Marutani, Ilú-tepui, Macizo del Chimantá, Ptari-tepui, upper Río Caroní, upper Río Paragua, Sierra Pakaraima), Amazonas (Cerro Aracamuni, Cerro Duida, Sierra Unturán). Western Guyana, Brazil (Amazonas: Serra Aracá). ◆Fig. 73. Blepharandra hypoleuca is an extremely variable species; see discussion in Mem. New York Bot. Gard. 32: 53–55. 1981.

3. BUNCHOSIA H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 153. 1821 [1822]. Shrubs or trees. Leaves usually bearing impressed glands abaxially on the blade; stipules distinct, borne on base of petiole. Inflorescence a pseudoraceme, simple or less commonly ternate, axillary without vegetative leaves or terminating a lateral shoot with 1 pair of vegetative leaves; 1 or both bracteoles often bearing 1(2) abaxial glands. Calyx bearing 8–10 often decurrent glands; petals yellow or whitish, glabrous. Stamens 10, usually glabrous; anthers ± alike. Gynoecium 2- or 3-carpellate; ovary with carpels connate and locules as many as carpels, every locule fertile; styles as many as carpels, distinct or partially to completely connate and then apparently only 1, stout, the large terminal stigmas subpeltate or apparently capitate. Fruit an indehiscent “drupe” (actually a berry), yellow, orange, or red at maturity, with 2 or 3 distinct 1-seeded pyrenes in a common fleshy exocarp, each pyrene with a smooth, brittle, cartilaginous wall. Mexico, Central America, West Indies, South America (all countries except Chile and Uruguay); ca. 65 species, 11 in Venezuela, 6 of these in the flora area. Key to the Species of Bunchosia 1.

1.

2(1).

2.

Inflorescence terminating a short lateral branch bearing a pair of sterile vegetative leaves (these sometimes deciduous, but leaving large leaf scars), or occasionally terminating a leaf-bearing main shoot; ovary glabrous or rarely sericeous, 2- or 3-carpellate .................................... 2 Inflorescence axillary, with small bracts only, every bract subtending 1 uniflorous peduncle (or sometimes 2 in B. glandulifera); ovary sericeous, 2-carpellate .................................................................................. 3 Leaf blades abaxially densely velutinous or tomentose, the hairs less dense on fully expanded leaves than on young leaves but almost always persistent and nearly uniformly distributed; fully expanded leaf blades 6–18(–22) cm long; inflorescence 4–12 cm long, bearing 10–30 flowers ........................................................................................................ B. mollis Leaf blades from the beginning densely velutinous on margin and abaxial midrib but bearing only a few hairs on surface between, glabrescent in age(?) or with hairs persistent on margin; leaf blades up to 4 cm long; inflorescence up to 3 cm long (in the only known collection, in which the

Bunchosia 101

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

5.

inflorescences are probably not fully elongated), bearing 8–12 flowers ...................................................................................................... B. petraea Styles 2/3 to completely distinct .................................................................. 4 Styles connate, the stigmas distinct or connate ........................................ 5 Leaves persistently very densely silvery- or golden-sericeous abaxially, the epidermis completely concealed by hairs ........................... B. argentea Leaves sparsely sericeous abaxially to apparently glabrate .... B. armeniaca Leaves thinly but persistently sericeous abaxially, undulate and crispate at margin; pseudoracemes bearing 10–20 flowers, sometimes 2 in the axil of 1 bract; peduncles 2.5–5 mm long; style sericeous; dried fruits 20–28 × 15–20 mm, the wall smooth .................................. B. glandulifera Leaves nearly or quite glabrate, plane or slightly revolute at margin, entire or slightly indented near glands; pseudoracemes usually bearing 20–60 flowers, 1 per bract; peduncles 0.5–2 mm long; style glabrous; dried fruits up to 15 × 16 mm, the wall granulate ............. B. decussiflora

Bunchosia argentea (Jacq.) DC., Prodr. 1: 582. 1824. —Malpighia argentea Jacq., Fragm. Bot. 57, pl. 83. 1800–1809. Shrub or tree (2–)3–20(–25) m tall; blade of larger leaves 10–23 × (5–)7–14 cm, abaxially densely and persistently silvery- or golden-sericeous; pseudoracemes 7–13 cm long, with 15–35(–50) flowers; anther connectives brown to red; ovary 2-carpellate, sericeous; styles distinct; fruits orange to red, 12–30 × 12–25 mm (dried), ± persistently sericeous. Evergreen lowland forests, 200–400 m; Bolívar (Río Nichare). Anzoátegui, Aragua, Barinas, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira, Yaracuy; Costa Rica, Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 76. There is great diversity in the plants to which this name has been applied; they may represent more than one species. Bunchosia armeniaca (Cav.) DC., Prodr. 1: 582. 1824. —Malpighia armeniaca Cav., Diss. 8: 410, pl. 238. 1789. —Palo de colmena. Tree 3–15 m tall; blade of larger leaves 10–27 × 5–10 cm, abaxially sparsely sericeous to nearly glabrate; pseudoracemes 6– 17 cm long, with 8–60 flowers; anther connectives dark red to black; ovary 2-carpellate, densely sericeous; styles at least 2/3 distinct, glabrous or proximally sericeous; fruits reddish, 12–19 × 14–17 mm (dried), ± persistently sericeous. Wet forests, 50–300 m; Delta Amacuro (west of Caño Guayo and east

of Caño Sacupana, Río Cuyubiní, town of Sierra Imataca east-southeast of Los Castillos), northern Bolívar (El Dorado, El Palmar, Pijiguaos, Río Caura). Colombia, Ecuador, Peru, Brazil, Bolivia. It is unlikely that everything passing under this name represents the same species. The styles are not always distinct in plants from Ecuador, but in Venezuela the character seems to be consistent. Bunchosia decussiflora W.R. Anderson, Mem. New York Bot. Gard. 32: 279. 1981. Shrub or tree 3–25 m tall; blade of larger leaves 12–19 × 4.5–10 cm, glabrate at maturity; pseudoracemes 8–15 cm long, with 20– 60 flowers; peduncles 0.5–2 mm long; anther connectives brown or reddish; ovary 2-carpellate, sericeous; style (from 2 connate) glabrous; fruits yellow or reddish, 9–15 × 9–16 mm (dried), glabrate and granulate at maturity. Evergreen lowland to upper montane moist forests, 100–1300 m; southern Amazonas (Raudal de los Guaharibos on upper Río Orinoco, Río Ocamo, Río Ugueto, Sierra de la Neblina). Guyana, French Guiana, Brazil. Bunchosia glandulifera (Jacq.) H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 154. 1821 [1822]. —Malpighia glandulifera Jacq., Collectanea 4: 207. 1790 [1791]. —Ciruela, Ciruela de fraile. Shrub or small tree 2–8 m tall; blade of larger leaves 8.5–18 × (4.5–)6–10(–12) cm, undulate and crispate at margin, abaxially

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Fig. 76. Bunchosia argentea

Burdachia 103

thinly but persistently sericeous; pseudoracemes 5–11 cm long, with 10–20 flowers, the bracts sometimes subtending 2 flowers; peduncles 2.5–5 mm long; anther connectives yellow or light brown; ovary 2-carpellate, sericeous; style (from 2 connate) 3– 3.5 mm long, sericeous; fruits orange to red, 20–28 × 15–20 mm (dried), beaked, the wall smooth, glabrate at maturity. Secondary woods, edges of forests, 50–100 m; Amazonas (La Esmeralda, Pimichín, Puerto Ayacucho, San Antonio del Sipapo, San Carlos de Río Negro). Anzoátegui, Aragua, Barinas, Carabobo, Distrito Federal, Guárico, Miranda, Portuguesa; West Indies, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia. Bunchosia glandulifera has a very large fleshy red-orange fruit. It is widely cultivated for its handsome foliage and showy edible fruits. It is known only as an introduced tree in the West Indies, Peru, Brazil, and Bolivia, and seems likely to be native only in Colombia and Venezuela. Its wide distribution in northern Venezuela suggests that those may be natural populations. None of the collections I have seen from northern Venezuela were described as cultivated, but the possibility remains that in a long-settled area like the Distrito Federal (the type locality) the species was introduced early (perhaps before the arrival of Europeans) and then became naturalized through dispersal by birds. Most, but not all, of the collections from Amazonas were described by the collectors as cultivated. It seems unlikely to me that this showy species is indigenous to southern Venezuela; I would expect it to be better collected there if that were the case.

Bunchosia mollis Benth., London J. Bot. 7: 127. 1848. —Ciruela de fraile, Ciruela montañera. Bunchosia rhombifolia Turcz., Bull. Soc. Imp. Naturalistes Moscou 36: 582. 1863. Bunchosia schomburgkiana Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 5: 45. 1914. Shrub or small tree 1–5 m tall; blade of larger leaves 6–18(–22) × 3–12 cm, abaxially ± uniformly densely velutinous or tomentose, the T- or Y-shaped hairs stellate at base of stalk; inflorescence terminating a lateral shoot with 1 pair of full-sized leaves, 4–12 cm long beyond leaves, with 10–30 flowers; ovary glabrous or very rarely sericeous, 2- or 3-carpellate; styles distinct or up to 1/2 connate; fruits orange to red, 6–11 mm diameter (dried), glabrous. Savannas, scrub forests, gallery forests, near sea level to 300 m; common in northeastern Bolívar. Anzoátegui, Barinas, Guárico, Miranda, Monagas, Nueva Esparta, Sucre; Guyana, Brazil (Roraima). Bunchosia petraea W.R. Anderson, Contr. Univ. Michigan Herb. 21: 40. 1997. Shrub 1.5 m tall; blade of larger leaves (not fully expanded?) 3–4 × 1.6–2 cm, velutinous on margin and abaxial midrib, some hairs with sharp spurs at base of very short stalk; inflorescence terminating a lateral shoot with 1 pair of full-sized leaves, up to 3 cm long (not fully elongated?), with 8–12 flowers; ovary glabrous, 2- or 3-carpellate; styles distinct; fruits unknown. Granitic outcrops, 50–100 m; Amazonas (Puerto Ayacucho–El Burro). Endemic.

4. BURDACHIA A. Juss. in Endl., Gen. Pl. 1064. 1840. Tetrapodenia Gleason, Bull. Torrey Bot. Club 53: 289. 1926. Shrubs or trees. Leaves bearing abaxial glands; stipules intra- and epipetiolar, completely connate, the pair 3–11 mm long, coriaceous, persistent on petiole. Inflorescence terminal, single or 2 or 3 together, each usually divided near base into 3(–5) axes, each axis a raceme of short cincinni; lowest bracteole and alternate subsequent bracteoles bearing 1 large eccentric abaxial gland. Flower buds spheroidal. Sepals all biglandular; petals pink or white; posterior petal bearing 2–4 large glands on each side of base of limb and several smaller glands distally. Stamens 10, glabrous; anthers rounded at apex, often exceeded at apex by the thick, fleshy connective. Ovary of 3 completely connate carpels, 3-locular but 1 of the posterior locules empty and smaller; styles 3, slender and subulate; stigma slightly internal and decurrent. Fruit an indehiscent fibrous or aerenchymatous nut, dry at maturity and without a stony endocarp, usually containing only 1 locule completely filled by 1 large seed.

104

M ALPIGHIACEAE

Amazonian Colombia, Venezuela, Guyana, Peru, Brazil; 2 or 3 species, 2 in Venezuela, both in the flora area. The third species of Burdachia, if it is recognized, is B. duckei Steyerm., which occurs along the Rio Negro in Amazonian Brazil between Barcelos and the Rio Urubú. It is a segregate from B. prismatocarpa. Key to the Species of Burdachia 1.

1.

Fruits pyramidal, bearing 8 or 9 longitudinal ribs, these extended at base into knobs or spurs; leaves minutely sericeous or tomentose abaxially to glabrate, with some hairs usually persisting on midrib; vegetative stems sericeous to glabrate; stipules abaxially sericeous to glabrate; peduncles sparsely to densely sericeous; ovary conoidal ... B. prismatocarpa Fruits conoidal to spheroidal, round in cross section, the wall smooth; leaves and vegetative stems glabrous; stipules abaxially glabrous; peduncles glabrous or with a few hairs in a line; ovary depressed-globose ............................................................................................ B. sphaerocarpa

Burdachia prismatocarpa A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 330. 1840. Burdachia williamsii Steyerm., Fieldiana, Bot. 28: 285. 1952. Shrub or tree 2–15 m tall; blade of larger leaves 10–21(–25) × (3–)4–12(–14) cm; fruits 9–20 × 7–17 mm. Along lowland rivers, 50– 200 m; Bolívar (Cerro Marimarota, Río Horeda near Ciudad Bolívar, Río Parhueña), Amazonas (basins of upper Río Negro and upper Río Orinoco). Apure (Río Meta); Amazonian Colombia, Peru, Brazil. ◆Fig. 77. Plants from Amazonas between San Fernando de Atabapo and Piedra Cocuy tend to have leaves < 6 cm wide, stipules only 3–5 mm long, a glabrous ovary, and fruits < 13 mm long, whereas elsewhere the leaves are mostly wider, the stipules are 6–10 mm long, the ovary is densely tomentose, and the fruits are mostly 14–20 mm long. The narrow-leaved form has been named as Burdachia williamsii, which I recognized in 1981. However, specimens that recombine the character states of these two taxa continue to accumulate. For example, Davidse 27754 (MICH, MO) and 27833 (MICH, MO, VEN) are B. williamsii with wide leaves; Liesner 8619 (MICH, MO, VEN), Stergios & P. Stergios 11293 (MICH, MO, NY, VEN), and Stergios et al. 11641 (MICH, VEN) are B. williamsii with a hairy ovary; Maguire & C. Maguire 35522 (MICH, NY, US, VEN) and Davidse & González 14086 from Apure (MICH) are B. prismatocarpa with short

stipules and fruits. Because of this variation I have decided to abandon B. williamsii as indefensible. I have given the name Burdachia prismatocarpa var. loretoënsis W.R. Anderson to plants from Loreto, Peru, and adjacent Brazil. Given the variation discussed above, it may be that that variety will not continue to merit recognition. If it is recognized, the plants of Venezuela should be called B. prismatocarpa var. prismatocarpa. Burdachia sphaerocarpa A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 330. 1840. —Burdachia prismatocarpa var. sphaerocarpa (A. Juss.) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 5: 60. 1914. Burdachia atractoides Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 5: 59. 1914. Tetrapodenia glandifera Gleason, Bull. Torrey Bot. Club 53: 289. 1926. —Burdachia sphaerocarpa var. glandifera (Gleason) W.R. Anderson, Mem. New York Bot. Gard. 32: 139. 1981. Shrub or tree 3–15(–20) m tall; blade of larger leaves (8–)10–24 × 5–11(–13) cm; fruits 18–28 × 13–18 mm. Lowland primary and secondary forests, usually on the banks of rivers or streams or in periodically flooded areas, 50–100 m; Amazonas (Río Casiquiare, San Carlos de Río Negro). Guyana, Brazil (Amazonas, Pará). ◆Fig. 78.

Burdachia 105

Fig. 77. Burdachia prismatocarpa

Fig. 78. Burdachia sphaerocarpa

106

M ALPIGHIACEAE

The type of Tetrapodenia glandifera came from the Amakura River, on the border between Guyana and Delta Amacuro, so the species must surely occur in Delta Amacuro too. In 1981 I recognized the plants of western Amazonia as Burdachia sphaerocarpa var. sphaerocarpa and those of Guyana as B.

sphaerocarpa var. glandifera, on the basis of an apparent difference in petal color, pink in the west and white in the east. Subsequently collectors have described the petals of some western populations as white, so there seems to be no justification for recognizing two taxa within this species.

5. BYRSONIMA H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 147. 1821 [1822]. Alcoceratothrix Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 1: 45. 1901. Trees, shrubs, or subshrubs. Leaves eglandular; stipules intra- and epipetiolar, distinct or partially to completely connate, persistent on petiole in most species. Inflorescence terminal, a raceme of few-flowered cincinni or a pseudoraceme (i.e., a raceme of 1-flowered cincinni); floriferous bracts and bracteoles eglandular; pedicels sessile or sometimes raised on a short peduncle. Sepals all biglandular or all eglandular, connate as far as tips of glands, the glands green, yellow, white, or pink; petals yellow, white, pink, or red, glabrous in most species; lateral 4 petals with slender recurved claws, the anterior pair with deeply cup-shaped limbs, the posterior pair shallower; posterior petal with a stout, erect claw and the limb smaller, flat or crumpled and often reflexed. Stamens 10, the anthers ± alike. Ovary with the 3 carpels completely connate, 3-locular, all locules fertile or the anterior sterile in some species; styles 3, slender and subulate, the stigmas minute and terminal or slightly internal. Fruit a drupe, the thin flesh green turning yellow, orange, red, purple, blue, or blue-black at maturity, the stone with a hard wall, 3-locular. Mexico, Central America, West Indies, South America (all countries except Chile, Argentina, and Uruguay); at least 125 species, 44 known or expected in Venezuela, 40 of these in the flora area. Key to the Species of Byrsonima 1. 1.

2(1).

2.

3(2).

Pedicels long-pedunculate, the primary peduncle 5–15 mm long; petals yellow; anthers glabrous ........................................................... B. maguirei Pedicels sessile or short-pedunculate, the peduncle 0–3 mm long; petals yellow, white, pink, or red, if yellow the anthers mostly sericeous or tomentose between or on sides of locules ................................................. 2 Stipules distinct (Caution: This refers to the pair of stipules borne on the adaxial face of a single petiole, best viewed by removing the petiole, to which the stipules will remain attached. In a few species two stipules from opposite petioles are short-connate across the node, but that is not the connation to which this couplet refers); petals white, pink, or red, often changing from white to pink or red with age; anthers glabrous; ovary and fruit glabrous; bracts and/or bracteoles persistent to or past maturity of fruit ..................................................................................... 3 Stipules 1/2 to completely connate beyond petiole; petals white, pink, red, or yellow; anthers glabrous or hairy; ovary and fruit glabrous or hairy; bracts and bracteoles persistent or deciduous .................................... 15 Vegetative internodes, abaxial surface of stipules, petioles, and abaxial surface of leaf blades at least initially hairy, the hairs persistent or deciduous ................................................................................................... 4

Byrsonima 107

3.

Vegetative internodes, abaxial surface of stipules, and leaves quite glabrous from the beginning except for short-hirsute axils of stipules (Caution: Do not confuse axis of inflorescence with vegetative internodes) ..................................................................................................... 9 4(3). Leaf blades with lateral veins very numerous and fine, not or hardly distinguishable from parallel veinlets and reticulum; flowers borne 1 per bract ........................................................................................................ 5 4. Leaf blades with principal lateral veins easily distinguished from finer veins and reticulum, usually 7–13 pairs, sometimes more in blades over 10 cm long; flowers borne 1–3 per bract ....................................... 7 5(4). Anther locules 1.8–3.1 mm long, cylindrical and unwinged or rarely bearing very narrow longitudinal wings < 0.1 mm wide, the inner and outer thecae about the same length; pedicels straight or slightly nodding in fruit; leaf blades mostly obtuse or slightly acuminate at apex, sometimes nearly rounded ............................................................ B. coniophylla 5. Anther locules 1–1.4 mm long, dorsiventrally flattened and bearing narrow membranous longitudinal wings, the outer thecae longer than the inner; pedicels usually decurved and sometimes twisted in fruit; leaf blades rounded or broadly obtuse at apex ............................................ 6 6(5). Leaf blades narrowly elliptic, 1.5–3(–3.6) cm wide, abaxially often persistently glaucous, the margin green; sepals revolute in anthesis; anther connective exceeding locules by 0.5–0.8 mm .................... B. bronweniana 6. Leaf blades elliptic or obovate, 3–5 cm wide, not glaucous abaxially, the margin yellow; sepals appressed in anthesis to eventually revolute; anther connective exceeding locules by 0.2–0.5 mm .................... B. duidana 7(4). Leaf blades abaxially persistently velutinous, the hairs on tissue between veins erect, ± straight, basifixed, the hairs on veins denser, twisted, sub-basifixed ................................................................................. B. cuprea 7. Leaf blades abaxially glabrous or sparsely tomentose to glabrate between veins, tomentose to glabrate on principal veins, most densely so on midrib ..................................................................................................... 8 8(7). Blade of larger leaves 8.5–14.5 × 4–8 cm, the petiole 10–19 mm long; stipules 3.5–7(–8) mm long, often acuminate; reticulum usually ± concolorous with areolar tissue; inflorescence 9–18 cm long ................................................................................................ B. punctulata 8. Blade of larger leaves 5–9(–10.5) × 3–4.5(–5) cm, the petiole 5–11 mm long; stipules 1.5–2.5(–3) mm long, acute or obtuse; fine reticulum visible (in dried leaves) abaxially or, usually, on both sides as a white mesh against darker areoles; inflorescence 5–10(–12.5) cm long ............................................................................................. B. leucophlebia 9(3). Petioles slightly shorter than stipules to slightly longer, never twice as long ....................................................................................................... 10 9. Petioles at least twice as long as stipules, often longer .......................... 11 10(9). Pedicels distally thickened, 2–3 mm diameter at apex, straight in bud; anther locules 1.6–2.2 mm long, the connective not or hardly exceeding them, by up to 0.1 mm; easternmost Bolívar ........................ B. pachypoda 10. Pedicels up to 1 mm diameter in flower, 1.5 mm in fruit, circinate in bud; anther locules 1–1.3 mm long, the connective exceeding them by 0.1– 0.5 mm; westernmost Bolívar and eastern Amazonas ...... B. steyermarkii

108

M ALPIGHIACEAE

11(9). Pedicels decurved to eventually twisted in old flowers and fruits ......... 12 11. Pedicels ascending in old flowers and fruits ........................................... 13 12(11). Leaf blade with the abaxial epidermis deeply pitted, at maturity ± densely and persistently glaucous ..................................... B. luetzelburgii 12. Leaf blade abaxially nearly or quite smooth and not or hardly glaucous ........................................................................................................ B. laevis 13(11). Pedicels becoming sigmoid in maturing fruits; petioles (18–)22–33 mm long; blade of larger leaves (11–)12.5–17.5 × (5–)6.5–8.5 cm ......... B. sp. A 13. Pedicels straight or curved but not sigmoid in fruits; petioles 5–17 mm long; blade of larger leaves 3–11(–13.5) × 1.5–6.5 cm ........................ 14 14(13). Pedicels straight in bud, 2.5–3 mm long in flower, 5–7 mm long and 1.5– 2 mm diameter in fruit; anther locules cylindrical and unwinged; sepals appressed in anthesis, not or only slightly revolute in fruit; stone of fruit apparently smooth; blade of larger leaves 3–5.5 × 1.5–3.5 cm ....................................................................................................... B. baccae 14. Pedicels circinate in bud, 5–11 mm long in flower, 7–12 mm long and 0.7– 1(–1.2) mm diameter in fruit; anther locules with the outer thecae dorsiventrally flattened and bearing narrow membranous longitudinal wings; sepals revolute at apex in anthesis, reflexed and revolute in fruit; stone of fruit rugose; blade of larger leaves (4.5–)6–11(–13.5) × (2.5–)3–6.5 cm ........................................................................... B. concinna 15(2). Specimens with flowers ........................................................................... 16 15. Specimens with fruits .............................................................................. 42 16(15). Petals yellow, sometimes turning orange or red with age ...................... 17 16. Petals white, pink, or red, often changing from white to pink or red with age ........................................................................................................ 25 17(16). Gnarled shrubs up to 60 cm tall; leaves mostly in dense clusters without measurable internodes ....................................................... B. verbascifolia 17. Shrubs or trees (0.8–)1–35 m tall; internodes usually > 5 mm long ...... 18 18(17). Leaf blades abaxially thinly to densely velutinous, the hairs with a straight, erect stalk, the branches mostly ≤ stalk .............................. 19 18. Leaf blades abaxially tomentose, sericeous, or glabrate, the hairs (if any) sessile, subsessile, or with a stalk much shorter than crosspiece or branches ............................................................................................... 21 19(18). All or many hairs of abaxial surface of leaf blades stellate, i.e., with > 2 branches; stipules (8–)10–25 mm long, deciduous .......... B. stipulacea 19. All hairs of leaf blades bifurcate, i.e., Y-shaped with only 2 branches; stipules 4–8 mm long, persistent on petiole ....................................... 20 20(19). Leaf blades ± strongly revolute at margin and persistently velutinous adaxially over entire surface; sepals adaxially tomentose; ovary and fruit very densely and persistently hairy over entire surface, the hairs subappressed to suberect; bracts and bracteoles mostly persistent during maturation of fruit ...................................................... B. linguifera 20. Leaf blades flat or only slightly revolute at margin, soon glabrescent adaxially or persistently velutinous or tomentose on midrib and sometimes on lateral veins; sepals adaxially glabrous; ovary and fruit glabrous or sparsely sericeous at apex; bracts and bracteoles mostly deciduous before maturity of fruit ...................................... B. poeppigiana

Byrsonima 109

21(18). Posterior petal with 2 or more glands at apex of claw or occasionally on base of limb .......................................................................................... 22 21. Posterior petal eglandular ....................................................................... 23 22(21). Ovary glabrous or very sparsely sericeous at apex; leaf blades abaxially ± persistently subsericeous or appressed-tomentose, the hairs distinctly stalked, with slightly twisted, nonparallel crosspieces > 0.5 mm long; leaf blades with 8–12 pairs of lateral veins strongly raised abaxially, parallel and anastomosing near margin, alternating with weaker, shorter, parallel veins; shrubs or small trees 2–5(–9) m tall ............................................................................................. B. chrysophylla 22. Ovary sericeous; leaf blades abaxially appressed-sericeous to glabrate, the hairs sessile or subsessile, with short, straight, parallel crosspieces up to 0.5 mm long; leaf blades with 15–20 or more pairs of fine lateral veins parallel and anastomosing near margin, none very prominent abaxially; trees 3–25 m tall ......................................................... B. spicata 23(21). Leaf blades abaxially tomentose to glabrate, the hairs ± twisted, not parallel, not or only moderately appressed; petiole of larger leaves 5–13 (–19) mm long ......................................................................... B. crassifolia 23. Leaf blades abaxially densely to sparsely sericeous or nearly glabrate, the hairs straight, appressed, and parallel; petiole of larger leaves (15–)20– 40 mm long ........................................................................................... 24 24(23). Leaf blades abaxially densely and persistently rusty brown-sericeous, occasionally belatedly glabrescent, the hairs 0.2–0.5 mm long; Delta Amacuro and eastern Bolívar ....................................................... B. aerugo 24. Leaf blades abaxially sparsely sericeous to nearly glabrate, the hairs 0.1– 0.2 mm long, never dense enough to hide epidermis; northwestern Bolívar and Amazonas ................................................................... B. crispa 25(16). Anthers hairy ........................................................................................... 26 25. Anthers glabrous ...................................................................................... 31 26(25). Leaves sessile or subsessile, the petiole up to 2 mm long .... B. coccolobifolia 26. Leaves petiolate, the petiole of larger leaves at least 5 mm long .......... 27 27(26). Anther locules rounded or acute at apex; bracts at least 3 mm long, mostly deciduous before maturity of fruit ...................................................... 28 27. Anther locules extended at apex into slender, sterile projections; bracts up to 1.6(–2) mm long, persistent in fruit or some persistent and some deciduous in the same inflorescence ....................................................... 29 28(27). Larger leaves with petiole 5–10 mm long, densely and ± persistently tomentose, the blade (5–)7.5–15 × (3–)4–7 cm; anthers 1.8–2.8 mm long, the locules sparsely to moderately tomentose with ± spreading and often twisted hairs that do not reach bulbous apex of connective; ovary glabrous; styles 2.7–3.7 mm long ................................ B. schomburgkiana 28. Larger leaves with petiole (15–)23–48 mm long and glabrous, the blade (13–)15–23.5 × (6–)7–16 cm; anthers (2.5–)3–3.8 mm long, the locules very densely hirsute their whole length with straight, appressed, parallel hairs that often reach as high as tapered apex of connective or beyond; ovary densely sericeous on distal half; styles 4.5–6.5 mm long ............................................................................................... B. fernandezii 29(27). Blade of larger leaves 5–8 × 3–4 cm; lateral veins and reticulum barely or

110

M ALPIGHIACEAE

29.

30(29).

30.

31(25). 31. 32(31).

32.

33(32).

33.

34(33).

34.

35(31).

35. 36(35).

36.

37(36). 37.

not visible in dried leaf blades; Gran Sabana, Bolívar ................. B. dubia Blade of larger leaves 9–18.5 × 3.5–7.5(–8) cm; lateral veins and reticulum generally visible on one or both sides of dried leaf blades; basins of middle Río Orinoco west and south to upper Río Negro .................... 30 Leaves glabrous or thinly sericeous to glabrate on petiole and abaxial surface of blade, the hairs (if present) quite straight and very strongly appressed; peduncles 0–0.5 mm long; bracts 0.5–1(–1.5) mm long ................................................................................................. B. japurensis Leaves persistently appressed-tomentose or loosely subsericeous on petiole and abaxial surface of blade, or patchily glabrescent in age, the hairs sinuous to twisted, appressed to erect; peduncles 0–3 mm long in the same inflorescence, mostly 0.5–2 mm long; bracts 1.2–1.5(–2) mm long ........................................................................................... B. basiliana Ovary sericeous ........................................................................................ 32 Ovary glabrous ......................................................................................... 35 Bracts and bracteoles persistent past maturity of fruit; bracts 0.7–2(–2.5) mm long, ≤ bracteoles; flowers often borne 2(3) per bract; all 3 locules of ovary fertile .......................................................................... B. christianeae Bracts and bracteoles all or most deciduous before or during anthesis; bracts 2.5–7 mm long, usually longer than bracteoles; flowers borne 1 per bract; only 2 locules of ovary fertile ........................................... 33 Leaf blades thinly to moderately sericeous abaxially with hairs not dense enough to completely conceal epidermis, sometimes eventually deciduous ................................................................................................. B. carraoana Leaf blades very densely and persistently sericeous or subsericeous abaxially, the hairs so dense as to completely conceal epidermis, even on older leaves ..................................................................................... 34 Inflorescence 5–16 cm long; stipules 2.5–3.8 mm long; petioles 13–20 mm long; blade of larger leaves 6.5–12.5 × 3.3–7.5 cm; eastern Bolívar ............................................................................................. B. chalcophylla Inflorescence 18–26 cm long; stipules 4.5–9 mm long; petioles 20–28 mm long; blade of larger leaves 11–17 × 5.5–9 cm; Amazonas ........................................................................................... B. macrostachya Lateral petals adaxially pilose on claw, abaxially pilose on limb; stipules 6–11 mm long, completely and smoothly connate, the pair rounded at apex .............................................................................................. B. tillettii Petals glabrous; stipules 1.5–7.5 mm long, if ≥ 6 mm the pair triangular, acute at apex, sulcate in middle .......................................................... 36 Leaf blades abaxially persistently dark brown-tomentose at maturity, the hairs very strongly twisted and so dense as to completely hide epidermis ....................................................................................................... B. huberi Leaf blades abaxially glabrous or glabrate at maturity or, if persistently hairy, the hairs with nearly straight branches and never so dense as to completely hide epidermis ................................................................... 37 Anthers with apex of connective about even with locules or exceeding them by no more than 0.3 mm ............................................................ 38 Anthers with connective much enlarged at apex, exceeding locules by 0.7– 1.6 mm .................................................................................................. 39

Byrsonima 111

38(37). Leaf blades with lateral veins very numerous and fine, not or hardly distinguishable from parallel veinlets and reticulum; petioles 8–16(–23) mm long; sepals abaxially glabrous or ciliate on margin and rarely bearing scattered hairs; hairs on filaments straight; fruit superior throughout development ...................................................... B. coniophylla 38. Leaf blades with principal lateral veins easily distinguished from finer veins and reticulum, usually 5–8 pairs; petioles 2–6 mm long; sepals abaxially sericeous; hairs on filaments kinky; fruit developing halfimmersed in enlarged, disk-like receptacle ......................... B. nitidissima 39(37). Bracts and bracteoles persistent through anthesis and past maturity of fruit .......................................................................................... B. kariniana 39. Bracts and bracteoles all or mostly deciduous before or during anthesis .................................................................................................................. 40 40(39). Bracts subtending only 1 flower, very rarely 2 ............................ B. frondosa 40. Bracts (all or at least some in every inflorescence) subtending 2–4 flowers .............................................................................................................. 41 41(40). Pedicels decurved in old flowers and fruit; styles strongly bent in buds, ± straightening during anthesis; stipules 1.7–4(–5) mm long; calyx glands mostly absent or rudimentary, rarely well developed ........... B. wurdackii 41. Pedicels straight or bent upward in old flowers and fruit; styles nearly or quite straight in buds; stipules 4–7.5 mm long; calyx glands well developed ..................................................................................... B. cowanii 42(15). Bracts and/or bracteoles persistent to or past maturity of fruit ............ 43 42. Bracts and bracteoles all or most deciduous before maturity of fruit .... 58 43(42). Leaves sessile or subsessile, the petiole up to 2 mm long ...................... 44 43. Leaves petiolate, the petiole at least 4 mm long ..................................... 45 44(43). Leaf blades abaxially glabrous or very soon quite glabrate; stipules 1–2 mm long; fruits quite superior throughout development ............................................................................................ B. coccolobifolia 44. Leaf blades abaxially sericeous or velutinous to glabrescent; stipules 2–5 mm long; fruits developing half-immersed in enlarged, disk-like receptacle .............................................................................. B. nitidissima 45(43). Bracts 3–5 times as long as bracteoles, strongly reflexed or revolute .............................................................................................................. 46 45. Bracts up to twice as long as bracteoles, sometimes the same length or shorter, appressed or spreading or somewhat reflexed ...................... 48 46(45). Leaf blades densely velutinous on both sides, the hairs Y-shaped with a straight, erect stalk, the branches mostly shorter than stalk; leaf blades ± strongly revolute at margin; fruits persistently hairy over entire surface with hairs subappressed to suberect .............. B. linguifera 46. Leaf blades sericeous or glabrate, the hairs (if any) sessile or subsessile, straight, appressed; leaf blades flat at margin; fruits proximally glabrescent, distally sericeous to glabrate ............................................... 47 47(46). Leaf blades densely and persistently rusty brown-sericeous abaxially, occasionally belatedly glabrescent, the hairs 0.2–0.5 mm long; Delta Amacuro and eastern Bolívar ....................................................... B. aerugo 47. Leaf blades sparsely sericeous to nearly glabrate abaxially, the hairs 0.1–0.2 mm long, never dense enough to hide epidermis; northwestern

112

M ALPIGHIACEAE

Bolívar and Amazonas ................................................................... B. crispa 48(45). Leaf blades persistently dark brown-tomentose abaxially at maturity, the hairs very strongly twisted and so dense as to completely hide epidermis ....................................................................................... B. huberi 48. Leaf blades glabrous or glabrate abaxially at maturity or, if persistently hairy, the hairs not dense enough to completely hide epidermis ....... 49 49(48). Fruits developing half-immersed in enlarged, disk-like receptacle ............................................................................................... B. nitidissima 49. Fruits nearly to quite superior throughout development ....................... 50 50(49). Sepals membranous in fruit, the portion beyond glands elongating to form a lingulate process at least twice as long as wide ....... B. schomburgkiana 50. Sepals ± coriaceous in fruit, the portion beyond glands often somewhat accrescent but triangular, about as wide as long, often auriculate at base ....................................................................................................... 51 51(50). Pedicels straight or slightly nodding in fruit .......................................... 52 51. Pedicels strongly decurved and/or twisted in fruit ................................. 55 52(51). Dried fruits 4–6 mm diameter; leaf blades with lateral veins very numerous and fine, not or hardly distinguishable from parallel veinlets and reticulum ............................................................................... B. coniophylla 52. Dried fruits (8–)9–20 mm diameter; leaf blades with principal lateral veins ± easily distinguished, usually 7–13 pairs ................................ 53 53(52). Blade of larger leaves 6.5–10.7 cm long; pedicels 2.5–6 mm long; bracts 2.5–7 mm long; fruits probably dark green or blue at maturity; 1450– 2000 m ..................................................................................... B. kariniana 53. Blade of larger leaves mostly 9–32 cm long; pedicels 6–10(–13) mm long; bracts 0.5–2(–2.5) mm long; fruits yellow, orange, or red at maturity; 30–950 m .............................................................................................. 54 54(53). Blade of larger leaves 3.5–7.5(–8) cm wide; stipules 1.5–3(–4.5) mm long; fruits red at maturity, 10–16 × 10–12 mm (dried); flowers 1 per bract; basins of middle Río Orinoco west and south to upper Río Negro ................................................................................................. B. japurensis 54. Blade of larger leaves (6–)7–15 cm wide; stipules 3–5 mm long; fruits yellow or orange at maturity, 15–20 mm diameter (dried); flowers 1 per bract or more often a cluster of 2(3); expected in easternmost Bolívar .............................................................................................. B. christianeae 55(51). Bracts (2.5–)3–5(–7) mm long, bracteoles 2–3 mm long; stipules 6–11 mm long, generally as long as petiole or longer, occasionally slightly shorter; flowers borne 1 or 2(3?) per bract .................................. B. tillettii 55. Bracts and bracteoles 0.5–2 mm long; stipules 1.5–4.5 mm long, always much shorter than petiole, never as much as half as long; flowers borne 1 per bract. ........................................................................................... 56 56(55). Blade of larger leaves 5–8 × 3–4 cm; lateral veins and reticulum barely or not visible in dried leaf blades; Gran Sabana, Bolívar ................. B. dubia 56. Blade of larger leaves 9–18.5 × 3.5–7.5(–8) cm; lateral veins and reticulum generally visible on one or both sides of dried leaf blades; basins of middle Río Orinoco west and south to upper Río Negro .................... 57 57(56). Leaves glabrous or thinly sericeous to glabrate on petiole and abaxial sur-

Byrsonima 113

57.

58(42). 58. 59(58). 59.

60(59). 60. 61(59). 61. 62(61). 62.

63(62). 63. 64(63).

64.

65(64).

65.

66(64). 66.

face of blade, the hairs (if present) quite straight and very strongly appressed; peduncles 0–0.5 mm long; bracts 0.5–1(–1.5) mm long ................................................................................................. B. japurensis Leaves persistently appressed-tomentose or loosely subsericeous on petiole and abaxial surface of blade, or patchily glabrescent in age, the hairs sinuous to twisted, appressed to erect; peduncles 0–3 mm long in the same inflorescence, mostly 0.5–2 mm long; bracts 1.2–1.5(–2) mm long ........................................................................................... B. basiliana Gnarled shrubs up to 60 cm tall; leaves mostly in dense clusters without measurable internodes ....................................................... B. verbascifolia Shrubs or trees 1–35 m tall; internodes mostly > 5 mm long, if shorter the plant at least 2 m tall .......................................................................... 59 Leaf blades thinly to densely velutinous abaxially, the hairs with a straight, erect stalk, the branches mostly ≤ stalk .............................. 60 Leaf blades tomentose, sericeous, glabrate, or glabrous abaxially, the hairs if present sessile, subsessile, or with a stalk much shorter than branches ............................................................................................... 61 All or many hairs of abaxial surface of leaf blades stellate, i.e., with > 2 branches; stipules (8–)10–25 mm long, deciduous .......... B. stipulacea All hairs of leaf blades bifurcate, i.e., Y-shaped with only 2 branches; stipules 4–8 mm long, persistent on petiole ....................... B. poeppigiana Leaves sessile or subsessile, the petiole up to 2 mm long; leaf blades usually rounded or cordate at base ................................... B. coccolobifolia Leaves petiolate, the petiole at least 5 mm long; leaf blades tapered, cuneate, or rounded at base .................................................................... 62 Sepals membranous in fruit, the portion beyond glands elongating to form a lingulate process at least twice as long as wide ....... B. schomburgkiana Sepals ± coriaceous in fruit, the portion beyond glands often somewhat accrescent but triangular, about as wide as long, often auriculate at base ....................................................................................................... 63 Pedicels nearly or quite straight, or ascending, in fruit ......................... 64 Pedicels strongly decurved or twisted in fruit ........................................ 69 Leaf blades abaxially nearly or quite glabrate as soon as expanded, with at most some appressed hairs persistent on midrib; fruit quite glabrous .............................................................................................................. 65 Leaf blades abaxially thinly to densely sericeous or subsericeous, the hairs persistent or eventually deciduous; fruit derived from a densely sericeous ovary and often bearing some hairs at maturity, especially at apex ...................................................................................................... 66 Bracts subtending only 1 flower, very rarely 2; inflorescence 6–15 cm long; stipules 1.5–2.5 mm long; blade of larger leaves 8–15 × 3–7 cm ................................................................................................... B. frondosa Bracts (all or at least some in every inflorescence) subtending 2–4 flowers; inflorescence 15–35 cm long; stipules 4–7.5 mm long; blade of larger leaves 15–25 × 7–13 cm .............................................................. B. cowanii Stipules completely and smoothly connate ...................................... B. aerugo Stipules mostly incompletely connate, the pair sulcate and shallowly to

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deeply bidentate at apex ...................................................................... 67 67(66). Leaf blades thinly to moderately sericeous abaxially with hairs not dense enough to completely conceal epidermis, sometimes eventually deciduous ................................................................................. B. carraoana 67. Leaf blades very densely and persistently sericeous or subsericeous abaxially, the hairs so dense as to completely conceal epidermis, even on older leaves ..................................................................................... 68 68(67). Inflorescence 5–16 cm long; stipules 2.5–3.8 mm long; petioles 13–20 mm long; blade of larger leaves 6.5–12.5 × 3.3–7.5 cm; eastern Bolívar ............................................................................................. B. chalcophylla 68. Inflorescence 18–26 cm long; stipules 4.5–9 mm long; petioles 20–28 mm long; blade of larger leaves 11–17 × 5.5–9 cm; Amazonas ... B. macrostachya 69(63). Leaf blades with 15–20 or more pairs of fine lateral veins, none very prominent ..................................................................................... B. spicata 69. Leaf blades with 5–12 pairs of principal lateral veins ............................ 70 70(69). Leaf blades glabrous from the beginning or very soon nearly glabrate; petiole of larger leaves 15–48 mm long ............................................... 71 70. Leaf blades subsericeous to densely tomentose with hairs persistent or eventually mostly deciduous; petiole of larger leaves 5–15 mm long .............................................................................................................. 72 71(70). Fruits (dried) 12–17 × 10–16 mm, persistently sericeous at apex; calyx glands well developed ........................................................... B. fernandezii 71. Fruits (dried) 6–7 mm diameter, quite glabrous; calyx glands mostly absent or rudimentary, rarely well developed ....................... B. wurdackii 72(70). Blade of larger leaves 5–6.8 cm long, broadly obtuse to rounded at apex, adaxially rugose, abaxially persistently tomentose with hairs so dense as to completely hide epidermis, even on older leaves. ............... B. huberi 72. Blade of larger leaves 6.5–17(–19) cm long, usually acute or acuminate at apex, sometimes narrowly obtuse, adaxially smooth or with veins slightly raised, abaxially tomentose or subsericeous to glabrescent, the hairs seldom so dense as to completely hide epidermis, especially on older leaves. ......................................................................................... 73 73(72). Fruits red at maturity, ovoid with a prominent apical beak ...... B. basiliana 73. Fruits yellow at maturity, somewhat ovoid when young but nearly globose at maturity ........................................................................................... 74 74(73). Sepals loosely sericeous adaxially; leaf blades elliptic, 2–3(–3.4) times as long as wide, persistently loosely subsericeous or appressed-tomentose abaxially, only rarely glabrescent in age ............................ B. chrysophylla 74. Sepals usually glabrous adaxially; leaf blades mostly broadly elliptic to subrotund, 1.4–2 times as long as wide, mostly glabrescent abaxially in age, often patchily glabrescent, occasionally persistently tomentose ................................................................................................. B. crassifolia Byrsonima aerugo Sagot, Ann. Sci. Nat. Bot. sér. 6, 12: 178. 1881. Byrsonima ferruginea var. macrophylla Benth., London J. Bot. 7: 119. 1848. Tree (4–)15–45 m tall; stipules 2.5–4.5 mm long, completely connate; petioles (15–)20–40 mm long; blade of larger leaves

11–22(–25) × 4–9(–11) cm, densely and ± persistently rusty brown-sericeous abaxially, occasionally belatedly glabrescent; bracts and bracteoles persistent or deciduous in fruit; bracts 3.5–5 mm long, strongly reflexed or revolute; bracteoles 0.6–1.5 mm long; pedicels nearly or quite straight, or ascending,

Byrsonima 115

in fruit; petals yellow, the posterior petal eglandular; anthers sericeous, especially between locules, the connective equaling locules or exceeding them by up to 0.6 mm; ovary densely sericeous; fruit yellow, 10–13 mm diameter (dried), sericeous to glabrate. Evergreen lowland to lower montane forests, 200–1300 m; Delta Amacuro (Caño Acoima, east-northeast of El Palmar, east-southeast of Imataca), widespread in eastern Bolívar to 63°W. Guyana, Suriname, French Guiana, Brazil (Roraima). ◆Fig. 85. This species is close to Byrsonima crispa A. Juss., and when its leaf hairs are eventually deciduous instead of persistent (e.g., Cardona 2150 [MICH, MO, VEN] and 2164 [MICH, MO, US, VEN]) the distinction becomes difficult. Nevertheless, B. aerugo is a useful taxon and I am not yet ready to abandon it, especially because the two species seldom if ever occur together. Byrsonima baccae W.R. Anderson, Contr. Univ. Michigan Herb. 20: 19. 1995. Shrub or small tree 1–6 m tall; vegetative internodes glabrous except in axil of stipules; stipules 1.5–2.5 mm long, distinct; petioles 5–8 mm long; blade of larger leaves 3–5.5 × 1.5–3.5 cm, glabrous, rounded or broadly obtuse at apex; bracts and bracteoles persistent in fruit, 1.5–3.2 mm long, stiff; pedicels straight in bud, 2.5–3 mm long in flower, straight in fruit, becoming 5–7 mm long, 1.5– 2 mm diameter; sepals appressed in anthesis, not or only slightly revolute in fruit; petals pink; anther locules glabrous, cylindrical, the connective exceeding locules by 0.3–0.6 mm; ovary and fruit glabrous; fruit 6–9 mm diameter (dried), the stone apparently smooth. On rocky slopes, in elfin forests, along streams, 1700–2200 m; Amazonas (Sierra Maigualida). Endemic. Byrsonima basiliana W.R. Anderson, Contr. Univ. Michigan Herb. 21: 45. 1997. Shrub or small tree; stipules 1.5–2.5 mm long, connate, rounded; petioles 8–12 mm long; blade of larger leaves 9.5–13.5 × 3.5– 5.7 cm, abaxially appressed-tomentose or loosely subsericeous to patchily glabrescent; bracts and bracteoles persistent or deciduous in fruit; bracts 1.2–1.5(–2) mm long; peduncles (0–)0.5–2(–3) mm long; bracteoles 0.7–1 mm long; pedicels decurved or twisted

in fruit; petals white and pink; anther locules sericeous, drawn out at apex into slender sterile extensions 0.5–1 mm long, the connective exceeding fertile part of locules by 0.5 mm; ovary glabrous or tomentose; fruit red, 8–9.5 × 6.5 mm (dried), beaked at apex, glabrous or tomentose to glabrate. Evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, Río Emoni). Endemic. Byrsonima bronweniana W.R. Anderson, Mem. New York Bot. Gard. 32: 107. 1981. Shrub or tree 2–8 m tall; vegetative internodes sparsely sericeous to glabrate; stipules 1–2 mm long, distinct; petioles 6–13 mm long; blade of larger leaves 5–9.5 × 1.5–3 (–3.6) cm, narrowly elliptic, rounded or obtuse at apex, sparsely sericeous to glabrate, abaxially often persistently glaucous, the lateral veins numerous and fine; bracts and bracteoles persistent in fruit; pedicels decurved or twisted in fruit; sepals revolute in anthesis; petals white turning pink; anthers glabrous, the locules 1–1.3 mm long, dorsiventrally flattened and narrowly winged, the connective exceeding locules by 0.5–0.8 mm, globose; ovary and fruit glabrous; fruits red or purplish, 5.5–7 mm diameter (dried). Along rivers, 50–200 m; Amazonas (basins of upper Río Orinoco and upper Río Negro). Colombia (Guainía), Brazil (Amazonas: upper Rio Negro and Rio Uaupés). ◆Fig. 82. Byrsonima carraoana Steyerm., Fieldiana, Bot. 28: 287. 1952. —Byrsonima chalcophylla var. carraoana (Steyerm.) W.R. Anderson, Mem. New York Bot. Gard. 32: 117. 1981. Byrsonima bolivarana Steyerm., Fieldiana, Bot. 28: 287. 1952. Shrub or tree (1.5–)3–12 m tall; stipules (2–)3–5 mm long, 2/3- to almost completely connate; petioles 7–18(–22) mm long; blade of larger leaves 5–12(–18.5) × 3–7(–10) cm, abaxially sericeous, sometimes eventually glabrescent, with hairs not dense enough to completely conceal epidermis; inflorescence 5–15 cm long; bracts and bracteoles deciduous before or during anthesis, 3–7 mm long or bracteoles shorter; pedicels nearly or quite straight in fruit; petals pink or pink in center and otherwise white; anther locules glabrous, cylindrical, the connective exceeding

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locules by 0.8–1.7 mm; ovary densely sericeous; fruits 8–10 mm long (dried), sericeous to glabrate. Montane, often dwarf forests, 1200–2000 m; eastern Bolívar (Auyán-tepui and Sierra de Lema south and east to Roraima). Guyana (Pakaraima Mountains, upper Mazaruni River basin). ◆Fig. 81. Since I treated this as a variety of Byrsonima chalcophylla, in 1981, accumulating collections have eroded the morphological differences between B. carraoana and B. macrostachya, and between B. macrostachya and B. chalcophylla. I now feel that B. carraoana, B. chalcophylla, and B. macrostachya should all have the same taxonomic status, and have elected to consider them a complex of three closely related species. One could also make the argument for treating them as three varieties of one species. B. carraoana is far better collected than the other two. Byrsonima chalcophylla Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 5: 57. 1914. Tree 3–12 m tall; stipules 2.6–3.8 mm long, 1/2–3/4-connate; petioles 13–20 mm long; blade of larger leaves 6.5–12.5 × 3.3– 7.5 cm, abaxially very densely and persistently sericeous; inflorescence 5–16 cm long; bracts and bracteoles deciduous before or during anthesis, 3–7 mm long or bracteoles shorter; pedicels nearly or quite straight in fruit; petals pink; anther locules glabrous, cylindrical, the connective exceeding locules by 0.6–1 mm; ovary densely sericeous; immature fruits 13 mm long (dried), sericeous to glabrescent. Tepui slope forests, ca. 1900 m; eastern Bolívar (Ilú-tepui, Roraima-tepui). Guyana. Byrsonima chalcophylla is known from only three collections, the two in the flora area and one from ca. 1700 m on the slopes of Mount Roraima in Guyana. See comments under Byrsonima carraoana. Byrsonima christianeae W.R. Anderson, Mem. New York Bot. Gard. 32: 112. 1981. Tree (8–)12–25 m tall; stipules 3–5 mm long, completely connate or distinct at apex; petioles 20–35(–45) mm long; blade of larger leaves (16–)18–30 × (6–)7–15 cm, initially sparsely sericeous, soon glabrate; bracts and

bracteoles persistent in fruit, 0.7–2.5 mm long; pedicels straight in fruit; petals white turning red in age; anther locules glabrous, unwinged, the connective exceeding locules by 1.2–2.9 mm; ovary sericeous; fruits yellow or orange, 15–20 mm diameter (dried), sparsely sericeous or glabrate. Expected in eastern Bolívar. Byrsonima christianeae is known presently only from mixed evergreen forests, 70– 950 m, from Guyana (Mt. Ayanganna in the upper Mazaruni River basin) and Brazil (Amapá, Pará). Byrsonima chrysophylla H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 150. 1821 [1822]. —Manteco. Shrub or small tree 2–5(–9) m tall; stipules 2–3 mm long, completely and smoothly connate; petioles 5–15 mm long; blade of larger leaves 7–17 × 4–7(–9) cm, ± persistently subsericeous or appressed-tomentose abaxially, the hairs distinctly stalked, with slightly twisted, nonparallel crosspieces > 0.5 mm long, with 8–12 pairs of principal lateral veins strongly raised abaxially; bracts and bracteoles mostly deciduous before maturity of fruit; pedicels mostly decurved or twisted in fruit; sepals loosely sericeous adaxially; petals yellow, the posterior petal bearing 2 glands at apex of claw; anthers sericeous, especially between locules, the connective equaling or slightly exceeding locules; ovary glabrous or very sparsely sericeous at apex; fruits yellow, globose, 6–10 mm diameter (dried), glabrous. In or at the edges of savannas, or in woodlands along rivers, 100–200(–900) m; scattered in northwestern and southwestern Bolívar, common in Amazonas (basins of upper Río Orinoco and the Río Negro). Amazonian Colombia, Suriname, Peru, Brazil, Bolivia. This species is close to Byrsonima spicata, and the placement of some collections in one species or the other can be rather arbitrary. Examples are Bunting et al. 3476 (MY) and Gentry & Berry 14694 (MICH, MO), both collected in the vicinity of Puerto Ayacucho. Byrsonima coccolobifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 148. 1821 [1822]. —Chaparro, Manteco merey. Byrsonima sessilifolia Benth., London J. Bot. 7: 124. 1848.

Byrsonima 117

Shrub or small tree 1–5 m tall; stipules 1– 2 mm long, connate; petioles 0–2 mm long; blade of larger leaves 6–15.5 × 4–10 cm, usually rounded or cordate at base, glabrous or very soon quite glabrate; bracts and bracteoles mostly deciduous in fruit or rarely many persistent, 1.5–3.5(–5) mm long or bracteoles smaller; pedicels decurved or twisted in fruit; sepals lingulate-accrescent and membranous in fruit; petals white and/ or pink; anther locules sericeous, the connective exceeding locules by (0–)0.2–1.3 mm; ovary and fruit glabrous; fruits green (yellow at maturity?), 6–8 mm diameter (dried). Savannas, 50–400 m; Delta Amacuro (Los Castillos), common in middle and northern Bolívar. Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Falcón, Guárico, Monagas, Sucre, Zulia; Cuba, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 80. In the southwestern extreme of its range (Bolivia, Mato Grosso, Paraguay) this species may reach heights of 10 m, but in Venezuela and Guyana it seldom if ever exceeds 5 m.

Yutajé, and Sierra de la Neblina). Western Guyana, Brazil (Roraima, on the border with Venezuela). In Bolívar and adjacent Guyana this species is variable but reasonably homogeneous. However, in Amazonas most populations are somewhat different, with relatively large leaves, narrow anther wings, and short fruiting pedicels. One collection (Fernández et al. 6041, MICH, MO) has the pedicels apparently straight in bud. The complex containing Byrsonima concinna probably deserves re-evaluation in Amazonas. A related species that may eventually be found in eastern Bolívar is Byrsonima rubrobracteata W.R. Anderson, Mem. New York Bot. Gard. 32: 105. 1981. It has been collected three times on the Ayanganna Plateau of western Guyana; other species from that area have been found in adjacent Venezuela. Byrsonima rubrobracteata is distinguished by its short dense inflorescence up to 6 cm long, with red bracts and bracteoles and only one flower per bract; its stipules are 3.5–5 mm long.

Byrsonima concinna Benth., London J. Bot. 7: 122. 1848. Byrsonima bracteolaris Benth., London J. Bot. 7: 123. 1848. Shrub or small tree 1.2–8(–15) m tall; vegetative internodes glabrous except in axil of stipules; stipules 1.5–2.5(–3.5) mm long, distinct; petioles 7–17 mm long; blade of larger leaves (4.5–)6–11(–13.5) × (2.5–)3–6.5 cm, glabrous, acute to rounded and often apiculate at apex; bracts and bracteoles persistent in fruit, (1–)2–5 mm long, usually spreading and often revolute; pedicels circinate in bud, 5–11 mm long in flower, straight or curved upward in fruit, becoming 7–12 mm long, 0.7–1(–1.2) mm diameter; sepals revolute at apex in anthesis, reflexed and revolute in fruit; petals pink(?) or white turning pink; anther locules glabrous, dorsiventrally flattened and bearing narrow membranous longitudinal wings, the connective exceeding locules by (0.2–)0.4–0.8 mm; ovary and fruit glabrous; fruits 4–8 mm diameter (dried), the stone rugose. Savannas, slope forests, (100–)400–2100 m; Bolívar (common in Gran Sabana and on tepuis as far west as Cerro Camarón), Amazonas (isolated populations on Cerro Marahuaka, Cerro Sipapo, Cerro

Byrsonima coniophylla A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 335. 1840. Shrub or small tree 1–8 m tall, occasionally shorter; vegetative internodes hairy to glabrescent; stipules 2.5–4(–5) mm long, distinct to nearly connate; petioles 8–16(–23) mm long; blade of larger leaves 7–12(–15.5) × 2.5–4.7(–5.2) cm, mostly obtuse, broadly obtuse, or slightly acuminate at apex, sometimes rounded, thinly sericeous to glabrate, often persistently glaucous abaxially, the lateral veins numerous and fine; bracts and bracteoles persistent in fruit, 2.5–4 mm long or bracteoles smaller; pedicels straight or slightly nodding in fruit; petals white turning red in age; anther locules 1.8–3.1 mm long, glabrous, cylindrical and unwinged or rarely bearing very narrow longitudinal wings < 0.1 mm wide, the connective equaling locules or exceeding them by up to 0.3 mm; ovary and fruit glabrous; fruits red, purplish, or black, 4–6 mm diameter (dried). Sandy seasonally flooded savannas, 100–200 m; common in Amazonas (upper Río Orinoco and Río Atabapo south to Río Casiquiare, Río Guainía, and Río Pasimoni). Colombia (Guainía, Vaupés), Brazil (northern Amazonas, southwestern Roraima).

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As I noted in my 1981 paper, plants from near the Río Atabapo sometimes have atypically small stature and leaves, which causes them to resemble their eastern relative Byrsonima eugeniifolia Sandwith. That species grows in sandy savannas of Guyana, Suriname, and Roraima and adjacent Amazonas, Brazil, including the tepui called Serra Aracá, but seems not to reach Venezuela. Byrsonima cowanii W.R. Anderson, Mem. New York Bot. Gard. 32: 123. 1981. Tree (2–)4–12 m tall; stipules 4–7.5 mm long, 1/2–4/5 connate, acute; petioles 11–21 (–25) mm long; blade of larger leaves 15–25 × 7–13 cm, sparsely sericeous to soon glabrate; inflorescence 15–35 cm long, the cincinni mostly 2–4-flowered; bracts and bracteoles deciduous during anthesis; bracts 3–8 mm long; bracteoles 1–3.5 mm long; pedicels straight or bent upward in fruit; calyx glands well developed; petals white and/or pink (white turning pink?); anther locules glabrous, unwinged, the connective exceeding locules by 0.8–1.5 mm; ovary and fruit glabrous; styles nearly or quite straight in bud; fruits dark blue, 7–10 mm diameter (dried). Forests, woodlands, often on granitic outcrops, 100–600 m; Amazonas (lowlands of the upper Río Orinoco and Río Atabapo from Río Cuao south to 3°12'N). Endemic? ◆Fig. 84. In recent years, specimens resembling this species have been collected in Roraima, Brazil, and Loreto, Peru, but they are so incomplete that accurate identification is not possible. Byrsonima crassifolia (L.) H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 149. 1821 [1822]. —Malpighia crassifolia L., Sp. Pl. 426. 1753. —Chaparro, Chaparro manteco. Byrsonima ferruginea H.B.K., Nov. Gen. Sp. 5: pl. 446. 1821. Byrsonima laurifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 150. 1821 [1822]. Shrub or small tree (0.8–)1–8(–10) m tall; stipules 2–3(–4) mm long, completely and smoothly connate; petioles 5–13(–19) mm long; blade of larger leaves 6.5–11(–15.5) × 3–6.5(–8) cm, broadly elliptic to subrotund, densely tomentose to glabrate on both sides; bracts and bracteoles deciduous in fruit; pedicels decurved in fruit; sepals adaxially glabrous or rarely sparsely tomentose; petals

yellow, the posterior petal eglandular; anthers pilose with few to many spreading hairs, the connective equaling locules or exceeding them by up to 0.2(–0.4) mm; ovary glabrous or sparsely to densely tomentosesericeous; fruits yellow, 8–10 mm diameter (dried), glabrous or sparsely tomentose to glabrate. Savannas, 50–1300 m; Delta Amacuro (near Los Castillos), throughout Bolívar and Amazonas. Common in savannas across northern Venezuela; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Byrsonima crassifolia is exceedingly variable through its range and may eventually yield defensible segregates, but within our area it is reasonably homogeneous. In my 1981 paper I recognized B. laurifolia, but subsequent study of the type has suggested that it is simply atypical material of this species. See discussion of probable hybrids under B. verbascifolia. Byrsonima crispa A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 335. 1840. —Manteco de agua. Byrsonima carmeniana Cuatrec., Webbia 13: 615. 1958. Tree (2.5–)6–30 m tall; stipules 3–4.5 mm long, completely connate; petioles 12–35(– 40) mm long; blade of larger leaves (8–)11– 20(–22) × (3–)4.5–8(–9) cm, sparsely sericeous to nearly glabrate abaxially; bracts (2.5–)3–4.5(–6) mm long, strongly reflexed or revolute, persistent or deciduous in fruit; bracteoles 0.5–1(–1.5) mm long, mostly persistent in fruit; pedicels mostly straight and ascending in fruit; petals yellow, the posterior petal eglandular; anthers sericeous, especially between locules, the connective equaling locules or exceeding them by up to 0.6 mm; ovary densely sericeous; fruits yellow, 10–13 mm diameter (dried), sericeous to glabrate. Nonflooded forests, 50–600 m; northwestern Bolívar (southeast of Pijiguaos, Túriba), Amazonas (Isla Ratón, Puerto Ayacucho, Río Cuao, Río Mawarinuma, San Antonio del Sipapo, Sierra Parima). Scattered in Costa Rica, Panama, Colombia, and Amazonian Peru, and common in Amazonian Brazil from Manaus east to Belém and south to Bolivia and Espírito Santo.

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Byrsonima cuprea Griseb. in Mart., Fl. Bras. 12(1): 19. 1858. Shrub or tree (2–)3–8(–16) m tall; vegetative internodes velutinous; stipules 5–9 mm long, distinct, acuminate; petioles 9–20 mm long; blade of larger leaves 8–16 × 4–8 cm, abaxially persistently velutinous; bracts and bracteoles persistent in fruit, 1.5–2.5(–3.5) mm long or bracteoles smaller; pedicels decurved in fruit; petals white turning pink or red in age; anther locules glabrous, linear, the connective exceeding locules by 0.5–0.9 mm; ovary and fruit glabrous; fruits red, 5–6 mm diameter (dried). Riparian forests, 100– 200 m; Amazonas (Río Atabapo, Río Atacavi, Río Casiquiare, Río Cuao, Río Negro, Río Orinoco, Río Pasimoni, Río Sipapo, Río Ventuari, Río Yatua). Brazil (Amazonas: upper Rio Negro). See discussion under Byrsonima punctulata. Byrsonima dubia W.R. Anderson, Contr. Univ. Michigan Herb. 17: 42. 1990. Shrub or small tree 3–4 m tall; stipules 2– 3 mm long, 3/4–5/6 connate, the apical lobes obtuse to rounded; petioles 15–18 mm long; blade of larger leaves 5–8 × 3–4 cm, obovate to nearly elliptic, obtuse to rounded and apiculate at apex, thinly sericeous to glabrate, the lateral veins and reticulum obscure; bracts and bracteoles persistent in fruit, 1– 1.6 mm long; pedicels decurved and/or twisted in fruit; petals probably pink or white; anther locules appressed-hirsute, with slender sterile extensions at apex, the connective exceeding fertile part of locules by 0.8–1 mm; ovary glabrous; fruits unknown. Thorn scrub in dry areas, 1400–1500 m; Bolívar (northwest of Cerro El Sol). Guyana (upper Potaro River). Byrsonima dubia is known from only two collections, and although these lack petals, it is keyed with some confidence as having pink or white petals because it is apparently related to B. gymnocalycina A. Juss., of Guyana, and B. laevigata (Poir. in Lam.) DC., a tall tree of lowland forests in Suriname, French Guiana, and northeastern Brazil (Amapá, Bahia, Maranhão, and Pará). Byrsonima duidana W.R. Anderson, Contr. Univ. Michigan Herb. 19: 361. 1993. Shrub or small tree 2–4 m tall; vegetative

internodes initially sericeous, soon or eventually glabrate; stipules 2–3 mm long, distinct; petioles 8–15 mm long; blade of larger leaves 5.5–10 × 3–5 cm, elliptic or obovate, rounded or obtuse and sometimes apiculate or retuse at apex, initially sparsely sericeous but soon glabrate, the lateral veins numerous and fine, the margin yellow; bracts and bracteoles persistent in fruit; pedicels decurved or twisted in fruit; sepals appressed in anthesis to eventually revolute, accrescent and reddish in fruit; petals white turning pink; anthers glabrous, the locules 1.1–1.3 mm long, dorsiventrally flattened and narrowly winged, the connective exceeding locules by 0.2–0.5 mm, globose; ovary and fruit glabrous; fruits 4.5–7 mm diameter (dried). Tepui scrub, edges of savannas, along valley forests, 50–2300 m; Amazonas (hills south and southeast of Cerro Camani, Cerro Duida, Cerro Marahuaka, Cerro Sipapo). Endemic. ◆Fig. 79. This is the species that I treated as Byrsonima bracteolaris Benth. in my 1981 paper on the Malpighiaceae of the Guayana Highland. That name has fallen into synonymy under B. concinna. Byrsonima duidana belongs to a difficult complex that contains B. laevis and B. luetzelburgii, and the taxonomy of that group is not entirely satisfactory; see comments under B. laevis. Byrsonima fernandezii Cuatrec., Webbia 13: 612. 1958. Tree (5–)10–25 m tall; stipules 1.5–3(–4) mm long, completely connate, obtuse or rounded at apex; petioles (15–)23–48 mm long; blade of larger leaves (13–)15–23.5 × (6–)7–16 cm, rounded and often apiculate at apex, glabrous; bracts and bracteoles deciduous before or during anthesis, or at the latest before maturation of fruit; bracts 3–5.5 mm long; bracteoles 1.5–2 mm long; pedicels decurved or twisted in fruit; petals white, perhaps turning pink in age(?); stamens densely hirsute; anthers (2.5–)3–3.8 mm long, the locules rounded at apex, the connective tapered, exceeding locules by 0.9–1.7 mm; ovary densely sericeous on distal half; styles 4.5–6.5 mm long; fruits 12–17 × 10–16 mm (dried), sericeous at apex. Lowland to lower montane forests, 100–800 m; Amazonas (Cerro Huachamacari, basin of upper Río Cuao, southeast of San Fernando de

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Atabapo). Scattered in Colombia (Vaupés), Brazil (Amazonas: north of Manaus, vicinity of Humaitá, São Paulo de Olivença). The type was said to have the flowers “moradas,” i.e., purple, a color otherwise unknown in the genus and all but unknown in the family. More recent collections state that the petals are white, as I would have expected, given the other characteristics of the species. Byrsonima frondosa A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 334. 1840. Tree 4–20 m tall; stipules 1.5–2.5 mm long, completely connate or the pair bidentate, triangular, acute or obtuse at apex; petioles 8–15(–18) mm long; blade of larger leaves 8–15 × 3–7 cm, initially sericeous but glabrate at maturity; inflorescence 6–15 cm long, the flowers borne 1(2) per bract; bracts and bracteoles deciduous before or during anthesis; bracts 3.5–5 mm long; bracteoles 1.5–2.5 mm long; pedicels straight in old flowers and fruit; calyx glands well developed; petals pink; anther locules glabrous, unwinged, the massive connective exceeding locules by 0.7–1.2 mm; ovary and fruit glabrous; styles nearly or quite straight in bud; mature fruits unknown. Wet forests along lowland rivers, ca. 100 m; Amazonas (Río Baría, Río Yatúa). Brazil (Amazonas: Rio Negro, Rio Uaupés). Byrsonima huberi W.R. Anderson, Contr. Univ. Michigan Herb. 21: 48. 1997. Tree 3 m tall; stems densely tomentose; stipules 3–4 mm long, completely and smoothly connate; petioles 8–12 mm long; blade of larger leaves 5–6.8 × 2.6–3.8 cm, broadly obtuse to rounded at apex, deeply rugose adaxially, abaxially very densely and persistently dark brown-tomentose; bracts and bracteoles persistent in fruit (? at least at nodes with maturing fruits); bracts 2.5– 3.5 mm long, bracteoles similar but smaller; petals probably white or pink; anther locules glabrous, cylindrical, the connective not or hardly exceeding locules; ovary and fruit glabrous; fruits orange, 10 mm diameter (dried). Upper montane shrubby woodland/meadow ecotone, ca. 2000 m; Amazonas (Sierra Maigualida). Endemic. Byrsonima huberi is known only from the type collection, which bears only extremely

immature flower buds and one detached fruit, so many of the character-states needed for placing this species in a key are not known with confidence, but the leaves are unlike those of any other species in the Guayana area. Byrsonima japurensis A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 335. 1840. Byrsonima inundata Benth., London J. Bot. 7: 122. 1848. Byrsonima uvulifera Spruce ex Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 1: 37. 1901. Byrsonima fluminensis Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 1: 43. 1901. —Byrsonima japurensis subsp. fluminensis (Nied.) Cuatrec., Webbia 13: 627. 1958. Byrsonima japurensis subsp. silvatica Cuatrec., Webbia 13: 626. 1958. Shrub or tree (2–)3–21 m tall; stipules 1.5–3(–4.5) mm long, connate, the pair acute to rounded; petioles (8–)12–20(–25) mm long; blade of larger leaves 9–18.5 × 3.5– 7.5(–8) cm, abaxially sparsely sericeous to usually glabrate; bracts and bracteoles persistent in fruit (at least the bracts), 0.5–1 (–1.5) mm long; pedicels straight or decurved/twisted in fruit; petals white or pink; anther locules sparsely to densely sericeous, drawn out at apex into slender, sterile extensions 0.3–1.2 mm long, the connective exceeding fertile part of locules by 0.3–1.2 mm; ovary glabrous or sericeous; fruits red, 10–16 × 9–12 mm (dried), often beaked at apex, glabrous or glabrate. Riparian forests, 50–300 m; Bolívar (basins of Río Caura, Río Maniapure, Río Paragua, Río Parguaza, Río Orinoco, and Río Suapure), Amazonas (basins of Río Asisa, Río Casiquiare, Río Mavaca, Río Orinoco, Río Pamoni, Río Pasiba, Río Ventuari, and Río Siapa). Anzoátegui, Apure, Guárico; Amazonian Colombia, Ecuador, Peru, and Brazil. Byrsonima kariniana W.R. Anderson, Mem. New York Bot. Gard. 32: 127. 1981. Shrub to 4 m tall; stipules 3.4–5 mm long, 1 /2–4/5 connate, acute; petioles 12–19 mm long; blade of larger leaves 6.5–10.7 × 3–5.5 cm, glabrous (initially sericeous but very soon quite glabrate); inflorescence 6–13 cm

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long; bracts and bracteoles persistent past maturity of fruit; bracts 2.5–7 mm long; bracteoles 2–3.5 mm long; pedicels 2.5–6 mm long, straight (or slightly decurved?) in fruit; petals pink (white turning pink?); anther locules glabrous, unwinged, the connective exceeding locules by 0.7–1(–1.5) mm; ovary and fruit glabrous; fruits blue(?) or dark green(?), ca. 10 mm diameter (dried). Tepui slope forests and savannas, 1400–2000 m; Amazonas (Cerro Sipapo). Apparently endemic. Related species occur at high elevations in northern Venezuela and in Trinidad, but none of those populations seems to be conspecific with the plant of Cerro Sipapo. Byrsonima laevis Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 1: 34. 1901. Shrub or tree 1.5–12 m tall; vegetative internodes glabrous except in axil of stipules; stipules 1–2 mm long, distinct; petioles (3–) 5–22 mm long; blade of larger leaves 6– 10.5(–13) × 3–7 cm, glabrous; bracts and bracteoles persistent in fruit, 0.5–1.5 mm long; pedicels decurved or eventually twisted in old flowers and fruit; petals white turning pink; anther locules glabrous, dorsiventrally flattened and longitudinally winged, the connective exceeding locules by 0.1–0.3 mm; ovary and fruit glabrous; fruits 4–12 mm diameter (dried). Forests and savannas on white sand, 100–1500 m; southern Amazonas (Cerro Aracamuni, Maroa/Yavita, Río Atacavi, Río Guainía/Casiquiare, Río Manapiare, Río Mawarinuma, San Fernando de Atabapo, Sierra de la Neblina). Brazil (northwestern Amazonas). The collections included here are a diverse lot, certainly closely related but perhaps not all one species. In my 1981 paper I recognized both Byrsonima laevis and B. amoena Cuatrec. as being in Amazonas, Venezuela. They were distinguished on the basis of size of leaves and fruits. Subsequent collections have eroded that distinction, and I can no longer justify recognizing both species in our area. I am using the older name, which happens to have been based on a type from near the mouth of Río Casiquiare. It remains to be seen whether B. amoena will stand as distinct elsewhere in western Amazonia.

Byrsonima leucophlebia Griseb. in Mart., Fl. Bras. 12(1): 15. 1858. Shrub or small tree 2–5(–8) m tall; vegetative internodes hairy to glabrescent; stipules 1.5–2.5(–3) mm long, distinct, acute or obtuse; petioles 5–11 mm long; blade of larger leaves 5–9(–10.5) × 3–4.5(–5) cm, sparsely tomentose to glabrate except midrib, the reticulum white against dark areoles; inflorescence 5–10(–12.5) cm long; bracts and bracteoles persistent in fruit, 1–2 mm long or bracteoles smaller; pedicels usually decurved in fruit; petals white to pink or red in age; anther locules glabrous, linear, the connective exceeding locules by 0.3–0.7 mm; ovary and fruit glabrous; fruits red to black, 4–5 mm diameter (dried). Shrubby associations on sand, often near rivers, 300–500 m; Bolívar (Río Acanán, Canaima, Río Carrao, Salto Angel). Widespread in Amazonian Brazil (eastern Pará and Maranhão west and south into Amazonas and Rondônia) and Bolivia (Beni). See discussion under Byrsonima punctulata. Byrsonima linguifera Cuatrec., Webbia 13: 613. 1958. Shrub or small tree 1.5–5(–6) m tall; stipules 4–7 mm long, smoothly and completely connate, the pair rounded at apex; petioles 4–10 mm long; blade of larger leaves 9.5–18 × 5–10 cm, obtuse or rounded at apex, ± revolute at margin, densely and persistently velutinous on both sides with Yshaped hairs; many bracts and bracteoles persistent in fruit; bracts 5–7 mm long, linear, strongly reflexed, several times as long as short triangular bracteoles; pedicels strongly reflexed and twisted in fruit; sepals adaxially tomentose; petals yellow; anthers loosely sericeous, especially between locules, the connective equaling locules or exceeding them by up to 1 mm; ovary and fruit densely hairy; mature fruits not seen. Grassy savannas, 50–1100 m; Amazonas (Cerro Parú, Río Asisa). Colombia (Vichada), Brazil (southwestern Amazonas, Rondônia), Bolivia (Beni). Byrsonima luetzelburgii Steyerm., Fieldiana, Bot. 28: 288. 1952. Byrsonima cretacea Nied., Repert. Spec. Nov. Regni Veg. 33: 71. 1933, non Gleason 1931.

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Shrub or tree 2–8 m tall; vegetative internodes glabrous except in axil of stipules; stipules 1.5–3 mm long, distinct; petioles (10–)15–25 mm long; blade of larger leaves 8–14 × (3.7–)4.5–8.3 cm, glabrous, the abaxial epidermis deeply pitted and ± densely and persistently glaucous at maturity; bracts and bracteoles persistent in fruit, 0.6–1.5 mm long; pedicels circinate in bud, decurved and eventually twisted in old flowers and fruit; petals pink or white and pink; anther locules glabrous, dorsiventrally flattened and longitudinally winged, the connective exceeding locules by 0.2–0.4 mm; ovary and fruit glabrous; fruits red at maturity, 4.5–6 mm diameter (dried). Riverbanks and lowland white-sand savannas, 100–200 m; Amazonas (Brazo Casiquiare, Caño Yagua, Cerro Yapacana, La Esmeralda, Río Autana, Río Manapiare, Río Ventuari). Brazil (northwestern Amazonas). Byrsonima macrostachya W.R. Anderson, Mem. New York Bot. Gard. 32: 118. 1981. Tree 14–20 m tall; stipules 4.5–9 mm long, 1/2-connate; petioles 20–28 mm long; blade of larger leaves 11–17 × 5.5–9 cm, abaxially very densely and persistently sericeous; inflorescence 18–26 cm long; bracts and bracteoles deciduous before or during anthesis, 2.5–4 mm long or bracteoles smaller; pedicels nearly or quite straight in fruit; petals pink; anther locules glabrous, cylindrical, the connective exceeding locules by 0.8–1.3 mm; ovary densely sericeous; immature fruits 12 mm long (dried), sericeous to glabrescent. Tepui slope forests, 1100– 2000 m; Amazonas (Cerro Marahuaka, Cerro Parú). Endemic. See comments under Byrsonima carraoana. Byrsonima maguirei W.R. Anderson, Mem. New York Bot. Gard. 32: 81. 1981. Shrub or tree 2–8 m tall; stipules 4–7 mm long, distinct or up to 1/2-connate, triangular; petioles 10–30 mm long; leaf blades 5–17 × 2.2–9.5 cm, obtuse at apex, abaxially tomentose to glabrate; bracts and bracteoles persistent in fruit; bracts 4–7 mm long; bracteoles 2.5–4 mm long; pedicels raised on peduncles 5–15 mm long, straight in fruit; petals yellow, turning pinkish in age?; anther locules

glabrous, linear, the connective not or hardly exceeding locules; ovary and fruit glabrous; fruits up to 20 mm diameter (dried). Scrub forests, tepui slopes, 1200–2200 m; Amazonas (Sierra de la Neblina). Endemic. Byrsonima nitidissima H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 152, pl. 448. 1821 [1822]. Shrub or tree 1.5–5(–10) m tall; stipules 2–5 mm long, usually completely and smoothly connate, rarely distinct at apex; petioles 2–6 mm long; blade of larger leaves 3.5–8.5 × 2–4.5 cm, rounded or broadly obtuse at apex, adaxially shining, abaxially sericeous or velutinous to glabrescent; bracts and bracteoles persistent in fruit; bracts 1.5– 3 mm long, bracteoles slightly smaller; pedicels decurved in fruit; petals white or pinkish; filaments bearded with kinky rusty brown hairs; anther locules cylindrical, glabrous, the connective not or hardly exceeding locules; ovary and fruit glabrous; fruits dark purplish, developing half-immersed in enlarged, disk-like receptacle, 4–5 mm diameter (dried). Scrub forests on and beside granitic outcrops, 50–200 m; Bolívar (upper Río Orinoco south of Maniapure), Amazonas (upper Río Orinoco north of Río Sipapo). Apure; Colombia (Río Orinoco). Byrsonima pachypoda W.R. Anderson, Mem. New York Bot. Gard. 32: 123. 1981. Shrub 2–4 m or tree to 8(–10) m tall; vegetative internodes glabrous except in axil of stipules; stipules 3–6 mm long, distinct, acute or obtuse at apex; petioles 3–5(–7) mm long, glabrous; blade of larger leaves 6–14.5 × 3–8.3 cm, obovate, broadly obtuse or rounded at apex, glabrous; bracts and bracteoles persistent in fruit; bracts 2–3 mm long, bracteoles slightly shorter; pedicels distally thickened, straight in bud and fruit; petals white to pink in age; anthers glabrous, the locules linear and unwinged, the connective swollen but not or hardly exceeding locules; ovary and fruit glabrous; fruits red to black, 7–8 mm diameter (dried). Low scrub, low forests (8–10 m), low Clusia-Magnolia woodlands, 1400–1900 m; Bolívar (Ilútepui). Guyana (upper Mazaruni River basin, Mount Ayanganna, Mount Wokomung).

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Byrsonima poeppigiana A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 335. 1840. Tree 6–25 m tall; stipules 4–8 mm long, completely connate, the pair rounded at apex; petioles 6–15(–18) mm long; blade of larger leaves 10–19(–24) × 4.5–10(–14.5) cm, abaxially thinly to densely velutinous with erect, bifurcate hairs; bracts and bracteoles mostly deciduous before maturation of fruit; bracts 1.5–4 mm long; bracteoles 0.5–1.5(–2) mm long; pedicels decurved or twisted in fruit; sepals glabrous on adaxial face; petals yellow; anthers loosely sericeous, at least between locules, the connective equaling or slightly exceeding locules; ovary glabrous or sparsely sericeous distally; fruits yellow, 8– 10 × 6–8 mm (dried), glabrous or with a few hairs at apex. Nonflooded evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, between San Carlos de Río Negro and Solano, Santa Rosa de Amanadona). Brazil (western Amazonas, Rondônia), Amazonian Peru (Loreto, Madre de Dios). Byrsonima punctulata A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 335. 1840. Byrsonima barkleyana Cuatrec., Webbia 13: 620. 1958. Small tree 2.5–9 m tall, rarely a shrub only 1 m; vegetative internodes hairy to glabrescent; stipules 3.5–7(–8) mm long, distinct, often acuminate; petioles 10–19 mm long; blade of larger leaves 8.5–14.5 × 4–8 cm, tomentose to glabrate at maturity except abaxial veins and midrib, the reticulum and areoles ± concolorous; inflorescence 9–18 cm long; bracts and bracteoles persistent in fruit, 1.5–3 mm long or bracteoles smaller; pedicels usually decurved in fruit; petals white turning pink or red in age; anther locules glabrous, linear, the connective exceeding locules by 0.3–1 mm; ovary and fruit glabrous; fruits red to blue, 4–6 mm diameter (dried). On sandy soil beside rivers, sometimes in seasonally flooded savannas, 100– 200 m; Amazonas (Río Atabapo, Río Sipapo). Colombia (Vaupés, Vichada), Brazil (Amazonas: basin of upper Rio Negro, southern Roraima). As noted in my 1981 paper, this species fills the morphological and geographical gap between Byrsonima cuprea and B. leuco-

phlebia, and may well have originated through hybridization between those two now-disjunct entities. Backcrossing to the putative parents probably continues, especially in the area of Manaus, where B. punctulata and B. leucophlebia co-occur, and the large-leaved plants of B. punctulata found in the region of San Fernando de Atabapo hardly differ from B. cuprea except in their hairs. Specimens of “B. cuprea” from the Atabapo area, in which the abaxial leaf hairs are sparser than usual, e.g., Velazco 244 (MO, PORT), are probably part of that hybrid complex, and call into question the feasibility of maintaining B. cuprea as a species. The name B. punctulata is the oldest in the complex. Byrsonima schomburgkiana Benth., London J. Bot. 7: 123. 1848. —Chaparro manteco. Tree 4–18 m tall, rarely a shrub only 1–2 m; stipules 2–4 mm long, smoothly connate; petioles 5–10 mm long, densely and ± persistently tomentose; blade of larger leaves (5–) 7.5–17 × (3–)4–9 cm, elliptic to obovate, tomentose but soon ± glabrate; bracts and bracteoles mostly deciduous in fruit, occasionally persistent; bracts (2–)3–5(–9) mm long, narrowly lingulate, bracteoles similar but shorter; pedicels straight or decurved in fruit; sepals lingulate-accrescent and membranous in fruit; petals pink or pink and white; anthers tomentose, the bulbous connective exceeding locules by (0.3–)0.5–1.1 mm; ovary and fruit glabrous; fruits yellow, 6–10 mm diameter (dried). Dry forests and savannas, 100–800 m; Bolívar (Caicara south to Picachos de Sabana Nueva, Represa Guri, between Río Perro de Agua and Río Juasjualito west of San Antonio, San Félix to Puerto Ordaz, Sierra de Maigualida), Amazonas (Río Ocamo). Guyana (Rupununi area), Brazil (Roraima). Byrsonima spicata (Cav.) DC., Prodr. 1: 580. 1824. —Malpighia spicata Cav., Diss. 8: 409, pl. 237. 1789. —Byrsonima coriacea var. spicata (Cav.) Nied. in Engl., Pflanzenr. IV. 141: 700. 1928, nom. superfl. —Manteco. Byrsonima propinqua Benth., London J. Bot. 7: 120. 1848.

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Tree 3–25 m tall; stipules 1–3 mm long, completely and smoothly connate; petioles 5–15(–20) mm long; blade of larger leaves 6.5–15(–16) × 1.7–5(–6) cm, tightly sericeous abaxially to usually eventually glabrate, the hairs sessile or subsessile with short, straight, parallel crosspieces up to 0.5 mm long, with 15–20 or more pairs of fine lateral veins, none very prominent; bracts and bracteoles deciduous in fruit; pedicels mostly decurved or twisted in fruit; petals yellow, the posterior petal bearing 2 or more glands at apex of claw or on base of limb; anthers sericeous, at least between locules, the connective equaling or slightly exceeding locules; ovary sericeous; fruits yellow-orange, 10–12 mm diameter (dried), sericeous to glabrate. Dry scrubland to wet forests, near sea level to 700(–1300) m; Delta Amacuro (near village of Morichito), common in Bolívar, Amazonas (Puerto Ayacucho to El Burro, Río Mawarinuma, Yutajé). Anzoátegui, Aragua, Barinas, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Panama, West Indies, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. The leaves of Byrsonima spicata are usually glabrescent at maturity. When the hairs persist (e.g., Davidse & Miller 27179, MICH, MO, VEN, Huber et al. 12911, MICH, MYF, VEN, Steyermark et al. 113966, MICH, VEN), the leaves have a different aspect, but I see no pattern suggesting that those plants deserve taxonomic recognition. In my 1981 paper (p. 96) I stated that B. propinqua is a name for such plants, but that was incorrect; the holotype of B. propinqua (K!) has rather large leaves for the species, but they are glabrescent. Byrsonima steyermarkii W.R. Anderson, Mem. New York Bot. Gard. 32: 124. 1981. Tree 6–7 m tall; vegetative internodes glabrous except in axil of stipules; stipules 2–5 mm long, distinct; petioles 2.5–5 mm long; blade of larger leaves 6.5–12.2 × 4–7 cm, glabrous; bracts and bracteoles persistent in fruit, 1–2.5 mm long; pedicels straight in fruit; petals white; anther locules glabrous, unwinged, the connective exceeding locules by 0.1–0.5 mm; ovary and fruit glabrous; fruits 5–7.5 mm diameter (dried). Low forests on tepui slopes and summit, 1200–2100

m; Bolívar (Cerro Jaua), Amazonas (Cerro Parú). Endemic. ◆Fig. 83. Byrsonima stipulacea A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 332. 1840. —Alcoceratothrix stipulacea (A. Juss.) Nied. in Engl., Pflanzenr. IV. 141: 762. 1928. —Chaparro manteco, Manteco, Manteco de agua. Byrsonima rugosa Benth., London J. Bot. 7: 118. 1848. —Alcoceratothrix rugosa (Benth.) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 1: 45. 1901. Tree (5–)7–25(–35) m tall; stipules 8–25 mm long, amplexicaulous, completely connate, each pair deciduous independently of and often well before the leaf; petioles (6–)10–27 mm long; blade of larger leaves 12–27 × 6–13 cm, abaxially velutinous with a mixture of long, basifixed simple hairs, stalked stellate hairs, and sessile stellate hairs; bracts and bracteoles deciduous in fruit; bracts 5–10 mm long; bracteoles 2–5 mm long; pedicels straight or somewhat decurved in fruit; petals yellow; anthers loosely sericeous on both sides, the connective usually exceeding locules by 0.4–1.1 mm; ovary densely short-velutinous with an overlay of appressed hairs; fruits orange-yellow, 12–18 mm diameter (dried), tomentose to glabrate. Evergreen lowland to lower montane forests, 100–1300 m; Delta Amacuro (east-northeast of El Palmar), widespread in Bolívar to 63°W, rare farther west (Río Caura at 5°35'N). Guyana, Suriname, French Guiana, Brazil (Rondônia, Roraima, and Amapá south to Espírito Santo). Byrsonima tillettii W.R. Anderson, Mem. New York Bot. Gard. 32: 118. 1981. Shrub or tree 2–5 m tall; stipules 6–11 mm long, completely connate, the pair rounded at apex; petioles 5–10 mm long; blade of larger leaves 9–11 × 4.5–6.5(–8) cm, abaxially sinuous-sericeous with hairs not dense enough to hide epidermis at maturity; bracts and bracteoles persistent in fruit; bracts (2.5–)3–5(–7) mm long; bracteoles 2–3 mm long; pedicels decurved in fruit; petals pink, the lateral 4 adaxially pilose on claw, abaxially pilose on limb; anthers glabrous, the connective exceeding locules by 0.7–1.2 mm; ovary and fruit glabrous; fruits 5–6 mm diameter (dried). Dense

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scrub, low forests with trees to 8 m in savannas, ca. 1300 m; Bolívar (Wadakapiapué-tepui west of Yuruaní-tepui). Guyana (Mount Ayanganna, upper Mazaruni River basin). Byrsonima verbascifolia (L.) DC., Prodr. 1: 579. 1824. —Malpighia verbascifolia L., Sp. Pl. 426. 1753. Byrsonima verbascifolia var. denudata Cuatrec., Webbia 13: 605. 1958. Gnarled shrub up to 60 cm tall, the leaves borne in dense sessile clusters on the thick corky stem, the internodes essentially lacking; stipules 6–12 mm long, connate, the pair triangular; petioles 10–35(–80) mm long; blade of larger leaves 14–25(–33) × 5–13 (–15) cm, obovate or spatulate, attenuate at base, usually obtuse or rounded and apiculate at apex, densely and usually persistently villous adaxially and woolly abaxially, rarely eventually subglabrate adaxially or on both sides; bracts and bracteoles narrowly triangular or subulate, deciduous; bracts 5– 11 mm long, bracteoles about half as long; pedicels decurved in fruit; petals yellow, the posterior petal eglandular; anthers loosely tomentose to glabrate, the connective not or hardly exceeding locules; ovary densely appressed-villous; fruits yellow, ca. 15 mm diameter (dried), thinly villous. Open savannas, often on white sand, 50–1200 m; common in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Guárico, Mérida, Monagas; Honduras, Nicaragua, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. Occasional plants in Venezuela have elongated internodes, short leaves with unwinged petioles, and small bracts. These have probably resulted from hybridization between Byrsonima verbascifolia and B. crassifolia, which is common and widespread and occurs in the same habitat. This taxon is extremely variable through its range, and probably represents a complex of several related taxa rather than a single species. The descriptive notes above refer only to the form found in Venezuela and the Guianas. Byrsonima wurdackii W.R. Anderson, Mem. New York Bot. Gard. 32: 119. 1981. Tree 3–19 m tall; stipules 1.7–4(–5) mm long, connate or distinct at apex, the pair tri-

angular, acute; petioles 15–35 mm long; blade of larger leaves 9.5–19 × 5–12 cm, sparsely sericeous to soon glabrate; inflorescence 10–25 cm long, the sessile cincinni 1– 3(4)-flowered; bracts and bracteoles caducous; bracts 3–4 mm long; bracteoles 0.5–1.5 mm long; pedicels decurved in old flowers and fruits; calyx glands usually absent or rudimentary, rarely well developed; petals white turning pinkish; anther locules glabrous, unwinged, the connective exceeding locules by 0.8–1.6 mm; ovary and fruit glabrous; styles strongly bent at apex in bud; fruits 6–7 mm diameter (dried). Savannas and scrubby forests on white sand, 100–200 m; Amazonas (upper Río Atabapo and La Esmeralda south to Maroa and San Carlos). Brazil (Amazonas: upper Rio Negro, Rio Uaupés). Byrsonima sp. A Tree 3–15 m tall; vegetative internodes glabrous except in axil of stipules; stipules 2–2.5 mm long, distinct; petioles (18–)22–33 mm long; blade of larger leaves (11–)12.5– 17.5 × (5–)6.5–8.5 cm, glabrous; bracts and bracteoles persistent in fruit, 1–3.5 mm long; pedicels ascending and becoming sigmoid in fruit, 8–12 mm long, 0.8–1 mm diameter; sepals abaxially sparsely velutinous or sericeous, elongating to 5–7 mm in fruit, the accrescent sepals often closely investing the enlarging fruit; petals and stamens unknown; fruits 6–8 mm diameter (dried), glabrous. Along black-water rivers, 100–200 m; Amazonas (Caño Caname, Río Atacavi, Río Autana, Río Cuchakén in Río Atabapo drainage). Otherwise known from one collection in Caquetá, Colombia. I plan to describe this species in the near future, giving it the epithet “flexipes” in reference to the pedicels, which are peculiarly sigmoid-ascending in fruit. The species resembles Byrsonima rodriguesii W.R. Anderson, a plant from the vicinity of Manaus that differs in its longer stipules, glabrous sepals that are soon reflexed or revolute in the enlarging fruit, fruiting pedicels that are thicker and curved upward but not sigmoid, and larger fruits. The petals of the new species can be expected to be white turning red, as in B. rodriguesii, and for the same reason its anthers should have a large, glandular connective that extends well beyond flattened locules.

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Fig. 79. Byrsonima duidana Fig. 80. Byrsonima coccolobifolia

Fig. 81. Byrsonima carraoana

Fig. 82. Byrsonima bronweniana

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Fig. 83. Byrsonima steyermarkii

Fig. 84. Byrsonima cowanii

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Fig. 85. Byrsonima aerugo

6. CLONODIA Griseb. in Mart., Fl. Bras. 12(1): 26. 1858. Atopocarpus Cuatrec., Webbia 13: 454. 1958. Skoliopterys Cuatrec., Webbia 13: 451. 1958. Woody vines or shrubs; stems terete, with many tiny punctiform lenticels. Leaves decussate; stipules none(?) or minute (ca. 0.3 mm long), triangular, borne on stem at base of petiole. Inflorescence a pseudoraceme, simple or compound to form a panicle, terminal or terminal and lateral. Calyx bearing 8–10 glands; petals pink and/or white, abaxially winged. Stamens 10, glabrous; filaments basally to 1/2-connate, straight or the posterior 3 (especially the 1 opposite posterior petal) sigmoid. Ovary of 3 carpels, connate at base, distinct distally; styles 3, stout, strongly unequal, the anterior thinner and usually shorter than the other 2, bent toward posterior petal, the 2 posterior styles turned toward anterior sepal; stigmas large, internal, the styles dorsally truncate to prominently hooked at apex. Fruit a schizocarp comprising 3 (or fewer by abortion) dry indehiscent mericarps separating from a short pyramidal torus; mericarp with the wings reduced to winglets or rounded outgrowths. Amazonian Colombia, southern Venezuela, Brazil, Bolivia; 3 species, 1 in Venezuela. Clonodia complicata (H.B.K.) W.R. Anderson, Mem. New York Bot. Gard. 32: 206. 1981. —Hiraea complicata H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 171. 1821 [1822]. —Mascagnia complicata (H.B.K.) Nied., Arbeiten Bot. Inst. Königl. Lyceum

Hosianum Braunsberg 3: 4. 1908. Hiraea nitida H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 171. 1821 [1822]. —Mascagnia nitida (H.B.K.) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 3: 4. 1908.

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Fig. 86. Clonodia complicata

Mascagnia lehmanniana Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum. Braunsberg 8: 59. 1926. —Skoliopterys lehmanniana (Nied.) Cuatrec., Webbia 13: 452. 1958. Clonodia racemosa var. orinocensis Nied. in Engl., Pflanzenr. IV. 141: 580. 1928. Woody vine, occasionally described as a shrub; petioles (3–)5–8 mm long; blade of larger leaves (4.5–)7.5–11.5(–13.5) × (2.5–)4– 6(–7) cm, ovate, acute or obtuse at apex, loosely sericeous to ± glabrate at maturity; pseudoracemes comprising (15–)20–50 flow-

ers; petals, especially the posterior, sometimes papillose on adaxial face; styles dorsally truncate or with a very short hook at apex; mericarps bearing a dorsal winglet 0.5–2 mm wide and 3–5 winglets on each side, mostly at right angles to dorsal winglet, sometimes connate with it at apex. Usually near rivers or in marshy places, near sea level to 300 m; common along the Río Orinoco and its tributaries in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Trujillo, Zulia; eastern Colombia. ◆Fig. 86.

7. DIACIDIA Griseb. in Mart., Fl. Bras. 12(1): 119. 1858. Sipapoa Maguire, Mem. New York Bot. Gard. 8: 124. 1953. Trees, shrubs, or subshrubs, with mostly basifixed or sub-basifixed hairs. Leaves eglandular, often glaucous, especially abaxially; proximal portions of stipules and petioles fused to form an interpetiolar sheath; distal portions of stipules (the

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part extending beyond petiole) distinct or connate. Inflorescence a simple or compound thyrse (i.e., a raceme or panicle of cincinni) or a pseudoraceme (i.e., a raceme of 1-flowered cincinni); bracts and bracteoles eglandular, caducous or eventually deciduous. Sepals all biglandular, accrescent in fruit; petals yellow, often with red claws. Stamens 6–10; anthers with each half bearing at apex and angled forward 1 or occasionally 2 stout, basifixed, awn-like hairs. Ovary composed of 3 completely connate carpels but only 2-locular, the anterior carpel reduced to a ridge of tissue; styles 3, slender and subulate with minute slightly internal stigmas. Fruit a spheroidal or ovoidal, dry, indehiscent, nut-like capsule ca. 2.5 mm high and wide, with a thin exocarp and a bony, smooth or rugose endocarp and containing 2 seeds (or 1 due to abortion), subtended and enclosed by accrescent (in most species red, membranous, veiny, wing-like) sepals. Endemic to the Guayana Shield in Colombia, southwestern Venezuela, and Brazil; 11 species, 10 in Venezuela, all in the flora area. Diacidia aracaënsis W.R. Anderson was described from the Serra Aracá in Brazil, but may eventually be found in Venezuela. It is similar to D. ferruginea, but differs in having 8 fertile stamens, the anther connectives hardly enlarged, the sepals glabrous on the margin, and the leaves soon glabrescent adaxially. Key to the Species of Diacidia 1.

1.

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

4.

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

Stamens 10; sepals only slightly accrescent in fruit, up to 3.5 × 2.5 mm; interpetiolar stipular sheath 2–3(–6) mm long, obscure, lacking median seams; 50–400 m .............................................................. D. galphimioides Stamens 6–9; sepals greatly accrescent in fruit, forming membranous wings 7–13 × (2.5–)4–12 mm; interpetiolar stipular sheath 4–26 mm long, marked by median seams; 300–2300 m ....................................... 2 Stipule lobes beyond petiole completely connate to form a single intrapetiolar structure ........................................................................... 3 Stipule lobes beyond petiole nearly or quite distinct ................................ 6 Stipules 7–11 mm long beyond petiole, acute or obtuse at apex ................................................................................................ D. kunhardtii Stipules 28–90 mm long beyond petiole, broadly obtuse or rounded at apex ........................................................................................................ 4 Inflorescence a simple pseudoraceme, glabrous or very soon glabrate; bracts, bracteoles, and sepals glabrous; shrubs 0.5–3 m tall .................................................................................................. D. stipularis Inflorescence compound, with a terminal axis and few to many lateral axes, and with the cincinni several-flowered; inflorescence, bracts, bracteoles, and sepals densely and persistently hairy; shrubs or trees (1–)2–15 m tall ....................................................................................... 5 Leaves glabrate abaxially or hairy only on veins; 600–1600 m ................................................................................................ D. glaucifolia Leaves densely and persistently rufous-sericeous abaxially over the whole surface; 1500–1900 m ............................................................. D. rufa Inflorescence a simple thyrse with several-flowered cincinni, glabrous; leaves glabrous or loosely sericeous to glabrate abaxially ........ D. cordata Inflorescence a pseudoraceme, hairy; leaves hairy abaxially .................. 7

Diacidia 131

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

Vegetative internodes glabrous .................................................. D. hypoleuca Vegetative internodes hairy ....................................................................... 8 Leaf blades glabrous or soon glabrescent adaxially except for very base; stipules 1–3 mm long beyond petiole .................................... D. ferruginea Leaf blades densely and persistently hairy adaxially; stipules 5–15 mm long beyond petiole ................................................................................ 9 Leaf blades woolly abaxially, the hairs much twisted and intertwined ....................................................................................................... D. vestita Leaf blades sericeous abaxially, the hairs straight, ± appressed, and parallel ................................................................................ D. steyermarkii

Diacidia cordata (Maguire) W.R. Anderson, Mem. New York Bot. Gard. 32: 66. 1981. —Sipapoa cordata Maguire, Mem. New York Bot. Gard. 18: 46. 1969. Shrub 0.5–2(–3) m tall; vegetative internodes glabrous; stipules 7–10 mm long beyond petiole, distinct, acute at apex; petioles 5–10 mm long including petiole-stipule sheath; blade of larger leaves 3–6.5 × 1.5–4.5 cm, deeply cordate at base, glabrous adaxially, glabrous or loosely sericeous to glabrate abaxially; inflorescence a simple thyrse, the cincinni 2–6(–8)-flowered, glabrous; fertile stamens 8 or 9; sepals in fruit up to 7 × 4 mm. Brushy rocky tepui slopes, 300–1900 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 88. Diacidia ferruginea (Maguire & K.D. Phelps) W.R. Anderson, Mem. New York Bot. Gard. 32: 68. 1981. —Sipapoa ferruginea Maguire & K.D. Phelps, Mem. New York Bot. Gard. 8: 126. 1953. Shrub or small tree 1–4 m tall; vegetative internodes persistently sericeous; stipules 1– 3 mm long beyond petiole, distinct, acute at apex; petioles 5–9 mm long including petiolestipule sheath; blade of larger leaves 3–4.5 × 1–2 cm, adaxially glabrous or soon glabrescent except for sericeous base, abaxially densely and persistently rufous- or yellowish-sericeous; inflorescence a pseudoraceme, tomentose; stamens 6; sepals in fruit up to 13 × 12 mm. Tepui savannas, 1100–2000 m; Amazonas (Cerro Parú). Endemic. ◆Fig. 89. Diacidia galphimioides Griseb. in Mart., Fl. Bras. 12(1): 120. 1858. Diacidia duckeana Maguire, Mem. New York Bot. Gard. 8: 124. 1953. Diacidia parvifolia Cuatrec., Webbia 13: 632. 1958.

Shrub or small tree 0.5–2(–4) m tall; vegetative internodes sericeous; stipules 2–5 (–7) mm long beyond petiole, completely connate and rounded at apex; petioles 4–8 mm long including petiole-stipule sheath; blade of larger leaves (2–)4–6(–9) × (0.7–)1.2–2.7 (–5) cm, loosely sericeous to subtomentose on both sides, occasionally glabrescent in age; inflorescence a simple thyrse with cincinni (1)2–10(–13)-flowered, glabrous or proximally sparsely sericeous; stamens 10; sepals in fruit up to 3.5 × 2.5 mm. On or among granitic outcrops, 50–400 m; Bolívar (Río Caura at 6°14'N), Amazonas (basins of upper Río Orinoco and upper Río Negro from Río Cuao south to Piedra de Cocuy). Adjacent Colombia and Brazil. In my 1981 paper I recognized Diacidia parvifolia as distinct from D. galphimioides. However, the accumulation of intermediates has convinced me that it is simply a smallleaved form that may not even have any genetic basis, reflecting nothing more than the extremely harsh conditions sometimes found on the granitic outcrops where it grows. Most of the characters I used in my earlier key overlap too much to be diagnostic, and the Venezuelan plants with the small leaves of D. parvifolia (Berry et al. 5114 [MICH], Morillo & Ishikawa 3470 [MICH, VEN], Morillo 6881 [MICH, VEN]) have the larger bracteoles of D. galphimioides. Diacidia glaucifolia (Maguire) W.R. Anderson, Mem. New York Bot. Gard. 32: 63. 1981. —Sipapoa glaucifolia Maguire, Mem. New York Bot. Gard. 18: 48. 1969. Shrub or tree (1–)2–15 m tall; vegetative internodes glabrous; stipules 70–90 mm long beyond petiole, completely connate and rounded at apex; petioles 34–53 mm long including petiole-stipule sheath; blade of

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larger leaves 16–27 × 11–18 cm, glabrous adaxially, sericeous on midrib and veins to soon glabrate abaxially; inflorescence a compound thyrse, the cincinni mostly 4–10-flowered, laxly rufous-sericeous; stamens (8)9; sepals in fruit up to 9 × 4 mm. Tepui slope forests, 600–1600 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Brazil (Amazonas: Serra Pirapucú). Diacidia hypoleuca (Maguire) W.R. Anderson, Mem. New York Bot. Gard. 32: 67. 1981. —Sipapoa hypoleuca Maguire, Mem. New York Bot. Gard. 8: 125. 1953. Shrub or tree 3–10 m tall; vegetative internodes glabrous; stipules beyond petiole 10–18 mm long, connate for up to 3 mm and distally distinct, acute or acuminate at apex; petioles 16–25 mm long including petiolestipule sheath; blade of larger leaves 5.5– 10.5 × 3–6 cm, adaxially glabrous, abaxially densely and persistently white- or yellowishsericeous; inflorescence a pseudoraceme, loosely sericeous; stamens 9; sepals in fruit up to 10 × 8 mm. Upper tepui slopes, 500– 1200 m; Amazonas (Cerro Yapacana). Endemic. Diacidia kunhardtii (Maguire) W.R. Anderson, Mem. New York Bot. Gard. 32: 66. 1981. —Sipapoa kunhardtii Maguire, Mem. New York Bot. Gard. 8: 125. 1953. Shrub or small tree to 4(–8) m tall; vegetative internodes glabrous; stipules 7–11 mm long beyond petiole, completely connate, acute or obtuse at apex; petioles 17–26 mm long including petiole-stipule sheath; blade of larger leaves 4–7 × 2.3–5 cm, adaxially sparsely sericeous to glabrate, abaxially spreading-sericeous on midrib and lateral veins; inflorescence a pseudoraceme (the cincinni rarely 2-flowered), villous; fertile stamens 8; sepals in fruit up to 13 × 4 mm. Tepui meadows and low woodlands on sandy soil, 1400–1600 m; Amazonas (Cuao-Sipapo massif). Endemic. ◆Fig. 90. Diacidia rufa (Maguire) W.R. Anderson, Mem. New York Bot. Gard. 32: 64. 1981. —Sipapoa rufa Maguire, Mem. New York Bot. Gard. 18: 48. 1969. Shrub or tree 2–12 m tall; vegetative internodes glabrous; stipules 40–80 mm long

beyond petiole, completely connate and rounded at apex; petioles 20–40(–50) mm long including petiole-stipule sheath; blade of larger leaves 8.5–17(–22) × 5–10(–14) cm, glabrous adaxially or sericeous on midrib, laxly and persistently rufous-sericeous all over abaxial surface; inflorescence a compound thyrse, the cincinni mostly 2–6-flowered, densely rufous-sericeous; stamens (8)9; sepals in fruit up to 9 × 4 mm. Low tepui forests and bromeliad-scrub zone on escarpment slopes and ridges, 1500–1900 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra Pirapucú). Diacidia steyermarkii (Maguire) W.R. Anderson, Mem. New York Bot. Gard. 32: 71. 1981. —Sipapoa steyermarkii Maguire, Mem. New York Bot. Gard. 18: 54. 1969. Shrub or tree (1–)3–8 m tall; vegetative internodes loosely sericeous to villous; stipules 9–15 mm long beyond petiole, distinct, acuminate at apex; petioles 23–30 mm long including petiole-stipule sheath; blade of larger leaves 6–10 × 3–4.5 cm, adaxially villous, abaxially densely sericeous; inflorescence a pseudoraceme, villous; stamens 8; sepals in fruit up to 9 × 4 mm. Tepui slope and summit vegetation, 1300–2300 m; Bolívar (Jaua-Sarisariñama massif). Endemic. ◆Fig. 87. Diacidia stipularis (Maguire & K.D. Phelps) W.R. Anderson, Mem. New York Bot. Gard. 32: 65. 1981. —Sipapoa stipularis Maguire & K.D. Phelps, Mem. New York Bot. Gard. 8: 127. 1953. Shrub 0.5–3 m tall; vegetative internodes glabrous; stipules 28–62 mm long beyond petiole, completely connate, broadly obtuse or rounded at apex; petioles 12–24 mm long including petiole-stipule sheath; blade of larger leaves 4.5–13.5 × 2.5–9.5 cm, adaxially glabrous, abaxially densely and ± persistently white- or yellowish-sericeous; inflorescence a pseudoraceme, glabrous or very soon glabrate; stamens 6(7); sepals in fruit up to 12 × 5 mm. Tepui scrub, 1600– 2000 m; Amazonas (Cerro Parú). Endemic. Diacidia vestita (Benth.) B.D. Jacks., Index Kew. 1: 741. 1895. —Coleostachys

Diacidia 133

Fig. 87. Diacidia steyermarkii

Fig. 88. Diacidia cordata

Fig. 89. Diacidia ferruginea

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vestita Benth., London J. Bot. 7: 124. 1848. —Sipapoa vestita (Benth.) Maguire, Mem. New York Bot. Gard. 8: 127. 1953. Shrub or small tree (1–)2–9 m tall; vegetative internodes loosely sericeous; stipules 5–11 mm long beyond petiole, distinct, acute or acuminate at apex; petioles 19–32 mm long including petiole-stipule sheath; blade of larger leaves 6–10(–13.5) × 2.8–5.5 cm, adaxially villous or subsericeous or subtomentose, abaxially woolly; inflorescence a pseudoraceme, villous; fertile stamens (7)8; sepals in fruit up to 13 × 7 mm. Open tepui slopes and woodlands, 1100–1900 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Sierra Parima). Endemic.

Fig. 90. Diacidia kunhardtii

8. DICELLA Griseb., Linnaea 13: 249. 1839. Woody vines. Stipules small, interpetiolar. Inflorescence a decompound panicle, the flowers borne ultimately in short pseudoracemes with decussate bracts, each bract subtending a 1-flowered peduncle with apical bracteoles. Sepals appressed in anthesis, the anterior eglandular, the lateral 4 biglandular, all accrescent in fruit; petals yellow, abaxially densely sericeous, adaxially glabrous or tomentose. Stamens 10; filaments and (in our species) the locules hairy. Ovary formed from 3 completely connate carpels, the 2 posterior locules full-sized and fertile, the anterior locule smaller and empty; styles distinct and straight, the posterior 2 stout, obliquely truncate or short-hooked at apex and with large internal stigmas, the anterior style absent or short, slender, rudimentary. Fruit composed of a dry, hard, indehiscent, nutlike structure with a thick, fibrous wall, containing 1 or 2 seeds, most often only 1, subtended by 5 dry wings formed by enlargement of the sepals. Costa Rica, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 7 species, not yet known from Venezuela but 1 likely to occur there in the flora area. See Mark W. Chase, 1981 [A revision of Dicella (Malpighiaceae), Syst. Bot. 6: 159–171]. Dicella julianii (J.F. Macbr.) W.R. Anderson, Acta Amaz. 5: 279. 1975. —Tetrapterys julianii J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser., 13: 805. 1950. Dicella amazonica Pires, Bol. Técn. Inst. Agron. N. 38: 27. 1960. Woody vine climbing to 30 m; petioles 13–24 mm long; blade of larger leaves 10– 19(–22) × 5–11 cm, elliptic, acuminate at apex, abaxially persistently sericeous; bracts (4–)5–7 × 3–5 mm, bracteoles similar but smaller; fruit spherical, 13–18 mm diameter;

wings (enlarged sepals) subtending fruit 20–55 × 7–20 mm, unequal, the anterior eglandular one shortest, the anterior-lateral pair intermediate, the posterior-lateral pair longest. In forests near rivers, 200–400 m. Western Amazonia (Colombia, Ecuador, Peru, Brazil); known from Rio Maturacá, Rio Cauaburí, on the lower slopes of the Brazilian side of the Sierra de la Neblina, and therefore to be expected on the Venezuelan side. ◆Fig. 91.

Dicella 135

B

A

G

C

D

E

F

Fig. 91. Dicella julianii. —A. Flowering branch, ×0.5. —B. Decussate flower buds, ×2.1. —C. Flower, ×2.6. —D. Androecium, ×5.3. —E. Stamens, ×8.2. —F. Gynoecium, posterior view (left) and anterior view (right), ×8.2. —G. Fruit, ×5.3. ©New York Botanical Garden 1981.

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G

H C

D

B A

E

F

Fig. 92. Diplopterys cabrerana. —A. Flowering branch, ×0.5. —B. Leaf base with glands, ×1.6. —C. Umbel of 4 buds, ×2.1. —D. Abaxial view of posterior-lateral petal (above) and posterior petal (below), ×4.3. —E. Abaxial view of part of androecium, stamen to left opposite anterior sepal, ×10.7. —F. Gynoecium, anterior style in center, ×10.7. —G. Mericarp with short wings, abaxial view (left), adaxial view (middle), cross section (right), ×1.1. —H. Mericarp with long wings, abaxial view (left), adaxial view (right), cross section (below), ×1.1. ©New York Botanical Garden 1981.

Excentradenia 137

9. DIPLOPTERYS A. Juss. in Deless., Icon. Sel. Pl. 3: 20. 1837 [1838]. Woody vines. Stipules small, distinct, interpetiolar. Inflorescence axillary, shorter than the subtending leaves, of 1–several simple 4-flowered umbels or 1–several racemes of up to 7 4-flowered umbels; pedicels sessile; bracts and bracteoles similar, lingulate, eglandular, spreading, persistent, borne in a cluster of 12 at base of umbel. Anterior sepal usually eglandular, rarely bearing 1 abaxial gland; 4 lateral sepals biglandular; petals yellow, long-fimbriate, abaxially sparsely sericeous. Stamens 10, all fertile; anthers unequal. Gynoecium of 3 free carpels adnate to a common torus; styles 3, distinct, with capitate terminal stigmas. Fruit dry, breaking apart into 3 mericarps; nut of mericarp with a hard woody pericarp, bearing a dorsal crest and 2–several lateral winglets or crests parallel to areole and interconnected by ridges. Mexico, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; 4 species, 1 in Venezuela. See Bronwen Gates, 1982 [Banisteriopsis, Diplopterys (Malpighiaceae), Fl. Neotrop. Monogr. No. 30]. Diplopterys cabrerana (Cuatrec.) B. Gates, Brittonia 31: 109. 1979. —Banisteriopsis cabrerana Cuatrec., Webbia 13: 493. 1958. —Yagé. Woody vine; petioles (4–)8–15(–22) mm long, bearing 2 large convex glands on margin at apex; blade of larger leaves (8.5–)10– 21(–26) × (3–)4.1–9 cm, elliptic, long-acuminate at apex, abaxially sparsely sericeous; mericarp with nut up to 15 mm diameter, bearing a dorsal crest or winglet 1–5 mm wide and essentially 4 roughly parallel

ridges or winglets on each side 0.5–10 mm wide, these irregular, dissected, and interconnected with transverse ridges. Riparian forests, 100–200 m; Amazonas (Río Mawarinuma, Río Orinoco above San Fernando de Atabapo). Amazonian Colombia, Ecuador, Peru, Brazil. ◆Fig. 92. Diplopterys cabrerana is widely used by indigenous peoples in western Amazonia as an admixture to Banisteriopsis caapi in the preparation of hallucinogenic concoctions.

10. EXCENTRADENIA W.R. Anderson, Contr. Univ. Michigan Herb. 21: 29. 1997. Woody vines. Leaves with petiole biglandular; stipules small, triangular, borne on very base of petiole or on adjacent stem, or absent; blade eglandular or bearing small glands on margin, the tertiary veins strongly parallel (scalariform). Inflorescence a single axillary raceme of 3–7(–9) 4-flowered umbels, with 1 umbel terminal and the other 1–4 pairs axillary to bracts bearing stipules and often petiole glands; floriferous bracts small, persistent, eglandular; floriferous peduncle short or absent; bracteoles small, persistent, one of each pair bearing 1 bulging eccentric abaxial gland toward center of umbel; pedicels circinate in bud as far as known. Sepals all eglandular or the lateral 4 biglandular, revolute in anthesis; petals yellow, glabrous. Stamens 10, all fertile; anthers ± alike. Ovary with the 3 carpels nearly distinct, all fertile; styles 3, inserted low on ventral face of carpels, the apex with a large internal stigma and dorsally truncate, apiculate, or bearing a hook up to 0.3 mm long. Fruits breaking apart into 3 samaras, each with a large, membranous, subcircular lateral wing borne on upper edge of nut, incised to nut at apex, usually continuous at base; dorsal wing small; intermediate winglets absent. Venezuela, Guyana, Suriname, French Guiana, Brazil, Bolivia; 4 species, 1 in Venezuela. Three of the species of Excentradenia are from north of the Amazon and the other is from Bolivia. It is close to Hiraea but differs in the small basal stipules, racemose branching of the inflorescence, eccentric bracteole glands, and samara wing

138

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usually continuous at the base. Only the type species occurs in Venezuela and the flora area. Excentradenia adenophora (Sandwith) W.R. Anderson, Contr. Univ. Michigan Herb. 21: 31. 1997. —Hiraea adenophora Sandwith, Kew Bull. 1951: 33. 1951. Woody vine; leaves opposite, subopposite, or alternate; stipules 0.4–0.8 mm long, borne on petiole at very base; petioles 10–20 mm long; blade of larger leaves 8–15.5 × 4.5–10.3 cm, ovate or elliptic to orbicular, obtuse or rounded and abruptly short-acuminate at

apex; floriferous peduncles 1–2.5 mm long; pedicels 7–10 mm long; anterior sepal eglandular, the lateral 4 biglandular; styles bowed outward, dorsally apiculate at apex; samaras depressed-circular with the nut positioned below the center, 57–66 mm wide, ca. 50 mm high. Near streams, 100–200 m; Delta Amacuro (east side of Río Cuyubiní), Bolívar (southwest of El Manteco on road to San Pedro de las Dos Bocas). Guyana. ◆Fig. 93.

H C B

E

A

D

F

G

Fig. 93. Excentradenia adenophora A–G: —A. Large leaf, adaxial view, ×0.54, and base of petiole with stipules, ×2.7. —B. Flowering branch, ×0.54. —C. Umbel of 4 circinate buds, ×2.7. —D. Base of umbel enlarged to show eccentric glands on 4 bracteoles, ×5.4. —E. Flower with posterior petal uppermost, ×2.7. —F. Anthers, adaxial view (left), side view (right), ×10.8. —G. Distal portion of style, side view, ×10.8. —H. Abaxial view of samara of E. propinqua (similar to that of E. adenophora), ×0.54. ©University of Michigan Herbarium 1997.

Glandonia 139

11. GLANDONIA Griseb. in Mart., Fl. Bras. 12(1): 23. 1858. Shrubs or trees. Leaves bearing abaxial glands; stipules interpetiolar, linear, 9–24 mm long, the adjacent stipules from opposite leaves connate in pairs, the 2 pairs at each node conduplicate and equitant over the apical bud, caducous, the members of a pair often splitting apart before falling off. Inflorescence terminal, usually unbranched, a raceme of short cincinni, the bracts and bracteoles persistent, the lowest bracteole and alternate subsequent bracteoles bearing 1 large eccentric abaxial gland. Flower buds pyramidal, with the conical-galeiform outermost petal completely covering the others. Sepals all biglandular; petals usually described as white, occasionally “yellow.” Stamens 10; filaments densely hirsute; anthers unwinged, tapering distally into 2 sterile awn-like extensions exceeding the slender connective. Ovary of 3 completely connate carpels, with all 3 locules fertile; styles 3, subulate; stigma minute and terminal. Fruit an indehiscent fibrous nut, cylindrical or truncate-conoidal, dry at maturity and without a stony endocarp, containing only 1 locule completely filled by 1 large seed. Amazonian Colombia, Venezuela, and Brazil; 3 species, 1 in Venezuela.

Glandonia williamsii Steyerm., Fieldiana, Bot. 28: 288. 1952. Shrub or small tree 2–8(–15) m tall; blade of larger leaves 10–22 × 3.3–10.3 cm; posterior petal bearing ca. 10 glands on each side of base of limb and distally eglandular; fruits 14–22 × 12–14 mm, bearing 7–9 low, rounded, longitudinal ribs. Seasonally flooded riparian forests of black-water rivers, 100–200 m; Amazonas (Río Atacavi, Río Casiquiare, Río Cuao, Río Guainía, upper Río Orinoco, Río Pasimoni, Río Puruname, Río Sipapo, Río Temi, Río Yatua). Basin of upper Río Negro in Colombia and Brazil. ◆Fig. 94.

Fig. 94. Glandonia williamsii

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12. HETEROPTERYS H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 163. 1821 [1822], nom. cons. Woody vines, shrubs, or small trees. Leaves usually bearing glands; stipules very small, distinct, triangular, borne on or beside base of petiole, or absent. Flowers borne in umbels, corymbs, or pseudoracemes, these single or grouped in racemes or panicles, axillary or terminal; petals mostly yellow or pink. Stamens 10; anthers ± alike, the connective not or hardly exceeding locules (fertile stamens apparently only 6 or 7 in H. maguirei). Ovary with the 3 carpels partially connate, all fertile; styles 3, the apex with a large, usually internal stigma and dorsally rounded, truncate, acute, or hooked, the stigma very rarely terminal. Fruits breaking apart into 3 samaras, each samara having its largest wing dorsal, thickened on the abaxial (lower) edge and (in most species) bent upward, the veins terminating in the thinner adaxial edge; much shorter winglets or crests present on sides of nut in some species; dorsal wing rudimentary in a few species. Mexico, Central America, West Indies, South America (all countries except Chile), western Africa (1 species); at least 125 species, 26 known or expected in Venezuela, 20 of these in the flora area. Key to the Species of Heteropterys 1. 1. 2(1). 2. 3(2).

3.

4(2).

4.

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

Petioles biglandular at base ....................................................................... 2 Petioles eglandular or bearing glands near or above middle. .................. 5 Pedicels sessile ........................................................................................... 3 Pedicels pedunculate .................................................................................. 4 Woody vine; petioles 14–20 mm long; blade of larger leaves 10–15 × 6– 11 cm, tomentose abaxially with the hairs clearly stalked; inflorescence a raceme or panicle of umbels .............................................. H. neblinensis Shrub or slender tree 1–3 m tall; petioles 5–10 mm long; blade of larger leaves 3–9 × 2–5 cm, sericeous or appressed-tomentose or almost glabrous abaxially with the hairs nearly or quite sessile; inflorescence a single umbel terminating a leafy shoot .............................. H. steyermarkii Inflorescence compound, paniculate, with the flowers borne ultimately in umbels or tight corymbs of 4–10; bracteoles eglandular; petals pink or pink and white, with a prominent abaxial wing on the limb; sepals appressed .................................................................................... H. cristata Inflorescence simple, an elongated pseudoraceme of 20–60 flowers; 1 of each pair of bracteoles bearing 1 large eccentric abaxial gland; petals yellow, abaxially smooth; sepals revolute at apex ..................... H. molesta Sepals erect or appressed; petals exposed early and through enlargement of bud, yellow, bronze, brown-maroon, pink, or white .......................... 6 Sepals revolute at apex; petals concealed by sepals during enlargement of bud, yellow ........................................................................................... 10 Mature leaf blades glabrous or glabrate abaxially or only sparsely sericeous, the hairs not apparent without a lens ....................................... 7 Mature leaf blades persistently and obviously sericeous or tomentose abaxially ................................................................................................. 8 Petals yellow, abaxially smooth or with the midvein prominent; anthers loosely sericeous; petioles 12–22 mm long; blade of larger leaves 9–20 × 4.5–12 cm; samaras 60–80 mm long, the nut smooth or rugose and

Heteropterys 141

7.

8(6).

8.

9(8).

9.

10(5). 10. 11(10).

11.

12(10).

12. 13(12). 13. 14(13). 14. 15(13).

without lateral crests or bearing a small crest 1.5 mm wide on each side. ........................................................................................ H. megaptera Petals pink or pink and white, abaxially prominently winged; anthers glabrous; petioles 4–12(–15) mm long; blade of larger leaves 6–12.5 × 3.3– 6.5 cm; samaras 28–37 mm long, the nut bearing on both sides a lateral crest or wing 1–5 mm wide, subentire or dissected into several winglets ..................................................................................................... H. cristata Leaf blades tightly and persistently sericeous abaxially; petals yellow, abaxially carinate; nut of samara with the sides quite smooth; blade of larger leaves 11–20 × 6–11 cm .......................................... H. macrostachya Leaf blades at least initially tomentose abaxially; petals pink or pink and white, 4 or all 5 abaxially prominently winged; nut of samara usually bearing short lateral crests or winglets; blade of larger leaves 4.5– 12.5 × 3–6.5 cm ...................................................................................... 9 Leaf blades eventually glabrate abaxially, soon glabrate and shining adaxially; pedicels 5–9 mm long; bracteoles appressed; stigmas laterally compressed, higher than wide ............................................. H. cristata Leaf blades persistently tomentose abaxially, ± persistently tomentose adaxially and not shining; pedicels 2–3 mm long; bracteoles spreading; stigmas laterally expanded, at least twice as wide as high .......... H. alata Bracts 4.5–8 mm long; bracteoles 3–5 mm long ...................................... 11 Bracts (0.5–)1–3(–4) mm long; bracteoles 0.5–2.5(–3) mm long ............. 12 Blade of larger leaves 23–33(–36) × 8–14(–19) cm, abruptly acuminate at apex; pseudoracemes 6–20 cm long; bracts 3–6 mm wide; bracteoles 3– 4 mm wide; fertile stamens 10; styles 3.8–4.5 mm long, with a well-developed dorsal hook 0.3–0.6 mm long at apex; samara wing 22–37 mm wide ................................................................................................. H. leona Blade of larger leaves 8–10.5 × 4.7–6.7 cm, very broadly obtuse or rounded and sometimes retuse and apiculate at apex; pseudoracemes 2–6(–9) cm long; bracts 1.3–3 mm wide; bracteoles 1–2.5 mm wide; fertile stamens 6 or 7 (as far as known); styles ca. 2.3 mm long, dorsally truncate or rounded at apex; samara wing 11–13 mm wide .................................................................................................. H. maguirei Inflorescence composed of umbels or corymbs of (3)4–6(–8) flowers or of short crowded pseudoracemes with 1–3 pairs of flowers and then a terminal umbel, the axis (excluding the floriferous peduncles and pedicels) up to 1.5 cm long (sometimes to 2.5 cm in H. nervosa) .............................................................................................................. 13 Inflorescence composed of elongated pseudoracemes not terminating in umbels, these mostly over 2.5 cm long, often much longer ................ 18 Leaf blades glabrous abaxially ................................................................ 14 Leaf blades sparsely to densely sericeous or tomentose-sericeous abaxially (older leaves sometimes eventually glabrescent) ............... 15 Branches of inflorescence usually with 1–3 pairs of flowers below terminal umbel; samaras 19–40 mm long........................................... H. nervosa Branches of inflorescence usually without flowers developed below terminal umbel; samaras 40–50 mm long.......................................... H. siderosa Bracts (and often bracteoles) reflexed or revolute; leaf blades sparsely sericeous abaxially, the hairs visible only with a lens .............. H. siderosa

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15.

16(15). 16. 17(16).

17.

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

19.

20(18).

20.

21(20).

21.

22(21).

22.

23(21).

Bracts and bracteoles ascending; leaf blades moderately to densely sericeous or tomentose-sericeous abaxially, the hairs visible without a lens .............................................................................................................. 16 Sepals proximally rusty brown, distally white; petals abaxially persistently sericeous; blade of larger leaves 5–7 cm wide ..... H. dichromocalyx Sepals uniformly dark brown; petals sparsely pilose, especially on margins, to glabrate; blade of larger leaves 2–4.5 cm wide ...................... 17 Woody vine; leaf blades acuminate at apex, the reticulum nearly or quite invisible on adaxial side; petiole of larger leaves 5–11 mm long; inflorescences short-stalked and crowded, mostly shorter than subtending leaves; anther connectives uniformly dark ....................... H. cuatrecasasii Shrub to 4 m tall; leaf blades obtuse or rounded and sometimes emarginate at apex, the reticulum readily visible on both sides; petioles 1– 4 mm long; inflorescences long-stalked and open, often longer than subtending leaves; anther connectives with a dark red spot just above insertion of filament, otherwise yellow ........................................ H. huberi Woody vines .............................................................................................. 19 Shrublets, shrubs, or small trees ............................................................. 20 Bracteoles (1 or both) usually bearing 1 or 2 prominent abaxial glands; petioles sometimes bearing 1 or 2 glands on distal half; samaras usually ca. 3 times as long as nut, the wing usually flabelliform, its abaxial edge often recurved ............................................................... H. orinocensis Bracteoles usually eglandular, rarely with minute marginal glands; petioles eglandular; samaras usually 4–5 times as long as nut or longer, the wing elongated, obovate or falcate, its abaxial edge usually curved or bent abruptly upward ........................................................... H. macradena Leaf blades densely and persistently tomentose to subsericeous abaxially, the hairs short-stalked, moderately sinuous to twisted, their crosspieces 0.5–1 mm long ............................................................ H. quetepensis Leaf blades originally sparsely to moderately sericeous with sessile, straight, tightly appressed hairs 0.15–0.4 mm long but usually soon glabrescent to quite glabrate ............................................................... 21 Leaves obtuse or rounded at apex, opposite, alternate, or whorled, the arrangement often variable on the same stem, seldom strictly decussate; petioles 0–5 mm long, eglandular ....................................................... 22 Leaves mostly narrowly acute or acuminate at apex, strictly decussate (if somewhat obtuse at apex, the petioles mostly bearing 2 sunken glands near middle); petioles 4 mm long or longer, eglandular or bearing 1 or 2 pairs of glands between middle and apex ........................................ 23 Leaf blades moderately to strongly revolute, with the fine reticulum much more prominent adaxially than abaxially; vegetative internodes nearly or quite glabrous; abaxial leaf epidermis papillose (due to protruding starch grains in guard cells), often glaucous ...................... H. oblongifolia Leaf blades nearly or quite flat, with the reticulum about as prominent abaxially as adaxially or more prominent abaxially; vegetative internodes minutely sericeous to glabrate; abaxial leaf epidermis smooth or obscurely papillose, not glaucous ........................................ H. atabapensis Leaf blades with the very fine reticulum equally visible on both sides in dried leaves, the finest subdivisions almost as prominent as the lateral

Heteropterys 143

23.

veins; most petioles bearing 2 sunken glands near middle, and occasionally 2 more near apex; bracts (and at least some bracteoles) deciduous before maturation of fruits; samaras 17–20 × 6–8 mm ............................................................................................ H. ayacuchensis Leaf blades with the reticulum more visible abaxially than adaxially, comprising 3 or 4 orders of prominence from lateral veins to finest and substantially less prominent veinlets; petioles eglandular; bracts and bracteoles mostly persistent; samaras usually 25 × 10 mm or larger ............................................................................................... H. macradena

Heteropterys alata (W.R. Anderson) W.R. Anderson, Contr. Univ. Michigan Herb. 16: 75. 1987. —Heteropterys beecheyana var. alata W.R. Anderson, Mem. New York Bot. Gard. 32: 184. 1981. Woody vine or shrub to 8 m tall; blade of larger leaves 4.5–7.5 × 3–5.2 cm, ovate, rounded or retuse and often mucronate at apex, densely and persistently tomentose on both sides, especially abaxially; flowers borne ultimately in umbels, corymbs, or short crowded pseudoracemes of 4–14; sepals erect or appressed in anthesis; petals pink or pink and white, the lateral 4 abaxially winged, the posterior carinate or with a narrow wing; samaras 18–30 × 7–12 mm, the nut usually bearing on both sides 1–several crests or winglets 0.5–2 mm wide. Open places in lowlands, especially in savannas and on granitic outcrops, 100–300 m; Bolívar (Cerro San Borja along lower Río Suapure), Amazonas (Cerro Parú, vicinity of Puerto Ayacucho). Cojedes, Miranda, Zulia; Colombia east of the Andes, especially common in the Llanos Orientales. Heteropterys atabapensis W.R. Anderson, Mem. New York Bot. Gard. 32: 198. 1981. Shrublet, shrub, or small tree 10 cm to 4 m tall; leaves alternate, opposite, or whorled, the arrangement often variable on the same stem; petioles 0–2 mm long, eglandular; blade of larger leaves 3.5–10(–12) × (0.5–)1– 6.5(–7.2) cm, ovate or elliptic or rarely linear, obtuse or rounded at apex, soon glabrate; flowers borne in pseudoracemes; sepals revolute at apex; petals yellow; samaras 10–23 × 5–10 mm. White-sand savannas, 100–200 m; Amazonas (upper Río Guainía north to Río Atabapo, Río Orinoco, and Río Ventuari, found as far north as Cerro Cuao). Colombia (Vaupés).

For a discussion of the parallel variation found in this species and its morphological and geographical sister Heteropterys oblongifolia, see p. 196 in the 1981 paper where it was described. Heteropterys ayacuchensis W.R. Anderson, Mem. New York Bot. Gard. 64: 227. 1990. Shrub 1–2 m tall; petioles 4–7 mm long, mostly biglandular near middle and occasionally near apex; blade of larger leaves 5– 8.8 × 2–4 cm, narrowly ovate, acute or acuminate or somewhat obtuse at apex, abaxially sericeous to soon ± glabrate, the very fine reticulum equally visible on both sides; flowers borne in pseudoracemes; sepals revolute at apex; petals yellow; samaras 17–20 × 6–8 mm. On and near granitic outcrops, 50–100 m; Amazonas (Puerto Ayacucho south to Morganito). Endemic. Heteropterys cristata Benth., London J. Bot. 7: 131. 1848. Heteropterys carinata Benth., London J. Bot. 7: 133. 1848. Woody vine; petioles (4–)6–12(–15) mm long, usually biglandular near base; blade of larger leaves 6–12.5 × 3.3–6.5 cm, ovate, acute or acuminate at apex, initially sericeous or tomentose, soon glabrate and shining adaxially, soon to eventually glabrate abaxially; inflorescence with the flowers borne ultimately in umbels or tight corymbs of 4–10; sepals erect or appressed in anthesis; petals pink or pink and white, abaxially winged; samaras 28–39 × 13–15 mm, the nut bearing on both sides a crest or wing 1–5 mm wide, subentire or dissected into several winglets. Lowland to lower montane forests, often along rivers, 100–900 m; Bolívar (southwest of Roraima-tepui), Amazonas (Isla Ratón, Puerto Ayacucho, Río Ventuari,

144

M ALPIGHIACEAE

San Juan de Manapiare, Sierra de la Neblina). Colombia, Guyana, Amazonian Peru, Brazil. Heteropterys cuatrecasasii W.R. Anderson, Mem. New York Bot. Gard. 32: 191. 1981. Woody vine; petiole of larger leaves 5–11 mm long; blade of larger leaves 4–9 × 2–4 cm, ovate or elliptic, acuminate at apex, abaxially persistently tomentose-sericeous or eventually glabrescent; inflorescence short-stalked and crowded, the flowers borne ultimately in umbels or corymbs of 4–6(–8); sepals revolute at apex; petals yellow; samaras 21–33 × 9–16 mm. Sabanitas and tepuislope forests, 1100–2000 m; Amazonas (Cerro Huachamacari, Cerro Marahuaka, Cerro Parú). Endemic. Heteropterys dichromocalyx W.R. Anderson, Mem. New York Bot. Gard. 32: 193. 1981. Woody vine; blade of larger leaves 11–12 × 5–7 cm, elliptic, acuminate at apex, abaxially thinly but ± persistently sericeous; flowers borne ultimately in umbels of 3–6; sepals revolute at apex, proximally rusty brown, distally white; petals yellow, abaxially persistently sericeous; samaras unknown. Montane forests, 1300–2000 m; Amazonas (expected on the slopes of Sierra de la Neblina). Brazil (Amazonas: Pico da Neblina). Heteropterys huberi W.R. Anderson, Contr. Univ. Michigan Herb. 16: 78. 1987. Shrub to 4 m tall; petioles 1–4 mm long; blade of larger leaves 5–8 × 2.5–4.5 cm, elliptic, obtuse or rounded and sometimes emarginate at apex, densely and ± persistently sericeous abaxially; inflorescence longstalked and open, the flowers borne ultimately in umbels of 4, often with an additional proximal pair of flowers; sepals revolute at apex; petals yellow, sparsely pilose on margin or glabrate; immature samaras 12– 14 × 5–5.5 mm. Open rocky shrubby places on tepui slopes, 600–800 m; Amazonas (Cerro Aracamuni). Endemic. Heteropterys leona (Cav.) Exell, Cat. Vasc. Pl. S. Tomé 123. 1944. —Banisteria leona Cav., Diss. 9: 424, pl. 247. 1790.

Malpighia reticulata Poir. in Lam., Encycl. Suppl. 4: 8. 1816. —Byrsonima reticulata (Poir.) DC., Prodr. 1: 581. 1824. —Heteropterys reticulata (Poir.) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 2: 54. 1903, non Griseb. in Mart. 1858. —Banisteria reticulata (Poir.) C.B. Rob. in Small, N. Amer. Fl. 25: 138. 1910. Banisteria multiflora DC., Prodr. 1: 589. 1824. —Heteropterys multiflora (DC.) Hochr., Bull. New York Bot. Gard. 6: 277. 1910. Heteropterys africana A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 276. 1840, nom. superfl. Woody vine; blade of larger leaves 23– 33(–36) × 8–14(–19) cm, often smaller near inflorescence, ovate or elliptic to obovate, abruptly acuminate at apex, abaxially thinly sericeous to glabrate; pseudoracemes 6–20 cm long; bracts 5–8 × 3–6 mm; bracteoles 4–5 × 3–4 mm; sepals revolute at apex; petals yellow; styles with a dorsal hook at apex 0.3– 0.6 mm long; samaras 35–50 × 22–37 mm, flabelliform to semicircular, the abaxial edge somewhat to strongly recurved, the nut flattened, trapezoidal, 17–25 mm diameter. Lowland riparian forests, near sea level to 100 m; Delta Amacuro (Caño Arature, Güiniquina, Río Cuyubiní). Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana and adjacent Brazil (Amapá, Pará), west Africa. Heteropterys leona has long been called H. multiflora in South America, but recent critical study by Christiane Anderson has shown that it is indistinguishable from the plant of west Africa, whose name is older and must be taken up for our species. I am grateful to C. Anderson for allowing me to cite her work here prior to its publication. Heteropterys macradena (DC.) W.R. Anderson, Mem. New York Bot. Gard. 32: 202. 1981. —Banisteria macradena DC., Prodr. 1: 590. 1824. Usually a woody vine, occasionally a shrub or small tree to 6 m tall; petioles eglandular; blade of larger leaves (5–)6.5– 16(–18) × (2.5–)3.5–7(–8.5) cm, ovate or elliptic, occasionally obtuse or rounded at apex but mostly acuminate, originally sericeous on both sides, soon glabrate adaxially, the

Heteropterys 145

hairs deciduous or sometimes ± persistent abaxially; bracteoles usually eglandular, rarely with minute marginal glands; flowers borne in a pseudoraceme; sepals revolute at apex; petals yellow; apex of styles usually extended dorsally into a hook up to 0.2 mm long; samaras (20–)25–42(–48) × (8–)10– 15(–18) mm. Mostly in or near forests, often along streams, 50–500(–1300) m; throughout Bolívar and Amazonas. Elsewhere in Venezuela this species, interpreted in a broad sense, occurs in all or most states; Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil. Heteropterys macrostachya A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 275. 1840. Woody vine, rarely a spreading shrub; petioles bearing 2–4 glands at or above middle; blade of larger leaves 11–22 × 6–11 cm, elliptic or slightly ovate or obovate, obtuse, short-acuminate, rounded, or retuse and mucronate at apex, soon glabrate adaxially, densely and persistently metallicsericeous abaxially; inflorescence a raceme or panicle of 4–6-flowered umbels; sepals erect or appressed in anthesis; petals yellow, abaxially carinate; samaras 25–60 × 12–27 mm, the nut smooth-sided. Forests and thickets, especially along streams and rivers, 50–900 m; Bolívar (Río Caura, Río Orinoco, Río Paragua, Río Suapure), Amazonas (Río Cuao, Río Yawarui affluent of Río Matacuni). Apure, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Nueva Esparta, Sucre, Táchira, Yaracuy; rare from Chiapas, Mexico, and Belize south to Nicaragua, ± common in Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Heteropterys maguirei W.R. Anderson, Mem. New York Bot. Gard. 64: 225. 1990. Woody vine or trailing shrub; blade of larger leaves 8–10.5 × 4.7–6.7 cm, elliptic, very broadly obtuse or rounded and sometimes retuse and apiculate at apex, abaxially ± persistently sericeous; pseudoracemes 2– 6(–9) cm long; bracts 4.5–8 × 1.3–3 mm; bracteoles 3–5 × 1–2.5 mm; sepals revolute at apex; petals yellow; styles dorsally truncate or rounded at apex; samaras 26–32 × 11–13 mm. Upland white-sand savannas,

1000–1100 m; Bolívar (Gran Sabana, 194 km north of Santa Elena on road to El Dorado). Western Guyana. The type collection has only 6 or 7 stamens fertile in each flower, whereas all other species of Heteropterys have all 10 stamens present and fertile. It remains to be seen whether that peculiarity will prove to be true of subsequent collections of this species; the type may have been abnormal and atypical. The second known collection is in fruit and sheds no light on this problem. Heteropterys megaptera A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 277. 1840. Heteropterys lasseri W.R. Anderson, Mem. New York Bot. Gard. 32: 180. 1981. Woody vine; petioles 12–22 mm long; blade of larger leaves 9–20 × 4.5–12 cm, broadly elliptic, rounded and abruptly shortacuminate at apex, abaxially sparsely sericeous or glabrate at maturity; inflorescence a panicle of 4–6-flowered umbels or 4-flowered umbels with another pair of flowers below; sepals erect or appressed in anthesis; petals yellow; anthers loosely sericeous; samaras 60–80 × 17–20 mm. Lower montane moist forests, 600–1200 m; Bolívar (La Escalera, Kavanayén, Río Venamo). Sucre (Península de Paria); Guyana, Atlantic Brazil (Bahia, Rio de Janeiro). Heteropterys molesta W.R. Anderson, Contr. Univ. Michigan Herb. 21: 59. 1997. Vine; petioles biglandular at base; blade of larger leaves 7.8–11.3 × 4–5.6 cm, elliptic or ovate, mostly obtuse and mucronulate at apex, adaxially loosely sericeous to glabrescent, abaxially tightly and persistently sericeous; inflorescence an unbranched axillary pseudoraceme of 20–60 flowers; 1 bracteole of each pair with 1 large eccentric abaxial gland; sepals revolute in anthesis; petals yellow; samaras unknown. Roadside thickets?, 50–100 m; Bolívar (between Upata and San Félix). Endemic. This species is isolated in the genus, and until it is collected with fruits we will not even be certain it belongs in Heteropterys. See discussion in the protologue. Heteropterys neblinensis W.R. Anderson, Mem. New York Bot. Gard. 32: 176. 1981.

146

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Woody vine; petioles 14–20 mm long, biglandular at base; blade of larger leaves 10–15 × 6–11 cm, subcircular or broadly ovate or elliptic, rounded and apiculate at apex or abruptly short-acuminate, abaxially densely and persistently sulfur-tomentose; flowers borne ultimately in umbels of 5–15 (–20?); pedicels sessile; sepals appressed in anthesis; petals “bronze” or “pale yellow tinged with bronze”; samaras 32–35 × 11–12 mm. Amazonas (known only from escarpment overlooking Cañón Grande, Sierra de la Neblina, 1700–2000 m). Endemic. Heteropterys nervosa A. Juss. in A. St.Hil., Fl. Bras. Merid. 3: 26. 1832 [1833]. Heteropterys suberosa Griseb., Linnaea 13: 229. 1839. Banisteria mossii C.V. Morton, Proc. Biol. Soc. Wash. 43: 157. 1930. —Heteropterys mossii (C.V. Morton) Cuatrec., Webbia 13: 478. 1958. Woody vine; blade of larger leaves 7–13 (–17) × (2–)3–6.5(–8) cm, elliptic, ovate, or somewhat obovate, acuminate at apex, glabrous; ultimate branches of inflorescence up to 1.5(–2.5) cm long, terminating in umbels of 4–6 flowers usually subtended by 1–3 proximal pairs of flowers; sepals revolute at apex; petals yellow; samaras 19–40 × 8–13 (–16) mm. In lowlands, along rivers and in ± open savannas, often on white sand, 50–300 m; southern Amazonas (Caño Caname, La Esmeralda south to Río Negro and Río Baría, San Antonio). Eastern Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian and central Brazil, Bolivia, Paraguay. The plants to which I have applied this name are rather diverse through their range, and may represent more than one species. Heteropterys oblongifolia Gleason, Bull. Torrey Bot. Club 58: 377. 1931. Shrublet, shrub, or small tree 20 cm to 4 m tall; leaves alternate, opposite, or whorled, the arrangement often variable on the same stem; petioles 0–5 mm long, eglandular; blade of larger leaves 6–12(–13.6) × 1–5 (–6.5) cm, narrowly to broadly elliptic or somewhat ovate or obovate, obtuse or rounded at apex, sparsely sericeous to soon glabrate; flowers borne in pseudoracemes; sepals revolute at apex; petals yellow; samaras 18–27 × 5–11 mm. Lowland savannas,

100–200 m; Amazonas (upper Río Orinoco and Río Guainía south to San Carlos de Río Negro and Río Pasimoni). Endemic. ◆Fig. 97. See note under Heteropterys atabapensis. Heteropterys orinocensis (H.B.K.) A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 276. 1840. —Banisteria orinocensis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 162. 1821 [1822]. Heteropterys acutifolia A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 276. 1840. Heteropterys helicina Griseb. in Mart., Fl. Bras. 12(1): 67. 1858. Woody vine; petioles often biglandular near apex; blade of larger leaves 9–18(–20) × 3.5–7(–8.5) cm, ovate or rarely elliptic, obtuse, acute, or acuminate at apex, thinly sericeous to glabrate; flowers borne in elongated pseudoracemes; one or both bracteoles usually bearing 1 or 2 prominent abaxial glands; sepals revolute at apex; petals yellow; samaras (16–)19–23 × (8–)11–13 mm, usually flabelliform, the abaxial edge often recurved. Common along rivers, 50–200 m; Bolívar (Río Orinoco and tributaries as far north as Caicara), Amazonas (Río Orinoco and Río Negro and their tributaries). Eastern Apure, southern Guárico; Amazonian Colombia, Peru, Brazil, Bolivia. ◆Fig. 96. Some of the larger-leaved plants from Venezuela to which I have applied this name could pass for Heteropterys riparia Cuatrec., Webbia 13: 483. 1958, a species of Amazonian Colombia, Peru, and Brazil. I see no discontinuity that would allow me to recognize two species in Venezuela, which leads me to wonder whether H. riparia will survive careful study elsewhere. Heteropterys quetepensis Steyerm., Fieldiana, Bot. 28: 292. 1952. Shrub or small tree 2–3 m tall; blade of larger leaves 6–11.2 × 3–6 cm, elliptic, mostly obtuse or rounded at apex, sometimes abruptly short-acuminate, abaxially persistently tomentose to subsericeous; flowers borne in pseudoracemes; sepals revolute at apex; petals yellow; samaras 20–28 × 8–11 mm. Dry rocky places with shrubby vegetation, near sea level to 200 m; Bolívar (between Caicara and Los Pijiguaos), Amazonas (northeast of Puerto Ayacucho). Anzoátegui, Apure, Cojedes, Guárico, Sucre.

Heteropterys 147

Heteropterys siderosa Cuatrec., Webbia 13: 476. 1958. Woody vine; blade of larger leaves 10–22 × 4–9 cm, elliptic or slightly obovate, acuminate to rounded at apex, abaxially sparsely sericeous to glabrate; ultimate branches of inflorescence terminating in umbels of 4–6 (–8) flowers; bracts (and often bracteoles) reflexed or revolute; sepals revolute at apex; petals yellow; samaras 40–50 × 12–21 mm. Evergreen lowland forests, 50–200 m; Amazonas (Caño Yagua, Río Casiquiare, Río Cataniapo, Río Cunucunuma, Río Yatua, Tamatama). Amazonian Colombia, Guyana, French Guiana, Amazonian Brazil.

Heteropterys steyermarkii W.R. Anderson, Mem. New York Bot. Gard. 32: 178. 1981. Shrub or slender tree 1–3 m tall; petioles 5–10 mm long, biglandular at base; blade of larger leaves 3–9 × 2–5 cm, elliptic or slightly obovate, mostly obtuse or rounded and often apiculate at apex, rarely acute, sericeous or appressed-tomentose or almost glabrous abaxially with the hairs nearly or quite sessile; inflorescence an umbel of 3–11 flowers terminating a leafy shoot; pedicels sessile; sepals appressed in anthesis; 4 lateral petals brown-maroon, posterior petal white or pale yellow; samaras 18–30 × 8–14 mm. Rocky tepui slopes and marshy meadows, 1200– 1900 m; Amazonas (Cerro Autana, Cerro Cuao, Cerro Sipapo). Endemic. ◆Fig. 95.

Fig. 95. Heteropterys steyermarkii

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Fig. 96. Heteropterys orinocencis

Fig. 97. Heteropterys oblongifolia

Hiraea 149

13. HIRAEA Jacq., Enum. Syst. Pl. 4. 1760. Woody vines, sometimes shrubby. Leaves usually bearing glands distally on petiole or against abaxial base of midrib, and often on margin of blade, the tertiary veins often strongly parallel; stipules borne on petiole, from slightly above base to near apex, usually long and subulate. Inflorescences axillary, usually 1–several umbels of 4–many flowers, the umbels when 4-flowered often borne in a cyme; bracts and bracteoles eglandular; pedicels sessile. Anterior sepal eglandular, 4 lateral sepals biglandular or eglandular, with both forms present in some species; petals yellow, rarely red at maturity, glabrous. Stamens 10, all fertile; anthers ± alike. Ovary with the 3 carpels nearly distinct, all fertile; styles 3, inserted low on ventral face of carpels, the apex with a large internal stigma and dorsally rounded to prominently hooked. Fruit breaking apart into 3 samaras, each butterfly-shaped with 2 discrete lateral wings, these coriaceous and reduced in a few species; dorsal wing small, sometimes reduced to a crest or lost; intermediate winglets or slender projections rarely present. Mexico, Central America, West Indies, and South America (all countries except Chile and Uruguay); at least 47 species, ca. 20 in Venezuela, 10 of these in the flora area. Key to the Species of Hiraea 1.

1. 2(1).

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

4.

5(4).

5.

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

Abaxial leaf hairs stalked, erect, Y- or T-shaped, occasionally with an admixture of V-shaped hairs (sessile with the 2 branches stiff and ascending) .................................................................................................. H. ternifolia Abaxial leaf hairs (if any) nearly or quite sessile and ± appressed .......... 2 Leaf blades abaxially very densely and persistently sericeous, the hairs usually completely concealing the epidermis and producing a golden or silvery-metallic sheen ................................................................. H. faginea Leaf blades abaxially thinly sericeous or subtomentose to glabrate ........ 3 Umbel with a globose or depressed-globose, often dark glandular cushion, 0.7–2 mm diameter, in the center between bracteoles ......................... 4 Umbel without a prominent central cushion, at most with a tiny obscure central body up to 0.3 mm diameter ..................................................... 6 Leaf blades at least shallowly cordate at base; limb of lateral petals 6.5– 8 mm long, 7.5–9 mm wide; petals long-fimbriate with the divisions 0.3–0.7 mm long ....................................................................... H. fimbriata Leaf blades cuneate or rounded at base; limb of lateral petals 4–7 mm long and wide; petals subentire or short-fimbriate with the divisions up to 0.3 mm long ........................................................................................ 5 Lateral veins prominent abaxially, hardly or not at all raised adaxially; limb of lateral petals 5–7 mm long and wide; nut of samara 3–4 mm diameter ............................................................................. H. apaporiensis Lateral veins not or only slightly raised abaxially, almost as prominulous adaxially as abaxially; limb of lateral petals 4–5 mm long and wide; nut of samara ca. 2 mm diameter ................................................. H. tepuiensis Leaf blades bearing obscure or prominent glands on margin, at least distally......................................................................................................... 7 Leaf blades with the margin eglandular. .................................................. 8 Petiole of larger leaves 14–20 mm long; pedicels 14–28 mm long; stipules borne 5 mm or more below apex of petiole ........................ H. steyermarkii

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7.

8(6). 8. 9(8). 9. 10(9).

10.

11(10).

11. 12(9). 12. 13(12). 13. 14(13).

14.

Petiole of larger leaves 7–11 mm long; pedicels 9–13 mm long (–15 mm in fruit); stipules borne at apex of petiole or up to 2 mm below .................................................................................................... H. fagifolia Lateral veins not or only slightly raised abaxially, almost as prominulous adaxially as abaxially; stipules 0.5–1 mm long ..................... H. tepuiensis Lateral veins quite prominent abaxially, hardly or not at all raised adaxially; stipules 1–5 mm long ............................................................ 9 Specimens with fruits (H. celiana is not known in fruit and is not keyed here) ..................................................................................................... 10 Specimens with flowers ........................................................................... 12 Samaras with the dorsal wing absent or represented by a rounded crest 0.2(–0.5) mm high; inflorescence an unbranched umbel, with 1–7 umbels in a vertical array in each axil .............................................. H. affinis Samaras with a small but definite dorsal wing; inflorescence usually a cyme of 3 umbels, with 1–3 cymes in a vertical array in each axil, the cyme occasionally reduced to 1 umbel ................................................ 11 Blade of larger leaves up to 7.5 cm wide; stipules borne at apex of petiole or up to 2 mm below; pedicels up to 13 mm long (–15 mm in fruit) .................................................................................................... H. fagifolia Blade of larger leaves 8.5–17 cm wide; stipules borne slightly above middle of petiole; pedicels 11–20 mm long ........................... H. neblinensis Posterior petal glandular-fimbriate all around margin ................ H. fagifolia Posterior petal with the margin dentate to fimbriate but the divisions not glandular-thickened ............................................................................. 13 Leaf blades cuneate at base; styles dorsally rounded or truncate at apex ......................................................................................................H. celiana Leaf blades rounded or cordate at base; styles with a short (ca. 0.2– 0.3 mm), rounded dorsal hook at apex ................................................ 14 Petals dentate to short-fimbriate, with the divisions up to 0.2 mm long; inflorescence an unbranched umbel, with 1–7 umbels in a vertical array in each axil; petioles up to 11 mm long .............................. H. affinis Petals long-fimbriate, with the divisions 0.5–1 mm long; inflorescence usually a cyme of 3 umbels, with 1–3 cymes in a vertical array in each axil, the cyme occasionally reduced to 1 umbel; petioles 9–19 mm long ............................................................................................... H. neblinensis

Hiraea affinis Miq., Linnaea 19: 133. 1847 [1846]. Large woody vine; stipules 1–2 mm long, borne somewhat below to somewhat above middle of petiole; petioles 4–11 mm long; blade of larger leaves 11–23 × (4–)5–12 cm, rounded or cordate at base, eglandular on margin, abaxially sparsely sericeous or glabrate at maturity; inflorescence a 4-flowered umbel, the umbels solitary in each axil or 2– 7 in a vertical array; pedicels (9–)11–18 mm long; sepals adaxially glabrous; petals pale yellow, dentate or short-fimbriate with the divisions nonglandular; styles with a short

rounded dorsal hook at apex; samaras with lateral wings 22–30 mm wide, 35–56 mm high, membranous, the dorsal wing absent or represented by a rounded crest 0.2(–0.5) mm high. Lowland wet forests along rivers, 50– 300 m; Delta Amacuro (Río Amacuro on the border with Guyana, east-northeast of El Palmar). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Hiraea apaporiensis Cuatrec., Webbia 13: 404. 1958. Woody vine; stipules 0.5–1(–1.5) mm long, borne between middle and apex of petiole;

Hiraea 151

petioles (4–)6–10(–12) mm long; blade of larger leaves 6–16(–21) × 3–6(–8) cm, cuneate or rounded at base, eglandular on margin, abaxially thinly sericeous at maturity, with the lateral veins prominent abaxially and hardly or not at all raised adaxially; inflorescence a 4-flowered umbel, the umbels solitary in each axil or 2 or 3 in a vertical array, each umbel with a dark glandular hemispherical or globose cushion in center between bracteoles; pedicels 8–13 mm long; petals subentire or short-fimbriate with the divisions nonglandular, the limb of lateral petals 5–7 mm long and wide; samaras with nut 3–4 mm diameter, lateral wings 7–11 mm wide, (6–)11–15 mm high. Evergreen lowland forests near rivers, 100–200 m; Amazonas (between San Fernando de Atabapo and the Río Orinoco). Amazonian Colombia, Brazil (western Amazonas). Hiraea celiana W.R. Anderson, Mem. New York Bot. Gard. 32: 249. 1981. Woody vine; stipules 1–1.5 mm long, borne between middle and apex of petiole; petioles 10–18 mm long; blade of larger leaves 12.5–28 × 5–12 cm, cuneate at base, eglandular on margin, abaxially thinly sericeous to glabrate at maturity; inflorescence a 4-flowered umbel, the umbels solitary in each axil or 2–4 in a vertical array; pedicels 16–22 mm long; sepals adaxially glabrous; petals all fimbriate with the divisions nonglandular; styles dorsally rounded or truncate at apex; samaras unknown. Mixed wet forests and scrub savannas, 100–1500 m; Amazonas (Cerro Cuao, Cerro Sipapo). Endemic. Hiraea fagifolia (DC.) A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 258. 1840. —Banisteria fagifolia DC., Prodr. 1: 590. 1824. Woody vine, occasionally described as a shrub; stipules 2.5–5 mm long, borne within 2 mm of apex of petiole; petioles 7–11 mm long; blade of larger leaves 8.5–18.5 × 2.5– 7.5 cm, rounded or subcordate at base, usually bearing several small glands on distal margin or occasionally eglandular, abaxially mostly glabrate at maturity except for sericeous midrib and sometimes the lateral veins; inflorescence a single axillary cyme of 3 4-flowered umbels; pedicels 9–13 mm long

(–15 mm in fruit); posterior petal glandularfimbriate all around margin; samaras with lateral wings 13–20 mm wide, 20–30 mm high, membranous. Moist lower montane forests, 200–300 m; Bolívar (Altiplanicie de Nuria). Lara, Sucre; southeastern Mexico, Central America, Trinidad, all countries of South America except Chile and Uruguay. See discussion below under Hiraea faginea. The specimens that I have called H. fagifolia sens. lat. came from Delta Amacuro (Río Amacuro) and Bolívar (upper Río Caura, Río Venamo). Hiraea faginea (Sw.) Nied., Verz. Vorles. Königl. Lyceums Hosianum Braunsberg 1906/07: 16. 1906. —Malpighia faginea Sw., Prodr. 74. 1788. —Hiraea swartziana A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 258. 1840, nom. superfl. Hiraea chrysophylla A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 258. 1840. Hiraea fulgens A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 258. 1840. Hiraea fulgens var. demerarensis A. Juss., Arch. Mus. Hist. Nat. 3: 572. 1843. —Hiraea demerarensis (A. Juss.) Nied., Verz. Vorles. Königl. Lyceum Hosianum Braunsberg 1906/07: 15. 1906. Woody vine, sometimes shrubby; stipules 2.5–5 mm long, borne between middle and apex of petiole; petioles 4–9(–11) mm long; blade of larger leaves (6–)8–15(–20) × 3–6 (–8) cm, rounded or subcordate at base, usually bearing several small glands on distal margin, abaxially densely and usually persistently sericeous, the straight, appressed hairs giving lamina a golden or silvery-metallic sheen; inflorescence a single axillary cyme of 3 4-flowered umbels, sometimes reduced to a single umbel; pedicels (7–)8–11 (–15) mm long; posterior petal glandularfimbriate all around margin; samaras with lateral wings 5–15 mm wide and high, trapezoidal or rectangular, entire or lobed, usually coriaceous, often irregularly reduced. Riparian forests, near sea level to rarely as high as 200 m; Delta Amacuro (Caño Araguao, Caño Güiniquina, Curiapo), Bolívar (Río Caura, Río Cuyuní, Río Paragua), Amazonas (Isla Ratón, Río Casiquiare, Río Mavaca, Río Metacuni, Río Ocamo, upper Río Orinoco, Río Siapa). Ap-

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parently not known from northern Venezuela; Nicaragua, Costa Rica, Panama, Lesser Antilles, Colombia, Guyana, Suriname, French Guiana, Brazil (north of the Amazon). ◆Fig. 98. Hiraea faginea and H. fagifolia are very similar, and presumably closely related. Most specimens can immediately and unequivocally be assigned to one or the other, because in H. faginea the leaf blades are abaxially very densely and persistently metallic-sericeous, with the hairs completely hiding the epidermis, and in H. fagifolia the leaf blades are abaxially glabrous (or very early glabrate) except for the persistently sericeous midrib and sometimes the lateral veins. Moreover, the samara in H. faginea is usually ± coriaceous and reduced, often very irregularly so, suggesting adaptation for dispersal by water; most fruiting collections of H. fagifolia have membranous, butterflyshaped samaras obviously adapted for dispersal by wind. The two species have overlapping ecological requirements, but H. faginea seldom ventures much above sea level or away from rivers while H. fagifolia is much more likely to be found in upland forests, just as one would expect of a plant that has retained dispersal by wind. H. fagifolia has a much wider geographic range than H. faginea, and through much of their ranges the two species are allopatric, e.g., H. fagifolia is common in Peru, Bolivia, and Paraguay, while H. faginea is common in southern Venezuela and adjacent Brazil where H. fagifolia is hardly known. Their ranges do overlap, especially in the Venezuelan Guayana and the Guianas, and in those areas they seem to hybridize. I do not consider that sufficient reason to combine two taxa that are mostly distinct and distinctive, but the occasional intermediate plants do pose a problem for the users of this flora, because they will not key well. The putative hybrids have been found on the Río Caura and near the Río Cuyuní in Bolívar, in areas where “typical” H. faginea occurs. When the leaf blades are abaxially very thinly sericeous, so as to resemble most closely H. fagifolia, I am calling the intermediates H. fagifolia sens. lat. When the leaf blades are more densely sericeous, but with hairs not dense enough to hide the epidermis, I am using the name H. faginea sens. lat.

Hiraea fimbriata W.R. Anderson, Mem. New York Bot. Gard. 32: 245. 1981. Woody vine, occasionally described as a shrub 2–4 m tall; stipules 1–2.5 mm long, borne between middle and apex of petiole; petioles 4–10 mm long; blade of larger leaves 8–22 × 4–10.5 cm, cordate at base, eglandular on margin, glabrate at maturity or with the midrib persistently sericeous; inflorescence a 4-flowered umbel, the umbels solitary in each axil or 2–4 in a vertical array, each umbel with a dark glandular hemispherical cushion in center between bracteoles; pedicels 7–14 mm long; petals longfimbriate with the divisions nonglandular, the limb of lateral petals 6.5–8 mm long, 7.5– 9 mm wide; mature samaras unknown. Riparian, seasonally flooded forests, 50–100 m; Bolívar (middle Río Orinoco and tributaries south of Río Caura), Amazonas (middle Río Orinoco and tributaries north of Río Sipapo). Anzoátegui, Apure; Colombia (Vichada: Río Meta). Hiraea neblinensis W.R. Anderson, Contr. Univ. Michigan Herb. 21: 66. 1997. Large woody vine; stipules 2–3 mm long, borne slightly above middle of petiole; petioles 9–19 mm long; blade of larger leaves 13.5–31 × 8.5–17 cm, shallowly cordate at base, eglandular on margin, abaxially mostly glabrate at maturity or with scattered hairs especially on midrib; inflorescence a cyme of 3 4-flowered umbels, sometimes reduced to a single umbel, the cymes solitary in each axil or 2 or 3 in a vertical array; pedicels 11–20 mm long; sepals adaxially sparsely sericeous; petals all long-fimbriate with the divisions 0.5–1 mm long and nonglandular; styles with a rounded dorsal hook ca. 0.2–0.3 mm long at apex; immature samaras with lateral wings 18 mm wide, 30 mm high. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma). Endemic. Hiraea steyermarkii W.R. Anderson, Contr. Univ. Michigan Herb. 21: 69. 1997. Woody vine; stipules 4.5–6 mm long, borne between base and middle of petiole or slightly higher; petioles 14–20 mm long; blade of larger leaves 14–23.5 × 7–13.7 cm, cuneate to obtuse at base, bearing a series of large glands on distal 1/2–2/3 of margin, abaxially nearly glabrate at maturity except

Hiraea 153

for sericeous midrib; inflorescence a cyme of 3–7 4-flowered umbels, the cymes solitary in each axil or 2, one above the other; pedicels 14–28 mm long; posterior petal fimbriate all around margin with the divisions ± glandular-thickened, especially proximally; mature samaras unknown. Lowland forests, 100– 300 m; Bolívar (east of Las Chicharras, Río Asa). Endemic. The collection from near Las Chicharras is sterile and has much larger leaves than the type, and may represent a different species; more collections are needed to shed light on this problem. Hiraea tepuiensis Steyerm., Fieldiana, Bot. 28: 293. 1952. Woody vine, rarely described as a small tree; stipules 0.5–1 mm long, borne at or above middle of petiole; petioles 6–15 mm long; blade of larger leaves 6–15 × 3–6(–7) cm, cuneate or rounded at base, eglandular on margin, loosely sericeous or subtomentose to glabrescent except persistently sericeous on abaxial midrib, with lateral veins not or only slightly raised abaxially, almost as prominulous adaxially as abaxially; inflorescence a 4-flowered umbel, the umbels solitary in each axil or 2–5 in a vertical array, each umbel without a prominent cushion in center between bracteoles, rarely with a small glandular knob; pedicels 8–15 mm long; petals erose or short-fimbriate with the divisions nonglandular, the limb of lateral petals 4–5 mm long and wide; samaras with nut ca. 2 mm diameter, the lateral wings 6– 10 mm wide, 13–18 mm high. Moist montane to lower montane forests, often gallery forests in areas of savannas, (500–)700–1800 m; common in southeastern Bolívar (Gran Sabana and Macizo del Chimantá), Amazonas (Cerro Aratitiyope, Cerro Parú). Sucre. ◆Fig. 99. Hiraea ternifolia (H.B.K.) A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 257. 1840. —Malpighia ternifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 146. 1821 [1822]. Hiraea platytriphylla Hochr., Bull. New York Bot. Gard. 6: 276. 1910. Hiraea bifurcata W.R. Anderson, Mem. New York Bot. Gard. 32: 242. 1981. Woody vine forming thickets, shrub to 2.5 m tall, or small tree to 5 m tall; leaves ternate or decussate; stipules (1.5–)2–6 mm

long, borne from above base to near apex of petiole; petioles 5–11(–19) mm long; blade of larger leaves 7–16(–18.5) × (3.5–)5–12 cm, broadly cuneate, rounded, or cordate at base, bearing several small glands on distal margin, adaxially densely velutinous with Yshaped hairs to glabrescent in age, abaxially densely and persistently velutinous over whole surface with Y-shaped hairs; inflorescence usually an axillary cyme of 3 or 7 4flowered umbels, sometimes a single umbel, sometimes 2 cymes, one above the other; pedicels 13–30(–35) mm long; posterior petal usually glandular-fimbriate all around margin; samara with the lateral wings 13–20 mm wide, 22–30 mm high, membranous. Open, dry forests on granitic outcrops, 50– 400 m; Bolívar (Río Orinoco lowlands southward from Caicara), Amazonas (Río Orinoco lowlands northward from Samariapo). Mérida; Panama, Colombia. Hiraea ternifolia is the oldest name in a difficult complex of species that extends from Mexico to eastern Brazil; species limits and the correct application of names in that complex are not at all clear. The description above applies to H. ternifolia sens. str. as I understand it. A second taxon in the complex, rather easily distinguished from H. ternifolia sens. str., also occurs in the flora area. It differs from the above description in these ways: adaxial surface of leaf blade initially subsericeous, the hairs very short-stalked with a sinuous, ± horizontal crosspiece often 1–1.5 mm long, glabrescent in age; abaxial surface of blade initially velutinous with Y- or T-shaped hairs but glabrescent in age with some hairs usually persistent on midrib and principal lateral veins. In addition, the leaf blade tends to be more deeply cordate at base and is often notably bullate. Such plants have been collected in forests, savannas, and thickets, 100–200 m, northwestern Bolívar (Caicara to El Burro) and northeastern Bolívar (El Palmar, El Miamo, north of Tumeremo, Reserva Forestal Imataca). I have seen similar plants from Zulia. For the present I am calling such plants Hiraea aff. ternifolia (H.B.K.) A. Juss. There are several published names that may eventually prove to be available for them, but I do not understand this group well enough to be willing either to apply one of those names to this taxon or to propose a new name in the complex at this time.

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Fig. 98. Hiraea faginea

Fig. 99. Hiraea tepuiensis

14. JUBELINA A. Juss. in Deless., Icon. Sel. Pl. 3: 19, pl. 32. 1837 [1838]. Woody vines. Leaves with the petiole eglandular; stipules small, borne on base of petiole; blade bearing impressed abaxial glands, the lateral veins interconnected by ± parallel tertiary veins. Inflorescences axillary and terminal, decompound, thyrsiform, containing much-reduced bract-like leaves below the floriferous bracts, the flowers borne ultimately in umbels of 4 or corymbs of 6; bracts and bracteoles large, pubescent on both sides, persistent. Sepals narrowly ovate, oblong, or obovate, the anterior eglandular, the lateral 4 each biglandular or (in our species) bearing 1 large abaxial gland; petals yellow or (in our species) pink or pink and white. Stamens 10, glabrous. Ovary of 3 carpels adnate to a common axis; styles 3, the apex with a large internal stigma and dorsally truncate or short-hooked. Fruits breaking apart into 3 samaras, each bearing a semicircular dorsal wing and 2 large lateral wings confluent at base, each lateral wing with a sterile cavity in its base parallel to the fertile locule. Central America, Colombia, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil; 6 species, 2 in Venezuela, both in the flora area. See W.R. Anderson, 1990 [The taxonomy of Jubelina (Malpighiaceae), Contr. Univ. Michigan Herb. 17: 21–37].

Jubelina 155

Key to the Species of Jubelina 1.

1.

Stems subsericeous or appressed-tomentose, the limb of hairs at right angles to the stalk; leaf blades bearing abaxially on each side 1 or 2 or occasionally 3 glands at base and 1–4 glands distally in a single row; calyx glands revolute at apex; samaras with lateral wings flat and lacking wings or winglets between them and dorsal wing, or at most a single flat crest or winglet 1–12 mm wide parallel to dorsal wing ........................................................................................... J. grisebachiana Stems velutinous, the hairs with a long fusiform stalk and very short erect branches; leaf blades bearing abaxially (4)5–15 scattered glands on each side; calyx glands attached for their whole length; samaras with strongly plicate or corrugated lateral wings consisting of an outer membranous wing, an inner dentate wing parallel to and hiding the central dorsal wing, and transverse winglets between them .................................................................................................. J. magnifica

Jubelina grisebachiana W.R. Anderson, Contr. Univ. Michigan Herb. 17: 30. 1990. —Bejuco de iguana, Bejuco de sapo. Jubelina bracteosa auctt. non Mascagnia bracteosa Griseb. in Mart., Fl. Bras. 12(1): 97. 1858.

Fig. 100. Jubelina grisebachiana

Woody vine; stems subsericeous or appressed-tomentose; petioles 14–23 mm long; blade of larger leaves 13–18 × 7–12 cm, broadly elliptic, rounded or abruptly shortacuminate at apex, persistently velutinous to tomentose on both sides or sometimes glabrescent adaxially; bracts and bracteoles

156

M ALPIGHIACEAE

pink, 4–6.5 × 1.5–2(–2.5) mm, narrowly obovate or spatulate; pedicels 5–7 mm long; sepals 4.5–6.5 × 1–1.5 mm, narrowly obovate, reflexed, distally inflated, the lateral 4 all bearing 1 large abaxial gland; petals pink, the lateral 4 sparsely to densely sericeous abaxially, the posterior glabrous; samaras elliptic, 60–90 × 25–40 mm, with the lateral wings flat. Open areas and secondary forests, 100–200 m; Amazonas (between Solano and San Carlos de Río Negro and San Simón de Cocuy). Basins of upper Río Negro and Río Vaupés in Colombia and Brazil. ◆Fig. 100. Jubelina magnifica W.R. Anderson, Mem. New York Bot. Gard. 32: 228. 1981. Woody vine; stems velutinous; petioles 9–

20(–25) mm long; blade of larger leaves 18– 28 × 10–18 cm, broadly ovate or elliptic, abruptly acuminate at apex, persistently velutinous on both sides; bracts and bracteoles pink, 6–8 × 3–4 mm, obovate or elliptic; pedicels 8–15 mm long; sepals 6.5–7 × 1.8–2.5 mm, narrowly obovate or oblong, reflexed, distally inflated, the lateral 4 all bearing 1 large abaxial gland; lateral petals pink, abaxially sericeous, posterior petal white with pink tints, glabrous; samaras subcircular, 40–50(–70) mm diameter, with the lateral wings strongly plicate or corrugated. Nonflooded riparian forests, 100–200 m; Amazonas (Río Baría, Río Mawarinuma, Río Siapa, Río Yaciba, Río Yatúa, San Carlos de Río Negro). Endemic.

15. LOPHANTHERA A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 328. 1840. Shrubs or trees, the stems often containing white latex. Leaves bearing glands on petiole or abaxial surface of blade or both; stipules intra- and slightly epipetiolar,

Fig. 101. Lophanthera longifolia

Lophopterys 157

2 3

/ to completely connate. Flowers borne in a usually terminal thyrse composed of few-flowered cincinni or dichasia or a pseudoraceme; 1 or both bracteoles usually bearing 1 large gland; petals pink or yellow, entire or minutely denticulate. Stamens 10, glabrous; anthers alike, their outer locules bearing dark longitudinal wings. Ovary of 3 fertile carpels connate along a central axis; styles 3, slender and subulate with minute terminal stigmas. Fruits breaking apart into 3 dry, unwinged, 1-seeded cocci, the mericarps indehiscent or slightly dehiscent along the keel but not enough so to release the spheroidal seed. Costa Rica, Venezuela, Brazil, Bolivia; 5 species, 1 in Venezuela. Lophanthera longifolia (H.B.K.) Griseb. in Mart., Fl. Bras. 12(1): 25. 1858. —Galphimia longifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 173. 1821 [1822]. Shrub or small tree 1–6(–10) m tall; stipules (6–)7–10 mm long; petioles (10–)14– 30 mm long, usually bearing 2(–4) glands near middle; blade of larger leaves 12–30 × 4–10 cm, usually bearing few to many small abaxial glands; inflorescence 12–35 cm long, usually pendulous, the cincinni comprising

1–4(–10) flowers, 1 bracteole of each pair terminating in a sessile or stalked gland; sepals all biglandular; petals yellow; ovary glabrous; cocci 7–9 × 3–4 mm, glabrous, subcylindrical, the proximal half filled with aerenchyma, the distal half containing the seed. Along rivers, 100–200 m; Amazonas (from Isla Ratón and La Esmeralda on the upper Río Orinoco to San Simón de Cocuy on the Río Negro). Amazonian Brazil and Bolivia. ◆Fig. 101.

16. LOPHOPTERYS A. Juss. in Deless., Icon. Sel. Pl. 3: 18, pl. 29. 1837 [1838]. Woody vines or shrubs. Leaves densely and persistently sericeous abaxially; stipules minute or absent. Inflorescence paniculate (simple in 1 species not found in the flora area), the flowers borne ultimately in pseudoracemes. Anterior sepal eglandular, the 4 lateral sepals rarely eglandular, usually each bearing a single, very large, circular or elliptic, radially lineate gland; petals yellow, glabrous or very sparsely sericeous abaxially. Stamens 10. Ovary of 3 free carpels borne on a short pyramidal torus; styles 3, stout, with large internal stigmas. Fruits breaking apart into 3 samaras, each bearing a relatively short, flabellate or trapezoidal dorsal wing and 2 much longer, narrow, forward-pointing lateral wings 3 or more times as long as wide (lateral wings lacking in 1 species not found in the flora area). Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 8 species, 2 in Venezuela, both in the flora area. Key to the Species of Lophopterys 1.

1.

Peduncles none or up to 1 mm long in fruit; blade of larger leaves 16–35 × 10–23 cm, obovate; petiole of larger leaves 20–60 mm long; pseudoracemes containing 10–50 flowers; samaras with nut 9–11 mm diameter, lateral wings 45–62 × 10–18 mm .................................... L. euryptera Peduncles well developed, 1.7–9 mm long; blade of larger leaves 8.9– 19.8 × 3.4–10 cm, elliptic or ovate (i.e., widest at or below the middle); petiole of larger leaves 10–18(–22) mm long; pseudoracemes containing (2–)4–28 flowers; samaras with nut 3.4–4.5(–6) mm diameter, lateral wings 15–26 × 4–7 mm ................................................................ L. inpana

Lophopterys euryptera Sandwith, Kew Bull. 1951: 34. 1951. Woody liana up to 35 m long. Evergreen lowland to lower montane forests, 50–400 m;

Delta Amacuro (Río Acure, Río Grande east of El Palmar), Bolívar (upper Río Cuyuní). Northwestern Guyana. ◆Fig. 102.

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M ALPIGHIACEAE

Fig. 102. Lophopterys euryptera

Lophopterys inpana W.R. Anderson, Contr. Univ. Michigan Herb. 17: 46. 1990. Woody vine or shrub up to 4 m tall. Evergreen lowland forests, roadside thickets, and

open savannas among sandstone outcrops, 100–200 m; Amazonas (southeast of San Fernando de Atabapo). Amazonian Peru, Brazil, and Bolivia.

17. MALPIGHIA L., Sp. Pl. 425. 1753. Shrubs or small trees. Leaves usually bearing 2(–6) glands abaxially on blade, the petiole eglandular; stipules small, borne on stem between petioles. Inflorescence an axillary pseudoraceme congested into a dense corymb or umbel; bracts and bracteoles eglandular; pedicels pedunculate; petals pink, pale purple, or white. Sta-

Malpighia 159

mens 10, all fertile, glabrous; anthers alike or the 2 opposite the posterior-lateral petals larger. Ovary with the 3 carpels completely connate, the 3 locules all fertile; styles 3, the apex with a large internal or subterminal stigma and dorsally rounded, truncate, or hooked. Fruit a fleshy red or orange drupe (or berry), with 3 pyrenes united in the center or free at maturity but then usually retained in a common fleshy exocarp. Mexico, West Indies, Central America, Colombia, Venezuela, Ecuador, Peru; 43 species, 2 in Venezuela, both in the flora area. Key to the Species of Malpighia 1.

1.

Leaves most often rounded or obtuse at apex and often emarginate and apiculate, but sometimes acute and rarely slightly acuminate, some pairs crowded in dense shoots with very short internodes, others separated by much longer internodes; umbels sessile or raised on a stalk 1– 3(–5) mm long and containing 2–4 flowers; petals cuneate at base of limb, often sparsely sericeous abaxially on claw and in middle of limb, the lateral 4 with a narrow abaxial keel; styles divergent from the base, the posterior 2 bowed outward and then ascending; filaments opposite posterior-lateral petals notably thicker than the other 8; calyx glands 6–10 ...................................................................................... M. emarginata Leaves usually short- to long-acuminate at apex, occasionally acute, evenly spaced, successive pairs separated by well-developed internodes; umbels or corymbs raised on a stalk (2–)3–12(–22) mm long and containing (3)4–10(–12) flowers; petals rounded or truncate at base of limb, abaxially glabrous, smooth; styles ± straight, parallel or divergent distally; filaments ± alike in thickness; calyx glands 6 ...... M. glabra

Malpighia emarginata DC., Prodr. 1: 578. 1824. —Acerola, Cereza, Semeruco. Malpighia punicifolia auctt. non L. 1762. Shrub or small tree 2–6 m tall; petioles (1–)2–4 mm long; blade of larger leaves 2.5–7 × 1.4–3.3 cm (up to 10 × 5 cm in cultivated plants), ovate, elliptic, or obovate, bearing 2(–4) abaxial glands in the proximal third, sparsely sericeous to glabrate; petals pink or purplish (in age?); styles with distinctly internal stigmas, dorsally truncate or apiculate at apex; fruits red, up to 17 × 22 mm. Dry areas with xerophytic vegetation, near sea level to 200 m; Delta Amacuro (Tucupita, cultivated), Bolívar (Ciudad Bolívar, Las Trincheras, Parque Cachamay near Río Caroní, Tumeremo, probably all cultivated or escaped from cultivation). Common in the coastal lowlands of northern Venezuela, where perhaps native, and elsewhere cultivated; apparently native from Mexico to Honduras, elsewhere in Central America and the West Indies probably escaped from cultivation, apparently native in Colombia(?) and coastal Ecuador.

Widely cultivated for the edible fruit, rich in vitamin C, and readily naturalized. Malpighia glabra L., Sp. Pl. 425. 1753. —Cereza, Guayabito rojo, Semeruco. Malpighia punicifolia L., Sp. Pl. ed. 2, 609. 1762. Shrub or small tree 1–6 m tall; petioles 1– 3 mm long; blade of larger leaves 3–10 × 1.5– 5 cm, narrowly to broadly elliptic or ovate, bearing (0–)2–4 abaxial glands in proximal fourth, glabrous or very sparsely sericeous; petals pink or pale purple; styles truncate at apex with the stigma slightly internal or apparently terminal; fruits red, 7–10 × 10–13 mm. Relatively dry deciduous or semideciduous forests, occasionally in the understory of lowland evergreen forest, 200–300 m; Bolívar (vicinity of La Paragua). Northwestern Venezuela from Zulia to Miranda; U.S.A. (southern Texas), Mexico, Central America, Greater Antilles, Colombia, Ecuador, Peru (cultivated, native too?). ◆Fig. 103.

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Fig. 103. Malpighia glabra

18. MASCAGNIA (Bertero ex DC.) Colla, Hortus Ripul. 85. 1824. —Hiraea sect. Mascagnia Bertero ex DC., Prodr. 1: 585. 1824. Vines, mostly woody (a few species from outside the flora area are shrubs or small trees). Leaves usually bearing glands; stipules small, distinct, triangular, borne between petioles or on base of petiole. Inflorescence mostly axillary or terminal pseudoracemes, sometimes congested and reduced to form corymbs or umbels, single or grouped in panicles; floriferous peduncles usually well developed. Petals yellow or yellow and orange or yellow turning red, or pink, lilac, or white. Stamens 10; anthers ± alike. Ovary with the 3 carpels connate along a central axis, all fertile; styles 3, the apex usually with a large internal stigma and dorsally rounded, truncate, acute, or short-hooked; stigma nearly or quite terminal in a few species. Fruits breaking apart into 3 samaras, each samara having its largest wings lateral, 2 discrete wings or a single wing continuous at the base or at both base and apex; dorsal wing small, sometimes reduced to a crest or lost; intermediate winglets present or absent; wings reduced or rudimentary in a few species. Central America, West Indies, Mexico, South America (all countries except Chile and Uruguay); ca. 60 species, 16 in Venezuela, 15 of these in the flora area. As I have noted in previous publications, Mascagnia as recognized here is a heterogeneous and probably unnatural genus, some of whose species will very likely have to be removed eventually to segregate genera. The species in our flora that will surely remain in Mascagnia even after that segregation are M. cynanchifolia, M. dissimilis, M. divaricata, M. eggersiana, M. ovatifolia, M. schunkei, and M. sepium.

Mascagnia 161

Key to the Species of Mascagnia based on flowering material 1. 1. 2(1). 2. 3(2).

3. 4(2).

4.

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

6.

7(6).

7.

8(6).

8. 9(8).

9.

Petals abaxially ± densely and persistently tomentose or sericeous ....... 2 Petals glabrous or bearing a few abaxial hairs ......................................... 5 Margin of bracteoles and sepals with a row of stalked clavate or capitate glands ..................................................................................................... 3 Margin of bracteoles and sepals eglandular ............................................. 4 Leaf blades originally sericeous on both sides with sessile appressed hairs, glabrate at maturity or the hairs persistent abaxially and often on adaxial midrib ............................................................... M. glandulifera Leaf blades velutinous to glabrate adaxially, persistently velutinous abaxially with the prominently stalked hairs T- or Y-shaped ..... M. surinamensis Petals lilac or pink; leaf blades abaxially persistently sericeous and bearing (1–)several small glands in a row parallel to but set in from the margin; sepals completely concealing petals in bud, revolute in anthesis; ultimate units of inflorescence ± elongated pseudoracemes 2– 9(–15) cm long ...................................................................... M. macrodisca Petals yellow; leaf blades nearly or quite glabrate at maturity, bearing many tiny impressed glands on margin or on adaxial surface of revolute margin, abaxially eglandular between midrib and margin; sepals leaving outermost petal exposed during enlargement of bud, appressed in anthesis; ultimate units of inflorescence congested or corymbose pseudoracemes up to 1.5 cm long .................................... M. sinemariensis Petals pink, lilac, or white, or a combination of those colors ................... 6 Petals yellow or yellow turning red in age .............................................. 10 Inflorescence “simple,” i.e., each pseudoraceme axillary to a well-developed vegetative leaf; bracteoles eglandular or 1 or both bearing 1 or 2 small abaxial glands ........................................................................... 7 Inflorescence usually “compound,” i.e., pseudoracemes grouped in terminal or axillary panicles containing only tiny bracts or much-reduced leaves; bracteoles eglandular ................................................................ 8 Leaf blades initially sericeous adaxially, usually soon glabrate; anthers glabrous; flowers ± evenly distributed along axis of pseudoraceme; inflorescence sericeous.............................................................. M. schunkei Leaf blades persistently velutinous adaxially; anthers sericeous; flowers mostly congested in distal half of axis of pseudoraceme; inflorescence tomentose [petal color unknown; placed here on the basis of similarity in other characters to M. schunkei] .................................. M. cynanchifolia Anthers pilose; dried leaf blades smooth on adaxial surface, the reticulum not raised and hardly visible; petioles usually bearing 2–4 glands near middle, or eglandular ............................................................ M. divaricata Anthers glabrous (rarely very sparsely pilose); fine reticulum prominent on adaxial surface in dried leaf blades; petioles eglandular ................ 9 Leaf blades with a discolored band (0.5–)1–2 mm wide all around margin, the band reddish in dried leaves; pedicels quite glabrous or initially sericeous at very apex but usually soon glabrate; petioles 11–20 mm long; ovary and immature fruit sparsely to moderately sericeous with straight appressed hairs ......................................................... M. dissimilis Leaf blades uniformly green; pedicels initially loosely sericeous their whole length (sometimes only sparsely so), eventually ± glabrescent;

162

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10(5).

10.

11(10).

11.

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

13.

14(13).

14.

petioles 8–11(–18) mm long; ovary and immature fruit densely subtomentose with loose sinuous ± spreading hairs ...................... M. ovatifolia Mature, full-sized leaves persistently golden-, silvery-, or brown-sericeous abaxially, the hairs so dense and appressed as to completely conceal epidermis and usually give leaf a metallic sheen; glands of leaf blade, if present, borne on very margin ............................................................ 11 Mature, full-sized leaves abaxially glabrate or variously sericeous, velutinous, or tomentose, but the hairs never dense enough to completely conceal epidermis; glands of leaf blade, if present, borne on abaxial surface between midrib and margin ...................................... 13 Petioles biglandular near or above middle; anthers 0.5–0.7 mm long, glabrous; calyx glands (if present) 0.5–1.5 mm long; petals with limb up to 2.5 mm long; floriferous bracts eglandular ........................ M. poeppigiana Petioles biglandular at or just above base; anthers 1–1.5 mm long, usually sericeous between locules and bearing a few abaxial hairs; calyx glands 1.7–2.5 mm long; petals with limb 3.5–5 mm long; floriferous bracts often bearing 1 or 2 large glands, or eglandular ................................... 12 Leaf blades abaxially golden- or silvery-sericeous, with the reticulum ± prominent ................................................................................ M. stannea Leaf blades abaxially dark brown-sericeous, with the reticulum ± obscured by outer layer of hairs ................................................... M. castanea Sepals completely concealing petals until flower opens, revolute afterwards; calyx glands 10 (all 5 sepals biglandular); petals yellow turning red in age; inflorescence densely and persistently sericeous or tomentose-sericeous with mostly white hairs, the overall aspect gray ..................................................................................................... M. liesneri Sepals leaving outermost petal(s) exposed during enlargement of bud, appressed after flower opens; calyx glands 8 (lateral 4 sepals biglandular, anterior sepal eglandular); petals persistently yellow; inflorescence bearing yellowish, brownish, or reddish hairs, or glabrate, the overall aspect green or brown .......................................................................... 14 Leaf blades abaxially velutinous to subsericeous, occasionally glabrescent but the dark brown or reddish hairs persistent at least on midrib; stems velutinous to glabrescent in age; petiole of larger leaves 8– 18(–22) mm long; pedicels sericeous or subvelutinous, with some hairs persistent in fruit at least on distal half; ovary densely tomentose-sericeous; samaras sparsely sericeous ............................................. M. sepium Leaf blades abaxially glabrous or soon glabrate; stems soon glabrate; petioles 3–8 mm long; pedicels glabrous; ovary sparsely sericeous; samaras soon quite glabrate ............................................................... M. eggersiana Key to the Species of Mascagnia based on fruiting material

1. 1. 2(1).

Lateral wings of samaras discrete, i.e., divided to nut at base and apex ................................................................................................................ 2 Lateral wing of samaras continuous at base, continuous at apex or incised part way or completely to nut ............................................................... 6 Mature, full-sized leaves thinly sericeous to glabrate abaxially, the hairs never dense enough to completely conceal epidermis .......................... 3

Mascagnia 163

2.

Mature, full-sized leaves persistently golden-, silvery-, or brown-sericeous abaxially, the hairs so dense and appressed as to completely conceal epidermis, usually giving leaf a metallic sheen .................................... 4 3(2). Leaf blades abaxially persistently sericeous and bearing (1–)several small glands in a row parallel to but set in from margin; ultimate units of inflorescence ± elongated pseudoracemes 2–9(–15) cm long .... M. macrodisca 3. Leaf blades nearly or quite glabrate at maturity, bearing many tiny impressed glands on margin or on adaxial surface of revolute margin, abaxially eglandular between midrib and margin; ultimate units of inflorescence congested or corymbose pseudoracemes up to 1.5 cm long .......................................................................................... M. sinemariensis 4(2). Petioles biglandular near or above middle; calyx glands (if present) 0.5– 1.5 mm long; floriferous bracts eglandular ........................ M. poeppigiana 4. Petioles biglandular at or just above base; calyx glands 1.7–2.5 mm long; floriferous bracts often bearing 1 or 2 large glands, or eglandular ..... 5 5(4). Leaf blades abaxially golden- or silvery-sericeous, with the reticulum ± prominent ................................................................................ M. stannea 5. Leaf blades abaxially dark brown-sericeous, with the reticulum ± obscured by outer layer of hairs ................................................... M. castanea 6(1). Margin of bracteoles and sepals with a row of stalked clavate or capitate glands ..................................................................................................... 7 6. Margin of bracteoles and sepals eglandular ............................................. 8 7(6). Leaf blades originally sericeous on both sides with sessile appressed hairs, glabrate at maturity or the hairs persistent abaxially and often on adaxial midrib ............................................................... M. glandulifera 7. Leaf blades velutinous to glabrate adaxially, persistently velutinous abaxially with the prominently stalked hairs T- or Y-shaped ..... M. surinamensis 8(6). Calyx glands 10 (all 5 sepals biglandular) [species unknown in fruit, but placed here on the basis of fruits known in closest relatives] ... M. liesneri 8. Calyx glands 8 (lateral 4 sepals biglandular, anterior eglandular) .......... 9 9(8). Samaras with lateral wing entire at apex or emarginate or incised partway or almost to nut and connate with dorsal wing; torus after fall of samaras surrounded by a 3-lobed disciform outgrowth of the receptacle; sepals appressed in anthesis ..................................................... 10 9. Samaras with lateral wing incised to nut at apex and free from dorsal wing; torus not surrounded by disciform structure; sepals revolute in anthesis ................................................................................ M. macrodisca 10(9). Inflorescence “simple,” i.e., each pseudoraceme axillary to a well-developed vegetative leaf; bracteoles eglandular or 1 or both bearing 1 or 2 small abaxial glands ......................................................................... 11 10. Inflorescence usually “compound,” i.e., the pseudoracemes grouped in terminal or axillary panicles containing only tiny bracts or reduced leaves; bracteoles eglandular ........................................................................... 12 11(10). Leaf blades initially sericeous adaxially, usually soon glabrate; flowers ± evenly distributed along axis of pseudoraceme; inflorescence sericeous ................................................................................................... M. schunkei 11. Leaf blades persistently velutinous adaxially; flowers mostly congested in distal half of axis of pseudoraceme; inflorescence tomentose ........................................................................................... M. cynanchifolia

164

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12(10). Disciform structure surrounding fruiting torus pilose; samaras usually distinctly wider than high ...................................................... M. ovatifolia 12. Disciform structure surrounding fruiting torus glabrous; samaras higher than wide, about as high as wide, or a little wider than high ............ 13 13(12). Pedicels glabrous or initially sericeous at very apex but usually soon glabrate ................................................................................................ 14 13. Pedicels at least initially sericeous, subvelutinous, or velutinous, eventually glabrescent in some species but with some hairs persisting, especially on distal half ............................................................................... 15 14(13). Leaf blades cordate at base, green and often crispate at margin; petioles 3–8 mm long .......................................................................... M. eggersiana 14. Leaf blades cuneate to rounded at base, smooth at margin and with a discolored band (0.5–)1–2 mm wide, the band reddish in dried leaves; petioles 11–20 mm long ................................................................. M. dissimilis 15(13). Leaf blades ± persistently velutinous or tomentose to subsericeous abaxially, the hairs dark brown or reddish (occasionally glabrescent but hairs persistent on midrib); hairs of inflorescence dark brown or reddish; petioles eglandular; leaf blades rounded or cordate at base ..................................................................................................... M. sepium 15. Leaf blades at maturity sparsely sericeous to glabrate, the hairs (if present) white or yellowish; hairs of inflorescence white or gray; petioles usually bearing 2–4 glands near middle, occasionally eglandular; leaf blades cuneate, truncate, or rounded at base, seldom cordate ................................................................................................ M. divaricata Mascagnia castanea (Cuatrec.) W.R. Anderson, Mem. New York Bot. Gard. 32: 218. 1981. —Heteropterys castanea Cuatrec., Webbia 13: 475. 1958. Woody vine; leaf blades abaxially dark brown-sericeous, with the reticulum ± obscured by outer layer of hairs; in other known characters this taxon (which has not been collected with fruits) is indistinguishable from Mascagnia stannea, which see. Evergreen lowland forests, 100–200 m; Amazonas (Caño Yagua). Brazil (Amazonas: Rio Dimití). This striking plant may not merit recognition, although its chestnut-brown leaves distinguish it immediately from M. stannea. I have decided to maintain it for now in order to draw attention to it as worthy of future study, which may reveal it to be nothing more than a trivial variant. It is worth noting that Davidse et al. 17463 (MICH, MO), which is M. castanea, and Davidse et al. 17374 (MO, VEN), which is M. stannea, were collected in the same area on the same day. Mascagnia cynanchifolia Griseb. Mart., Fl. Bras. 12(1): 95. 1858.

in

Woody vine; petioles 3–6 mm long, eglandular; blade of larger leaves 5–8.5 × 2– 4 cm, adaxially persistently velutinous, abaxially persistently sericeous; inflorescence a simple axillary pseudoraceme with flowers congested in distal half of axis; bracteoles eglandular or 1 or both bearing 1 or 2 abaxial glands; lateral 4 sepals biglandular; petals probably pink, glabrous; anthers sericeous; samaras suborbicular or broadly elliptic, 15–36 mm diameter, the lateral wing continuous at base, incised at apex ca. halfway to nut. Evergreen lowland forests, 100–200 m; Amazonas (area of San Fernando de Atabapo). Brazil (Amazonas: vicinity of Manaus). No collection I have seen of this species records the color of the petals. I am assuming that they are probably pink, on the basis of the similarity of this species to Mascagnia schunkei, but that is a questionable assumption; see discussion under M. schunkei. Mascagnia dissimilis C.V. Morton & Moldenke, Phytologia 1: 19. 1933. Woody vine; petioles 11–20 mm long, eglandular; blade of larger leaves 11–25 × 6–

Mascagnia 165

11 cm, soon glabrate, broadly obtuse or rounded at base, with a discolored band (0.5–)1–2 mm wide at margin; inflorescence paniculate; pedicels glabrous or initially sericeous at very apex but usually soon glabrate; lateral 4 sepals biglandular; petals white or pink, glabrous; anthers glabrous; samaras ovate or orbicular, (20–)25–53 mm diameter, glabrate at maturity or with a few straight appressed hairs on nut, the lateral wing continuous at base, emarginate at apex. Evergreen lowland and lower montane forests, 100–300 m; Amazonas (Río Cuao, Río Cunucunuma, Tamatama). Western Amazonia (Colombia, Ecuador, Peru, Brazil, Bolivia). Mascagnia divaricata (H.B.K.) Nied. in Engl. & Prantl, Nat. Pflanzenfam. III. 4: 55. 1890. —Hiraea divaricata H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 169. 1821 [1822]. Hiraea oblongifolia DC., Prodr. 1: 585. 1824. Woody vine; petioles 10–19(–24) mm long, eglandular or, usually, bearing 2–4 glands near middle; blade of larger leaves 6–13 × 3.5–6.5(–8) cm, thinly sericeous to glabrate at maturity; inflorescence paniculate, the axes of ultimate pseudoracemes velutinous with very short gray hairs; 4 lateral sepals biglandular; petals lilac, pink, or pinkish lilac, glabrous; anthers pilose; samaras suborbicular, (18–)20–30 mm diameter, the lateral wing continuous at base, incised up to halfway to nut at apex. Evergreen lowland forests, 50–100 m; Delta Amacuro (Río Acure between La Margarita and Puerto Miranda). Fairly common in northern Venezuela (Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Portuguesa, Sucre); Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Mascagnia divaricata is widely but erroneously known as M. ovatifolia; see discussions in Contr. Univ. Michigan Herb. 19: 380–383. 1993. Mascagnia eggersiana (Nied.) W.R. Anderson, Mem. New York Bot. Gard. 32: 224. 1981. —Mascagnia tenuifolia var. eggersiana Nied., Arbeiten Bot. Inst. Königl.

Lyceum Hosianum Braunsberg 3: 10. 1908. Woody vine; stems soon glabrate; petioles 3–8 mm long, eglandular; blade of larger leaves 8–19.5 × 4–10.5 cm, cordate at base, often crispate at margin, soon glabrate; inflorescence paniculate, sparsely velutinous; pedicels glabrous; 4 lateral sepals biglandular; petals yellow, glabrous; samaras ovate, 20–35 mm diameter, soon glabrate, the lateral wing continuous at base, emarginate at apex or incised up to halfway to nut. Mostly in evergreen forests, also at edges of savannas and cultivated areas, 100– 600 m; northwestern Bolívar (southeast of Pijiguaos, mouth of Río Parguaza), Amazonas (Parhueña). Aragua, Distrito Federal, Falcón, Táchira, Trujillo, Yaracuy; Ecuador, Peru. ◆Fig. 105. Mascagnia glandulifera Cuatrec., Webbia 13: 365. 1958. Woody vine; stems sericeous to glabrate; petioles 8–13 mm long, bearing (2–)4–10 small glands in 2 rows; blade of larger leaves 11–14.5 × 4.5–8 cm, sericeous to glabrate on both sides or the hairs persistent abaxially and on adaxial midrib; bracteoles and sepals bearing a row of small stalked glands on margin; sepals completely concealing petals until flower opens, the lateral 4 abaxially biglandular; petals yellow, abaxially densely tomentose; samaras 45–60 mm wide, the lateral wing continuous at base, divided to nut at apex. Evergreen lowland forests, ca. 100 m; Amazonas (northeast of San Carlos). Northwestern Amazonia (Colombia, Peru, Brazil). Mascagnia liesneri W.R. Anderson, Contr. Univ. Michigan Herb. 17: 51. 1990. Woody vine; petioles 10–20 mm long, eglandular or biglandular on distal half; blade of larger leaves 7–11.5 × 3.8–5.8 cm, adaxially soon glabrate, abaxially thinly sericeous to glabrate, eglandular or with a row of tiny glands between midrib and margin; inflorescence paniculate, the flowers borne ultimately in pairs, 2 or 3 pairs usually congested to form corymbs or umbels; sepals all biglandular, completely concealing petals in bud, revolute in anthesis; petals yellow turning red in age, glabrous; stigmas terminal or nearly so; fruits unknown. Wet disturbed lowland forests, 100–200 m;

166

M ALPIGHIACEAE

Amazonas (San Carlos de Río Negro north to the Brazo Casiquiare). Endemic. The fruit of Mascagnia liesneri is not known, but it can be expected to resemble that of Mascagnia leucanthele Griseb. in Mart., in which the samaras are suborbicular with the membranous lateral wing continuous at the base and incised to the nut at the apex. Mascagnia macrodisca (Triana & Planch.) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 3: 16. 1908. —Hiraea macrodisca Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 326. 1862. Woody vine; petioles (8–)10–25 mm long, usually bearing 2–8 glands in 2 rows; blade of larger leaves 8–14.5(–18) × 4–9(–12) cm, abaxially persistently sericeous, bearing glands between midrib and margin; pseudoracemes 2–12(–16) cm long; sepals completely concealing petals in bud, revolute in anthesis, the lateral 4 biglandular; petals lilac or pink, abaxially densely sericeous or tomentose; samaras 20–62 × 25–70 mm, the lateral wing continuous at base or rarely divided nearly or quite to nut, divided to nut at apex. Evergreen lowland forests and secondary forests, 100–500 m; fairly common in Bolívar, rare in Amazonas (Mavaca). Apure, Barinas; Colombia, Guyana, Suriname, Amazonian Ecuador, Peru, Brazil (Acre, Amazonas, Pará), and Bolivia. ◆Fig. 106. Mascagnia ovatifolia (H.B.K.) Griseb., Fl. Brit. W.I. 121. 1860. —Hiraea ovatifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 170. 1821 [1822]. Mascagnia nervosa Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 3: 12. 1908. Woody vine; petioles 8–11(–18) mm long, eglandular; blade of larger leaves 9.5–16.5 × 5–8.2 cm, soon glabrate, rounded at base; inflorescence paniculate; pedicels initially loosely sericeous, glabrescent in age; petals white or pink, glabrous; anthers glabrous or very sparsely pilose; samaras broadly elliptic, 17–30 × 25–37 mm, usually distinctly wider than high, the lateral wing continuous at base, emarginate or shallowly incised at apex; disciform structure surrounding fruiting torus pilose. Semideciduous forests, ca. 200 m; Bolívar (near El Manteco). Apure, Aragua, Barinas, Carabobo, Distrito Fe-

deral, Falcón, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Panama, Colombia, Trinidad. This is not the species that has been called Mascagnia ovatifolia in all recent floras, which have continued the misapplication of that name by Niedenzu in Das Pflanzenreich. That species is treated here as M. divaricata. The type of Hiraea ovatifolia (P-HBK!), which Niedenzu never saw, is conspecific with his M. nervosa. See discussions in Contr. Univ. Michigan Herb. 19: 380–383. 1993. Mascagnia poeppigiana (A. Juss.) W.R. Anderson, Mem. New York Bot. Gard. 32: 219. 1981. —Hiraea poeppigiana A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 260. 1840. —Tetrapterys poeppigiana (A. Juss.) Griseb. in Mart., Fl. Bras. 12(1): 87. 1858. Mascagnia subsericea Cuatrec., Webbia 13: 370. 1958. Mascagnia materdei Cuatrec., Brittonia 11: 167. 1959. Woody vine; petioles 7–9(–11) mm long, biglandular near or above middle; blade of larger leaves 8.5–15(–19) × 4.2–9 cm, abaxially very densely and persistently metallic-sericeous; floriferous bracts eglandular; all sepals eglandular or the lateral 4 biglandular, the glands 0.5–1.5 mm long; petals yellow, glabrous, the limb up to 2.5 mm long; anthers 0.5–0.7 mm long, glabrous; samaras 9–18 × 16–22 mm, butterflyshaped, the lateral wing divided to nut at base and apex. Evergreen lowland forests, 100–200 m; Amazonas (Río Cunucunuma, Río Yureba). Colombia, Guyana, Peru, Brazil, Bolivia. Mascagnia schunkei W.R. Anderson, Mem. New York Bot. Gard. 32: 222. 1981. Mascagnia cordifolia var. peruviana J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13 : 787. 1950. Woody vine; petioles 5–15 mm long, eglandular; blade of larger leaves 5.5–17 × 2.3–7 cm, adaxially sericeous to soon glabrate, abaxially persistently sericeous; inflorescence a simple axillary pseudoraceme with flowers ± evenly distributed along axis; bracteoles eglandular or 1 (rarely both) bearing 1 abaxial gland; lateral 4 sepals biglandular; petals pink or lilac(?), glabrous;

Mascagnia 167

samaras suborbicular, 25–36 mm diameter, the lateral wing continuous at base, notched at apex or incised partway or almost all the way to nut. Evergreen lowland forests, 50– 300 m; Amazonas (Mavaca, Puerto Ayacucho). Widespread in Amazonia [French Guiana, Peru, Brazil (Amapá, Amazonas, Mato Grosso, Pará), Bolivia]. When I described Mascagnia schunkei I admitted uncertainty as to whether the petals are yellow or pink. I now have available 2 collections from Peru for which the petals were described as yellow, and 7 from Peru, Brazil, Bolivia (Amapá and Pará), and French Guiana for which they were described as pink or lilac. In most other characters these collections are alike, so I still cannot say with confidence which form gets the name (the Peruvian type was in fruit). I am tentatively assuming the petals in the “true” M. schunkei to be pink, but that may prove to be wrong. Mascagnia sepium (A. Juss. in A. St.-Hil.) Griseb. in Mart. —Hiraea sepium A. Juss. in A. St.-Hil., Fl. Bras. Merid. 3: 19, pl. 165. 1832 [1833]. Woody vine; stems velutinous to glabrescent in age; petioles 8–18(–22) mm long, eglandular; blade of larger leaves 8–16 × 5– 10.2 cm, rounded or cordate at base, abaxially velutinous or rarely tomentose to subsericeous with dark brown or reddish hairs, occasionally glabrescent; inflorescence paniculate, velutinous with dark brown or reddish hairs; pedicels at least sparsely sericeous or subvelutinous; 4 lateral sepals biglandular; petals yellow, glabrous; samaras ovate to orbicular, 16–34 mm diameter, sparsely and loosely sericeous, the lateral wing continuous at base and apex, often shallowly notched at apex. Evergreen lowland to lower montane forests, 50–1200 m; widespread in Bolívar and Amazonas. Apure, Distrito Federal, Falcón, Miranda, Sucre, Táchira, Yaracuy, Zulia; Colombia, Ecuador, Peru, Brazil. ◆Fig. 104. Our plant is probably separable from true Mascagnia sepium, an exceedingly variable complex based on a type from Rio de Janeiro. Mascagnia sinemariensis (Aubl.) Griseb. in Mart., Fl. Bras. 12(1): 93. 1858. —Banisteria sinemariensis Aubl., Hist. Pl. Guiane 1: 462, pl. 185. 1775.

Malpighia volubilis Sims, Bot. Mag. 21: 809. 1805. —Mascagnia volubilis (Sims) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 3: 22. 1908. Hiraea schizoptera Turcz., Bull. Soc. Imp. Naturalistes Moscou 36: 584. 1863. —Mascagnia schizoptera (Turcz.) Cuatrec., Webbia 13: 373. 1958. Mascagnia hondensis C.V. Morton, Proc. Biol. Soc. Wash. 45: 52. 1932. Woody vine; petioles (5–)8–15 mm long, usually bearing 2–6 glands in 2 rows; blade of larger leaves (6–)8–16.5 × (2.5–)4–9 cm, abaxially sparsely sericeous to glabrate, bearing impressed glands on margin; pseudoracemes up to 1.5 cm long, corymbose; sepals leaving petals exposed in enlarging bud, appressed in anthesis, the lateral 4 biglandular; petals yellow, abaxially densely sericeous; samaras 12–30 × 20–50 mm, butterfly-shaped with lateral wing divided to nut at base and apex, sometimes reduced and corky. Evergreen lowland forests, 100– 300 m; Bolívar (50 km south of Caicara, El Dorado to Santa Elena de Uairén road, El Palmar, Isla Anacoco, Upata), Amazonas (Caño Yureba on lower Río Ventuari, Río Sipapo, Río Mavaca, Río Metacuni). Apure, Falcón, Mérida, Táchira, Zulia; Mexico, Central America, Lesser Antilles, Colombia, Guyana, French Guiana, Amazonian Ecuador, Peru, and Brazil. Mascagnia stannea (Griseb.) Nied., Verz. Vorles. Königl. Lyceum Hosianum Braunsberg 1912/13: 12. 1912. —Heteropterys stannea Griseb., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1853: 46. 1854. Hiraea laurifolia A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 260. 1840, non Mascagnia laurifolia Nied. in Chodat & Hassl. 1907. Tetrapterys benthamiana Griseb. in Mart., Fl. Bras. 12(1): 88. 1858. —Mascagnia benthamiana (Griseb.) W.R. Anderson, Mem. New York Bot. Gard. 32: 217. 1981. Mascagnia sericans Nied. in Chodat & Hassl., Bull. Herb. Boissier sér. 2, 7: 284. 1907. Woody vine, or a shrub with young branches twining when support is available; petioles 4–9 mm long, biglandular at or slightly above base; blade of larger leaves 10–19.5(–26) × 6.5–12(–14) cm, abaxially

168

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Fig. 104. Mascagnia sepium Fig. 105. Mascagnia eggersiana

Fig. 106. Mascagnia macrodisca

Mezia 169

very densely and persistently golden- or silvery-metallic-sericeous; floriferous bracts mostly bearing 1 or 2 large abaxial glands, or eglandular; lateral 4 sepals biglandular, the glands 1.7–2.5 mm long; petals yellow, glabrous, the limb 3.5–5 mm long; anthers 1– 1.5 mm long, usually sericeous between locules and bearing a few abaxial hairs; samaras 15–30 × 20–40 mm, butterfly-shaped, the lateral wing divided to nut at base and apex, sometimes much dissected. Evergreen lowland forests, often on riverbanks or in seasonally flooded places, 100–200 m; Amazonas (Caño Yagua, Cariche, Río Cuao). Costa Rica, Panama, Colombia, Peru, Brazil, Bolivia, Paraguay. Mascagnia stannea is exceedingly variable in the size and shape of its leaves, but relatively consistent in its reproductive structures. The small-leaved populations in southern Brazil and Paraguay probably deserve segregation, but those in northern South America are not easily separable from the type of Heteropterys stannea, which came from Costa Rica. The descriptive notes given above apply only to the plants of southern Venezuela and nearby, and do not attempt to convey the variation throughout the taxon’s range. I am recognizing here two segregates from this complex, M. poeppigiana and M. castanea. The former is fairly well supported by several characters and a distribution throughout western Amazonia. The latter has a much weaker claim to recognition; see the discussion under M. castanea.

Note that the oldest name for this species as it is defined here is Hiraea laurifolia A. Juss., but the combination cannot be made in Mascagnia because that would create a later homonym. However, if/when this species is transferred to a different genus, the epithet laurifolia should be taken up for it. On the other hand, if M. poeppigiana is combined with M. stannea, the epithet poeppigiana, which was published at the same time as laurifolia, can be used for it immediately, in Mascagnia or any other genus. Mascagnia surinamensis (Kosterm.) W.R. Anderson, Contr. Univ. Michigan Herb. 17: 53. 1990. —Mascagnia multiglandulosa var. surinamensis Kosterm., Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 25: 5. 1936. Woody vine; petioles 4–12 mm long, eglandular or biglandular in middle third; blade of larger leaves 6.5–12 × 3–7 cm, velutinous to glabrate adaxially, persistently velutinous abaxially with T- or Y-shaped hairs; bracteoles and sepals bearing a row of small stalked glands on margin; sepals leaving outermost petal(s) exposed in bud, the lateral 4 abaxially biglandular; petals yellow, abaxially densely tomentose; samaras 22–30(–40) mm wide, the lateral wing continuous at base, divided to nut at apex. Evergreen lowland forests, roadside thickets, 100–200 m; Amazonas (Puerto Ayacucho– Gavilán). Guyana, Suriname, Amazonian Brazil, Bolivia.

19. MEZIA Schwacke ex Nied. in Engl. & Prantl, Nat. Pflanzenfam. III. 4: 58. 1890. —Diplopterys sect. Mezia (Schwacke ex Nied.) Nied., Arbeiten Bot. Inst. Königl. Lyceum Hosianum Braunsberg 4: 16. 1912. Woody vines, shrubs, or small trees. Leaves with the petiole eglandular; stipules minute, interpetiolar, caducous; blade bearing impressed abaxial glands or eglandular. Inflorescence tightly reddish- or brown-sericeous throughout, axillary and terminal, often decompound, containing much-reduced bract-like leaves, the flowers borne ultimately in umbels of 4; bracts and bracteoles abaxially sericeous, adaxially glabrous or sparsely sericeous; bracts smaller than bracteoles, deciduous before maturation of fruits; peduncles well developed; bracteoles borne just below flower, large, globose-cymbiform, the inner enclosing bud until flower opens, the outer enclosing bud and inner bracteole, persistent or deciduous before maturation of fruits; pedicels absent or very short, up to 5 mm long in fruit; old flowers (not setting fruit) deciduous at base of peduncle, not at joint between peduncle and pedicel. Sepals narrowly oblong or spatulate, the anterior eglandular, the lateral 4 each bearing 2 large compressed glands, these distinct or partially to completely connate;

170

M ALPIGHIACEAE

petals yellow. Stamens 10, dimorphic, the 5 opposite sepals differing from the 5 opposite petals in size and shape, and sometimes in pubescence. Ovary of 3 carpels adnate to a common axis; styles 3, the apex with a large internal stigma and dorsally truncate or short-hooked or pedaliform (i.e., with a short broad extension resembling from above the sole of a shoe), the anterior style shorter and often more slender than the 2 posterior styles. Fruits breaking apart into 3 samaras, each bearing 2 large lateral wings that are distinct or more often confluent at base, a smaller dorsal wing, and often additional wings, winglets, or crests between them or outside the lateral wings. Panama, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 12 species, 4 in Venezuela, all in the flora area. Key to the Species of Mezia 1. 1. 2(1).

2.

3(1). 3.

Leaves very densely and persistently sericeous abaxially, the hairs producing a reddish or rusty brown metallic sheen .................................. 2 Leaves glabrous or thinly sericeous to glabrate abaxially, the hairs on mature leaves not dense enough to completely conceal epidermis ...... 3 Shrub or small tree 2–8 m tall; blade of larger leaves 9–17 × 5–10 cm; petioles 10–15 mm long; samaras 30–42 mm diameter, the central dorsal wing and 2 parallel winglets flat, the latter not connected by transverse winglets to lateral wing; lateral wing of samara nearly flat, tomentose, the hairs sinuous and spreading ................................................. M. huberi Woody vine; blade of larger leaves 19–29 × 10–18.5 cm; petioles 20–40 mm long; samaras 45–60 mm diameter, the central dorsal wing and 2 parallel winglets strongly corrugated and the latter connected to lateral wing by several transverse winglets; lateral wing of samara wrinkled or corrugated, sericeous, the hairs straight and appressed ........... M. rufa Anthers opposite sepals densely tomentose, those opposite petals sparsely tomentose to glabrous ............................................................... M. curranii Anthers all glabrous .................................................................... M. includens

Mezia curranii W.R. Anderson, Mem. New York Bot. Gard. 32: 236. 1981. Diplopterys involuta var. ovata Nied. in Engl., Pflanzenr. IV. 141: 227. 1928. Woody vine; petioles 12–24 mm long; blade of larger leaves 12.5–23 × 4.5–9.5 cm, abaxially very sparsely sericeous to glabrate; anthers opposite sepals ± densely tomentose, those opposite petals sparsely tomentose to glabrous; samaras 55–65 mm diameter, the lateral wing continuous at base, flat to wrinkled, thinly and loosely sericeous, the central dorsal wing and 2 parallel winglets wavy or corrugated, the latter connected to lateral wing by several mostly transverse winglets. Evergreen lowland forests, Amazonas (Culebra). Amazonian Peru. Mezia curranii is apparently little more than M. includens with hairy anthers, and it

remains to be seen whether that character is sufficient basis to justify its continued recognition, but it is true that the widespread and variable M. includens is generally consistent in its glabrous anthers. Mezia huberi W.R. Anderson, Contr. Univ. Michigan Herb. 19: 384. 1993. Shrub or small tree 2–8 m tall; petioles 10–15 mm long; blade of larger leaves 9–17 × 5–10 cm, abaxially very densely and persistently reddish- or dark brown-sericeous; anthers glabrous; samaras suborbicular, 30–42 mm diameter, the lateral wing continuous at base, flat, tomentose with sinuous and spreading hairs, the dorsal wing and 2 intermediate winglets flat and parallel. Sandy savannas and adjacent gallery forests, 100–600 m; Amazonas (west of San Juan de Manapiare). Endemic. ◆Fig. 107.

Mezia 171

B

C

A F E

H

D I G

Fig. 107. Mezia huberi. —A. Fruiting branch, ×0.53. —B. Samara, abaxial view (left), side view looking into apical notch (right), ×1.1. —C. Umbel of 4 flower buds with 2 cut off, ×2.7. —D. Flower, side view with 1 posterior-lateral petal removed, ×2.7. —E. Lateral petal, abaxial view, ×5.3. —F. Posterior petal, abaxial view, ×5.3. —G. Androecium laid out, adaxial view, the stamen opposite anterior sepal to left, ×5.3.—H. Anthers, side view, from opposite a sepal (left) and opposite a posterior-lateral petal (right), ×10.6. —I. Gynoecium, side view, anterior style to left, ×8. ©University of Michigan Herbarium 1993.

172

M ALPIGHIACEAE

Mezia includens (Benth.) Cuatrec., Webbia 13: 450. 1958. —Tetrapterys includens Benth., London J. Bot. 7: 133. 1848. —Diplopterys includens (Benth.) Nied. in Engl., Pflanzenr. IV. 141: 226. 1928. Stenocalyx involutus Turcz., Bull. Soc. Imp. Naturalistes Moscou 31: 394. 1858. —Diplopterys involuta (Turcz.) Nied. in Engl., Pflanzenr. IV. 141: 226. 1928. Woody high-climbing vine; petioles 14– 21(–25) mm long; blade of larger leaves 13– 20(–21) × 6–9.5(–10.5) cm, abaxially initially ± densely sericeous, mostly glabrescent and thinly sericeous to glabrate at maturity; anthers glabrous; samaras suborbicular or wider than high, 70–110 mm across, the lateral wing continuous at base, wrinkled or corrugated, thinly sericeous, the central dorsal wing and 2 parallel winglets strongly corrugated, the latter connected to lateral wing by several transverse winglets. Evergreen lower montane forests, especially along rivers, 200–1300 m; Bolívar (east of 63°W, from Upata and Río Grande south to Río Icabarú, and upper Río Caura at 64°W), Amazonas (scattered localities from Puerto Ayacucho to Sierra Parima). Carabobo; Panama, Colombia, Guyana, French Guiana, Amazonian Ecuador, Peru, Brazil. See discussion of Bernardi 7170 under Mezia rufa. Mezia rufa W.R. Anderson, Mem. New York Bot. Gard. 32: 234. 1981. Woody vine; petioles 20–40 mm long; blade of larger leaves 19–29 × 10–18.5 cm, abaxially densely and persistently metallicsericeous with the hairs usually giving blade

a rusty brown or reddish appearance; anthers glabrous; samaras 45–60 mm diameter, the lateral wing continuous at base or incised to nut, wrinkled or corrugated, sericeous with hairs straight and tightly appressed, the central dorsal wing and 2 parallel winglets strongly corrugated, the latter connected to lateral wing by several corrugated transverse winglets. Evergreen lowland forests, 100–300 m; Amazonas (Maroa to Yavita, base of Sierra de la Neblina). Brazil (Amazonas: Rio Maturacá, Rio Negro), Peru (Loreto). Mezia rufa was described to accommodate plants with very large, persistently sericeous leaves. In characters of their flowers they resemble M. includens. The fruits of M. rufa are not well known; they seem to be mostly smaller than in M. includens, probably reflecting a shift toward dispersal by water, but they have complicated intermediate ruffles as in M. includens. The leaves of M. includens are often densely sericeous at first, but usually soon glabrescent and at most thinly sericeous abaxially at maturity, although occasionally the smaller leaves near the inflorescence are persistently sericeous. In one collection from Río Grande in northeastern Bolívar (Bernardi 7170 [G, MER, MICH]) the few leaves present, subtending branches of the inflorescence, are small but abaxially densely and persistently sericeous. Its samaras resemble those of M. includens. Given the small size of the leaves and the location, I suspect that Bernardi 7170 represents M. includens rather than M. rufa; if larger leaves had been collected they probably would have been glabrate.

20. PTERANDRA A. Juss. in A. St.-Hil., Fl. Bras. Merid. 3: 72. 1832 [1833]. Shrubs or trees. Leaves often crowded at tips of branchlets, eglandular (except for tiny angular translucent dots often present in blade); stipules intra- and epipetiolar, basally to completely connate. Inflorescence reduced to fasciculate clusters in axils of leaves or bracts or above leaf scars; bracts and bracteoles eglandular; pedicels sessile; petals abaxially sparsely to densely sericeous on claw and in center of limb, often persistent in fruit. Stamens 10; anthers glabrous, the outer locules bearing introrse longitudinal wings. Ovary of 3 distinct carpels; styles 3, attached ventrally or subapically, slender and subulate with minute terminal stigmas. Fruits comprising up to 3 dry indehiscent cocci with a papery exocarp and a moderately thick, corneous but not bony endocarp. Panama, Colombia, Venezuela, Guyana, Brazil, Bolivia; 14 species, 3 known or expected in Venezuela, all of these in the flora area.

Pterandra 173

See Christiane Anderson, 1997 [Revision of Pterandra (Malpighiaceae), Contr. Univ. Michigan Herb. 21: 1–27]. Key to the Species of Pterandra 1.

1.

2(1).

2.

Stipules abaxially glabrous even in bud (but often with a sericeous patch at point of attachment to petiole); blade of larger leaves 9.5–14 cm long; bracts and bracteoles triangular, 1.2–1.5 × 1–1.5 mm; sepals strongly revolute ................................................................................... P. guianensis Stipules initially densely sericeous abaxially, the hairs persistent or deciduous at maturity; blade of larger leaves up to 9.8 cm long; bracts and bracteoles linear or narrowly triangular, 1.7–3 × 0.4–1.3 mm; sepals appressed or slightly recurved. ............................................................. 2 Leaf blades white or yellow abaxially, bearing scattered, dark red or brown, short (up to 0.7 mm), straight, sessile hairs; lateral veins of leaf blade flush with abaxial surface or prominulous; petioles 15–27 mm long; anther wings 0.4 mm wide ............................................. P. flavescens Leaf blades light green abaxially, densely brown- or white-sericeous, the hairs up to 1.2 mm long, straight to somewhat serpentine, sessile or short-stalked; lateral veins of leaf blade prominently raised abaxially; petioles 3–13 mm long; anther wings 0.2 mm wide..................... P. sericea

Fig. 108. Pterandra flavescens

174

M ALPIGHIACEAE

Pterandra flavescens Maguire, Mem. New York Bot. Gard. 8: 128. 1953. Shrub or small tree 2–10 m tall; stipules 3.5–4.5 mm long, 2/3–completely connate, abaxially densely sericeous; petioles 15–27 mm long; leaf blades 5–9.8 × 2–5 cm, abaxially white or yellow and very sparsely sericeous; bracts and bracteoles linear or narrowly triangular, 1.7–3 × 0.6–1.3 mm; sepals all biglandular, appressed or slightly recurved; petals cream or pale yellow; anther wings 0.4 mm wide. Tepui meadows and boggy areas along river, ca. 1500 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 108. Pterandra guianensis W.R. Anderson, Mem. New York Bot. Gard. 32: 35. 1981. Tree up to 20 m tall; stipules 3–3.7 mm long, nearly to completely connate, abaxially glabrous or with a sericeous patch at base; petioles 8–15 mm long; blade of larger leaves 9.5–14 × 3.5–6 cm; bracts and bracteoles triangular, 1.2–1.5 × 1–1.5 mm; sepals all

biglandular, strongly revolute; petals greenish yellow; anther wings 0.2 mm wide. Known only from the type, collected in mixed evergreen forest below 762 m on Mt. Ayanganna in western Guyana; to be expected in La Gran Sabana of southeastern Bolívar. Pterandra sericea W.R. Anderson, Brittonia 28: 407. 1976 [1977]. Shrub or tree 2.5–10(–15) m tall; stipules 3–4 mm long, completely connate or notched at apex, abaxially densely sericeous to glabrescent; petioles 3–13 mm long; leaf blades 3–8.5 × 1.6–4.3 cm, abaxially light green and densely sericeous; bracts and bracteoles linear, 1.8–3 × 0.7–1 mm; sepals all biglandular or all eglandular, appressed; petals white to greenish white becoming pale yellow in age; anther wings 0.2 mm wide. By rivers in forests, 100–800 m; northwestern and eastern Bolívar (Apacará-tepui, Auyán-tepui, Cerro Bolívar, Río Ambutuir, Río Carrao, Río Caura). Western Guyana.

21. SPACHEA A. Juss. in Deless., Icon. Sel. Pl. 3: 19. 1837 [1838]. Shrubs or trees, the stems often containing white latex. Leaves bearing impressed glands in blade, abaxially at base and sometimes distally, adaxially near apex; stipules intra- and epipetiolar, connate. Inflorescence terminal, a raceme of short 1–several-flowered cincinni, 1 bracteole often bearing 1 large gland; petals pink or white. Stamens 10, the anthers unwinged, the connective shorter than the locules. Carpels 2 or 3, connate along a central axis, each bearing a stout, untapered style with the broad terminal stigma often becoming subpeltate or bilobed in anthesis. Plants morphologically gynodioecious but functionally dioecious, the pistillate flowers bearing flat unopened anthers with aborted pollen, the apparently bisexual flowers bearing large polleniferous anthers and a small ovary that does not mature into a fruit. Fruits breaking apart into 2 or 3 dry, unwinged, smooth, indehiscent, 1seeded cocci bearing the persistent styles. West Indies, Central America, and northern South America (Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil); ca. 6 species, 1 in Venezuela. Spachea elegans (G. Mey.) A. Juss. in Deless., Icon. Sel. Pl. 3: 19, pl. 31. 1837 [1838]. —Malpighia elegans G. Mey., Prim. Fl. Esseq. 178. 1818. Tree 4–15 m tall; stipules 3–6 mm long, completely connate; petioles 7–10 mm long; blade of larger leaves 6–12(–18) × 3–6(–7.5) cm, abaxially sparsely reddish-tomentose or subsericeous at least on midrib, bearing 2–4 impressed glands abaxially near base and often several distally, plus 2–4 glands on adaxial surface near apex; inflorescence a

pseudoraceme (i.e., all cincinni 1-flowered); bracts eglandular; some plants with 1 bracteole terminating in a ± stalked gland; calyx bearing 8 or 9 glands; petals pink; carpels 2(3); cocci of fruits 4.5–6 × 3–4 mm. Riparian forests, near sea level to 200 m; Delta Amacuro (Río Amacuro), Bolívar (Puerto Ordaz, Río Botanamo, lower Río Caroní, San Félix). West Indies (probably introduced), Trinidad, Guyana, Suriname, French Guiana, Brazil (Roraima). ◆Fig. 109.

Spachea 175

C B L

K A

D

I

F

E

H G

J

Fig. 109. Spachea elegans. —A. Flowering branch, ×0.6. —B. Adaxial view of leaf apex enlarged to show glands, ×1.1. —C. Node with stipules, ×5.5. —D. Bract, peduncle, and bracteoles (twisted peduncle makes bract appear above rather than below peduncle), ×5.5. —E–G. Flower, stamen, and gynoecium of functionally pistillate plant: E ×4.4, F ×11, G ×8.3. —H–J. Flower, stamen, and gynoecium of functionally staminate plant: H ×4.4, I ×11, J ×8.3. —K. Intact fruit, ×5.5. —L. Coccus, adaxial view, ×5.5. ©New York Botanical Garden 1981.

176

M ALPIGHIACEAE

22. STIGMAPHYLLON A. Juss. in A. St.-Hil., Fl. Bras. Merid. 3: 48. 1832 [1833]. Brachypterys A. Juss. in Deless., Icon. Sel. Pl. 3: 20. 1837 [1838]. Woody or herbaceous vines, a few species shrubby. Leaves with the blade entire or lobed; stipules small, distinct, interpetiolar; petioles often long, usually bearing 2 large glands at apex, these sometimes just above petiole on base of blade. Inflorescence unbranched or more commonly a dichasium (or occasionally a small thyrse) of congested pseudoracemes, these usually corymbose or umbellate. Lateral 4 sepals biglandular, the anterior usually eglandular; sepals erect or appressed in anthesis; petals yellow or yellow and red, glabrous or rarely pubescent abaxially, the limb largest in anterior-lateral pair, smaller in posterior-lateral pair, and smallest in posterior petal. Stamens 10, the filaments usually unequal in length and thickness; anthers very unequal in most species, the 4 opposite the lateral sepals often with reduced locules or sometimes sterile and the 1 opposite the posterior petal often small; anthers subequal in several species, including S. bannisterioides in our area. Ovary with the 3 carpels partially connate, all fertile; styles 3, the apex with an internal stigma and dorsally truncate, hooked, or bearing a foliaceous lateral appendage (foliole), the foliole symmetrical on the anterior style, 1-sided on the posterior styles. Fruits breaking apart into 3 samaras, each samara having its largest wing dorsal, thickened on the adaxial (upper) edge, the veins terminating in the thinner abaxial edge; much shorter winglets or crests present on sides of nut in some species; dorsal wing much reduced in a few species. Mexico, Central America, West Indies, South America (all countries except Chile); 1 species, S. bannisterioides, also occurs in West Africa; 91 species, 11 known or expected in Venezuela, 5 of these in the flora area. See Christiane Anderson, 1997 [Monograph of Stigmaphyllon (Malpighiaceae), Syst. Bot. Monogr. 51]. Key to the Species of Stigmaphyllon 1.

1.

2(1).

2.

3(2).

Flowers (3)4(–6) per umbel; all 3 styles dorsally hooked but lacking folioles; “samaras” with dorsal wing reduced to a triangular apical crest 4–9 mm high ................................................................... S. bannisterioides Flowers 8–40 per umbel or pseudoraceme; all 3 styles or at least the posterior 2 bearing well-developed folioles; samaras with an elongated dorsal wing 25–55 mm long or with dorsal wing partially encircling nut and 30–44 mm long measured from base of nut ................................... 2 Leaf blades abaxially pubescent with T-shaped hairs and bearing stipitate (nail-like) or peltate marginal glands 0.2–0.6 mm long and 0.2–0.4 mm diameter at the apex; samaras with nut 12–19 mm diameter, the locule surrounded by air chambers, and dorsal wing partially encircling nut .................................................................................................. S. adenodon Leaf blades abaxially glabrous or sparsely to densely sericeous with sessile or subsessile hairs and bearing sessile marginal glands 0.2– 1.5 mm diameter and sometimes filiform glands up to 1.6 mm long; samaras with nut 2.8–7 mm diameter and dorsal wing not encircling nut ................................................................................................................ 3 Flowers 8–15 per umbel; petals fimbriate with fimbriae up to 0.6 mm long; anterior style and its opposing stamen longer than posterior styles and their opposing stamens; samaras with nut smooth-sided or with 1–5 prominent ribs, the wing widest at base .................. S. puberum

Stigmaphyllon 177

3.

4(3). 4.

Flowers 15–40 per umbel or pseudoraceme; petals erose to denticulatefimbriate with teeth or fimbriae up to 0.2 mm long; anterior style and its opposing stamen mostly shorter than posterior styles and their opposing stamens; samaras with nut usually bearing 1–3 coarsely toothed to lacerate lateral winglets and/or spurs and crests, rarely smooth, the wing narrowed beyond nut, widest distally ...................... 4 Leaf blades abaxially sparsely to very densely sericeous, the hairs (0.2–)0.3– 0.5(–0.7) mm long, usually touching to overlapping .............. S. sinuatum Leaf blades abaxially appearing glabrous to the naked eye, but usually very sparsely sericeous, the hairs ca. 0.1(–0.2) mm long and widely spaced, never touching ................................................ S. convolvulifolium

Stigmaphyllon adenodon A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 288. 1840. Stigmaphyllon grenadense Nied., Ind. Lect. Lyc. Reg. Hos. Brunsberg. p. aest. 1900: 26. 1900. Woody vine; petioles up to 115 mm long; blade of larger leaves 7–15 × 4–15 cm, cordate or ovate, cordate at base in larger leaves, abaxially pubescent with T-shaped hairs, the margin with stipitate nail-like or peltate glands; flowers 15–30(–40) in each umbel or congested pseudoraceme; peduncles 3.5–17.5 mm long; limb of petals erose or erose-denticulate; anthers pubescent; anterior style shorter than posterior styles, all 3 bearing folioles. Grenada, Trinidad, Amazonian Colombia, Venezuela, Ecuador, Amazonian Peru and Brazil; 2 varieties, 1 in Venezuela. The second variety, var. macropterum C.E. Anderson, has a long samara wing similar to that of most other species of the genus. It is found in Ecuador and Peru. S. adenodon var. adenodon Samaras with nut 12–19 mm diameter including air chambers, the dorsal wing partially encircling nut, 30–44 mm long from base of nut. Wet areas along rivers, near sea level; Delta Amacuro (La Margarita–Puerto Miranda, Teima, Tucupita–La Horqueta). Sucre; Grenada, Trinidad, Amazonian Colombia, Peru, and Brazil. Stigmaphyllon bannisterioides (L.) C.E. Anderson, Taxon 41: 328. 1992. —Malpighia bannisterioides L., Pl. Surin. 9. 1775. Banisteria ovata Cav., Diss. 9: 429, pl. 257. 1790. —Brachypterys borealis A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 291. 1840, nom. superfl. —Stigmaphyllon

ovatum (Cav.) Nied., Ind. Lect. Lyc. Reg. Hos. Brunsberg. p. aest. 1900: 31. 1900. —Brachypterys ovata (Cav.) Small, N. Amer. Fl. 25: 138. 1910. Vine or twining shrub to 3 m tall; petioles up to 18 mm long; blade of larger leaves 5– 11(–13) × 2–4.5(–5.5) cm, narrowly elliptic to ovate, attenuate or truncate at base, sparsely sericeous abaxially, the margin eglandular; flowers (3)4(–6) in each umbel; peduncles 0.2–2.5 mm long; limb of petals erose; anthers glabrous; styles subequal, all with a dorsal hook at apex and without folioles; mericarps with nut 8–11 mm diameter, bearing 4–6 ribs or crests on each side radiating from areole, without air chambers, and dorsal wing reduced to a triangular crest 4–9 × 5.5–7.5 mm. Seashores, mangrove swamps, salt marshes, sea level to 50 m; Delta Amacuro (Atoiba, Caño Güinipa, Caño Güiniquina, Curiapo, region of Pedernales). Miranda, Sucre, Yaracuy; Atlantic coast from southern Mexico through Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil as far south as Ceará, Atlantic coast of western Africa. Stigmaphyllon convolvulifolium A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 289. 1840. Vine to 15 m; petioles up to 100 mm long; blade of larger leaves 6–16 × 3–12 cm, ovate or cordate, cordate at base, abaxially very sparsely and minutely sericeous (apparently glabrous), the margin with sessile glands 0.2–0.5 mm diameter and filiform glands up to 1.6 mm long; flowers 15–40 in each congested pseudoraceme; peduncles 4–12.5 mm long; limb of petals erose to denticulate-fimbriate; anthers glabrous; anterior style shorter than posterior styles, all 3 bearing folioles; samaras with nut bearing short winglets, crests, or spurs on sides and dorsal

178

M ALPIGHIACEAE

wing 34–42 × 12–20 mm, widest distally. Moist forests, along rivers, secondary growth, roadsides, near sea level to 300 m; expected in Delta Amacuro adjacent to Guyana. Guyana, Suriname, French Guiana, northeastern Brazil. Stigmaphyllon puberum (Rich.) A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 289. 1840. —Banisteria pubera Rich., Actes Soc. Hist. Nat. Paris 1: 109. 1792. Woody vine; petioles up to 72 mm long; blade of larger leaves 9–18(–20) × 4.5–12.5 cm, elliptic to ovate, attenuate or truncate or sometimes cordate at base, ± densely sericeous abaxially, the margin with sessile glands 0.3–0.4 mm diameter; flowers 8–15 in each umbel; peduncles (0.8–)1.5–4.8 mm long; limb of petals digitate-fimbriate, the fimbriae up to 0.6(–0.8) mm long; anthers glabrous or pubescent; anterior style longer than posterior styles, all 3 bearing folioles; samaras with nut smooth-sided or with 1–5 prominent ribs and dorsal wing 25–41 × 9–

15 mm, widest at base. Evergreen lowland forests, gallery forests, river banks, mangrove swamps, near sea level to 100 m; Delta Amacuro (La Margarita–Puerto Miranda, Misión del Guayo, Río Amacuro, Río Cuyubiní). Monagas; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil.

Fig. 110. Stigmaphyllon sinuatum

Tetrapterys 179

Stigmaphyllon sinuatum (DC.) A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 288. 1840. —Banisteria sinuata DC., Prodr. 1: 588. 1824. Banisteria heterophylla Willd., Sp. Pl. ed. 4, 2: 742. 1799, non Stigmaphyllon heterophyllum Hook. 1843. —Banisteria splendens DC., Prodr. 1: 588. 1824, nom. superfl. —Stigmaphyllon fulgens A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 289. 1840, nom. superfl. —Stigmaphyllon splendens Cuatrec., Webbia 13: 531. 1958, nom. superfl. Stigmaphyllon martianum A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 289. 1840. Stigmaphyllon hypoleucum Miq., Linnaea 18: 51. 1844. Stigmaphyllon brachiatum Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 18: 316. 1862. Stigmaphyllon monancistrum Nied., Ind. Lect. Lyc. Reg. Hos. Brunsberg. p. hiem. 1899–1900: 13. 1899. Woody vine; petioles up to 130 mm long; leaf blades 6–21 × 4.5–20 cm, triangular or ovate to orbicular, acute to deeply auriculate at base, abaxially sparsely to very densely

silver-sericeous, the margin with sessile glands 0.4–1.5 mm diameter and sometimes with filiform glands up to 1.5 mm long; flowers 15–35(–40) in each congested pseudoraceme; peduncles 2.5–11 mm long; limb of petals erose to denticulate; anthers glabrous; anterior style ≤ posterior styles, all 3 bearing folioles; samaras with nut usually bearing short winglets, crests, or spurs on sides and dorsal wing 35–55 × 10–18 mm, widest distally. Primary and secondary forests, especially wet forests, but also in vegetation on white sand, along rivers, at roadsides, and in thickets, 50–900 m; Delta Amacuro (Río Grande, Sierra Imataca), widespread in lowlands of Bolívar and Amazonas. Present in most states of northern Venezuela; Colombia, Guyana, Suriname, French Guiana, Ecuador, northern Peru, northern Brazil, Amazonian Bolivia. ◆Fig. 110. Stigmaphyllon sinuatum is common throughout much of Amazonia and excessively polymorphic. For a thorough discussion of its variability and comparison to similar species, see C. Anderson, Contr. Univ. Michigan Herb. 19: 393–413. 1993.

23. TETRAPTERYS Cav., Diss. 9: 433. 1790, nom. cons. Woody vines or shrubs, occasionally described as small trees. Leaves usually bearing glands; stipules small, interpetiolar or epipetiolar, or absent. Flowers borne in umbels, corymbs, or pseudoracemes, these often grouped in panicles. Calyx usually bearing 8 or 10 glands, eglandular in a few species; petals yellow or pink. Stamens 10, all fertile; anthers ± alike. Ovary with the 3 carpels centrally connate, all fertile; styles 3, the apex with an internal to terminal stigma and dorsally smooth, truncate, or short-hooked. Fruits breaking apart into 3 samaras, each samara having its largest wings lateral, usually 4 discrete wings; dorsal wing smaller, sometimes reduced to a crest or lost; intermediate winglets or projections sometimes present; all wings reduced to rudimentary outgrowths in a few species. Mexico, Central America, West Indies, South America (all countries except Chile and Uruguay); at least 70 species, ca. 18 in Venezuela, 11 of these in the flora area. The taxonomy of this genus is far from satisfactory; as it is studied and better resolved in coming years the number of species recognized will probably increase substantially. Key to the Species of Tetrapterys 1.

Inflorescence simple or compound, terminating in umbels of 4(–6) flowers; bracteoles smaller than floriferous bracts or the same size, eglandular or abaxially callose ................................................................................. 2

180

1.

2(1).

2.

3(2).

3.

4(1).

4. 5(4).

5.

6(5).

6. 7(6).

M ALPIGHIACEAE

Inflorescence an axillary or terminal pseudoraceme, usually unbranched, rarely basally ternate, the flowers sometimes reduced to 1 pair or crowded distally to form a few-flowered corymb or umbel; bracteoles mostly larger than floriferous bracts (wider and often at least as long), often bearing marginal or abaxial glands ............................................. 4 Stipules distinct; styles slender, with small, discrete, nearly terminal stigmas; nonfloriferous bracts of the inflorescence inconspicuous, 5 mm long or less, lanceolate; bracteoles longer than wide; calyx glands (if present) becoming stalked in older flowers and fruits ......... T. mucronata Stipules connate in interpetiolar pairs, often caducous but then leaving a prominent interpetiolar scar; styles stout, stigmatic on internal angle of apex with the stigmas decurrent; inflorescence containing conspicuous, often orbicular, foliaceous bracts 4–20 mm long, much smaller and thinner than vegetative leaves but much larger than floriferous bracts, the large bracts deciduous and usually absent from fruiting specimens; bracteoles about as wide as long or wider; calyx glands sessile ........... 3 Samaras with winglets or aculeate outgrowths between dorsal and lateral wings, the upper lateral wings 12–22(–25) mm long, the lower 4– 10 mm long; stipule pair 1–2 mm wide, the scar stretched to 2.5 mm at older nodes; leaf blades (7.5–)9–14.5 × (3–)4–6.5 cm; petioles 7–12 mm long .............................................................................................. T. discolor Samaras usually devoid of outgrowths between dorsal and lateral wings, the upper lateral wings 22–35 mm long, the lower 11–15(–20) mm long; stipule pair 2.5–4 mm wide, the scar stretched to 5.5 mm at older nodes; leaf blades (11–)13–20 × (5–)6–10.5 cm; petioles (12–)14–20(–27) mm long .......................................................................................... T. crispa Calyx bearing 8 glands, i.e., lateral 4 sepals biglandular, anterior eglandular; petals abaxially ± abundantly appressed-tomentose or subsericeous ................................................................... T. maranhamensis Calyx bearing 10 glands, i.e., all 5 sepals biglandular; petals glabrous or bearing a few appressed hairs abaxially ............................................... 5 Limb of lateral petals denticulate or entire; filaments densely to sparsely sericeous or rarely glabrous; glands on leaf blade marginal or none; nut of samara usually bearing aculeate outgrowths between dorsal and lateral wings ............................................................................................... 6 Limb of lateral petals fimbriate; filaments glabrous; glands present on abaxial surface of leaf blade in most species, between midrib and margin; nut of samara usually devoid of outgrowths between dorsal and lateral wings (all wings reduced to thick, irregular outgrowths in T. oleifolia). ................................................................................................. 8 Blade of larger leaves up to 6 cm long, persistently appressed-tomentose abaxially or eventually glabrescent, the hairs ± serpentine and loose ............................................................................................. T. aristeguietae Blade of larger leaves 3.5–15 cm long, thinly sericeous to glabrate abaxially, the hairs (if any) straight and appressed ............................. 7 Woody vine or robust shrub or small tree 2–3 m tall, usually growing by rivers; leaf blades (2.5–)3.5–7 cm wide; calyx glands (2.5–)3–4 mm long; lateral wings of samara (7–)8–14 mm long, 4, subequal; anthers (1.3–)1.5–1.8 mm long; bracteoles 2–5 mm long ..................... T. styloptera

Tetrapterys 181

7.

Wiry-stemmed shrub 0.2–0.8(–1.5) m tall, in savannas; leaf blades 0.7–2.8 cm wide; calyx glands 1.2–2 mm long; lateral wings of samara 1–3(–5) mm long, variable in number, irregular and often unequal; anthers 0.9–1.1(–1.5) mm long; bracteoles up to 2 mm long ................... T. gracilis 8(5). Stems, inflorescence, and leaves very soon nearly or quite glabrate, except for golden-sericeous axillary buds; leaf blades with margin notably thickened and veins usually obscure or invisible; virgate shrubs 0.2– 2(–3) m tall .................................................................................... T. pusilla 8. Stems and usually the inflorescence densely and persistently sericeous; leaves sericeous to glabrate; leaf blades with margin sometimes revolute but usually not or only slightly thickened, the veins visible on one or both sides; shrubs or vines. ............................................................... 9 9(8). Leaf blades abaxially thinly sericeous to glabrate, the epidermis and glands easily seen ................................................................ T. fimbripetala 9. Leaf blades abaxially densely and persistently sericeous, the hairs nearly or completely concealing the epidermis and often hiding the glands, if any ........................................................................................................ 10 10(9). Leaf blades 2.5–5.1 cm wide, with a row of 9–20 abaxial glands between midrib and margin; lateral wings of samara normal, i.e., all 4 well developed, subequal, thin .................................................... T. rhodopteron 10. Leaf blades up to 2.4 cm wide, eglandular or with 2(–4) abaxial glands near base; lateral wings of samara reduced to rounded crests or thick or aculeate irregular outgrowths ............................................... T. oleifolia Tetrapterys aristeguietae W.R. Anderson, Mem. New York Bot. Gard. 32: 255. 1981. Woody vine(?) or shrub or treelet 1–3.5 m tall; petioles 2–4 mm long; blade of larger leaves 2.3–6 × 1.3–3 cm, abaxially persistently appressed-tomentose to glabrescent, eglandular or with minute glands on margin; inflorescence a short axillary pseudoraceme; bracteoles larger than bracts, eglandular; sepals all biglandular; petals yellow, abaxially very sparsely sericeous, denticulate; filaments sericeous; samaras with lateral wings 2.5–10 mm long, irregular and unequal, with several long narrow projections between lateral wings and dorsal crest. Sandy savannas, semideciduous forests, rocky outcrops, 50– 100 m; Bolívar near Río Orinoco (Cerro Carichana, Cerro El Jobito, Ciudad Bolívar, San Félix). Anzoátegui, Apure.

tiny abaxial glands between midrib and margin; flowers borne ultimately in umbels of 4; 4 lateral sepals biglandular, anterior eglandular; petals yellow, glabrous, entire or sinuate; stigmas internal, decurrent; upper lateral wings of samara 22–35 mm long, lower pair 11–15(–20) mm long, aculeate outgrowths usually not present between dorsal and lateral wings. Evergreen lowland to lower montane forests, usually near rivers, 100–700 m; Delta Amacuro (east-northeast of El Palmar, town of Sierra Imataca eastsoutheast of Los Castillos, Río Cuyubiní), northeastern Bolívar (near San Felipe), southern Amazonas (Río Ocamo). Apure, Carabobo, Cojedes, Falcón, Lara, Miranda, Portuguesa, Yaracuy, Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, and Bolivia.

Tetrapterys crispa A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 265. 1840. Woody vine; stipules connate in interpetiolar pairs, leaving a scar 2.5–5.5 mm wide; petioles (12–)14–20(–27) mm long; blade of larger leaves (11–)13–20 × (5–)6– 10.5 cm, glabrate at maturity, with a row of

Tetrapterys discolor (G. Mey.) DC., Prodr. 1: 587. 1824. —Triopterys discolor G. Mey., Prim. Fl. Esseq. 182. 1818. Woody vine; stipules connate in interpetiolar pairs, leaving a scar 1–2.5 mm wide; petioles 7–12 mm long; blade of larger leaves (7.5–)9–14.5 × (3–)4–6.5 cm, soon glabrate,

182

M ALPIGHIACEAE

with a row of tiny abaxial glands between midrib and margin; flowers borne ultimately in umbels of 4; 4 lateral sepals biglandular, anterior eglandular; petals yellow, glabrous, entire; stigma internal, decurrent; upper lateral wings of samara 12–22(–25) mm long, lower pair 4–10 mm long, several narrow winglets or rounded or aculeate outgrowths present between dorsal and lateral wings. Along rivers, in forests, at shrubby roadsides, 50–700 m; Delta Amacuro (east-northeast of El Palmar, southeast of Piacoa, Río Amacuro, upstream from San Victor), Bolívar (common north of 6°N, Río Karún). Anzoátegui, Apure, Falcón, Lara, Monagas, Sucre, Táchira; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, and Bolivia. Tetrapterys fimbripetala A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 263. 1840. Tetrapterys huachamacariensis W.R. Anderson, Mem. New York Bot. Gard. 32: 261. 1981. Woody vine, sometimes a shrub 1–2.5 m tall; stems ± persistently sericeous; petioles (1–)2–11 mm long; blade of larger leaves 4– 11(–15) × (1.5–)2.3–5(–6) cm, abaxially thinly sericeous to glabrate, with a row of 6– 20 abaxial glands on each side between midrib and margin; inflorescence an axillary pseudoraceme of 4–16 flowers; bracteoles larger than bracts; sepals all biglandular; petals yellow, sometimes tinged with red, glabrous, fimbriate; filaments glabrous; samaras with lateral wings 12–20(–23) mm long, lacking outgrowths between dorsal and lateral wings. Along streams, edge of moist forests, occasionally in savannas, 50–2000 m; widespread in southern Bolívar and Amazonas. Guyana, Suriname, French Guiana, Brazil (Amazonas: Serra Aracá, Roraima), Peru (Huánuco). ◆Fig. 112. Tetrapterys huachamacariensis was based on shrubby plants from Cerro Duida and Cerro Huachamacari with leaves that are smaller than usual for the species. The additional collections of this complex that have accumulated over the last 20 years reveal that it is not rare for T. fimbripetala to be shrubby in various parts of its range, especially at higher elevations, and plants with small leaves also occur at scattered locali-

ties, in both shrubby and climbing plants. It therefore seems best to treat all of that variation as a single variable species. Tetrapterys gracilis W.R. Anderson, Mem. New York Bot. Gard. 32: 257. 1981. Wiry-stemmed shrub 0.2–0.8(–1.5) m tall; petioles 3–6(–9) mm long; blade of larger leaves 3.5–6.5(–10) × 0.7–2.8 cm, soon glabrate, eglandular except for small impressed marginal glands; inflorescence an axillary pseudoraceme of 4–12(–26) flowers; bracteoles larger than bracts, often bearing abaxial glands; sepals all biglandular; petals yellow, glabrous or bearing a few abaxial hairs, entire or denticulate; filaments sericeous to subglabrous; samaras with lateral wings 1–3(–5) mm long, variable in number, irregular and often unequal. Sandy savannas, 100–200 m; Bolivar? (see discussion below), Amazonas (Río Cuao to Río Pasimoni). Colombia (Vaupés). Tetrapterys gracilis was described to accommodate plants that are obviously closely related to T. styloptera but differ from it in their habitat, habit, size of various parts, and reduced, irregular samaras. Most plants have very narrow leaves and low stature and look quite different from the riparian vines of T. styloptera, but occasional plants have higher stature and wider leaves and then resemble T. styloptera more strongly. One very anomalous plant is Stergios et al. 6266 (MICH, MO, PORT), which was found in rain forest on the Río Cuyuní in northeastern Bolívar, far from all populations of T. gracilis and in the wrong habitat. It was a shrub with leaves that are large for T. gracilis and toward the small end of the range for T. styloptera. Its fruits bear much-dissected, rudimentary, rounded outgrowths instead of wings, and therefore fit T. gracilis well. Assuming that T. gracilis deserves recognition, it may be that we should consider Stergios et al. 6266 to be that species, but it may also be that it represents a population of T. styloptera in which the fruits have become reduced independently of and in parallel to the same tendency in T. gracilis. Tetrapterys maranhamensis A. Juss., Arch. Mus. Hist. Nat. 3: 537. 1843. Hiraea gracilis Benth., London J. Bot. 7: 135. 1848.

Tetrapterys 183

Woody vine or shrub 1–2 m tall; stems soon glabrate; blade of larger leaves 4–8.5 × 1.5–3.8 cm, abaxially loosely sericeous to glabrate, with 1–5 glands in a row between midrib and margin; inflorescence an axillary or terminal pseudoraceme of 8–30 flowers; bracteoles wider than bracts but often not as long, one of each pair bearing 1 large eccentric abaxial gland; 4 lateral sepals biglandular, anterior eglandular; petals yellow, abaxially ± abundantly appressed-tomentose or subsericeous, erose; filaments glabrous; samaras with lateral wings 9–14 mm long, bearing irregular outgrowths between dorsal and lateral wings. Evergreen lowland and lower montane forests and thickets, 50–300 m; Bolívar (south of Altiplanicie de Nuria and southeast of Campamento CVG-Las Flores, El Dorado). Guyana, Amazonian Brazil and Bolivia. Tetrapterys mucronata Cav., Diss. 9: 434, pl. 262. 1790. Tetrapterys crebriflora A. Juss. in A. St.Hil., Fl. Bras. Merid. 3: 9. 1832 [1833]. —Tetrapterys mucronata subsp. crebriflora (A. Juss.) Kosterm. in Pulle, Fl. Surinam. 2: 182. 1936. —Tetrapterys mucronata var. crebriflora (A. Juss.) J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser., 13: 807. 1950. Tetrapterys glaberrima Benth., London J. Bot. 7: 134. 1848. Tetrapterys silvatica Cuatrec., Webbia 13: 425. 1958. Woody vine, rarely described as a shrub or small tree; stipules minute, distinct; petioles 5–15(–20) mm long; blade of larger leaves (4–)6–15(–17) × (2–)3–8(–10.5) cm, abaxially glabrous or very thinly sericeous to glabrate, with 2 glands at base and usually a distal row between midrib and margin; flowers borne ultimately in umbels of 4–6; bracteoles narrowly triangular, smaller than bracts; sepals all eglandular or the lateral 4 biglandular, the glands becoming stalked in age; petals yellow, often with reddish flecks, glabrous, entire or denticulate; stigmas internal or nearly terminal; lateral wings of samara 5–22 mm long, the upper pair ca. twice as long as the lower, aculeate outgrowths usually present between dorsal and lateral wings. Evergreen lowland forests, along streams, in roadside shrubs, 50–400

m; common in Bolívar and Amazonas. Apure; Costa Rica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (from Minas Gerais north), Bolivia. Tetrapterys mucronata is an exceedingly variable species, especially in the size and shape of its leaves, but it hangs together as a monophyletic taxon. Whether that should be considered one variable species or a complex of closely related taxa I shall leave to a future monographer to decide. Tetrapterys oleifolia (Benth.) Griseb. in Mart., Fl. Bras. 12(1): 86. 1858. —Hiraea oleifolia Benth., London J. Bot. 7: 136. 1848. —Heteropterys oleifolia (Benth.) Griseb., Linnaea 22: 19. 1849. Shrub 0.5–1 m tall; stems densely and persistently golden-sericeous; petioles 4–13 mm long; blade of larger leaves 5.5–10 × 0.9– 2.4 cm, abaxially densely and persistently sericeous to belatedly glabrescent, eglandular or with 1(2) pairs of impressed glands (often hidden under hairs) near base; inflorescence an axillary pseudoraceme of 2–6 flowers; bracteoles larger than bracts; sepals all biglandular; petals yellow, sometimes tinged with red, glabrous, fimbriate; filaments glabrous; mericarp bearing a dorsal crest up to 2 mm wide and several short, rounded or aculeate lateral outgrowths. Upland areas along streams or in sandy savannas, 1000–1400 m; Bolívar (Cerro La Danta northwest of Cerro Venamo, Gran Sabana, Río Aponguao, southwest of Roraima-tepui). Western Guyana. ◆Fig. 113. As I noted in 1981, the fruit of Tetrapterys oleifolia, which apparently has only rudimentary wings, is known only from Steyermark et al. 105495 (NY), in which the fruits are immature. No other fruiting collections have come to my attention in the last 19 years. Tetrapterys pusilla Steyerm., Fieldiana, Bot. 28: 294. 1952. Virgate subshrubs or shrubs 0.2–2(–3) m tall; vegetative parts glabrous or soon glabrate except for golden-sericeous axillary buds; petioles 1.5–3(–5) mm long; blade of larger leaves 2.5–6(–7) × 1.3–3(–3.5) cm, with several abaxial glands between midrib and thickened margin; inflorescence an axillary pseudoraceme; bracteoles larger than

184

M ALPIGHIACEAE

bracts, bearing small abaxial glands; sepals all biglandular; petals yellow, glabrous, fimbriate; filaments glabrous; styles with stigma internal to apparently terminal; samaras with lateral wings 5–7 mm long, subequal, lacking outgrowths between lateral wings and dorsal crest. Sandy upland savannas and tepui meadows, rocky slopes, 500– 1400(–2600) m; Bolívar (common in the Gran Sabana, Ilú-tepui). Western Guyana. ◆Fig. 111. Tetrapterys rhodopteron Oliv., Timehri 5: 190. 1886. Tetrapterys phylladenophora Maguire & Steyerm., Mem. New York Bot. Gard. 9: 481. 1957. Woody vine, sometimes a shrub 0.5–2.5 m tall; stems persistently sericeous; petioles 6– 12 mm long; blade of larger leaves 6–10.5 × 2.5–5.1 cm, abaxially densely and persistently sericeous, with a row of 9–20 glands on each side between midrib and margin; inflorescence an axillary pseudoraceme of 4–16 flowers; bracteoles larger than bracts; sepals all biglandular; petals yellow, sometimes tinged with red, glabrous, fimbriate; filaments glabrous; samaras with lateral wings 8–17 mm long, lacking outgrowths between dorsal and lateral wings or rarely with 1 aculeate outgrowth. Near streams, apparently often in open rocky or sandy places but also reported from gallery forests, 400–1700 m; Bolívar (common in the Gran Sabana from Auyán-tepui east and south). Western Guyana. Tetrapterys rhodopteron is essentially identical to T. fimbripetala except for the fact that the leaf blades are densely and persistently sericeous abaxially in T. rhodopteron, thinly sericeous to glabrate in T. fimbripetala. Distinguishing between the two taxa is generally easy, but it may be that T. rhodopteron simply represents the extreme of a cline of variation and does not merit recognition. In two Bolívar collections that I am calling T. rhodopteron, Holst & Liesner 2419 (MICH, MO) from Río Samay and Liesner & Holst 18822 (MICH, MO) from near El Paují, the hairs are of intermediate density and the distinction is somewhat arbitrary. Tetrapterys styloptera A. Juss., Ann. Sci. Nat. Bot. sér. 2, 13: 262. 1840.

Bunchosia squarrosa Griseb., Linnaea 22: 11. 1849. —Tetrapterys squarrosa (Griseb.) Griseb. in Mart., Fl. Bras. 12(1): 87. 1858. Tetrapterys boliviensis Nied., Verz. Vorles. Königl. Lyceum Hosianum Braunsberg 1909/10: 17. 1909. Woody vine, rarely described as shrub or tree 2–3 m tall; petioles (3–)4–9(–11) mm long; blade of larger leaves (4.5–)7–15 × (2.5–)3.5–7 cm, abaxially thinly sericeous to glabrate, eglandular except for small impressed marginal glands; inflorescence an axillary or terminal pseudoraceme of 8–24 or more flowers; bracteoles larger than bracts, often bearing 1 or 2 abaxial glands; sepals all biglandular; petals yellow, glabrous or abaxially very sparsely sericeous, denticulate; filaments usually sericeous; samaras with 4 subequal lateral wings (7–)8–15 mm long, bearing several slender outgrowths between dorsal and lateral wings. Along rivers, often at edge of gallery forests, occasionally in open savannas, near sea level to 1200 m; common throughout much of Bolívar and Amazonas. Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Táchira, Trujillo, Yaracuy, Zulia; Nicaragua, Panama, Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil, and Bolivia. ◆Fig. 114. See discussion of Stergios et al. 6266 under Tetrapterys gracilis.

Fig. 111. Tetrapterys pusilla

Tetrapterys 185

Fig. 112. Tetrapterys fimbripetala Fig. 113. Tetrapterys oleifolia

Fig. 114. Tetrapterys styloptera

186

M ALVACEAE

MALVACEAE by Paul A. Fryxell Herbs, shrubs, or trees, often stellate-pubescent; roots fibrous or woody, sometimes fleshy in perennial herbs; stems erect or procumbent, sometimes repent. Leaves alternate, stipulate (the stipules rarely suppressed), ovate, lanceolate, sometimes lobed or dissected, with hairs that are stellate, simple, sometimes prickly, sometimes glandular, or rarely lepidote. Flowers solitary or fasciculate in the axils of the leaves or grouped in inflorescences (usually racemes or panicles, sometimes spikes or scorpioid cymes, rarely umbels or heads). Involucel (or epicalyx, made up of a whorl of 3–many bractlets at the base of the calyx) present or absent. Calyx gamosepalous, truncate, 5-toothed, 5-lobed, or 5-parted; petals 5, distinct, usually clawed, adnate to the staminal column at base. Androecium monadelphous; anthers reniform, numerous (rarely reduced to 5); pollen spheroidal, echinate. Gynoecium superior, 3–40-carpellate; styles 1–40; stigmas truncate, capitate, or decurrent. Fruit schizocarpic or capsular, sometimes a berry. Seeds reniform or turbinate, pubescent or glabrous, rarely arillate. Principally tropics and subtropics, also with a few temperate-zone genera in the Americas, the West Indies, Europe, Africa, southern Asia, the East Indies, Pacific Islands, and Australia; ca. 110 genera and ca. 1800 species, 15 genera and 55 species in the flora area. The genus Uladendron Marc.-Berti, with a single species U. codesuri Marc.Berti (Pittieria 3: 10. 1971), was originally described in the Malvaceae and placed in the tribe Hibisceae. Further consideration, however, suggests that a more likely placement is in the Sterculiaceae tribe Eriolaeneae, currently known from the single Asiatic genus Eriolaena (ca. 17 species). Uladendron is evidently not congeneric with Eriolaena, although further studies are clearly needed. Placement as a second genus in tribe Eriolaeneae may be appropriate or perhaps some other disposition. Uladendron does not conform to the Malvaceae most notably in its dense, heavy wood and its distinctively winged seeds. Key to the Genera of Malvaceae 1. 1.

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

Fruits capsular, 3–5-locular; involucel present; staminal column surmounted by 5 sterile teeth (tribe Hibisceae) ......................................... 2 Fruits schizocarpic (or if pseudocapsular, involucel absent), 3–many-carpellate; involucel present or absent; staminal column surmounted by 5 sterile teeth or antheriferous at summit ........................................... 5 Calyx asymmetrical, splitting laterally at anthesis, deciduous in fruit; fruits elongate (fusiform); foliar nectaries absent ............. 1. Abelmoschus Calyx symmetrical, not splitting laterally, persistent; fruits ± ovoid; foliar nectaries present or absent ................................................................... 3 Stigmas capitate; fruits 5-locular .................................................. 7. Hibiscus Stigmatic lobes decurrent; fruits 3–5-locular (tribe Gossypieae) ................................................................................................................ 4 Involucel of 3 ovate-laciniate bracts enclosing the bud; plants cultivated for seed hairs (cotton); foliar nectaries present .................... 5. Gossypium

M ALVACEAE 187

4. 5(1).

5.

6(5).

6.

7(6).

7. 8(7).

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

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

14(13).

14.

Involucel of ca. 9 subulate or lanceolate bracts not enclosing the bud; seeds ± pubescent, but not cultivated; foliar nectaries absent .. 3. Cienfuegosia Styles and stigmas 10, twice as many as carpels of fruit (5); staminal column surmounted by 5 sterile teeth; foliar nectaries present (in Urena) or absent; involucel usually present (tribe Malvavisceae) ................... 6 Styles and stigmas 3–many, of same number as carpels of fruit; staminal column antheriferous at summit; foliar nectaries absent; involucel absent (tribe Malveae) ............................................................................... 9 Leaves with prominent abaxial nectaries; bractlets of involucel 5, alternate with calyx lobes; mericarps with numerous glochidiate spines; corolla lavender ............................................................................ 14. Urena Leaves lacking nectaries; bractlets of involucel usually > 5, sometimes (in Malachra) absent; mericarps with 3 retrorsely barbed spines or without spines; corolla of various colors ....................................................... 7 Flowers solitary or aggregated into inflorescences, involucellate but not subtended by specialized bracts; mericarps 3-spined, smooth or viscid ..................................................................................................... 9. Pavonia Flowers aggregated into capituliform inflorescences and subtended by specialized, ± ovate bracts; mericarps smooth, spineless ..................... 8 Plants usually setose with urticating hairs; floral bracts often basally dissected or setose, entire; involucellar bracts absent or subulate .................................................................................................. 8. Malachra Plants stellate-pubescent; floral bracts stellate-pubescent, serrate; involucellar bracts often bifurcate ................................................... 10. Peltaea Mericarps 1-seeded .................................................................................. 10 Mericarps usually 3-seeded ..................................................................... 12 Seeds encircled by a pectinate endoglossum within the carpel; fruits somewhat inflated; flowers solitary in the leaf axils, sometimes nodding .......................................................................................................... 4. Gaya Endoglossum lacking; fruits not inflated; flowers solitary in the leaf axils or in inflorescences, usually erect ....................................................... 11 Calyx basally rounded, without costae (ribs); mericarps fragile walled; inflorescences congested ....................................................... 13. Sidastrum Calyx basally 10-costate; mericarps usually indurate; inflorescences various ............................................................................................. 12. Sida Fruits inflated, subglobose, setose, pendent; corolla white ..... 6. Herissantia Fruits not inflated, variously shaped, stellate-pubescent to glabrate, erect; corolla yellow or white ............................................................... 13 Mericarps apically spinose, not differentiated into upper and lower cells, the seeds uniseriate; calyx subequal to fruit ................. 11. Pseudabutilon Mericarps apically bulbous-apiculate, differentiated into 1-seeded lower cell and 2-seeded upper cell, the 2 upper seeds collateral; calyx shorter than fruit .............................................................................................. 14 Cells of mericarp divided by an endoglossum; lower leaves petiolate, uppermost leaves sessile and amplexicaul; inflorescence spiciform ................................................................................................... 2. Briquetia Endoglossum lacking; all leaves petiolate; inflorescence paniculate ............................................................................................... 15. Wissadula

188

M ALVACEAE

Fig. 115. Abelmoschus moschatus

1. ABELMOSCHUS Medik., Malvenfam. 45. 1787. Robust herbs or subshrubs, annual or perennial, often tomentose or hispid. Leaves large, long-petiolate, palmately lobed, crenate or dentate, lacking foliar nectaries. Flowers solitary in the leaf axils. Involucel of 4–16 bractlets, usually caducous. Calyx spathaceous, splitting asymmetrically at anthesis and falling with the corolla; corolla large and showy, yellow with a dark red center. Androecium included in corolla, apically 5-dentate. Style single with 5 sessile capitate stigmas. Capsule elongate, 5-carpellate, pubescent or hispid, dehiscent. Seeds numerous, reniform, pubescent or squamose. Asia, introduced elsewhere in cultivation and sometimes escaped and naturalized; 6 or more species, 3 in Venezuela, 1 of these in the flora area. Abelmoschus moschatus Medik., Malvenfam. 46. 1787. —Hibiscus moschatus (Medik.) Salisb., Prodr. Stirp. Chap. Allerton 387. 1796. —Hibiscus abelmoschus L., Sp. Pl. 696. 1753. Erect herb 1–3 m tall; stems retrorsely

hispid. Cultivated and occasionally naturalized in wet habitats, near sea level to 300 m; Delta Amacuro (Capure), Bolívar (Río Erebato). Widespread in the rest of Venezuela; widely cultivated in the lowland tropics of both hemispheres. ◆Fig. 115.

2. BRIQUETIA Hochr., Annuaire Conserv. Jard. Bot. Genève 6: 11. 1902. Herbs or subshrubs, usually with a single erect stem, branching only in the inflorescence, ± soft-pubescent with stellate and simple hairs. Lower leaves long-petiolate grading to sessile and amplexicaul immediately below the inflorescence, the

Briquetia 189

blades broadly ovate, sometimes lobed, deeply cordate, serrate to undulate (even subentire), acute or acuminate. Inflorescence a leafless terminal spike or panicle, the flowers generally subtended by small trifid bractlets. Involucel absent. Calyx small; petals yellow. Androecium included, antheriferous at apex. Styles 5–14 with capitate stigmas. Fruits schizocarpic but pseudocapsular, strigose or glabrate; mericarps 5–14, 2-celled (the upper cell vestigial in the type species), the cells divided by an endoglossum, the lower cell 1-seeded, the upper cell 1- or 2-seeded. Seeds ± pubescent. Mexico, Central America, Cuba, northern South America south to Brazil and Paraguay; 5 species, 1 in Venezuela. Briquetia spicata (H.B.K.) Fryxell, Brittonia 28: 231. 1976. —Abutilon spicatum H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 271. 1821 [1822]. —Wissadula spicata (H.B.K.) C. Presl, Reliq. Haenk. 2: 117. 1835. —Pseudabutilon spicatum (H.B.K.) R.E. Fries, Kongl. Svenska Vetenskapsakad. Handl. n.s. 43(4): 98. 1908. —Cadillo. Subshrub 1–1.5 m tall; inflorescences spiciform. Forests and disturbed vegetation, commonly growing in shaded habitats, 50– 1000 m; Bolívar (Santa Elena de Uairén), Amazonas (Capibara, Maroa, east of San Fernando de Atabapo). Widespread in northern Venezuela; other distribution as in genus. ◆Fig. 116.

Fig. 116. Briquetia spicata

190

M ALVACEAE

3. CIENFUEGOSIA Cav., Diss. 3: 174. 1787. Redutea Vent., Descr. Pl. Nouv. t. 11. 1800. —Redoutea H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 293. 1821 [1822], spelling variation. Perennial herbs or subshrubs (sometimes shrubs) with procumbent, ascending, or erect stems, glabrous or pubescent, ± gland-dotted throughout. Leaves linear, elliptic, digitately divided, or reniform, entire (in the flora area) or serrate, pubescent to glabrous, with or without (in the flora area) adaxial foliar nectaries; stipules subulate to foliaceous. Pedicels usually solitary in the leaf axils, with or without trimerous involucellar nectaries at summit. Involucel of ca. 9 bractlets (sometimes absent). Calyx often prominently gland-dotted; petals yellow (in the flora area), pink, or purple. Androecium included, apically 5-dentate. Style single; stigmas usually 3, capitate or decurrent. Capsules usually 3-locular, glabrous or pubescent. Seeds several, densely comose to subglabrous. U.S.A. (southernmost Florida and Texas), Mexico, Central America, West Indies, Colombia, Guyana, Brazil, Bolivia, Paraguay, South America, Africa, Arabian Peninsula; 26 species, 2 in Venezuela, both in the flora area. Key to the Species of Cienfuegosia 1.

1.

Plants hirsute; stigmas capitate; leaves ovate to elliptic, pinnately veined; involucel subequal to calyx; petals 2.5–5 cm long; fruits antrorsely villous externally; seeds 2–3 mm long, subglabrous ................... C. affinis Plants glabrate; stigmas decurrent; leaves broadly ovate or rhomboid to moderately 3-lobed, palmately veined; involucel shorter than calyx; petals 1–2.5 cm long; fruits glabrous externally (copiously hairy on internal suture margins); seeds 3–4 mm long, densely comose ............................................................................................. C. heterophylla

Fig. 117. Cienfuegosia affinis

Fig. 118. Cienfuegosia heterophylla

Gaya 191

Fig. 119. Gaya gaudichaudiana

192

M ALVACEAE

Cienfuegosia affinis (H.B.K.) Hochr., Annuaire Conserv. Jard. Bot. Genève 6: 54. 1902. —Hibiscus affinis H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 289. 1821 [1822]. Fugosia campestris Benth., J. Bot. (Hooker) 4: 120. 1842. Fugosia guianensis Klotzsch ex Schomb., Reis. Br.-Guiana 3: 1171. 1848. Fugosia retusa Turcz., Bull. Soc. Imp. Naturalistes Moscou 31: 197. 1858. Cienfuegosia phlomidifolia (A. St.-Hil.) Garcke, Bonplandia 8: 150. 1860. —Fugosia phlomidifolia A. St.-Hil. in A. St.Hil. et al., Fl. Bras. Merid. 1: 197. 1827. Erect shrub 1–2 m tall. Savannas, 100– 300 m; Bolívar (between Ciudad Bolívar and

El Pao, northwest of El Manteco, La Paragua, Río Aro). Widespread in northern Venezuela; Colombia, Guyana, Brazil, Bolivia, Paraguay. ◆Fig. 117. Cienfuegosia heterophylla (Vent.) Garcke, Bonplandia 8: 150. 1860. —Redutea heterophylla Vent., Descr. Pl. Nouv. t. 11. 1800. Subshrub < 1 m tall. Savannas, near sea level to 300 m; Delta Amacuro (Sacupana), Bolívar (between Río Caroní and Ciudad Bolívar). Anzoátegui, Guárico, Monagas, Miranda, Nueva Esparta, Sucre; Virgin Islands, Aruba, northern Colombia, Trinidad, Brazil (Bahia). ◆Fig. 118.

4. GAYA H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 266. 1821 [1822]. Erect herbs or subshrubs, sparingly branched, minutely puberulent, sometimes also pilose. Leaves sometimes distichous, ovate to oblong, cordate, crenate, serrate, or subentire, acute, minutely puberulent. Pedicels solitary in the leaf axils, the flowers sometimes nodding. Involucel absent. Calyx small, basally rounded, not prominently veined; petals 0.5–1.5 cm long, usually yellow. Androecium included, antheriferous at apex; anthers yellow or orange. Styles 8–30, slender; stigmas capitate. Fruits schizocarpic, subglobose to broadly conical, somewhat inflated, sparsely pubescent to glabrate; mericarps 8–30, dehiscent, 1-seeded, usually with a specialized pectinate endoglossum arising ventrally and encircling the seed. Seeds ± puberulent. Neotropics; 33 species, 2 in Venezuela, 1 of these in the flora area. Gaya gaudichaudiana A. St.-Hil., Fl. Bras. Merid. 1: 151. 1827 [1825]. Erect subshrub ca. 1 m tall. Savannas, 100–400 m; Bolívar (Laguna Larga 8 km

southeast of Upata). Distrito Federal, Lara, Miranda, Sucre; Colombia, Ecuador, Brazil, Bolivia, Uruguay. ◆Fig. 119.

5. GOSSYPIUM L., Sp. Pl. 693. 1753; Gen. Pl. ed. 5, 309. 1754. Shrubs or trees, erect (in the flora area) or rarely procumbent, stellate-pubescent or glabrate, ± gland-dotted throughout. Leaves ovate, weakly lobed, or deeply parted, entire, with one or more abaxial foliar nectaries (rarely absent); stipules subulate to falcate, caducous or persistent. Flowers in the leaf axils or in sympodial inflorescences. Pedicels usually surmounted by trimerous involucellar nectaries. Involucel of 3 whorled bracts that are often (in the flora area) cordate, foliaceous, laciniate or dentate, enclosing the bud and persisting in fruit, the bracts sometimes reduced to linear straps, small scales, or deciduous at anthesis. Calyx truncate or 5dentate (rarely 5-lobed); petals cream, yellow, or rose, sometimes with a dark spot at base, often gland-dotted. Androecium included, apically 5-dentate. Style solitary, with 3–5 decurrent stigmatic lobes. Capsule glabrous (rarely pubescent), often prominently gland-dotted, ovoid, dehiscent, 3–5-loculate. Seeds densely lanate, rarely subglabrous. Tropical and subtropical (often arid) regions of North America, South America, Africa, the Middle East, and Australia; 50 species, 2 in Venezuela, both in the flora area.

Gossypium 193

Several species are cultivated commercially for the cotton crop and are widely distributed. Key to the Species of Gossypium 1.

1.

Capsules 3-celled, narrowly ovoid to elongate, pitted; calyx to 10 mm long, usually truncate; stipules 1–5 cm long; leaves 3–7-lobed, the central lobe ovate to lanceolate, usually > 1.5 times as long as broad .... G. barbadense Capsules 3–5-celled, ovoid or subglobose, smooth; calyx usually < 6 mm (excluding teeth), truncate or with acute lobes or acuminate teeth; stipules 0.5–1.5(–2) cm long; leaves 3- or 5-lobed, the central lobe triangular to ovate, usually 1–1.5 times as long as broad .............. G. hirsutum

Gossypium barbadense L., Sp. Pl. 693. 1753. —Algodón. Gossypium vitifolium Lam., Encycl. 2: 135. 1786. Gossypium peruvianum Cav., Diss. 6: 313, pl. 168. 1788. Shrub 1–3 m tall. Usually in houseyard

Fig. 120. Gossypium barbadense

cultivation, rarely naturalized, near sea level to 600 m; Delta Amacuro (Río Acure, Sacupana), scattered in Bolívar and Amazonas. Native to South America but now cultivated in many parts of the world, especially the U.S.A., Peru, Egypt, Uzbekistan, and Tadzhikistan. ◆Fig. 120.

194

M ALVACEAE

Gossypium hirsutum L., Sp. Pl. ed. 2, 975. 1763. —Algodón. Gossypium religiosum L., Syst. Nat. ed. 12, 462. 1767. Gossypium latifolium Murr., Novi Comment. Soc. Regiae Sci. Gott. 7: 22. 1776. Gossypium punctatum Schumach. & Thonn. in Schumach., Beskr. Guin. Pl. 309. 1827.

Shrub 1–2 m tall. In houseyard cultivation or naturalized on roadsides or other disturbed habitats, 50–100 m; Bolívar (between Ciudad Guayana and Puerto Ordaz, between Upata and El Dorado), Amazonas (Río Cunucunuma). Native to Mexico and the West Indies, now widely distributed in cultivation to many parts of the world.

6. HERISSANTIA Medik., Vorles. Churpfälz. Phys.-Ökon. Ges. 4(1): 244. 1788. Bastardia sect. Gayoides Endl., Gen. Pl. 986. 1840. —Gayoides (Endl.) Small, Fl. S.E. U.S. 764. 1903. Bogenhardia Rchb., Repert. Herb. 1: 200. 1841; 2: 48. 1841. Herbs, shrubs, or subshrubs, erect or decumbent, pubescent or hirsute, sometimes viscid. Leaves ovate, cordate, dentate; stipules subulate, caducous. Flowers solitary or several in the leaf axils. Involucel absent. Calyx lobes lanceolate or ovate; petals white. Androecium included, antheriferous at apex. Styles 10–14; stigmas capitate. Fruits schizocarpic, oblate, inflated, pendulous; mericarps 10–14, pubescent or setose, dehiscent, with fragile walls. Seeds 1–3, glabrous or minutely scabridulous. Neotropics; 5 or more species, 1 in Venezuela. Herissantia crispa (L.) Brizicky, J. Arnold Arbor. 49: 279. 1968. —Sida crispa L., Sp. Pl. 685. 1753. —Abutilon crispum (L.) Medik., Malvenfam. 29. 1787. —Gayoides crispum (L.) Small, Fl. S.E. U.S. 764. 1903. —Bogenhardia crispa (L.) Kearney, Leafl. W. Bot. 7: 120. 1954. Trailing to semi-erect perennial herb or subshrub; fruits usually setose. A common, almost weedy plant occurring in many habitats, 100–500 m; Bolívar (El Miamo, Tumeremo, Upata). Widespread in northern Venezuela; from southern U.S.A. to northern Argentina, as well as at a few localities in the Old World. ◆Fig. 121.

Fig. 121. Herissantia crispa

Hibiscus 195

7. HIBISCUS L., Sp. Pl. 693. 1753; Gen. Pl. ed. 5, 310. 1754. Subshrubs, shrubs, or trees (rarely herbs), glabrescent, pubescent, or hispid. Leaves elliptic, lanceolate, ovate, or cordiform, sometimes lobed or parted, dentate or less commonly subentire; abaxial foliar nectaries present or absent. Pedicels solitary or fasciculate in the leaf axils, sometimes aggregated apically. Involucel present (rarely suppressed), the elements 8–10(–20), distinct or basally connate. Calyx 5lobed; petals yellow, lavender, red, or other colors, sometimes large and showy. Staminal column usually included, apically 5-dentate. Styles 5, distally free, with capitate stigmas. Capsules 5-locular, dehiscent, glabrous or pubescent, ovoid or oblong. Seeds numerous, glabrescent or hirsute. Americas, Africa, Asia, Australia, and elsewhere, mostly in tropical and subtropical, but also in temperate climates; ca. 200 species, 12 in Venezuela, 10 of these in the flora area. Many species are in cultivation as ornamentals, such as Hibiscus rosa-sinensis L., which is found cultivated in many towns in the Venezuelan Guayana. Key to the Species of Hibiscus 1.

1.

2(1). 2.

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

6.

Involucel gamophyllous, a 8–20-lobed cup; stipules prominent (15–25 mm long, 7–10 mm wide), enclosing the developing shoot, deciduous; corolla yellow fading to orange, 4–6 cm long; leaves coriaceous, discolorous, broadly ovate and deeply cordate, subentire; trees or large shrubs in mangrove and littoral habitats .................... H. pernambucensis Involucellar bracts distinct or nearly so; stipules usually < 15 mm long, subulate, not enclosing the shoot; corolla of various colors and sizes; leaves various, usually serrate; herbs or shrubs in nonlittoral habitats ................................................................................................................ 2 Calyx lobes ecostate or costae not marginal, lacking nectary on midrib; involucellar bracts various but not bifurcate ........................................ 3 Calyx lobes with prominent marginal costae, often (except H. radiatus) with nectary on midrib of each calyx lobe; involucellar bracts often bifurcate .................................................................................................... 6 Bracts of involucel abruptly expanded distally to a reniform blade .................................................................................................... H. sororius Bracts of involucel linear to lanceolate, not expanded distally ................ 4 Stems pubescent but lacking aculei; petals 6–10 cm long; involucellar bracts 4(5), ovate, 8–12 mm wide ......................................... H. dimidiatus Stems aculeate; petals 9–11 cm long; involucellar bracts ca. 10, narrowly linear ...................................................................................................... 5 Aculei 2–4 mm long, abundant; leaves usually 3-lobed; fruits 3.5–4 cm long .............................................................................................. H. trilobus Aculei 1–2.5 mm long, sparse; leaves unlobed, narrowly lanceolate or sometimes subhastate; fruits 2 cm long ..................................... H. striatus Calyx becoming fleshy and bright red in fruit; foliage often reddish, essentially glabrous; corolla yellow with a red center; involucellar bracts obscurely bifurcate ................................................................. H. sabdariffa Calyx not becoming fleshy in fruit, green; foliage green, pubescent to glabrate; corolla lavender or red; involucellar bracts manifestly bifurcate ......................................................................................................... 7

196

7(6).

7.

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

M ALVACEAE

Nectaries absent on midribs of calyx lobes and on midrib of lower leaf surface; corolla dark red or purplish; leaves glabrate or sparsely aculeate, deeply palmately parted with narrow lobes; plants introduced for gardens, sometimes naturalized .................................................... H. radiatus Nectaries present on midribs of calyx lobes and on midrib of lower leaf surface; corolla lavender; leaves variously pubescent, simple or shallowly lobed (or lowermost sometimes deeply lobed); plants of swampy habitats .................................................................................................. 8 Stems densely stellate-pubescent, usually lacking aculei ........ H. furcellatus Stems sparsely pubescent, with at least scattered aculei ........................ 9 Involucel and calyx hirsute or hispid with long simple hairs; plants erect ................................................................................................. H. bifurcatus Involucel and calyx with sparse, very fine stellate hairs; plants often reclining or trailing ............................................................... H. peruvianus

Hibiscus bifurcatus Cav., Diss. 3: 146, pl. 51, fig. 1. 1787. —Sarsa hueca. Erect or often scandent shrub 1–3(–5) m tall; stems subglabrous. On river banks and in swampy habitats, near sea level to 400 m; Delta Amacuro (Río Amacuro, Caño Araguao, Río Cuyubini), Bolívar (widespread), Amazonas (middle Río Casiquiare, Río Orinoco near mouth of Río Cunucunuma, Samariapo). U.S.A. (Florida Keys), Central America (Honduras, Nicaragua, Panama), West Indies, Colombia, Ecuador, Brazil. Hibiscus dimidiatus Schrank, Denkschr. Bayer. Bot. Ges. Regensburg 2: 61. 1822. Pavonia ulbrichiana Kearney, Madroño 12: 116. 1953. Perennial herb or subshrub 1–2 m tall with densely pubescent stems and broad involucellar bracts. Granitic hills along rivers, 100–200 m; Amazonas (Raudal de Atures). Guyana, Brazil (Amazonas, Maranhão, Pará). ◆Fig. 122. Hibiscus furcellatus Desr. in Lam., Encycl. 3: 358. 1789. —Cayena, Malva. Erect shrub 1–2 m tall with pubescent stems, lowermost leaves sometimes deeply lobed. On river banks, swampy habitats, near sea level to 1000 m; Delta Amacuro (Paloma), Bolívar (Río Suapure, San Ignacio, near Santa Elena de Uairén), Amazonas (La Esmeralda, Río Casiquiare, Río Manapiare, upper Río Orinoco, near Puerto Ayacucho, San Carlos de Río Negro, San Fernando de Atabapo). Anzoátegui, Apure, Barinas, Carabobo, Monagas; U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Guyana, Peru, Brazil, Bolivia, Paraguay, Argentina.

Hibiscus pernambucensis Arruda, Diss. Pl. Brazil 44. 1810. Hibiscus bracteosus DC., Prodr. 1: 455. 1824. Shrub or tree 1–8 m tall with prominent but deciduous stipules; lower surface of leaves whitish. Estuaries, river banks (near their mouths), littoral habitats, near sea level to 50 m; Delta Amacuro (Caño Angosturita, Caño Güiniquina, Misión de Guayo). Aragua, Carabobo, Mérida, Táchira, Zulia; Mexico, West Indies, Central America, much of South America. ◆Fig. 124. This species is common along neotropical coastlines. It is often confused with, but is distinct from, the paleotropical Hibiscus tiliaceus L. Hibiscus peruvianus R.E. Fries, Kongl. Svenska Vetenskapsakad. Handl. ser. 3, 24(2): 31. 1947. —Algodoncillo. Shrub (often reclining or scandent) 2–3 m tall. River banks, associated flooded areas, wet savannas, 50–200 m; Bolívar (Río Caura), Amazonas (islands of Río Orinoco opposite mouth of Río Parhueña). Eastern Ecuador and Peru, Brazil (Acre, Amazonas, Roraima). Hibiscus radiatus Cav., Diss. 3: 150. 1787. Erect or sprawling herb or subshrub to 1.5 m tall with glabrescent stems and often with red flowers. Sometimes naturalized in disturbed habitats, 800–900 m; Bolívar (Santa Elena de Uairén). Apure, Carabobo, Cojedes, Distrito Federal, Guárico, Sucre, Zulia; native to southern and southeastern Asia, introduced as a garden plant in other tropical areas and sometimes naturalized. ◆Fig. 123.

Hibiscus 197

Hibiscus sabdariffa L., Sp. Pl. 695. 1753. —Flor de Jamaica, Rosa Jamaica. Hibiscus fraternus L., Pl. Surin. 12. 1775. Subshrub 1–3 m tall, most plant parts glabrate and ± red-pigmented. Cultivated and often naturalized in disturbed habitats, 100–300 m; Bolívar (San Martín de Turumbán southwest of Tumeremo). Apure, Monagas; probably of African origin, cultivated pantropically and often naturalized.

Hibiscus sororius L., Pl. Surin. 12. 1775. —Dau-joro. Shrub ca. 1 m tall, often reclining or scandent, the stems roughly pubescent. Marshy habitats, riparian forests, near sea level to 200 m; Delta Amacuro (Caño Araguao, Caño Guapoa near Caño Güiniquina). Monagas; Southern Mexico, Guatemala, Belize, Panama, West Indies, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 125.

Fig. 122. Hibiscus dimidiatus

Fig. 123. Hibiscus radiatus

198

M ALVACEAE

Fig. 124. Hibiscus pernambucensis

Fig. 125. Hibiscus sororius

Malachra 199

Hibiscus striatus Cav., Diss. 3: 146, t.54, fig. 1. 1787. Scattered in U.S.A. (Texas), eastern Mexico, Cuba, Jamaica, Honduras, Colombia, Trinidad, Suriname, Peru, Brazil, northern Bolivia, Paraguay, Uruguay, Argentina; 3 subspecies, 1 in Venezuela. H. striatus subsp. lambertianus (H.B.K.) O.J. Blanch. ex Proctor, J. Arnold Arbor. 63: 267. 1982. —Hibiscus lambertianus H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 291. 1821 [1822]. Hibiscus salviifolius A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 1: 249. 1827 [1828]. Hibiscus cubensis A. Rich. in Sagra, Hist. Phys. Cuba, Bot. Pl. Vasc. 140. 1845. Perennial herb or subshrub 1.5–3 m tall; stems densely tomentose. Swampy habitats, near sea level to 100 m; Delta Amacuro (Ca-

ño Jota-Sabuca and Laguna Travesía on Caño Mariusa). Scattered in U.S.A. (Texas), eastern Mexico, Honduras, Cuba, Jamaica, Colombia, Trinidad, Suriname, Peru, and northern Bolivia. Hibiscus trilobus Aubl., Hist. Pl. Guiane 708. 1775. Guatemala, Belize, Jamaica, Hispaniola, Puerto Rico, Venezuela, Suriname; 3 subspecies, 1 in Venezuela. H. trilobus subsp. ingratus (Miq.) O.J. Blanch. & Fryxell, Novon 10: 191. 2000. —Hibiscus ingratus Miq., Linnaea 19: 143. 1846. Erect subshrub or shrub ca. 1 m tall, prominently aculeate. Swamps, river banks, 50–100 m; Bolívar (middle Río Botanamo). Suriname.

8. MALACHRA L., Mant. Pl. 1: 13. 1767; Syst. Nat. ed. 12, 458. 1767. Erect herbs or subshrubs, sometimes puberulent, more commonly hispid, often with urticating hairs. Leaves suborbicular to broadly ovate, angular, palmately lobed to digitately divided, truncate, serrate, acute to obtuse, usually pubescent; stipules filiform, usually persistent. Flowers in condensed head-like racemes, sessile or pedunculate, subtended by broad foliar bracts, these cordate-ovate, often with white intercostal tissue at base. Involucel absent (except present in M. radiata). Calyx deeply 5-lobed; corolla white, yellow, or lavender. Androecium included, apically 5-dentate. Styles 10; stigmas capitate. Fruits schizocarpic, glabrous; mericarps 5, smooth or reticulate, 1-seeded. Seeds glabrous. Neotropics; 8–10 species, 4 in Venezuela, all in the flora area. Key to the Species of Malachra 1. 1.

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

Inflorescences axillary; stem hairs to 1.5 mm long; corolla yellow, 1–1.5 cm long; leaves weakly lobed or unlobed ...................... M. alceifolia Inflorescences terminal; stem hairs 2–4 mm long; corolla white or lavender, 1.5–2.5 cm long; leaves digitately 3–7-lobed below to ovate above ................................................................................................................ 2 Corolla lavender, 2–2.5 cm long; involucel present ........................ M. radiata Corolla white, 1.5–1.8 cm long; involucel absent ...................................... 3 Leaves shallowly lobed with acute sinuses ................................... M. fasciata Leaves manifestly lobed with rounded sinuses ........................... M. palmata

Malachra alceifolia Jacq., Collectanea 2: 350. 1788 [1789]. Herb or subshrub 0.5–2 m tall; flowers yellow. Deciduous forests, pastures, roadsides, secondary vegetation, often in wet soil, near sea level to 500 m; western Delta Amacuro, Bolívar (Ciudad Bolívar, El Dorado, Río Botanamo). Widespread in northern Venezue-

la; Mexico, Belize, Guatemala, Nicaragua, Costa Rica, Panama, West Indies, Colombia, Guyana, Ecuador, Peru. ◆Fig. 126. Malachra fasciata Jacq., Collectanea 2: 352. 1788 [1789]. Malachra kegeliana Garcke, Linnaea 22: 52. 1849.

200

M ALVACEAE

Herb or subshrub 0.5–1 m tall, prominently hispid; flowers white. Riparian forests, roadsides, often in wet soil, near sea level to 100 m; Delta Amacuro (Caño Manamo). Apure, Aragua, Carabobo, Falcón, Guárico, Portuguesa, Sucre, Yaracuy; widely distributed in the Neotropics, adventive in the Paleotropics. Malachra palmata Moench, Methodus 615. 1794. Malachra heptaphylla Fischer in Hornem., Hort. Bot. Hafn. 1: 78. 1813. Urena moenchii Gómez, Anales Soc. Esp. Hist. Nat. 19: 219. 1890, non Urena palmata Roxb. 1814.

Fig. 126. Malachra alceifolia

Herb or subshrub 0.5–1 m tall with palmately lobed leaves. Disturbed habitats, generally in wet soil, 200–300 m; Bolívar (northwest of El Manteco on edge of Lago Guri). Lara; Guyana, Brazil. ◆Fig. 128. Malachra radiata (L.) L., Syst. Nat. ed. 12, 459. 1767. —Sida radiata L., Sp. Pl. ed. 2, 965. 1762 [1763]. Herb or subshrub 1–2 m tall, prominently hispid; flowers lavender. Mud flats near beaches, swamps, near sea level to 50 m; Delta Amacuro (Caño Güiniquina). Apure; Mexico, Nicaragua, Panama, West Indies, Suriname, Peru, Brazil, Paraguay. ◆Fig. 127.

Pavonia 201

Fig. 127. Malachra radiata

Fig. 128. Malachra palmata

9. PAVONIA Cav., Diss. 2: (unpaged addendum). 1786; 3: 132. 1787, nom. cons. Prostrate perennial herbs, erect subshrubs, or shrubs (rarely arborescent), often stellate-pubescent, sometimes viscid, sometimes glabrate. Leaves ovate, elliptic, lanceolate, oblanceolate, deltoid, sometimes lobed, sometimes asymmetrical, dentate or crenate (rarely entire), lacking foliar nectaries. Flowers axillary and solitary or aggregated in racemes, panicles, or heads. Involucel 4–23-parted, the bractlets distinct or basally connate. Calyx 5-lobed; petals white, lavender, purple, or yellow, sometimes with a basal spot. Androecium included or exserted, surmounted by 5 apical teeth. Styles 10; stigmas capitate. Fruits schizocarpic, dry or viscid, pubescent or glabrous; mericarps 5, sometimes winged, spined, rugose, or otherwise ornamented. Seeds solitary, glabrous or slightly pubescent. Southern U.S.A., Mexico, Central America, West Indies, South America, Africa, southern Asia, Australia; ca. 250 species, 17 in Venezuela, 9 of these in the flora area. Key to the Species of Pavonia 1. 1. 2(1).

Leaves oblong-elliptic to obovate; inflorescences often capituliform; mericarps with 3 retrorsely barbed spines ................................................... 2 Leaves narrowly to broadly ovate; flowers solitary in the leaf axils or in terminal racemes; mericarps without spines ........................................ 3 Involucellar bracts 5–7, 2–4 mm wide; spines apically crowded, the 2 lateral spines < 1 mm apart at base .............................................. P. fruticosa

202

2. 3(1). 3.

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

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

M ALVACEAE

Involucellar bracts 8–11, < 2 mm wide; spines not crowded, the 2 lateral spines ca. 3 mm apart at base ............................................. P. castaneifolia Flowers in terminal racemes; corolla greenish yellow; fruits woody; plants of mangrove habitats .............................................................. P. paludicola Flowers solitary in the leaf axils; corolla lavender, rose, or yellow, sometimes with a dark red center; fruits with relatively fragile walls; plants of upland habitats .................................................................................. 4 Corolla yellow with a red center ................................................................ 5 Corolla lavender or rose ............................................................................. 6 Leaves broadly ovate, softly tomentose; involucellar bracts 6, broadly lanceolate (2.5–4 mm wide); plants ± erect or ascending ............... P. sidifolia Leaves asymmetrically deltoid, the lower surface stellate-pubescent, the upper surface with simple hairs; involucellar bracts ca. 15, arcuate, linear, ciliate; plants prostrate ................................................ P. cancellata Leaves narrowly ovate to lance-elliptic; involucellar bracts 7–9, shorter than calyx at anthesis ........................................................... P. angustifolia Leaves ovate to broadly elliptic; involucellar bracts 11–18, 3–6 times the length of calyx at anthesis ..................................................................... 7 Flowers solitary in the leaf axils on pedicels 2–7 cm long; corolla rose; filaments 3–5 mm long; fruits dry (nonviscid) ..................... P. imatacensis Flowers in crowded terminal panicles, the pedicels 1–3 cm long; corolla rose-pink or red; filaments 4–10 mm long; fruits viscid ....................... 8 Involucellar bracts 11–13; petals 3–4.5 cm long; androecium slightly exserted; leaves longer than wide .......................................... P. dasypetala Involucellar bracts 15–18; petals 2–3 cm long; androecium manifestly exserted; lower leaves about as wide as long .................... P. malacophylla

Pavonia angustifolia Benth., J. Bot. (Hooker) 4: 119. 1842. Subshrub 0.8–2 m tall. Savannas, 50–200 m; Bolívar (southwest of Caicara). Guyana, Brazil (Minas Gerais, Roraima), Bolivia. Pavonia cancellata (L.) Cav., Diss. 3: 135. 1787. —Hibiscus cancellatus L., Pl. Surin. 12. 1775. —Auyamilla. Pavonia guanacastensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 672. 1937. Procumbent trailing herb with asymmetrical leaves. Deciduous forests, savannas, roadsides, often in sand, near sea level to 500 m; northern Bolívar. Widespread in northern Venezuela; Nicaragua, Costa Rica, Brazil, Bolivia, Paraguay. ◆Fig. 130. Pavonia castaneifolia A. St.-Hil. & Naud., Ann. Sci. Nat. Bot. ser. 3, 18: 44. 1842, “castanaefolia.” Herb or subshrub 0.4–1.0 m tall; fruits spinose. Edges of paths in forests, along river beaches, 200–1000 m; Bolívar (Río Caura,

Río Paragua, Salto Pará, Santa Elena de Uairén). Falcón, Monagas; Central America, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 131. Pavonia dasypetala Turcz., Bull. Soc. Imp. Naturalistes Moscou 31: 189. 1858. —Lopimia dasypetala (Turcz.) Standl., Contr. U.S. Natl. Herb. 18: 114. 1916. Shrub or small tree 1–7 m tall. Wet forests, often along streams, 200–600 m; Amazonas (Cerro Guanay, Yutajé). Honduras, Nicaragua, Costa Rica, Panama, Colombia. ◆Fig. 132. Pavonia fruticosa (Mill.) Fawcett & Rendle, Fl. Jamaica 5: 130. 1926. —Sida fruticosa Mill., Gard. Dict. ed. 8, no. 18. 1768. Subshrub 0.5–1.5 m tall with spinose fruits. Evergreen lowland forests and forest edges, near sea level to 200 m; Delta Amacuro (Río Orocoima, between Tucupita and La Horqueta, between Tucupita and Los Güires), Bolívar (El Palmar, La Vergareña,

Pavonia 203

middle Río Caura). Widespread in northern Venezuela; Costa Rica, Panama, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. Pavonia imatacensis Steyerm., Acta Bot. Venez. 3: 181. 1968. Subscandent shrub 2–4 m tall, the axillary flowers with tubular rose-colored corollas and exserted genitalia. Evergreen lowland forests, 200–300 m; Bolívar (Río Toro). Endemic.

Pavonia malacophylla (Link & Otto) Garcke, Jahrb. Königl. Bot. Gart. Berlin 1: 221. 1881. —Sida malacophylla Link & Otto, Icon. Pl. Select. 67, t. 30. 1822. —Lopimia malacophylla (Link & Otto) Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11: 97. 1823. Shrub or small tree 1–5 m tall. Deciduous forests, savannas, wet forests, edges of swamps, 50–300 m; Bolívar (Corozal 6 km from Maniapure, El Pao), Amazonas (road

Fig. 129. Pavonia sidifolia

Fig. 130. Pavonia cancellata

204

M ALVACEAE

Fig. 131. Pavonia castaneifolia

Pavonia 205

Fig. 132. Pavonia dasypetala

Fig. 133. Pavonia paludicola

206

M ALVACEAE

Fig. 134. Pavonia malacophylla

Peltaea 207

between Puerto Ayacucho and Samariapo). Mexico, Belize, Nicaragua, Panama, Cuba, Brazil, Bolivia. ◆Fig. 134. Pavonia paludicola Nicolson ex Fryxell in R.A. Howard, Fl. Lesser Antilles 5: 241. 1989. —Malache scabra B. Vogel in Trew, Pl. Select. 50, t. 90. 1772. —Pavonia spicata Cav., Diss. 3: 136, pl. 46., fig. 1. 1787, nom. illeg. based on Malache scabra B. Vogel. —Pavonia racemosa (Sw.) Sw., Fl. Ind. Occid. 2: 1215. 1800, non Cav. 1787. —Althaea racemosa Sw., Prodr. 102. 1788. —Pavonia scabra (B. Vogel) Cif., Atti Inst. Bot. Univ. Pavia ser. 6, 8: 321. 1936, non C. Presl 1835. Shrub 1–3(–5) m tall; flowers greenish yellow, in terminal racemes. Riparian for-

ests, mangrove habitats, near sea level to 100 m; Delta Amacuro (widespread). U.S.A. (southern Florida), Belize, Honduras, Nicaragua, Costa Rica, Panama, West Indies. ◆Fig. 133. Pavonia sidifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 283. 1821 [1822], “sidaefolia.” Malache chiapensis Standl., Contr. U.S. Natl. Herb. 23: 772. 1923. —Pavonia chiapensis (Standl.) Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 231. 1929. Widely branching softly pubescent shrub or subshrub 1 m tall. Deciduous forests, savannas, disturbed vegetation, 50–300 m; Bolívar (Caruachi, southeast of Upata). Mexico, Panama, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 129.

10. PELTAEA (C. Presl) Standl., Contr. U.S. Natl. Herb. 18: 113. 1916, nom. cons. —Malachra sect. Peltaea C. Presl, Reliq. Haenk. 2(2): 125. 1835. Shrubs or subshrubs, erect or decumbent, stellate-pubescent. Leaves elliptic to ovate to suborbicular, crenate-serrate, acute or obtuse, lacking foliar nectaries. Flowers solitary in the leaf axils or more commonly grouped in dense head-like aggregations subtended by specialized floral bracts, the bracts subsessile, cordateovate, the heads subsessile to long-pedunculate. Involucellar bracts 8–13, bifurcate or spatulate. Calyx 5-lobed; petals yellow or lavender. Androecium included, apically 5-dentate, antheriferous ± along length of column. Styles 10; stigmas capitate. Fruits schizocarpic; mericarps 5, glabrous or pubescent, smooth. Seeds solitary, glabrous or pubescent. Central America, West Indies, South America; 19 species, 5 in Venezuela, 4 of these in the flora area. Key to the Species of Peltaea 1. 1.

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

3.

Corolla yellow; flowers in heads at the ends of branches, subtended by ovate floral bracts; bractlets of involucel clearly bifurcate .................. 2 Corolla lavender; flowers solitary in the leaf axils, the specialized floral bracts weakly if at all developed; bractlets of the involucel spatulate ................................................................................................................ 3 Mericarps glabrous; distal surface of bifurcate bractlets pubescent .....................................................................................................P. trinervis Mericarps hirsute; distal surface of bifurcate bractlets subglabrous, the margin ciliate .......................................................................... P. sessiliflora Mericarps laterally winged, apically hirsute; involucellar bractlets medially geniculate; stellate pubescence fine (hairs 0.5 mm diameter) ............................................................................................. P. surumuensis Mericarps unwinged, evenly stellate-pubescent; involucellar bractlets not geniculate; stellate pubescence coarse (hairs 1 mm diameter or more) ..................................................................................................... P. speciosa

208

M ALVACEAE

Fig. 135. Peltaea speciosa

Fig. 136. Peltaea trinervis

Fig. 137. Peltaea surumuensis

Pseudabutilon 209

Peltaea sessiliflora (H.B.K.) Standl., Contr. U.S. Natl. Herb. 18: 113. 1916. —Pavonia sessiliflora H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 281. 1821 [1822]. Shrub 1–2 m tall; flowers yellow; fruits pubescent. Roadsides, wet habitats, 50–100 m; Amazonas (north of Puerto Ayacucho). Guárico, Lara, Sucre, Táchira; Colombia. Peltaea speciosa (H.B.K.) Standl., Contr. U.S. Natl. Herb. 18: 113. 1916. —Pavonia speciosa H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 281. 1821 [1822]. Shrub to 1 m tall; flowers lavender, axillary. Savannas, open habitats, 50–1000 m; Bolívar (widespread), Amazonas (Auyántepui, base of Cerro Guanay, north of Cerro Parú, Río Ventuari). Guárico, Táchira; Cuba, Colombia, Guyana, Suriname, Brazil, Bolivia. ◆Fig. 135. Peltaea surumuensis (Ulbr.) Krapov. &

Cristóbal, Bonplandia 7: 55. 1993. —Pavonia surumuensis Ulbr., Notizbl. Bot. Gart. Mus. Berlin-Dahlem 6: 325. 1915. Peltaea aliculata Krapov. & Cristóbal, Kurtziana 2: 206. 1965. Shrub ca. 1 m tall; flowers lavender; carpels winged. Savannas, riparian forests, 400–1200 m; Bolívar (Canaima, Gran Sabana between Luepa and Santa Elena de Uairén, Río Aprada in Río Urimán basin). Endemic. ◆Fig. 137. Peltaea trinervis (C. Presl) Krapov. & Cristóbal, Kurtziana 2: 168. 1965. —Malachra trinervis C. Presl, Reliq. Haenk. 2: 126. 1835. Shrub 0.5–2 m tall; flowers yellow; fruits glabrous. Rocky outcrops, savannas, river banks, 50–800 m; Bolívar (widespread). Widespread in northern Venezuela; Mexico, Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Guyana, Brazil. ◆Fig. 136.

Fig. 138. Pseudabutilon umbellatum

210

M ALVACEAE

11. PSEUDABUTILON R.E. Fries, Kongl. Svenska Vetenskapsakad. Handl. n.s. 43(3): 96. 1908. Subshrubs or shrubs, glabrescent or pubescent, sometimes glandular-pubescent. Leaves elliptic, ovate, or cordiform, crenate or serrate. Flowers solitary or glomerate in the leaf axils or aggregated into inflorescences. Involucel absent. Calyx lobes lanceolate or ovate; petals often yellow or orange, less often white. Androecium included, the column filamentiferous at apex. Styles 5–10; stigmas capitate. Fruits schizocarpic (but sometimes pseudocapsular); mericarps 5–10, usually apically acute or acuminate. Seeds 3 per carpel. U.S.A. south to Argentina; 19 species, 1 in Venezuela. Pseudabutilon umbellatum (L.) Fryxell, Contr. Univ. Michigan Herb. 21: 190. 1997. —Sida umbellata L., Syst. Nat. ed. 10, 1145. 1759. Erect shrub 1–2 m tall; flowers small, yel-

low; fruits 6–8-carpellate. Disturbed areas, ca. 50 m; Bolívar (Ciudad Bolívar). Widely distributed in Mexico, West Indies, and elsewhere in northern and western South America. ◆Fig. 138.

12. SIDA L., Sp. Pl. 683. 1753; Gen. Pl. ed. 5, 306. 1754. Perennial herbs or subshrubs, erect or prostrate, glabrous or pubescent, sometimes viscid. Leaves ovate (sometimes lobed), elliptic, rhombic, or linear, usually dentate. Flowers solitary in the leaf axils, in axillary glomerules, or in dense or open terminal inflorescences. Pedicels shorter than to much longer than calyx. Involucel absent. Calyx 5-lobed, often basally 10-ribbed; corolla white, yellow, orangish, rose, or purplish, sometimes with a dark red center. Androecium included, antheriferous at apex. Styles 5–14; stigmas capitate. Fruits schizocarpic, glabrous or pubescent; mericarps 5–14, usually indurate, usually laterally reticulate, with 2 apical spines or spines suppressed. Seeds solitary, glabrous. Americas, Africa, Asia, Australia; ca. 150 species, 19 in Venezuela, 12 of these in the flora area. Key to the Species of Sida 1.

1.

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

Leaves entire, linear or narrowly lanceolate, short-petiolate to subsessile; inflorescences terminal, few-flowered, corymbiform, rising above the leaves; carpels 7–9, muticous; corolla whitish with a red center ..................................................................................................... S. linifolia Leaves manifestly serrate (at least apically so), linear, lanceolate, rhombic, or ovate, the petiole usually manifest; inflorescences various (not as above); carpels 5–14, muticous or spinose; corolla usually yellow or white, with or without a red center ...................................................... 2 Plants procumbent ........................................................................... S. ciliaris Plants erect ................................................................................................. 3 Leaves and branching pattern distichous; stipules usually 3(or more)veined, broadly falcate ........................................................................... 4 Leaves and branches spirally arranged; stipules 1(–3)-veined, subulate to narrowly falcate ..................................................................................... 6 Flowers white; leaves broadly elliptic, obtuse, softly tomentose; carpels 5 ............................................................................................. S. jamaicensis Flowers yellow (sometimes white); leaves lanceolate, acute, hirsute to glabrate; carpels 5–9 .............................................................................. 5

Sida 211

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

Carpels 5; calyx 4–5 mm long ..................................................... S. glomerata Carpels 7–9; calyx 6–8 mm long ......................................................... S. acuta Carpels with retrorsely barbed apical spines ........................................... 7 Carpels muticous or apically spinose or aristate, but the spines or aristae not retrorsely barbed ............................................................................. 8 7(6). Leaves narrowly linear; flowers axillary ............................... S. angustissima 7. Leaves ovate; flowers in terminal panicles ................................. S. cordifolia 8(6). Leaves ovate-cordate, serrate throughout; flowers subsessile, in terminal inflorescences; mericarps 5–8, muticous ................................. S. aggregata 8. Leaves linear-lanceolate or rhombic, serrate throughout or basally entire; mericarps 5–14, muticous, spinose, or aristate .................................... 9 9(8). Mericarps 10–12, aristate, the aristae as long as the mericarp body, filiform, curled; leaves often > 8 cm long, rhombic, acute; flowers axillary or in axillary umbels ...................................................................... S. setosa 9. Mericarps 5–14, muticous or spinose, the spines shorter than the mericarp body, straight; leaves < 8 cm long, variously shaped; flowers usually axillary .................................................................................... 10 10(9). Carpels 7–14 .............................................................................. S. rhombifolia 10. Carpels 5 ................................................................................................... 11 11(10). Calyx 4 mm long; leaves linear or narrowly lanceolate, concolorous; corolla yellow ................................................................................... S. serrata 11. Calyx 5–7 mm long; leaves lanceolate to oblanceolate to rhombic, discolorous; corolla white with red center ....................................... S. viarum Sida acuta Burm. f., Fl. Indica 147. 1768. —Escoba, Escobilla. Sida carpinifolia L. f., Suppl. Pl. 307. 1781, non Mill. 1768. Erect shrub or subshrub ca. 1 m tall with distichous branching. Common in disturbed habitats, roadsides, forests, near sea level to 500 m; Delta Amacuro (Caño Manamo, Santa Catalina, between Tucupita and Los Güires), Bolívar (between Moitaco and Río Aro), Amazonas (east of Cerro Marahuaka, Puerto Ayacucho, below San Juan de Manapiare). Widely scattered in northern Venezuela; Pantropics. ◆Fig. 140. Sida aggregata C. Presl, Reliq. Haenk. 2: 106. 1835. Sida setifera C. Presl, Reliq. Haenk. 2: 105. 1835. Sida savannarum K. Schum. in Mart., Fl. Bras. 12(3): 308. 1891. Erect shrub 1–1.5 m tall; leaves broadly ovate; flowers subsessile. Deciduous forests, savannas, roadsides, disturbed vegetation, ca. 50 m; Bolívar (Laguna Franco east of Ciudad Bolívar). Widespread in northern Venezuela; Mexico, Honduras, Nicaragua,

Costa Rica, Panama, West Indies, Colombia, Guyana. ◆Fig. 142. Sida angustissima A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 1: 179. 1827, non Miq. 1850. Erect subshrub ca. 1 m tall; leaves linear (almost filiform); fruits spinose. Savannas, cerrados, 50–300 m; Bolívar (east of El Miamo, La Grulla, Puerto Ordaz). Brazil (Bahia, Minas Gerais, Piauí). ◆Fig. 145. Sida ciliaris L., Syst. Nat. ed. 10, 1145. 1759. Procumbent perennial herb with the leaves, flowers, and fruits all crowded apically. Roadsides, pastures, savannas, disturbed sites, near sea level to 200 m; Delta Amacuro (San Antonio), northern Bolívar. Widespread in northern Venezuela; southern U.S.A., Mexico, Central America, West Indies, most of South America. ◆Fig. 143. Sida cordifolia L., Sp. Pl. 684. 1753. —Malva mansa. Erect shrub or subshrub to 1.5 m tall; fruits spinose. Forests, savannas, roadsides, disturbed sites, 50–900 m; Delta Amacuro (Caño Manamo), Bolívar (Ciudad Bolívar,

212

M ALVACEAE

Fig. 139. Sida cordifolia

Fig. 140. Sida acuta

Sida 213

Fig. 141. Sida serrata

Fig. 142. Sida aggregata

214

M ALVACEAE

Fig. 143. Sida ciliaris

Fig. 144. Sida linifolia

Fig. 145. Sida angustissima

Sidastrum

northeast of El Manteco, Santa Elena de Uairén). Widespread in northern Venezuela; Mexico, Guatemala, Honduras, El Salvador, Nicaragua, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa, Asia, Australia. ◆Fig. 139. Sida glomerata Cav., Diss. 1: 18, pl. 2. 1785. —Escoba, Escoba negra. Subshrub 0.5–1 m tall with distichous branching. Deciduous forests, roadsides, river banks, disturbed vegetation, 50–200 m; Bolívar (widespread), Amazonas (Isla Ratón, Puerto Ayacucho, below Quiratare, San Carlos de Río Negro, San Juan de Manapiare). Widespread in northern Venezuela; Nicaragua, Costa Rica, Panama, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay. Sida jamaicensis L., Syst. Nat. ed. 10, 1145. 1759. Subshrub 0.5–1 m tall with distichous branching; leaves softly pubescent; flowers white. Deciduous forests, disturbed vegetation, ca. 50 m; Bolívar (Ciudad Bolívar). Aragua, Carabobo, Falcón, Portuguesa, Zulia; Mexico, Honduras, Nicaragua, Costa Rica, Panama, West Indies, Colombia. Sida linifolia Juss. ex Cav., Diss. 1: 14, pl. 2, fig. 1, 1785. —Trebol sabanero. Erect to ascending subshrub to 1 m tall; leaves narrow, entire; inflorescences corymbiform. Savannas, forest margins, roadsides, pastures, 50–1000 m; Delta Amacuro (between Los Castillos de Guayana and Piacoa), Bolívar (widespread), Amazonas (Puerto Ayacucho). Widespread in northern Venezuela; Mexico, Central America, West Indies, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay, Uruguay, Africa, Fiji. ◆Fig. 144. Sida rhombifolia L., Sp. Pl. 684. 1753. —Escoba.

215

Subshrub ca. 1 m tall; flowers yellow; leaves ± rhombic. Roadsides, pastures, savannas, disturbed vegetation, riparian forests, near sea level to 500 m; Delta Amacuro (Isla Cocuina, Pedernales), Bolívar (southwest of El Dorado, El Paují, northwest of Kilómetro 88), Amazonas (Gavilán, Puerto Ayacucho, San Carlos de Río Negro, San Fernando de Atabapo). Widespread in northern Venezuela; almost cosmopolitan weed, reaching temperate climates as an annual. Sida serrata Willd. ex Spreng., Syst. Veg. 3: 111. 1826. Subshrub < 1 m tall with small leaves and flowers. Wet savannas, forest margins, 100– 200 m; Bolívar (road between Casa Blanca and Anacoco), Amazonas (Palomar, Raudal de Atures, Raudal de Maipures, San Juan de Manapiare). Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 141. Sida setosa Mart. ex Colla, Herb. Pedem. 1: 416. 1833. Sida surinamensis Miq., Linnaea 22: 469. 1849. Subshrub or shrub 1–2(–3) m tall; fruits with long, capillary setae. Evergreen lowland forests, river banks, disturbed sites, sometimes in periodically flooded sites, near sea level to 1500 m; Delta Amacuro (San Victor), widespread in Bolívar and Amazonas. Nicaragua, Panama, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil. Sida viarum A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 1: 182. 1827. —Conejilla. Subshrub 0.5–1 m tall; leaves narrow, rhombic; corollas with a red center. Savannas, 50–300 m; Bolívar (between Caicara and Ciudad Bolívar, El Manteco). Mexico, Central America, Brazil, Bolivia, Paraguay, Argentina.

13. SIDASTRUM Baker f., J. Bot. 30: 137. 1892. Erect subshrubs 1–2 m tall, ± stellate-pubescent. Leaves ovate to lanceolate or elliptic, serrate or crenate, acute or obtuse. Flowers solitary in the leaf axils, on short axillary racemes, or forming terminal racemes or panicles, the individual pedicels often subtended by 3 stipuliform bractlets. Involucel absent. Calyx small, cupuliform, ecostate; petals small. Androecium included, antheriferous at apex; anthers few (5–20). Styles 5–10; stigmas capitellate. Fruits schizocarpic, oblate to coni-

216

M ALVACEAE

cal, often stellate-pubescent, smooth or rugulose; mericarps 5–10, essentially indehiscent. Seeds solitary, sparsely pubescent to subglabrous. Neotropics; 7 species, 2 in Venezuela, 1 of these in the flora area. Sidastrum micranthum (A. St.-Hil.) Fryxell, Brittonia 30: 452. 1978. —Sida micrantha A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 1: 190. 1827; Pl. Usuel. Bras. t. 49. 1827. Subshrub 1–3 m tall with small flowers and fruits; inflorescences crowded. Deciduous forests, disturbed vegetation, roadsides, ca. 50 m; Bolívar (Ciudad Bolívar). Widespread in northern Venezuela; Costa Rica, West Indies, Colombia, Guyana, Brazil (Bahia, Minas Gerais), Bolivia. ◆Fig. 146.

Fig. 146. Sidastrum micranthum

Wissadula 217

14. URENA L., Sp. Pl. 692. 1753; Gen. Pl. ed. 5, 309. 1754. Shrubs 0.5–2 m tall, ± stellate-pubescent. Leaves often 3- or 5-angled, -lobed, or -parted, less often ovate, oblong, or lanceolate, crenate or serrate, with one or more prominent foliar nectaries on abaxial side. Flowers solitary or glomerulate in the axils or forming terminal racemes. Involucel gamophyllous, 5-lobed, the lobes alternate with the lobes of the calyx. Calyx 5-lobed; petals rose or lavender. Androecium included, apically 5-dentate; anthers subsessile. Styles 10; stigmas capitellate. Fruit schizocarpic, 5-lobed, the lobes convex; mericarps 5, prominently glochidiate (in the flora area) or smooth, essentially indehiscent. Seeds solitary. New World and Old World (southern Asia, Australia) tropics; 6–8 species, 2 in Venezuela, both in the flora area. Key to the Species of Urena 1.

1.

Leaves ovate, angulate, or shallowly 3-lobed, the sinuses acute to narrowly rounded; involucel 5–9 mm long, sometimes exceeding calyx ....................................................................................................... U. lobata Leaves palmately 5-parted, the sinuses broadly rounded; involucel 4–5 mm long, subequal to calyx ........................................................ U. sinuata

Urena lobata L., Sp. Pl. 692. 1753, s. lat. Shrub 1–1.5 m tall. Disturbed areas, 50–200 m; Amazonas (San Fernando de Atabapo, Yavita). Mostly Pantropics. ◆Fig. 147. This is a polymorphic species, treated by Gürke (Bot. Jahrb. Syst. 16: 361–385. 1892) as including 9 varieties and by Hochreutiner (Annuaire Conserv. Jard. Bot. Genève 5: 131–146. 1901) as comprising 14 varieties. In either case the varieties are poorly delimited. Some of the specimens from the Venezuelan Guayana appear to conform to Urena reticulata Cav., [= U. lobata var. reticulata (Cav.) Gürke], for example Guánchez 1825

(MO), but until this complex variation pattern can be more satisfactorily resolved, the species is here treated as U. lobata s. lat. Urena sinuata L., Sp. Pl. 692. 1753. —Urena lobata var. sinuata (L.) Hochr., Annuaire Conserv. Jard. Bot. Genève 5: 141. 1901. Shrub or subshrub 0.5–2 m tall. Abandoned cultivated areas, disturbed sites, near sea level to 200 m; Delta Amacuro (Pedernales, Sacupana, between Tucupita and La Horqueta), Bolívar (Caicara, between Moitaco and Río Aro, La Urbana), Amazonas (Río Padamo). Mostly Pantropics.

15. WISSADULA Medik., Malvenfam. 24. 1787. Herbs or subshrubs, usually erect, stellate-pubescent or sometimes glabrate. Leaves broadly ovate to narrowly triangular, entire or crenate-dentate. Flowers sometimes solitary in the leaf axils, usually in condensed or open terminal panicles. Involucel absent. Calyx 5-lobed, usually small; petals usually yellowish, sometimes white. Androecium included, antheriferous at apex. Styles 3–6; stigmas capitate. Fruits schizocarpic but septicidal dehiscence often suppressed; mericarps 3–6, divided into indehiscent lower cell and dehiscent upper cell by a constriction, apically bulbous-apiculate. Seeds 1 in the lower cell, usually 2 in the upper cell, the lower seed relatively densely pubescent, the upper seeds less so. U.S.A. (southern Texas, Louisiana), Mexico, Central America, West Indies, South America, Africa; ca. 25 species, 4 in Venezuela, all of these in the flora area.

218

M ALVACEAE

Fig. 147. Urena lobata

Fig. 148. Wissadula periplocifolia

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

Leaves basally truncate, ovate to elliptic, the herbage ferruginoustomentose .................................................................................. W. excelsior Leaves ± cordate, ovate or triangular; herbage with whitish to tan tomentum ......................................................................................................... 2 Leaves shallowly cordate, narrowly triangular, with straight margins; corolla sometimes with red center .................................... W. periplocifolia Leaves deeply cordate, broadly ovate, with curved margins; corolla without red center ......................................................................................... 3

M ARANTACEAE 219

3(2). 3.

Inflorescence an open panicle; corolla yellow; mericarps with pronounced medial constriction .......................................................... W. hernandioides Inflorescence a crowded, narrow, racemiform panicle; corolla white; mericarps with imperfectly developed medial constriction ........... W. contracta

Wissadula contracta (Link) R.E. Fries, Kongl. Svenska Vetenskapsakad. Handl. 43(4): 60. 1908. —Sida contracta Link, Enum. Hort. Berol. Alt. 2: 204. 1822. Subshrub to 2.5 m tall; panicles crowded, racemiform; flowers white. Roadsides, open habitats, ca. 900 m; Bolívar (Santa Elena de Uairén). Aragua, Distrito Federal, Lara, Miranda, Monagas, Yaracuy; Mexico (Chiapas), Guatemala, West Indies, Colombia, Guyana, Brazil, India. Wissadula excelsior (Cav.) C. Presl, Reliq. Haenk. 2: 118. 1835. —Sida excelsior Cav., Diss. 1: 27, pl. 5. 1785. Abutilon patens A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 1: 200. 1827. —Wissadula patens (A. St.-Hil.) Garcke, Z. Naturwiss. 63: 123. 1890. Shrub 1–2 m tall; leaves ± elliptic, with truncate bases. Disturbed forests, river banks, roadsides, 200–300 m; Bolívar (Represa Guri). Lara, Zulia; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay. Wissadula hernandioides (L’Hér.) Garcke, Z. Naturwiss. 63: 124. 1890. —Sida

hernandioides L’Hér., Stirp. Nov. 121, t. 58. 1785 [1789]. Wissadula amplissima (L.) R.E. Fries, Kongl. Svenska Vetenskapsakad. Handl. n.s. 43(4): 48. 1908, and many subsequent authors, non Sida amplissima L. Subshrub to 2 m tall with lax, terminal panicles. Forests, savannas, along streams, roadsides, disturbed vegetation, near sea level to 50 m; Delta Amacuro (Caño Manamo). Táchira; Southern U.S.A., Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Africa, India. Wissadula periplocifolia (L.) C. Presl ex Thwaites, Enum. Pl. Zeyl. 27. 1858. —Sida periplocifolia L., Sp. Pl. 684. 1753. Wissadula zeylanica Medik., Malvenfam. 25. 1787. Shrub 1–2 m tall with narrowly triangular leaves, very open inflorescences. Savannas, pastures, roadsides, secondary vegetation, 50–300 m; Bolívar (Ciudad Bolívar, northwest of El Manteco). Widespread in northern Venezuela; Mexico, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, West Indies, Colombia, Brazil, India, East Indies. ◆Fig. 148.

MARANTACEAE by Lennart L. Andersson, Helen Kennedy, and Mats Hagberg Rhizomatous herbs, small and rosulate, very tall and tree-like, or scandent; rhizome axis often starch-storing. Roots often with terminal (subterminal) starch-storing tubers. Leaves distichous in vegetative shoots, or spirodistichous (irregularly arranged due to secondary twisting of axis), differentiated into sheath, petiole (often missing), pulvinus, and blade; sheath not ligulate; blade with a prominent midrib and closely set, parallel, sigmoid lateral veins which fuse near margin, lateral veins interconnected by numerous parallel, regularly arranged, secondary veinlets. Inflorescence a simple thyrse or a ± complex synflorescence of aggregated thyrses (florescences); thyrse with 1 or a series of 1- or 2-flowered cymules (florescence components) in the axils of bracts (spathes of Andersson); cymule usually with a 2- or

220

M ARANTACEAE

2- or 3-keeled prophyll at base backing parental axis, often also with a scale-like or membranous interphyll opposite the prophyll, often also with 1 or 2 dorsal (medial) and/or occasionally 1 or 2 lateral bracteoles. Flowers epigynous, 3-merous, 5-cyclic, with the calyx and corolla clearly different, asymmetric (the 2 flowers of a cymule presenting mirror images). Ovary with 3 locules, 2 of which are often empty and compressed, septa with conspicuous septal nectaries; ovules 1 per locule, basal, initially anatropous. Sepals distinct; corolla, androecium, and style fused at base to form a tube, which is often very long and narrow, rarely indistinct. Outer staminal whorl of (0)1 or 2 staminodes, the outer staminodes usually petaloid and showy, inner staminal whorl of 3 members, 1 monothecial fertile stamen (often with a lateral petaloid appendage) and 2 staminodes, 1 hooded (cucullate staminode), the other firm and fleshy with ± marked callosities (callose staminode). Style with a subapical, flattened stamp on dorsal side, explosively released in pollination; stigma confined to an apical cavity, scoop-shaped or blunt. Fruit usually capsular, grading into ± achene-like, and tardily dehiscent or indehiscent in a few species, ± fleshy and indehiscent in some genera (outside the flora area). Seeds arillate in all members with dehiscent fruits, without endosperm but with copious starchy perisperm; embryo horseshoe-shaped; center of perisperm with a canal (perisperm canal) formed by disintegration of the central core of tissue, perisperm canal unbranched throughout or ± extensively branched. Pantropics, a few species extend into warm-temperate North America and South America; 31 genera and ca. 530 species, 9 genera and 59 species in the flora area. Key to the Genera of Marantaceae by Lennart Andersson 1. 1. 2(1). 2. 3(2).

3.

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

5.

Ovary with 3-ovulate locules; fruit commonly 2- or 3-seeded ..... 1. Calathea Ovary with 1-ovulate locule (the other 2 being empty and compressed); fruit 1-seeded ......................................................................................... 2 Cymules 1-flowered (i.e., flowers unpaired) ............................. 6. Monotagma Cymules 2-flowered (i.e., flowers in pairs) ................................................ 3 Inflorescence long and ± terete, ± tail-like with bracts (spathes) remaining rolled up and tightly imbricated throughout anthesis; corolla tube > 10 times longer than wide; seed with perisperm canal simple throughout .......................................................................... 4. Ischnosiphon Inflorescence usually ± capitate or diffuse in outline, never tail-like , with bracts (spathes) loosening up at anthesis and usually ± spreading; corolla tube usually < 10 times longer than wide; seed with perisperm canal branched distally or throughout .................................................. 4 Leaf sheath flanks prematurely decaying, leaving a network of persistent fibers ....................................................................................... 3. Hylaeanthe Leaf sheath flanks remaining fleshy throughout life span of the leaf, when eventually disintegrating, not leaving a network of fibers ........ 5 Herbs of marshes; outer staminode solitary; style with very long ventral process from rim of stigmatic orifice, helically twisted when triggered; seed with perisperm canal branched from base .......................... 9. Thalia Herbs of terra firme or in briefly flooded forests; outer staminodes paired or none; style without ventral process from rim of stigmatic orifice, circinnately curved when triggered; seed with perisperm canal branched

Calathea 221

6(5).

6.

7(6).

7.

8(6). 8.

distally .................................................................................................... 6 Leaf blades all of the same symmetry (homotropic), i.e., with the narrow half consistently to either the right or the left (leaf blades antitropic in Maranta ruiziana, but then < 7 cm long and with midrib hirsute above); when tall and shrubby, all leaves separated by distinct internodes ......................................................................................................... 7 Leaf blades of two alternating symmetry forms (antitropic), with the narrow side of the blade alternating between right and left following the sequence on a particular shoot; when tall and shrubby, leaves clustered at nodes .................................................................................................. 8 Inflorescences head-like, bracts (spathes) 6–16, white, all turned to one side, arising from both center and axils of a basal rosette of leaves; herbs with slender, nearly vertically descending rhizome branches bearing distal, corm-like swellings ........................................... 7. Myrosma Inflorescences cylindrical or diffuse in outline (due to long, spreading cymule axes), bracts (spathes) 1-ranked, arising only from center of basal rosette of leaves , or at tips of leafy branches; runners when present ± horizontal, filiform throughout or fleshy throughout ............... 5. Maranta Sepals acute at apex; callose staminode distally petaloid; corolla tube longer than wide; bracts (spathes) persistent ......................... 2. Ctenanthe Sepals obtuse and hooded at apex; callose staminode fleshy throughout; corolla tube wider than long; bracts (spathes) deciduous ... 8. Stromanthe

1. CALATHEA G. Mey., Prim. Fl. Esseq. 6. 1818. Goeppertia Nees, Linnaea 6: 337. 1831. Monostiche Körn., Gartenflora 7: 88. 1858. Thymocarpus Nicolson, Steyerm. & Sivad., Brittonia 33: 22. 1981. by Helen Kennedy Herbs 0.1–4 m tall, aerial shoots usually unbranched. Leaves mostly basal or basal and cauline, occasionally all cauline; blades homotropic, ovate to elliptic, rarely obovate, apex rounded, obtuse, or acuminate, base cuneate to rounded, glabrous to pubescent. Inflorescences terminal on leafy shoots, occasionally on a separate, nonleafy shoot, solitary or occasionally several, the latter axillary and often clustered; bracts spirally arranged or distichous, green, white, or colored, persistent, rarely disintegrating; secondary bracts usually present; bracteoles membranous or claviculate, rarely absent. Flowers paired, usually sessile, opening spontaneously or remaining closed, rarely cleistogamous. Corolla tube elongated, 0.7–5 cm long, 4–25 × as long as wide. Outer staminodes (0)1. Ovary with 3 fertile 1-ovulate locules. Capsule 3-locular, 3-seeded. Seeds with basal, white, rarely colored aril. Mexico, Central America, South America; ca. 255 species, ca. 29 in Venezuela, 24 of these in the flora area. Key to the Species of Calathea 1.

Bracts obviously dimorphic, the lower fertile ones (subtending flowers) emarginate at tip, the terminal sterile ones (not subtending flowers) acute to obtuse at tip; inflorescence l per aerial shoot, cylindric, length > 4.5 times width; leaves all basal, patterned with 2 submarginal darker bands and the midrib area pale green .............................. C. densa

222

1.

2(1).

2.

3(2). 3.

4(3).

4.

5(3).

5.

6(5).

6.

7(6).

7.

8(6).

M ARANTACEAE

Bracts all fertile and similar; inflorescence 1 or more per aerial shoot, not cylindric or length < 4 times width; leaves various, not patterned or, if patterned, without submarginal darker bands .................................... 2 Inflorescences 1–2.5 cm long; bracts 3–10, < 1.4 × 1.4 cm; plants 9–45 cm tall; corolla tube ≤ 10 mm long; bracteoles 2, membranous, lateral not medial ........................................................................................... C. micans Inflorescences 3.5–25 cm long; bracts 3–50 or more, 2–8 × 1–8.4 cm; plants 30–350 cm tall; corolla tube 11–45 mm long; bracteoles absent, or else 1 or 2 and triangular, claviculate (stiff, hardened, needle- or awl-shaped, and functioning as extrafloral nectaries), or 1–4 and membranous, but with at least 1 medial ............................................................................ 3 Leaf blades pubescent throughout on both surfaces at 10× magnification ................................................................................................................ 4 Leaf blades glabrous on one or both surfaces or if pubescent on upper surface, hairs restricted to midrib and adjacent area, the apical margin or the base, but not throughout ................................................................. 5 Pulvinus densely pubescent on front and back but glabrous on sides, hairs conspicuous to the naked eye, 1–2 mm long; bracts 3–5, conspicuously villous with hairs to 3.5 mm long, green; flowers bright yellow; leaf often patterned with subrectangular brownish to dark green blotches on both sides of the midrib ................................................................ C. villosa Pulvinus densely pubescent on front, glabrous to subglabrous on back; bracts 6–10, glabrous, lavender-tinged; flowers lavender; leaf solid green ............................................................................................. C. liesneri Bracts and prophylls deciduous, the point of attachment of the flower pair readily visible, the bracts 4 or 5; bracteoles triangular, thickened, ca. 3– 4 × 3 mm; leaves borne singly, stiff, coriaceous, blade glabrous; capsule tuberculate .............................................................................. C. cannoides Bracts persistent, though occasionally dying early and disintegrating in age, but attachment of flower pairs not exposed (except C. elliptica, which has 6–12 persistent bracts); bracteoles claviculate, membranous, or absent, but not triangular; leaves various; capsule not tuberculate (except C. elliptica) ................................................................................ 6 Bracts distichous or subdistichous (but often appearing distichous when pressed); basal leaves 2–several, cauline ones 1 or 2; plants 1.3–3.5 m tall; inflorescence 6–30 cm long ............................................................. 7 Bracts spirally arranged; leaves 1 or more, all basal, or if 2 or more cauline, then pulvinus minutely tomentose; plants 0.35–2 m tall; inflorescence 3–15 cm long ........................................................................... 8 Lower surface of leaves white, waxy, apex broadly obtuse, rounded or truncate, the apical margin densely tomentose on the upper surface; bracts 6–12, subdistichous, maroon to brown; petals purple; staminodes yellow .................................................................................... C. lutea Lower surface of leaves gray-green or purple-tinged, not waxy, apex obtuse with acumen, glabrous to sparsely tomentose; bracts 18–34, distichous, bright orange to red-orange; petals and staminodes orange .................................................................................................... C. casupito Bracteoles claviculate (stiff, hardened, needle- or awl-shaped, functioning

Calathea 223

8. 9(8).

9. 10(9). 10.

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

12. 13(12).

13.

14(11). 14. 15(14).

15.

16(8). 16. 17(16).

17.

as extrafloral nectaries) ......................................................................... 9 Bracteoles membranous or absent .......................................................... 16 Bracts 3 or 4; pulvinus < 6 mm long; leaves > 3.5 times as long as wide, narrowly cuneate at base, acuminate at tip, often patterned with a purple band along midrib ...................................................... C. acuminata Bracts 5–60; pulvinus > 10 mm long; leaves various, but not as proportionately narrow or apex and/or base obtuse ...................................... 10 Bracts 6–12, spreading; rachis visible in flowering inflorescences; leaves with white lines or plain green on upper surfaces ................... C. elliptica Bracts 5–60, imbricate at least at base; rachis not visible in flowering inflorescences; leaves solid green, patterned with a pale band along midrib, patterned with a dark green band along the midrib and a paler area between midrib and margin, or in juvenile foliage, patterned with pink lines .............................................................................................. 11 Pulvinus 5.5–13 cm long; bracts 30–60 ................................................... 12 Pulvinus 1–5.5 cm long; bracts 5–32 ....................................................... 14 Inflorescence < 7 × 7 cm; bracts rotting to fibers; leaves usually without pattern, rarely with a pale brushed pattern between midrib and margin .................................................................................................. C. altissima Inflorescence > 8 × 8 cm; bracts not rotting or else rotting to lace-like structure ............................................................................................... 13 Bracts crinkled, papery, not rotting to lace-like structure; sepals densely appressed-tomentose at base; peduncle with dense, short, matted hairs below the inflorescence; juvenile leaves with pink lines ........... C. grandis Bracts rotting to lace-like structure; sepals glabrous at base; peduncle minutely hispid, the hairs stiff, their bases enlarged, wart-like; juvenile leaves plain green or patterned dark green along the midrib with a lighter green or yellow-green band on either side ...................... C. fragilis Inflorescence on a separate leafless shoot; leaves < 2 times as long as wide; peduncle < 15 cm long ............................................. C. cataractarum Inflorescence terminal on the leafy shoot, leaves > 2 times as long as wide, or if less, then peduncle > 30 cm long ................................................. 15 Leaves 2.7–5 times as long as wide; leaf base cuneate; leaf not patterned; pulvinus articulate with a raised ridge at junction with petiole; bracts green to purple ...................................................................... C. neblinensis Leaves 1.8–2.6 times as long as wide; leaf base obtuse to rounded, abruptly attenuate; leaf often patterned with a pale band along midrib; pulvinus not articulate with a raised ridge at junction with petiole; bracts pink ............................................................................. C. cyclophora Bracts 6–10 cm long, acuminate-attenuate at tip, longer than wide ......................................................................................................... C. ovata Bracts < 6 cm long, various ...................................................................... 17 Leaves both basal and cauline; bracts wider than long, 1.8–3.2 cm long; flowers closed at anthesis, cream-colored or purple; plants deciduous, dying back to the rhizome .......................................................... C. latifolia Leaves all basal; bracts longer than wide; flowers open at anthesis, or if closed, then bright yellow; plants not dying back to rhizome (except C. panamensis) ......................................................................................... 18

224

M ARANTACEAE

18(17). Pulvinus 0.4–2 cm long; bracteoles membranous; plants 20–60 cm tall; peduncle short (to 10 cm) or absent; inflorescences terminal on a leafy shoot ..................................................................................................... 19 18. Pulvinus 2–11 cm long; bracteoles present or absent; plants 50–250 cm tall; peduncles > 5 cm long (except in C. zingiberina); inflorescences terminal on a leafy shoot or on a separate leafless shoot ................... 21 19(18). Upper surface of leaf minutely tomentose apically along margin of broader side; pulvinus 1.5–2 cm long, terete; bracts pubescent on outer surface with hairs much more dense at the margin, subglabrous within ................................................................................................ C. mishuyacu 19. Upper surface of leaf pubescent along midrib or sparsely pubescent at base; pulvinus 0.4–1 cm long, laterally flattened; bracts pubescent with evenly distributed hairs on outer surface and inner surface apically .............................................................................................................. 20 20(19). Pulvinus minutely pubescent along center back; lateral veins 11–14 per 3 cm and 20–22 cross veinlets per 5 mm (both measured near midpoint of broader side of leaf); corolla tube densely appressed-pilose, lobes sparsely pilose; internodes sparsely pubescent ..................... C. lasseriana 20. Pulvinus glabrous along center back; lateral veins 18–20 per 3 cm and 24– 29 cross veinlets per 5 mm; corolla tube and lobes densely appressedpilose; base of leaf sheath and internodes sericeous .......... C. panamensis 21(18). Bracts < 30; inflorescences borne on a separate leafless shoot; plants 0.5– 1.3 m tall; bracts and prophylls persistent; juvenile foliage unpatterned; pulvinus 2–6 cm long; peduncle 1.3–15 cm ............................ 22 21. Bracts > 30; inflorescences terminal on a leafy shoot; plants 1.5–2.5(–3) m tall; bracts and prophylls shredding into fibers or rotting off in fruit or earlier; juvenile foliage patterned with a broad yellow-green area on each side of the darker green midrib or plain green; pulvinus 3–11 cm; peduncles 30–100 cm long ................................................................... 23 22(21). Leaves > 2.5 times as long as wide; bracts 2–5; peduncle densely pubescent just below bracts; plant 60–130 cm tall; bracteoles absent ............................................................................................... C. zingiberina 22. Leaves < 2.5 times as long as wide; bracts > 8; peduncle glabrous just below bracts; plants 50–80 cm tall; bracteoles membranous ....... C. erecta 23(21). Bracts dying during flowering, shredding apically in late fruit, the individual bracts readily discernible, length > 3 times width; corolla ± glabrous ......................................................................................... C. variegata 23. Bracts dying and shredding apically before flowering, densely crowded, straw-like, the individual bracts often difficult to discern, length < 2.5 times width; corolla densely pilose ............................... C. inocephala Calathea acuminata Steyerm., Fieldiana, Bot. 28: 161. 1951. Herb 0.5–1.1 m tall; leaves all basal, often with dark purple band along midrib. Evergreen forests on terra firme or partially flooded forests, often on white sands, 100– 200 m; Amazonas (Río Atabapo basin, lower Río Barío, Río Negro basin). Brazil (northern Amazonas). ◆Fig. 149.

Calathea altissima (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 141. 1862. —Phrynium altissimum Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 20. 1838. Herb 0.8–2.1 m tall; leaves all basal, rarely 1 cauline. Evergreen lowland forests on terra firme, 100–300 m; Amazonas (Río Casiquiare, Río Negro, Río Orinoco, and Río

Calathea 225

Sipapo basins). Apure; widespread in Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Calathea cannoides (Nicolson, Steyerm. & Sivad.) H. Kenn., Phytologia 69: 375. 1990. —Thymocarpus cannoides Nicolson, Steyerm. & Sivad., Brittonia 33: 24. 1981. Herb to 1.8 m tall, 1 leaf per shoot, inflorescence on a separate, nonleafy shoot. Evergreen forests, 100–600 m; Bolívar (Cerro Pitón, upper Río Cuyuni). Brazil (Amazonas, Maranhão, Pará, Rondônia). ◆Fig. 150. Calathea casupito (Jacq.) Schult., Mantissa 1: 8. 1822. —Maranta casupito Jacq., Fragm. Bot. 51, t. 64, fig. 2. 1809. Calathea trinitensis Britton, Bull. Torrey Bot. Club 48: 329. 1921. Herb 1.5–2 m tall; leaves basal and 1 cauline; bracts orange to red-orange. Montane evergreen forests, 1200–1400 m; Bolívar (Sierra de Maigualida), Amazonas (Sierra de la Neblina). Aragua, Carabobo, Miranda, Sucre, Yaracuy; Colombia, Trinidad, Guyana. ◆Fig. 151. Calathea cataractarum K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 95. 1902. —Tirita. Herb 20–40 cm tall; leaves basal, 1 or 2 per shoot. Lowland forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Guyana, Brazil (Amazonas: upper Rio Negro). Calathea cyclophora Baker, Bull. Misc. Inform. Kew 1895: 17. 1895. Herb 0.5–1 m tall; leaves basal; bracts pinkish. Semideciduous to evergreen lowland or lower montane forests, shrub savannas, granitic outcrops, 50–700 m; Delta Amacuro (upper Caño Arature, Serranía de Imataca), Bolívar (Altiplanicie de Nuria), Amazonas (Puerto Ayacucho to Samariapo road, Río Atabapo basin, Río Casiquiare, upper Río Negro basin, Río Orinoco). Amazonian Colombia, Guyana, Brazil. The species cited as Calathea cyclophora by A. M. E. Jonker-Verhoff and F. P. Jonker 1968 (Flora of Suriname 1(2): 169) is C. maasiorum H. Kenn. Calathea densa (K. Koch) Regel, Gartenflora 18: 98. 1869. —Phrynium

densum K. Koch, Wochenschr. Vereines Beförd Gartenbaues Königl. Preuss. Staaten 7: 277. 1864. Herb 30–50 cm tall; leaves basal. Evergreen forests, 100–500 m; Amazonas (upper Río Orinoco). Suriname, Amazonian Brazil. Calathea elliptica (Roscoe) K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 75. 1902. —Phrynium ellipticum Roscoe, Monandr. Pl. Scitam. t. 44 (inflorescence only), 1828 [1827]; t. 42 leaves only. 1828 [1826]. Calathea vittata hort. ex Körn., Gartenflora 7: 88. 1858. Herb 0.7–1.5 m tall; leaves 1–4, basal, sometimes patterned with white lines. Evergreen forests, savannas, 50–900 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (El Paují). Colombia, Guyana, Suriname, French Guiana, northern Brazil. Calathea erecta L. Andersson & H. Kenn., Nord. J. Bot. 6: 448. 1986. Herb 0.5–0.8 m tall; leaves 1–4(–6), basal; inflorescence usually on a separate shoot. Expected in Amazonas (near Sierra de la Neblina). Amazonian French Guiana, Brazil (Amapá, Amazonas: near Pico Neblina, Manaus). Calathea fragilis Gleason, Bull. Torrey Bot. Club 56: 21. 1929. —Bijau. Calathea duidae Steyerm., Fieldiana, Bot. 28: 162. 1951. Herb 1.3–2.5 m tall; leaves 2–6, basal; bracts rotting, leaving a lace-like network. Evergreen to semievergreen forests, 100–900 m; Bolívar (Gran Sabana, south and east of Río Caroní), Amazonas (Cerro Duida, Sierra de la Neblina). Guyana, Suriname, northern Amazonian Brazil. The leaves of Calathea fragilis are used for wrapping hallacas (tamales). Calathea grandis Petersen in Mart., Fl. Bras. 3(3): 124. 1890. —No’samo adü. Herb 1.3–3 m tall; leaves all basal; bracts dying in early flowering, becoming papery. Semideciduous to moist forests, along streams, 100–400 m; Bolívar (west of El Dorado, Río Caura, Río Cuyuni). Monagas (near Cachipo); Trinidad, Guyana, Suriname, French Guiana. Calathea inocephala (Kuntze) H. Kenn. & Nicolson, Ann. Missouri Bot. Gard. 62:

226

M ARANTACEAE

501. 1975. —Phyllodes inocelphalum Kuntze, Revis. Gen. Pl. 2: 694. 1891. Calathea barbillana Cufod., Ann. Naturhist. Hofmus. 46: 235. 1933. Calathea altissima auct. non (Poepp. & Endl.) Körn. 1862: sensu Woodson and Sherry, Ann. Missouri Bot. Gard. 32: 94. 1943 [1945]; Standl. & Steyerm., Fieldiana, Bot. 24: 209. 1952; Standl., Pub. Field Mus. Nat. Hist., Bot. Ser. 18: 192. 1937. Herb 1.5–3 m tall; leaves 2–4, basal, rarely 1 cauline. Moist to semideciduous forests on terra firme, 200–800 m; Delta Amacuro (near Serranía de Imataca), Bolívar (northeast of Serranía Pia-zoi), Amazonas (upper Río Orinoco). Apure, Barinas, Mérida, Miranda, Portuguesa, Táchira, Zulia; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Trinidad, Ecuador, Peru, Brazil. Calathea lasseriana Steyerm., Fieldiana, Bot. 28: 163. 1951. Herb to 0.6 m tall; leaves all basal; inflorescence basal. Evergreen lowland forests, ca. 200–300 m; Bolívar (Río Caura near Salto Pará), Amazonas (southeast base of Cerro Duida). Endemic. When the floral structure is known, this species may prove to be conspecific with the vegetatively similar Calathea propinqua (Poepp. & Endl.) Körn. Calathea latifolia (Willd. ex Link) Klotzsch in R.M. Schomb., Reis. Br.-Guiana 3: 918. 1848. —Alpinia latifolia Willd. ex Link in Spreng. et al., Jahrb. Gewächsk. 1(3): 22. 1820. —Thalia latifolia (Willd. ex Link) Link ex Schult. in Roem. & Schult., Syst. Veg. 1: 10. 1848. —Konono. Phyllodes platyphyllum Kuntze, Revis. Gen. Pl. 2: 696. 1891. Calathea allouia var. violacea auct. non Calathea violacea Lindl. 1825: sensu Woodson, Ann. Missouri Bot. Gard. 29: 332. 1942. Deciduous herb 0.8–2 m tall; leaves basal and cauline. Semi-deciduous forests, ca. 100 m; Bolívar (Corozal Village near Río Maniapure, Río Ore in Río Parguaza basin), Amazonas (Río Manaviche). Apure, Aragua, Barinas, Cojedes, Guárico, Lara, Miranda, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; Panama, Colombia, Trinidad, western Ecuador.

Calathea liesneri H. Kenn., Novon 3: 49. 1993. Rosulate herb 0.4–0.8 m tall; leaves 2–4, basal. Submontane and montane evergreen forests, 600–1000 m; Amazonas (between Cerro Duida and Cerro Marahuaca, eastern slopes of Cerro Huachamacari). Endemic. Calathea lutea (Aubl.) Schult., Mantissa 1: 8. 1822. —Maranta lutea Aubl., Hist. Pl. Guiane 4. 1755. —Aru-maka, Kasujö. Maranta cachibou Jacq., Fragm. Bot. 52, t. 69, 70. 1809. Calathea discolor G. Mey., Prim. Fl. Esseq. 7. 1818. Calathea magnifica C.V. Morton & Skutch, J. Wash. Acad. Sci. 20: 372. 1930. Herb 1.6–4 m tall; leaves 3–7, basal, 1 cauline, the lower surface white-waxy. Evergreen and semievergreen forests, often in full sun or along stream banks, near sea level to 200 m; Delta Amacuro (Caño Araguao, Piacoa, Río Acure basin), Bolívar (Río Nichare, lower Río Caura basin), Amazonas (Río Mawarinuma). Apure, Aragua, Barinas, Falcón, Mérida, Miranda, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, western Ecuador, Peru, Brazil. ◆Fig. 154. The petioles of Calathea lutea are used for making baskets, hence the origin of the name Calathea. Calathea micans (Mathieu) Körn., Gartenflora 7: 87. 1858. —Maranta micans Mathieu, Cat. Pl. 1853. —Phrynium micans (Mathieu) Klotzsch, Allg. Gartenzeitung 22: 249. 1854. Phrynium pusillum K. Koch, Berliner Klin. Wochenschr 1863: 359. 1863. Maranta amabilis Linden, Cat. Pl. Exot. 22: 6. 1869. —Calathea micans var amabilis (Linden) Petersen in Mart., Fl. Bras. 3(3): 97. 1890. Calathea albicans Brongn. ex K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 112. 1902. Calathea klugii J.F. Macbr., Field Mus. Nat Hist., Bot. Ser. 11: 55. 1931. Herb to 30(–45) cm tall; bracts 3–8(–10) per inflorescence. Submontane evergreen and swampy mature forests, 200–300 m; Delta Amacuro (east of Río Grande), Bolívar (Serranía Imataca). Apure, Aragua, Cara-

Calathea 227

Fig. 149. Calathea acuminata

Fig. 150. Calathea cannoides

228

M ARANTACEAE

Fig. 151. Calathea casupito

Fig. 152. Calathea villosa

Calathea 229

Fig. 153. Calathea micans

bobo, Miranda, Táchira, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil, Bolivia. ◆Fig. 153. Calathea mishuyacu J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 54. 1931. Herb 0.8–1.2 m tall; leaves 2–5, basal. Shrub savannas, 200–300 m; Bolívar (southeast of El Manteco). Amazonian Ecuador, Peru. Calathea neblinensis H. Kenn., Phytologia. 69: 373. 1990. Herb 0.5–1.5 m tall; leaves 3–9, basal. Evergreen lowland forests on sandy ultisols, 50– 200 m; Amazonas (Culimacare, base of Sierra de la Neblina, near San Carlos de Río Negro). Colombia (Vaupés), Brazil (Amazonas). Calathea ovata (Nees & Mart.) Lindl., Bot Reg. 14: pl. 1210. 1828, in notes. —Phrynium ovatum Nees & Mart., Nova

Fig. 154. Calathea lutea

Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 11: 27. 1823. Caulescent. Evergreen lowland forests, 100–200 m; Amazonas (Río Yatúa at mouth of Río Yaciba). Southeastern Brazil. Calathea panamensis Rowlee ex Standl., J. Wash. Acad. Sci. 15: 4. 1925. Deciduous herb 0.3–0.6 m tall; leaves all basal; inflorescence basal. Semideciduous and riparian forests, shrub savannas, 50–300 m; Bolívar (Río Caura at Salto Pará, lower Río Parágua), Amazonas (near Puerto Ayacucho, San Juan de Manapiare). Apure, Barinas, Carabobo, Cojedes, Portuguesa; Nicaragua, Costa Rica, Panama, Colombia, Ecuador. Calathea variegata Linden ex Körn., Mitt. Russ. Gartenbauver. St. Petersburg 2: 93. 1860. —Casupo. Herb 1.3–3 m tall; leaves all basal; juvenile foliage with dark green leaflet-like pattern along the midrib. Semievergreen for-

230

M ARANTACEAE

ests, shrub savannas, 100–400 m; Bolívar (Río Asa, lower Río Paragua, San Francisco). Barinas, Cojedes, Distrito Federal, Monagas, Portuguesa; Amazonian Colombia, Trinidad, Suriname, Ecuador, Peru, Brazil. Calathea villosa Lindl., Edward’s Bot. Reg. 31: t. 14. 1845. —Casupo amarillo, Casupo. Calathea pardina Planch. & Linden in Linden, Cat. Pl. Exot. 2. 1855. —Calathea villosa var. pardina (Planch. & Linden) Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 141. 1862. Calathea hirsuta Standl., J. Wash. Acad. Sci. 15: 4. 1925. Deciduous herb 0.5–1.2 m tall; leaves all basal, villose throughout. Semideciduous forests, 200–700 m; Bolívar (Los Pijiguaos, near confluence of Río Parágua and Río Caroní,

Río Parguaza, near Upata), Amazonas (near Puerto Ayacucho). Carabobo, Cojedes, Portuguesa; Nicaragua, Costa Rica, Panama, northern Colombia, Guyana, Suriname, northeastern Brazil. ◆Fig. 152. A variety (Calathea villosa var. glabra Schum.) has been described from Ceará, Brazil, but it is not clear whether it is sufficiently distinct to warrant that rank. Calathea zingiberina Körn., Bull. Soc. Naturalistes Moscou 35: 122. 1862. Herb 0.7–1.7 m tall; leaves borne singly; inflorescence borne on a separate, nonleafy shoot; bracts 2–4. Evergreen and riparian forests, terra firme, 200–900 m; Bolívar (northeast of El Manteco, upper Río Caura, Río Parágua basin), Amazonas (near Puerto Ayacucho). Amazonian Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia.

Fig. 155. Ctenanthe compressa

Hylaeanthe 231

2. CTENANTHE Eichler, Abh. Königl. Akad. Wiss. Berlin 1883: 81. 1884. by Lennart Andersson Rosulate or caulescent herbs, without specialized rhizome branches. Leaves antitropic. Inflorescence simple or a few-branched synflorescence. Florescence usually ± markedly dorsiventral with all flowers exposed on 1 side; bracts (spathes) usually green, usually persistent, herbaceous to coriaceous; florescence component with several to many cymules, cymule with distinct but usually very short axes (a few mm or less), without interphyll, often with 1 or 2 scale-like, dorsal bracteoles. Sepals ovate to elliptic, obtuse and ± hooded at apex; corolla tube 1–2 times longer than wide. Outer staminodes 2, ± equal, petaloid and showy, callose staminode distally petaloid and showy, stamen with small and inconspicuous, narrowly obovate appendage. Ovary 1-ovulate. Fruit a rather thin-walled capsule; pericarp ± papery at maturity. Seed arillate; perisperm canal branched distally. Neotropics; ca. 12 species, 1 in Venezuela. Ctenanthe compressa (A. Dietr.) Eichler, Abh. Königl. Akad. Wiss. Berlin 1882: 83. 1883. —Maranta compressa A. Dietr., Sp. Pl. 1: 22. 1831. Caulescent herb 0.5–1 m tall. Mostly on boulders and granitic outcrops in forested regions, near sea level to 500 m; Delta Ama-

curo (eastern side of Río Cuyubiní), Bolívar (Altiplanicie de Nuria, south of El Dorado, road between Tumeremo and El Dorado, Río Grande/Río Toro), Amazonas (Río Yureba, San Juan de Manapiare). Brazil. ◆Fig. 155. Identification of this Guayanan taxon is highly tentative.

3. HYLAEANTHE A.M.E. Jonker & Jonker, Acta Bot. Neerl. 4: 172. 1955. Myrosma L. f., pro parte, excl. type: sensu K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 140. 1902. by Lennart Andersson Rosulate herbs with strikingly soft-textured foliage, with specialized, distally starch-storing rhizomes, and subterminal root tubers. Leaves homotropic; sheath flanks obovate, prematurely disintegrating, leaving a network of persistent fibers. Inflorescence a terminal synflorescence of (1)2–6 florescences; florescences markedly dorsiventral; bracts (spathes) softly herbaceous; florescence component of 2 or 3 cymules; cymule 2-flowered, with distinct and often rather long axes, interphyll and bracteoles absent. Sepals oblong to sublinear, acute to obtuse or ± truncate; corolla tube 2.5–4.5 times longer than wide. Outer staminodes 2 and markedly unequal to solitary, petaloid and showy, callose staminode petaloid and showy distally, fertile stamen with a petaloid appendage ± equaling the inner staminodes. Ovary 1-ovulate. Fruit an asymmetrical capsule with rather thick, parenchymatic wall. Seeds arillate; perisperm canal branched distally. Mainly Amazonian; ca. 6 species, 1 species in Venezuela. Hylaeanthe unilateralis (Poepp. & Endl.) A.M.E. Jonker & Jonker, Acta Bot. Neerl. 4: 175. 1955. —Thalia unilateralis Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 24, t. 133. 1838. —Myrosma unilateralis (Poepp. & Endl.) K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 144. 1902.

Rosulate herb 0.5–1 m tall, with runners and root tubers. Understory of wet forests, at low altitude, 100–400 m; Bolívar (Río Caura, Salto Pará), Amazonas (Río Asisa, Río Orinoco near the mouth of Río Ugueto). Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 156.

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M ARANTACEAE

Fig. 156. Hylaeanthe unilateralis

Ischnosiphon 233

4. ISCHNOSIPHON Körn., Nouv. Mém. Soc. Imp. Naturalistes Moscou 11: 346. 1859. by Lennart Andersson Low and rosulate to tall and shrub-, tree-, or liana-like herbs. Leaves homotropic. Inflorescences usually clustered into terminal synflorescences, but synflorescences grading into simple, lateral inflorescences; florescences long and tail-like with bracts (spathes) remaining imbricate at anthesis; bracts (spathes) ± sclerotic; florescence composed of 1–17 cymules; cymule 2-flowered, axes undifferentiated, interphyll occasionally present; bracteoles 1 or 2, small and scale-like to long and strap-shaped with terminal gland (nectary). Sepals linear or sublinear, acute; corolla tube 10–30 times longer than wide. Outer staminode solitary, petaloid and showy, sometimes precociously shed (sect. Bambusastrum), callose staminode firm and fleshy throughout or petaloid distally, fertile stamen with a usually small appendage. Ovary 1-ovulate. Fruit an obliquely ellipsoid capsule (sometimes indehiscent?) with thin sclerotic wall. Seed arillate; perisperm canal simple throughout. Central America, Antilles, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, 35 species, 12 in Venezuela, all in the flora area. Key to the Species of Ischnosiphon 1. 1. 2(1).

2.

3(2).

3.

4(2). 4.

5(4). 5.

Leaf blade with apex displaced > 1 cm to 1 side of an imaginary straight line continuing the midrib ..................................................................... 2 Leaf blade with apex displacement < 1 cm ............................................... 7 Herbs erect with leaves gathered into a fan-shaped cluster on top of an unbranched, cane-like stem; inflorescences clustered into a distinct synflorescence which is terminal on the main shoot; outer staminode wilting attached ..................................................................................... 3 Herbs scandent with leaves on lateral short shoots, or shrubby plants with leaves ± evenly distributed along branches; inflorescences on lateral branches, solitary or 2(–4) together; outer staminode usually shed shortly after the bud opens (except in I. ursinus) ................................ 4 Leaf blades papillose (25×), the lower surface usually distinctly whitish and waxy; inflorescences 6–12 mm diameter; outer staminodes bright yellow .......................................................................................... I. obliquus Lower surface of leaf blades neither papillose nor waxy, grayish green; inflorescence 4–6 mm diameter; outer staminodes usually purple, at least distally ................................................................................. I. arouma Indument on leaves and inflorescences reddish brown; leaf blades broadest above middle, ± cuneate at base; erect, shrubby plants ....... I. ursinus Indument on leaves and inflorescences, when present, grayish to pale yellowish; leaf blades broadest at or below middle, usually ± rounded at base; plants usually scandent ................................................................ 5 Leaf blades usually 3–4 cm wide, with apex displacement usually < 1.5 cm; inflorescence solitary, usually on a leafless shoot ........................ I. gracilis Leaf blades usually > 4 cm wide, with apex displacement usually > 1.5 cm; inflorescences usually on leafy shoots, often in fascicles of 2 .............. 6

234

M ARANTACEAE

6(5).

Lower surface of leaf blades papillose (25×), usually white and waxy; inflorescence 4.5–6 mm diameter; bracts (spathes) densely hairy, > 4 cm long ........................................................................................ I. enigmaticus 6. Lower surface of leaf blades neither papillose nor waxy, ± grayish green; inflorescence 2–5 mm diameter; bracts (spathes) glabrous or inconspicuously hairy, > 4 cm long ................................................... I. puberulus 7(1). Shrubby or scandent herbs, well over 1 m tall when flowering; major stems segmented and bamboo-like, dark green .................................... 8 7. Rosulate herbs, usually < 1 m tall (exceptionally to 2.5 m), ± leafy at base; stems ± herbaceous, pale green, drying straw-colored ....................... 11 8(7). Herbs growing in black-water swamps and rivers; leaf blades obtuse at the very tip; inflorescences ± clustered ................................. I. polyphyllus 8. Herbs on terra firme; leaf blades sharply pointed at apex; inflorescences solitary ................................................................................................... 9 9(8). Major stems usually ± scabrous, especially below nodes; inflorescences usually borne on short, leafless or few-leaved branches; outer staminode precociously shed, callose staminode firm and fleshy throughout, shorter than outer staminode .................................. I. gracilis 9. Major stems smooth; inflorescences borne on long, many-leaved branches; outer staminode persistent, callose staminode petaloid distally, ± equaling outer staminode .................................................................... 10 10(9). Pulvinus minutely hirtellous adaxially; bracts (spathes) rather distinctly appressed-pilose ...................................................................... I. longiflorus 10. Pulvinus glabrous adaxially; bracts (spathes) glabrous or minutely puberulous at apex .............................................................. I. surumuensis 11(7). Leaf blades papillose and waxy on lower surface; bracts (spathes) minutely and indistinctly puberulous; bracteoles 1 per cymule, with spatulate gland .................................................................................... 12 11. Leaf blades neither papillose nor waxy on lower surface; bracts (spathes) ± densely hirsute with coarse, spreading hairs; bracteoles 2 per cymule, with needle-like gland ................................................................ I. hirsutus 12(11). Leaf sheaths ± tightly enclosing the stem, rigid, ± conspicuously striate from raised fiber bundles; blade 2.7–5.6 times longer than broad ................................................................................................ I. cannoideus 12. Leaf sheaths ± spreading, leaving upper part of stem visible, of ordinary herbaceous texture (drying papyraceous), ± smooth; blade 1.3– 3.6 times longer than wide ................................................... I. leucophaeus Ischnosiphon arouma (Aubl.) Körn., Nouv. Mém. Soc. Imp. Naturalistes Moscou 11: 348. 1859. —Maranta arouma Aubl., Hist. Pl. Guiane 3. 1775. Herb to 3.5 m tall, with all leaves borne in a fan-like cluster on top of an unbranched, cane-like stem. Edges of swamp forests, along small streams, often also in secondary growth, 50–1600 m; Delta Amacuro (Caño Capurito in Caño Capure basin, east-southeast of Los Castillos de Guayana, southeast of Piacoa, Serranía Imataca), Bolívar (Icabarú, Maripa, Represa Guri, Río Asa, Río

Cataniapo, Río Caura, Río Uairén, northeast of San Carlos de Río Negro, Urimán), Amazonas (Puerto Ayacucho, Río Cuao, Río Siapa, Santa Bárbara). Apure, Miranda, Portuguesa, Táchira, Zulia; Panama, Lesser Antilles, Colombia, Guyana, Suriname, French Guiana, Peru. ◆Fig. 157. Ischnosiphon cannoideus L. Andersson, Opera Bot. 43: 81. 1977. Herb of zingiber-like growth habit, 0.5– 2.5 m tall. Savanna scrub and campina forests, on white sand and other nutrient-poor

Ischnosiphon 235

soils, 50–200 m; Amazonas (Cerro Duida, Chapazón, Río Baría, Río Pasimoni, San Carlos de Río Negro, San Juan de Ucata). Brazil (Amazonas, Pará, Rôndonia, Roraima). Ischnosiphon enigmaticus L. Andersson, Opera Bot. 43: 62. 1977. —Carutillo. Scandent herb of bamboo-like habit, usually several m tall. Light-gaps in rain forests, well-drained soils, 500–900 m; Bolívar (El Paují, San Félix to Upata road). Guyana, French Guiana. Ischnosiphon gracilis (Rudge) Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 94. 1862. —Maranta gracilis Rudge, Pl. Guian. 1: 8. 1805. Ischnosiphon lasseriana Steyerm., Fieldiana, Bot. 28: 164. 1951. Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil; 2 subspecies, 1 in Venezuela. I. gracilis subsp. gracilis Scandent, bamboo-like herb to 6 m tall. Rain forest understory on well-drained soils, 100–900 m; Bolívar (El Paují, Perai-tepui, Río Acanán, San Ignacio), Amazonas (upper Río Cuao, Río Mawarinuma). Guyana, Suriname, French Guiana, Ecuador, Brazil (along South Atlantic coast in Amapá, Bahia, and Pará). Ischnosiphon hirsutus Petersen in Mart., Fl. Bras. 3(3): 135. 1890. Rosulate herb to 1.5 m tall. Forest understory and small light-gaps in rain forests, ca. 400 m; Bolívar (Río Paramichí). Colombia, Ecuador, Peru, Brazil. Ischnosiphon leucophaeus (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 91. 1862. —Calathea leucophaea Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 21. 1838. Colombia, Venezuela, Suriname, French Guiana, Ecuador (Pacific coast), Peru, Brazil, Bolivia; 2 subspecies, 2 in Venezuela, 1 of these in the flora area. I. leucophaeus subsp. leucophaeus Rosulate herb 0.3–1.5(–2) m tall. Lightgaps in forests, often on poorly drained soils, 50–200 m; Bolívar (near Puerto Ordaz), Amazonas (base of Cerro Sipapo, Maroa). Monagas; southern Colombia, Suriname,

French Guiana (Pacific coast), Peru, Brazil, Bolivia. Ischnosiphon longiflorus K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 160. 1902. Colombia, Venezuela, Peru, Brazil; 2 subspecies, 1 in Venezuela. I. longiflorus subsp. angustifolius L. Andersson, Opera Bot. 43: 102. 1977. Tree-like or straggling or scandent herb to 7 m tall. Nonflooded forests, 100–200 m; Amazonas (San Carlos de Río Negro, San Fernando de Atabapo). Colombia, Peru, Brazil (Amazonas, Pará, Roraima). Ischnosiphon obliquus (Rudge) Körn., Nouv. Mém. Soc. Imp. Naturalistes Moscou 11: 348. 1859. —Maranta obliqua Rudge, Pl. Guian. 1: 8, tab. 2. 1805. Herb 2–4(–6) m tall with leaves in a fanlike cluster on top of an unbranched, canelike stem. River margins and light swamp forests, also common in secondary growth, 50–700 m; Delta Amacuro (Curiapo south to Sierra Imataca), Bolívar (Cerro Abismo, Río Caura, Rio Erebato, mouth of Río Paragua), Amazonas (Río Coro Coro, Río Coromoto, Río Cunucunuma, Río Mawarinuma). Mérida; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Ischnosiphon polyphyllus (Poepp. & Endl.) Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 93. 1862. —Calathea polyphylla Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 23. 1838. —Plananillita, Tirita cania grulla, Tirite. Shrubby or tree-like herb 1–3 m tall. Black-water rivers and swamps, igapó forests, 50–200 m; Amazonas (Caño Yagua, lower Río Baría, Río Emoni in lower Río Siapa basin, Río Sipapo, San Carlos de Río Negro, Río Yatúa, Tamatama). Adjacent Colombia, Brazil (Rio Negro basin and along the Amazon from the Japura to the mouth). The split stems of this species are used in basket-making. Ischnosiphon puberulus Loes., Notizbl. Königl. Bot. Gart. Berlin 6: 281. 1915. Ischnosiphon gracilis var. scabra Petersen in Mart., Fl. Bras. 3(3): 139. 1890. —Ischnosiphon puberulus var. scaber

236

M ARANTACEAE

(Petersen) L. Andersson, Opera Bot. 43: 59. 1977. Ischnosiphon verruculosous J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 56. 1931. —Ischnosiphon puberulus var. verruculosous (J.F. Macbr.) L. Andersson, Opera Bot. 43: 59. 1977. Shrubby or, usually, scandent herb 1–10 m tall. Clearings and light-gaps in rain forests, gallery forests, 100–1000 m;, Delta Amacuro (El Palmar), Bolívar (San Félix to Upata road), Amazonas (Cerro Aracamuni, Cerro Huachamacari, Río Coro Coro, Río Mawarinuma, Río Orinoco below mouth of Río Atabapo). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

Ischnosiphon surumuensis Loes., Notizbl. Königl. Bot. Gart. Berlin 6: 276. 1915. Straggling, shrubby to ± scandent herb, usually 1–3 m tall. Well-drained, nutrientpoor soil in rain forests, 100–600 m; Amazonas (slope of Cerro Aracamuni, San Carlos de Río Negro). Brazil (Río Negro, Río Branco, and Río Curuá basins). Ischnosiphon ursinus L. Andersson, Nord. J. Bot. 4: 28. 1984. Shrubby herb 1–1.5 m tall. In savannas on granitic outcrops and white sand, 100– 200 m; Amazonas (Río Mawarinuma). Suriname, French Guiana, Brazil (Amazonas: Rio Negro basin).

Fig. 157. Ischnosiphon arouma

Maranta 237

5. MARANTA L., Sp. Pl. 2. 1753. by Lennart Andersson Low rosulate or tall shrubby herbs. Leaves usually homotropic. Inflorescences clustered into synflorescences, or simple at tips of leafy branches; florescences usually few-spathed, spike-like or diffuse in outline, bracts (spathes) usually ± spreading at anthesis; bracts (spathes) herbaceous to membranous, persistent; florescence component of 2–6 cymules; cymules 2-flowered, interphyll absent; bracteoles absent or rudimentary. Sepals narrowly oblong or triangular to linear, ± acute at apex; corolla tube 2–13 times longer than wide. Outer staminodes usually 2, subequal, petaloid and showy, occasionally 1, callose staminode firm and fleshy throughout, or petaloid distally, fertile stamen with a small petaloid, oblong to obovate appendage. Ovary 1-ovulate. Fruit capsular, wall thick and parenchymatic to thin and submembranous. Seed arillate; perisperm canal branched distally. Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; ca. 30 species, 8 in Venezuela, all in the flora area. Several species, such as Maranta bicolor and M. leuconeura, are grown as ornamentals for their colorful foliage. Key to the Species of Maranta 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

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

6.

Inflorescences with numerous membranous bracts (spathes); cymule axes very short ........................................................................... M. humilis Inflorescences with 1–3 herbaceous bracts (spathes); cymule axes very long ......................................................................................................... 2 Herbs rosulate, most leaves basal; fleshy starch-storing rhizome branches present .................................................................................................... 3 Herbs caulescent, all leaves cauline, or occasionally 1 or 2 leaves basal; specialized rhizome branches absent or filiform .................................. 5 Leaf blade linear to very narrowly oblong, 1.5–1.9 cm broad in basal leaves; sepals 8–12 mm long ..................................................... M. linearis Leaf blade ovate to oblong or narrowly so, > 2.5 cm broad in basal leaves; sepals > 12 mm long .............................................................................. 4 Leaf blade distinctly pilose all over the upper surface and/or lower surface; proximal spathe usually < 40 mm long; sepals 12–16 mm long; plants (partly) autogamous and usually richly fruiting ... M. arundinacea Leaf blade glabrous on both sides, or hairy only near midrib; proximal spathe > 45 mm long; sepals 15–20 mm long; plants allogamous and rarely and sparsely fruiting ................................................... M. amplifolia Leaves antitropic, blades < 7 cm long; midrib on upper surface densely villous or hirsute ....................................................................... M. ruiziana Leaves homotropic, blades > 10 cm long, at least in some leaves of each plant; midrib on upper surface glabrous or subglabrous ..................... 6 Sepals reflexed in fruit, starting to reflex at late anthesis; abaxial side of pulvinus and lower part of midrib glabrous; leaf blade 1.5–3 times longer than broad ..................................................................... M. protracta Sepals erect in fruit; abaxial side of pulvinus and lower part of midrib ± conspicuously hairy; or, if glabrous, leaf blades > 3 times longer than wide ........................................................................................................ 7

238

7(6). 7.

M ARANTACEAE

Leaf blade < 3 times longer than wide; abaxial side of pulvinus and lower part of midrib shaggy-villous ........................................................ M. gibba Leaf blade > 3 times longer than wide; abaxial side of pulvinus and lower part of midrib glabrous or sparsely appressed-pilose .............. M. rupicola

Maranta amplifolia K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 128. 1902. Rosulate herb usually 1–2 m tall; rhizome branches starch-storing. White-sand savannas, expected to be found in Bolívar. Guyana, Suriname, French Guiana, Peru, Bolivia. Maranta arundinacea L., Sp. Pl. 2. 1753. Rosulate herb, usually 0.3–0.8 m tall; rhizome branches starch-storing. Mainly in dry tropical forests, along brooks, at pool margins, 100–300 m; Bolívar (Caicara, Coromoto), Amazonas (San Fernando de Atabapo, San Juan de Manapiare). Cojedes, Distrito Federal, Guárico, Portuguesa; West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Argentina. ◆Fig. 160. This species is cultivated for the highquality starch stored in its rhizomes. Maranta gibba Sm. in Rees, Cycl. 22. 1819. Maranta divaricata auct. non Roscoe 1828, K. Schum. 1902, A.M.E. Jonker & Jonker 1957. Caulescent, shrubby herb 1–4 m tall. Clearings and light-gaps in semideciduous forests, on deep, well-drained latosol, 50–400 m; Delta Amacuro (Los Castillos, southeast of Piacoa), Bolívar (east of Ciudad Piar, southeast of Represa Guri, middle Río Botanamo, Río Paragua, Tumeremo). Aragua, Distrito Federal, Falcón, Miranda, Portuguesa, Sucre, Zulia; Central America south to Nicaragua, Lesser Antilles, Colombia (upper Magdalena and Cauca valleys), TrinidadTobago, Guyana, Suriname, Ecuador (coastal plain), Peru. Maranta humilis Aubl., Hist. Pl. Guian. 4. 1775. Saranthe urceolata Petersen in Mart., Fl. Bras. 3(3): 167. 1890. Rosulate herb to 0.5 m tall, with filiform runners and strikingly soft-textured foliage. On deep, well-drained soil in rain forest understory, also in old secondary growth, 200– 400 m; Bolívar (Los Pijiguaos, Río Ariza), Amazonas (between Platanal and Raudal

Fig. 158. Maranta linearis

Maranta 239

Fig. 159. Maranta humilis

Fig. 160. Maranta arundinacea

240

M ARANTACEAE

Guaharibos, near San Juan de Manapiare). Suriname, French Guiana, Brazil (Acre, Amazonas, Pará, Roraima). ◆Fig. 159. Maranta linearis L. Andersson, Nord. J. Bot. 6: 741. 1986. Rosulate herb to 0.8 m tall, with fleshy starch-storing rhizome branches. Savanna woods, granitic outcrops, 50–300 m; Bolívar (southwest of Caicara, Los Pijiguaos), Amazonas (Puerto Ayacucho to Samariapo road). Colombia (Arauca). ◆Fig. 158. Maranta protracta Miq., Linnaea 18: 71. 1844. Caulescent, shrubby herb 1–3 m tall. Clearings and light-gaps in rain forests, deep, well-drained latosol, ca. 300 m; Bolívar (Ciudad Piar to La Paragua Road, middle Río Botanamo). Guyana, Suriname, Brazil

(Amazonas, Bahia, Pará, Roraima). Maranta ruiziana Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 45. 1862. Caulescent herb 0.3–1 m tall, with filiform, distally thickened and starch-storing rhizomes. Rain forest understory, 300–400 m; Amazonas (Río Manaviche). Lesser Antilles, western Colombia, Guyana, Suriname, French Guiana, western and southern Brazil, Ecuador, Peru. Maranta rupicola L. Andersson, Nord. J. Bot. 6: 751. 1986. Caulescent herb 1–2 m tall. Rock savannas and glades in white-sand forests, 200– 300 m; Bolívar (Río Toro). Falcón; Guyana, Suriname, French Guiana, Brazil (coastal Pará, and along Atlantic coast from Pernambuco to Bahia).

6. MONOTAGMA K. Schum. in Engl., Pflanzenr. IV. 48(Heft 11): 166. 1902. by Mats Hagberg Rosulate herbs. Leaves homotropic. Inflorescence usually a richly branched terminal synflorescence, sometimes congested; florescence ± spike-like, when young with the sclerotic (due to large fibre bundles) bracts (spathes) densely rolled up around the rachis and the enclosed florescence component, during anthesis bracts (spathes) spread in a comb-like manner but still enclose the florescence component; in a few species the bracts (spathes) remain imbricate throughout anthesis; florescence component of 2–8 1-flowered cymules, axis short (0.5–3 mm), interphylls present in a few species, bracteoles very rarely present. Sepals membranous, with obtuse to truncate apex; corolla tube long and narrow (length/width ratio ca. 5–30), lobes oblong with obtuse and hooded apex; outer staminode solitary, petaloid and showy, sometimes rudimentary or absent. Callose staminode usually firm and fleshy, sometimes with a petaloid rim, usually with a shelf-shaped callus. Ovary 1ovulate. Capsule with a thin sclerotic wall, opening along 1–3 longitudinal slits. Seed with a tubular aril at the base, with or without 2 lobes on one side. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 40 species, 9 in Venezuela, all in the flora area. Key to the Species of Monotagma 1. 1. 2(1).

2.

Inflorescence congested, head-like; aril tubular without lobes ................ 2 Inflorescence not head-like; aril with 2 lobes on one side ........................ 3 Scape (26–)35–81(–91) cm; inflorescence globular; peduncle of terminal florescence 0.2–0.4(–0.5) cm; bracts (spathes) bright red, 1.7–2.4 cm long; outer staminode 0.5–4.5 × 0.1–2 mm, rarely absent ... M. secundum Scape 15–52(–71) cm; inflorescence ovoid; peduncle of terminal florescence 0.4–0.8 cm; bracts (spathes) greenish to slightly reddish,

Monotagma 241

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

(2.1–)2.2–3.2 cm long; outer staminode 4–8 mm × (1–)2.5–6 mm ............................................................................................. M. tomentosum Bracts (spathes) red; flowers without an outer staminode ....................... 4 Bracts (spathes) greenish to brownish; flowers with an outer staminode ................................................................................................................ 5 Leaves with abaxial white waxy coating, midrib reaching only 2/3 of the leaf length; inflorescence subtended by a bract ................ M. rhodanthum Leaves without wax, midrib reaching apex of the leaf; inflorescence subtended by a bladed leaf ............................................... M. yapacanensis Inflorescence subtended by a bract ........................................................... 6 Inflorescence subtended by a bladed leaf .................................................. 7 Pulvinus glabrous; length from base to broadest part of the leaf blade 6–10 cm ....................................................................................... M. ovatum Pulvinus pilose throughout or at least adaxially; length from base to broadest part of the leaf blade 11–24 cm ................................ M. spicatum Petiole always absent in basal leaves; phyllotaxy spirodistichous, plants with (7–)11–25 basal leaves .................................................. M. vaginatum Petiole always present in basal leaves; phyllotaxy distichous or shoot axis irregularly twisted; 0–7(–15) basal leaves ............................................ 8 Lowermost spathe of terminal florescence 1.3–2.2 cm long; inflorescence nodes (4–)6–10; scape 12–56 cm ...................................... M. plurispicatum Lowermost spathe of terminal florescence 2.6–3.8 cm long; inflorescence nodes 3–5(–7); scape (10–)26–117 cm ........................................... M. laxum

Monotagma laxum (Poepp. & Endl.) K. Schum., Pflanzenr. IV. 48(Heft 11): 168. 1902. —Calathea laxa Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 22, pl. 130. 1838. —Lengua de vaca. Herb 0.5–2 m tall. In light gaps, along paths in primary forests, along roadsides, in secondary scrub, 50–600 m; southern Bolívar (El Paují, Icabarú), Amazonas (Caname, mouth of Caño Casiquiare, Puerto Ayacucho, Río Baría, Río Cataniapo, Río Coro Coro, Río Coromoto, Río Emoni, Río Mawarinuma, San Carlos de Río Negro, San Fernando de Atabapo). Apure; Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Monotagma ovatum Hagberg, Ann. Missouri Bot. Gard. 77: 421. 1990. Herb 0.5–0.7m tall, outer staminode orangish. Upland montane forests, in crevices or among boulders on rocky escarpments, 500–1200 m; Bolívar (Cerro Venamo, Km 105–107 on El Dorado to Santa Elena road). Adjacent Guyana. Monotagma plurispicatum (Körn.) K. Schum. in Engl., Pflanzenr. IV. 48(Heft

11): 169. 1902. —Ischnosiphon plurispicatus Körn., Bull. Soc. Imp. Naturalistes Moscou 35: 1. 1862. —Casupo, Tirita. Herb 0.3–1 m tall. Primary and slightly disturbed forests, 100–900 m; Bolívar (Cerro Camarón, Ciudad Piar, El Paují, Río Caura, middle Río Tonoro, Salto Pará, San Francisco), Amazonas (Caño Majagua off Caño Parucito, Cerro Yureba, Río Baría, Río Coro Coro, Río Mawarinuma, mouth of Río Paragua, Río Pasimoni, east-southeast of Santa Bárbara, Solano, Yavita). Mérida, Táchira, Zulia; Honduras, Nicaragua, Costa Rica, Panama, Colombia, Suriname, French Guiana, Peru, Brazil (Amazonas, Pará), Bolivia. ◆Fig. 163. Monotagma rhodanthum Maguire & Wurdack, Mem. New York Bot. Gard. 10(2): 15. 1960. Herb 1–1.5 m tall. Montane forests, ca. 800–1100 m; Amazonas (Cerro Duida, Cerro Imerí, Sierra de la Neblina). Endemic. Monotagma secundum (Petersen) K. Schum. in Engl., Pflanzenr. IV. 48(Heft

242

M ARANTACEAE

11): 167. 1902. —Ischnosiphon secundus Petersen in Mart., Fl. Bras. 3(3): 135. 1890. Monotagma surinamense Pulle, Enum. Vasc. Pl. Surinam 112, pl. 3. 1906. Herb 0.4–1 m tall. Forests on latosol, 100– 600 m; Bolívar (6 km southeast of Los Pijiguaos, Río Caura, Río Samay in Río Icabarú basin), Amazonas (Pimichín). Colombia, Suriname, Peru, Brazil. ◆Fig. 164. Monotagma spicatum (Aubl.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 14. 1931. —Maranta spicata Aubl., Hist. Pl. Guiane 4. 1775. —Casupo. Herb 0.4–1.3 m tall; outer staminode bluish. Wet places in forests, 50–1100 m; Delta Amacuro (Río Amacuro, El Palmar), Bolívar (west of Amaruay-tepui, Caño Pablo near Salto Pará, Cerro Guaiquinima, El Paují, Perai-tepui, San Ignacio), Amazonas (Sierra de la Neblina). Monagas; Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil (Amapá, Maranhão, Pará). ◆Fig. 162. Monotagma tomentosum K. Schum. ex Loes., Notizbl. Königl. Bot. Gart. Berlin 6: 286. 1915. Monotagma duidae Steyerm., Fieldiana, Bot. 28: 164. 1951. Herb 0.3–1 m tall. Confined to white-sand forests, 100–800 m; Bolívar (Icabarú, Río Apacará, Serranía Pia-Zoi), Amazonas (Cerro Duida, near San Carlos de Río Negro). Peru, Brazil. Monotagma vaginatum Hagberg, Ann. Missouri Bot. Gard. 77: 421. 990. —Lengua de vaca hoja fina. Herb 0.6–1.5 m. Confined to white-sand forests, 50–600 m; Amazonas (Cerro Aracamuni, lower Río Baría, Río Mawarinuma, San Carlos de Río Negro). French Guiana, Peru (Iquitos), Brazil (Manaus). ◆Fig. 165. Monotagma yapacanensis Steyerm. & Bunting, Acta Bot. Venez. 8: 114. 1973. Herb 1.5–2 m tall. Along streams in forests, 100–800 m; Amazonas (Caño Yutajé at base of Cerro Yutajé, Cerro Yapacana, Cerro Yutajé, Gavilán, near Puerto Ayacucho, Río Coro Coro, Río Coromoto). Endemic. ◆Fig. 161.

Fig. 161. Monotagma yapacanensis

Monotagma 243

Fig. 162. Monotagma spicatum

Fig. 163. Monotagma plurispicatum

244

M ARANTACEAE

Fig. 164. Monotagma secundum

Fig. 165. Monotagma vaginatum

Myrosma 245

7. MYROSMA L. f., Suppl. Pl. 8, 80. 1781. by Lennart Andersson Small rosulate herbs, with filiform rhizome branches distally thickened into corm-like structures. Leaves homotropic. Inflorescence simple or 2 or 3 clustered into a synflorescence, arising both from center of basal leaf rosette and from leaf axils; florescence compact, markedly dorsiventral with all flowers produced on 1 side; bracts (spathes) submembranous, persistent; florescence component with a single cymule; cymule 2-flowered, axes very short, interphyll and bracteoles absent. Sepals ± elliptical, obtuse to subacute; corolla tube about as long as wide. Outer staminodes 2, petaloid and showy, callose staminode petaloid distally, fertile stamen with a petaloid appendage ± equaling inner staminodes. Ovary 1-ovulate. Fruit capsular, thinwalled. Seeds arillate; perisperm canal branched distally. Antilles, South America; 1 species. Myrosma cannifolia L. f., Suppl. Pl. 80. 1781. Rosulate herb to 0.5 m tall, with filiform runners thickened distally into corm-like structures. Deciduous to semideciduous forests on well-drained ground, 50–100; Bolívar (74 km southwest of Caicara, Los Pijiguaos), Amazonas (south of Puerto Ayacucho, Río Sipapo). Apure, Cojedes, Guárico; Antilles, Guyana, Suriname, French Guiana, Brazil (Goiás, Maranhão, Mato Grosso, Pará, Roraima), Bolivia. ◆Fig. 166. This species is cultivated by the Amerindians for its starchy rhizome tubers. The present distribution was probably much influenced by cultivation.

Fig. 166. Myrosma cannifolia

246

M ARANTACEAE

8. STROMANTHE Sonder, Neue Allg. Deutsche Garten-Blumenzeitung 5: 225. 1849. by Lennart Andersson Caulescent, tree-like or shrubby herbs. Leaves antitropic, on major branches clustered at several-leaved nodes. Inflorescences clustered into distinct synflorescences or solitary at tips of leafy branches; inflorescence spike- or head-like to diffuse in outline, often ± distinctly zygomorphic. Bracts (spathes) herbaceous to coriaceous, often brightly colored, persistent or caducous; florescence component of 1–5 cymules, axes distinct, interphyll absent; bracteoles absent or rudimentary. Sepals ovate to elliptic, obtuse, ± concealing corolla and inner staminodes; corolla tube shorter than wide. Outer staminodes 2, petaloid and showy over rudimentary to absent, callose staminode firm and fleshy throughout, fertile stamen with a small, oblong to narrowly obovate, petaloid appendage. Ovary 1-ovulate. Fruit capsular, wall ± parenchymatic. Seed arillate; perisperm canal branched distally. Central America, West Indies, South America; ca. 15 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Stromanthe 1.

1.

Herbs with leaves clustered distally on an unbranched, cane-like stem; bracts (spathes) orange; outer staminodes absent or rudimentary; fruit greenish, with persistent calyx ................................................ S. jacquinii Herbs shrubby with long, straggling, leafy branches; bracts (spathes) green or purplish-tinged; outer staminodes petaloid and showy (violet to purplish); fruit orange to dark red, without calyx when mature ...................................................................................................... S. tonckat

Fig. 167. Stromanthe tonckat

Thalia 247

Stromanthe jacquinii (Roem. & Schult.) H. Kenn. & Nicolson, Ann. Missouri Bot. Gard. 62: 501. 1975. —Maranta jacquinii Roem. & Schult., Syst. Veg. 1: 558. 1817. Maranta lutea Jacq., Icon. Pl. Rar. 2(1): pl. 201. 1781 [1789], non Aubl. 1775. —Stromanthe lutea Eichler, Abh. Königl. Akad. Wiss. Berlin 1883: 81. 1884. Caulescent herb 1–2 m tall. Clearings and light-gaps in seasonal, evergreen forests, ca. 900 m; Bolívar (Matacuchillo west of Santa Elena). Widespread elsewhere in Venezuela; Central America, northern and

western Colombia, western Ecuador. Stromanthe tonckat (Aubl.) Eichler, Abh. Königl. Akad. Wiss. Berlin 1883: 80. 1884. —Maranta tonckat Aubl., Hist. Pl. Guiane 3. 1775. Caulescent, shrubby herb ca. 1–3 m tall. Light-gaps in rain forests, common in secondary growth, ca. 300–400 m; Bolívar (near El Palmar). Mérida, Miranda, Portuguesa, Sucre, Táchira, Yaracuy; Central America, West Indies except Greater Antilles, Colombia, Guyana, Suriname, French Guiana, eastern Brazil. ◆Fig. 167.

Fig. 168. Thalia geniculata

248

M ARANTACEAE

9. THALIA L., Sp. Pl. 1193. 1753. by Lennart Andersson Rosulate aquatic herbs usually with a tall and branched synflorescence. Inflorescences arranged in synflorescences, small and spike-like in active portion; bracts (spathes) shed with the mature fruit and leaving a long, scarred rachis; florescence component with a single cymule; cymule with very short axes, interphylls and bracteoles absent. Sepals very small, membranous, ± triangular; corolla tube obsolete. Outer staminode solitary, petaloid and showy, callose staminode largely firm and fleshy, with narrow petaloid rim, hooded staminode with double appendage, fertile stamen with an irregularly obovate, petaloid appendage. Ovary 1-ovulate; style with a very long ventral process from rim of stigmatic orifice, helically twisted when triggered. Fruit indehiscent; pericarp very thin. Seed arillate (but aril often very small); perisperm canal branched from base. Southern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, Ecuador, Brazil, Bolivia, Argentina, Paraguay, tropical Africa; ca. 5 species, 1 in Venezuela. Thalia geniculata L., Sp. Pl. 1193. 1753. Thalia trichocalyx Gagnep., Bull. Soc. Bot. France 51: 180. 1904. Emergent aquatic herb 1–3 m tall, with ample panicle-like synflorescence. Open, seasonal pools and swamps, ca. 50 m; Delta Amacuro (Caño Mánamo), Bolívar (between

Upata and La Grulla). Apure, Aragua, Carabobo, Falcón, Guárico, Portuguesa, Sucre; U.S.A. (Florida, southern Louisiana), Mexico, Central America, West Indies, Colombia, Trinidad, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, Argentina, Paraguay, Africa. ◆Fig. 168.

MARCGRAVIACEAE by Stefan Dressler Terrestrial, hemiepiphytic, or epiphytic lianas or shrubs, rarely small trees; hypophyllous glands (glands on the lower surface of the leaves), raphide cells and variously shaped sclereids frequent. Leaves alternate (Marcgravia) or spirally arranged (all other genera), simple, estipulate, glabrous; margins entire or minutely crenate. Inflorescences terminal, racemose, sometimes in pseudo-umbels or pseudospikes, erect or pendulous. Bracts transformed into variously shaped nectaries. Flowers bisexual, actinomorphic, hypogynous, 2-bracteolate. Bracteoles persistent, sepaloid, sometimes lacking. Sepals 4 or 5, unequal, free or nearly so, imbricate, persistent. Petals 3–5, imbricate, free or connate. Stamens 3–many; filaments free or basally connate, uniseriate; anthers basifixed or nearly so, dithecal, tetrasporangiate, introrse, longitudinally dehiscent. Ovary superior, completely or incompletely 2–20-locular; ovules numerous. Fruit a subglobose, apiculate capsule with persistent style and stigma, loculicidally and septifragously dehiscent from the base, or berry-like and irregularly dehiscent, pulpy inside. Seeds hemispheric to reniform, anatropous, few to numerous with a shiny reticulate testa. Endosperm scanty or lacking; embryo straight. Tropics and subtropics of Central America, West Indies, and South America; 7 genera and ca. 130 species, 5 genera and 16 species in the flora area.

Marcgravia 249

Key to the Genera of Marcgraviaceae 1.

1.

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

Leaves distichous; apical flowers abortive, only the nectaries developed, fertile flowers without nectaries; sepals 4; petals united into a deciduous cap; sterile and potentially fertile branches with differently shaped leaves ............................................................................................... 1. Marcgravia Leaves spirally arranged; all flowers fertile and provided with nectaries; sepals 5; petals 5, ± free or connate, reflexed at anthesis; sterile and potentially fertile branches with similarly shaped leaves ................... 2 Inflorescence umbellate or subumbellate ...................... 2. Marcgraviastrum Inflorescence spicate or racemose ............................................................. 3 Inflorescence spicate; nectaries inserted on the rachis next to the flowers .................................................................................................. 4. Sarcopera Inflorescence racemose; nectaries variously inserted on the pedicel ....... 4 Nectaries inserted at the base of the calyx, their limb auriculate; stamens < 10, generally 3 or 5; inflorescence mostly < 25 cm long ..... 5. Souroubea Nectaries inserted at various distances from the calyx, but never at its base, their limb never auriculate; stamens numerous (> 10); inflorescence mostly > 25 cm long ........................................................ 3. Norantea

1. MARCGRAVIA L., Sp. Pl. 503. 1753. Climbing shrubs or woody vines, with dimorphic branches: sterile, juvenile branches creeping, appressed to the substrate, and attached by roots; fertile ones free, mostly pendulous, often angular and provided with wart-like lenticels. Leaves distichous, those of juvenile branches small, sessile, thin, ovate, mostly asymmetrically cordate at the base, those of mature branches larger, subsessile to petiolate, thicker, often with a drip tip, the lower surface of the leaves with prominent midrib and variously shaped and numbered hypophyllous glands. Inflorescences umbellate or subumbellate, flowers pedicellate, the apical (central) ones abortive with only the bracts (nectaries) well developed. Nectaries hood-, pitcher-, or dipper-shaped, their stalk often lenticellate outside. Bracteoles on the upper half of the pedicel, sometimes lacking. Sepals 4, decussate; petals 4, connate into a deciduous cap. Stamens 6–many; filaments mostly filiform, flattened; anthers slender-ovate to triangular. Ovary 3–20-locular; ovules numerous per locule; stigma capitate to umbonate. Fruit a leathery capsule; calyx persistent; seeds tiny, embedded in a juicy pulp; testa reticulate, purple and black. Southern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 60 species, 13 in Venezuela, 9 of these in the flora area. Key to the Species of Marcgravia 1. 1. 2(1). 2. 3(2).

Flowers erect on pedicels; lower surface of leaves with a submarginal row of many poriform glands ..................................................... M. patellulifera Flowers angled on pedicels; lower surface of leaves without submarginal poriform glands ...................................................................................... 2 Lower surface of leaves punctate, ± densely covered with numerous black dots ......................................................................................................... 3 Lower surface of leaves epunctate, lacking numerous black dots ........... 5 Rachis of inflorescence elongated, with a section (interstice) between the

250

3. 4(3).

4.

5(2).

5. 6(5).

6.

7(6). 7.

8(7).

8.

M ARCGRAVIACEAE

fertile flowers and the nectaries that bears no pedicels; nectaries with conspicuously protruding orifices ......................................... M. eichleriana Rachis contracted to form a clavate or globose structure, not elongated; nectaries without conspicuously protruding orifices ............................ 4 Leaves < 12 cm long, coriaceous when dry, the base acute to obtuse, the blade slightly obovate-elliptic; inflorescence terminal on normal-leaved branches ................................................................................... M. maguirei Leaves 15–25 cm long, chartaceous when dry, the base round to cordate, the blade slightly ovate-elliptic; inflorescence “cauliflorous” on leafless side shoot ............................................................................... M. punctifolia Rachis of inflorescence elongated, with a section (interstice) between the fertile flowers and the nectaries that bears no pedicels; corolla cap large, usually > 10 mm long ...................................................... M. coriacea Rachis clavate or globose, not elongated; corolla cap smaller, usually < 10 mm long .......................................................................................... 6 Leaves thick-coriaceous; venation mostly not prominent when dry; petiole mostly < 5 mm long; leaf blades with 1–3 conspicuous glands on lower surface, the margins often with conspicuous dark marginal glands ......................................................................................... M. sororopaniana Leaves chartaceous to subcoriaceous; venation mostly prominent when dry; petiole mostly > 5 mm long; leaf blades with inconspicuous glands or glands lacking .................................................................................... 7 Nectaries longer than the pedicels; lower surface of leaf blades without glands; leaves often drying reddish brown; Bolívar state ...... M. purpurea Nectaries shorter than or as long as the pedicels; lower surface of leaf blades with 1 or 2 glands (sometimes lacking); leaves not drying conspicuously reddish brown, rather greenish brown or conspicuously bicolored; Amazonas state ........................................................................ 8 Nectaries > 1 cm long; leaf apex cuspidate; leaf base often obtuse to round; leaves drying brownish, both surfaces conspicuously different in color; inflorescence mostly on normal-leaved branches, peduncle usually short, 8–10(–16) cm long ............................................................. M. sprucei Nectaries < 1 cm long; leaf apex acuminate; leaf base acute to obtuse; leaves drying greenish, both surfaces similar in color; inflorescence often long-pedunculate, sometimes “cauliflorous” ............ M. pedunculosa

Marcgravia coriacea Vahl, Eclog. Amer. 2: 39. 1798. —Bejuco pantalla, Ero, Matekesashe, Tëpo po mën (Panare). Marcgravia roraimae Gilg & Werderm. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 105, fig. 57. 1925. Liana; leaves 6–15.5 × 2–5 cm, oblong to elliptic, the apex acuminate, the base cuneate, shortly attenuate, acute, or obtuse, the lower surface with or without 1–3(4) small, poriform glands per side; pedicels 2– 3.5 cm long; interstice 3–7 mm long; corolla caps 9–17 mm long; nectaries 25–60 mm long, bent outward, the stalk (5–)7–10 mm

long. Riparian, gallery, swamp, and cloud forests, near sea level to 1800 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Barinas, Carabobo, Falcón, Miranda, Sucre; Colombia (Amazonas?), Guyana, Suriname, French Guiana, Brazil (Amapá, Maranhão, Pará, Pernambuco). ◆Fig. 169. Marcgravia coriacea is reported to be used to treat rheumatism and amoebiasis. The type of Marcgravia roraimae has 1–4 poriform glands on the lower surface of the blade. This is often found in collections from the west of the distributional range (Venezuelan Guayana and adjacent areas). Addition-

Marcgravia 251

ally, the number of stamens seems to be fewer (< 20). Recognition as a subspecies was considered but abandoned since it was not found to be consistent within this geographical area. Moreover, even within one collection both patterns of glands sometimes occur (e.g., the collection at VEN of Tamayo 2960 has these glands, the duplicates at F and US lack them). Marcgravia eichleriana Wittm. in Mart., Fl. Bras. 12(1): 230. 1878. Liana; leaves 7–13.5 × 3–4 cm, elliptic, the apex acuminate, the base acute, the lower surface with a row of 1–3 small poriform glands; pedicels 2–2.5 cm long; interstice 2–3 mm long; corolla caps 6–8 mm long; nectaries 28–33 mm long, often bent outward, the stalk 10–12 mm long. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro). Colombia (Vaupés), Brazil (Amazonas). Marcgravia maguirei de Roon, Acta Bot. Neerl. 19: 799. 1970. Liana, very variable in leaf and nectary shape; leaves 5–12 × 3.5–5.8 cm, obovate to elliptic, the apex acuminate, acute to obtuse, rarely retuse, the base acute, cuneate, or obtuse, the lower surface of the leaf with 1–3 conspicuous poriform glands per side; pedicels 2–3(–3.5 in fruit) cm long; corolla caps 7–8 mm long; nectaries 12–32 mm long, the stalk 5–7 mm long. Montane rain and cloud forests, 1200–1600 m; Bolívar (Cerro Jaua, Cerro Sarisariñama, southwestern Cerro Venamo, kilómetro 88–126 south of El Dorado, Macizo del Chimantá [Abácapa-tepui]), Amazonas (Cerro Huachamacari, Cerro Sipapo, Cerro Yutajé, Río Iguapo). French Guiana, Brazil (Amazonas). Some collections have the glandular dots on the lower leaf surfaces only faintly or irregularly dispersed. Those are intermediate to Marcgravia sororopaniana. Both species are similar and only discernible by the punctate versus epunctate leaves. Marcgravia patellulifera de Roon, Acta Bot. Neerl. 19: 802. 1970. Liana; leaves 6–12 × 2.2–3.8 cm, narrowelliptic to elliptic, sometimes slightly obovate, coriaceous; pedicels 4–5(–6 in fruit) cm long; corolla caps (narrow) conical, 8–11 mm

long; nectaries 2–3 cm long, the stalk 4–6 mm long. Evergreen lowland forests, 50–200 m; Amazonas (Río Mawarinuma). Aragua, Carabobo. Marcgravia pedunculosa Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 371. 1862. —Bejuco de gallineta. Marcgravia parviflora Rich. ex Wittm. in Mart., Fl. Bras. 12(1): 227. 1878. Marcgravia parviflora var. macrophylla Wittm. in Mart., Fl. Bras. 12(1): 228. 1878. Liana; leaves 7–20 × 3–7 cm, narrow-elliptic, the apex acuminate, the base acute to obtuse, the lower surface of the leaves with 1 or 2 inconspicuous glands per side; pedicels 1–2.5 cm long; corolla caps 4–7 mm long; nectaries 5–10 mm long, the stalk 2–5 mm long. Evergreen lowland and riparian forests, 50– 200 m; Amazonas (Puerto Ayacucho, Río Mawarinuma, San Carlos de Río Negro, Yavita). Guyana, Suriname, French Guiana, Ecuador (Napo), Brazil (Amazonas). Marcgravia punctifolia S. Dressler, Novon 8: 137. 1998. Liana; leaves relatively large, 15–25 × 4– 11 cm, elliptic, the apex acuminate, the base round to cordate, the lower surface with numerous dark glandular dots; inflorescences “cauliflorous,” on rather long, leafless lateral shoots (peduncle); pedicels 2–3(–4 in fruit) cm long; nectary ca. 15 mm long, the stalk ca. 8 mm long. Forests along lower tepui slopes, 500–1000 m; Bolívar (Amaruay-tepui, Cerro Marutaní). Brazil (Amazonas: Río Uaupés; Roraima: Serra dos Surucucus). Marcgravia purpurea I.W. Bailey, Amer. J. Bot. 9: 378. 1922. Liana; leaves 10–18 × 4.5–8 cm, elliptic, the apex acuminate, the base obtuse, the lower surface without glands; venation conspicuously embossed when dry; pedicels 2– 3(–3.5 in fruit) cm long; corolla caps 7–9 mm long; nectaries 15–32 mm long, the stalk ca. 5 mm long. Evergreen lowland forests, 50–300 m; Bolívar (Río Caura), Amazonas (Río Mawarinuma). Guyana, Suriname, French Guiana, Brazil (Amazonas, Mato Grosso, Pará). Marcgravia sororopaniana Steyerm., Fieldiana, Bot. 28: 378. 1952.

252

M ARCGRAVIACEAE

mature inflorescence

young inflorescence

Fig. 169. Marcgravia coriacea

Marcgravia steyermarkii de Roon, Bol. Soc. Venez. Ci. Nat. 32: 355. 1976. Liana, variable in leaf and nectary shape; leaves 4–10 × 1.8–4.8 cm, obovate to elliptic, the apex acuminate, acute, obtuse, or retuse, the base acute, cuneate, or obtuse, the lower surface of the leaf with 1–3 conspicuous poriform glands per side; pedicels 2–2.7(–3 in fruit) cm long; corolla caps 5–7 mm long; nectaries 15–35 mm long, the stalk 4–9 mm long. Montane rain and cloud forests, (200–)700–1600(–2300) m; Bolívar (Cerro Sarisariñama, Cerro Uroi in Río Chicanán basin, Cerro Venamo, road between El Dorado and Santa Elena de Uiarén, Ilú-tepui, Río Cuyuní, Río Yudi, Sororopán-tepui). Monagas(?); Guyana, French Guiana(?), Brazil.

The type of Marcgravia steyermarkii is merely a smaller-leaved specimen of M. sororopaniana. Marcgravia sprucei (Wittm.) Gilg, Bot. Jahrb. Syst. 25(Beibl. 60): 28. 1898. —Marcgravia parviflora var. sprucei Wittm. in Mart., Fl. Bras. 12(1): 228. 1878. Liana; leaves 8–16 × 3–6.5 cm, elliptic to slightly ovate, the apex cuspidate to acuminate, the base obtuse to acute, the lower surface of the leaf without or with only 1 inconspicuous poriform gland per side, the upper leaf surface often light grayish green when dry; pedicels 15–25 mm long; corolla caps 5–7 mm long; nectaries 1.3–2 cm long, the stalk 3–5 mm long. Evergreen lowland forests, 50–

Marcgraviastrum 253

300 m; Amazonas (upper Río Baría, Río Casiquiare, Río Cuao, Río Mavaca, Río Mawarinuma, Río Yaciba above the mouth of

Río Casiquiare, Río Yatua). (Amazonas, Pará), Colombia, Peru.

Brazil

2. MARCGRAVIASTRUM (Wittm. ex Szyszyl.) de Roon & S. Dressler, Bot. Jahrb. Syst. 119: 332. 1997. —Norantea subsect. Marcgraviastrum Wittm. ex Szyszyl. in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 163. 1893. Sprawling terrestrial or (hemi-)epiphytic shrubs or lianas. Leaves spirally arranged, sessile or petiolate, coriaceous, producing a nonciliate or long-ciliate fracture when broken perpendicular to the midvein, with 2 basal glands, and either (1)2–6 or 12–18(–22) glands per side in 1(2) fairly uniform row(s), or numerous glands (25–160) scattered in a zone inside the loops of the secondary veins, the leaf

Fig. 170. Marcgraviastrum mixtum

254

M ARCGRAVIACEAE

apex mucronate or retuse to emarginate (due to loss of mucro); margins entire, sometimes revolute. Inflorescences umbelliformly contracted racemes with (2–)5– 14(–22) flowers. Flowers erect on straight, slender, or stout pedicels, subtended by sessile, rarely stalked, pendulous, saccate, tubular, or pouch-shaped nectariferous bracts attached to the lower 1/4–1/3 of the pedicel. Bracteoles sepaloid, appressed to the calyx, rarely 2–3 mm distant from the calyx. Sepals 5, quincuncial, coriaceous. Petals 5, free to variously connate, imbricate in bud, strongly reflexed at anthesis. Stamens 12–75; filaments mostly free with the outer whorl basally adnate to corolla, linear, flattened or somewhat triangular; anthers (sub)sagittate. Ovary 5–9-locular; ovules numerous per locule; stigma smooth or slightly capitate. Fruit a leathery capsule, globose. Seeds reniform to hemispheric; testa reticulate, red-black or shiny black. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 15 species, 2 species in Venezuela, both in the flora area. Key to the Species of Marcgraviastrum 1. 1.

Leaf apex mostly acute; stamens 40–55; southwestern Amazonas state .................................................................................................... M. mixtum Leaf apex obtuse; stamens 60–65; central eastern Bolívar state ................................................................................................ M. pendulum

Marcgraviastrum mixtum (Triana & Planch.) Bedell in de Roon & S. Dressler, Bot. Jahrb. Syst. 119: 332. 1997. —Norantea mixta Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 374. 1862. Norantea peduncularis Poepp. ex Wittm. in Mart., Fl. Bras. 12(1): 238. 1878. Sprawling shrub or liana; leaves elliptic to obovate, with 2–6(–35) small to mediumsized hypophyllous glands in a row or zone 3–5 mm from the margin, the leaf apex acute to bluntly acuminate, mucronate or emarginate through loss of mucro; inflorescence with mostly 13–22 flowers. Evergreen lowland to montane forests, ca. 200 m; Amazonas (Río Negro). Aragua, Mérida, Miranda, Táchira; Colombia, Ecuador, Peru, Brazil (Amazo-

nas), Bolivia. ◆Fig. 170. Marcgraviastrum pendulum (Lanj. & Heerdt) Bedell, Novon 9: 252. 1999. —Norantea pendula Lanj. & Heerdt, Recueil Trav. Bot. Néerl. 37: 282. 1940. Sprawling shrub or liana; leaves broadly oblong, often with 4–8 small to medium-sized hypophyllous glands per side, 5–9 mm from the margin, sometimes hardly visible, the apex obtuse to rounded, or retuse through loss of mucro; inflorescence with mostly 10–14 flowers. Lowland to montane forests, 100–800 m; Bolívar (Río Uiri-yuk on upper Río Cuyuní, Venezuela-Guyana border), Amazonas (Sierra de la Neblina). Mérida, Miranda; Guyana, Suriname, French Guiana, Brazil (Amapá).

3. NORANTEA Aubl., Hist. Pl. Guiane 554. 1775. (Hemi-)epiphytic or terrestrial woody lianas or sprawling shrubs. Leaves spirally arranged, the blade coriaceous, with 2 basal glands, petiolate, producing a nonciliate fracture when broken perpendicular to midvein, the margins entire, slightly revolute, the apex mucronate. Inflorescences dense multiflorous racemes with 120–350 flowers. Flowers pedicellate, subtended by a purple or bright orangered, tubular-saccate, pendulous, nectariferous bract on a slender petiole adnate to the upper half of the pedicel. Bracteoles located below the calyx. Sepals 5, quincuncial, coriaceous. Petals 5, free to slightly basally connate, imbricate in bud, reflexed at anthesis. Stamens 20–35; filaments basally adnate to corolla, linear in

Norantea 255

outline, triangular in cross section; anthers linear to subsagittate. Ovary 5-locular; ovules 10–20 per locule; stigma smooth or slightly 5-lobed. Fruit a berry-like, leathery capsule, globose. Seeds small; testa reticulate, black. Costa Rica, Lesser Antilles, northern South America, southern Brazil; 2 species, 1 in Venezuela. Norantea guianensis Aubl., Hist. Pl. Guiane 554, t. 220. 1775. (Hemi-)epiphytic liana or shrub; leaves obovate-elliptic to elliptic, the base cuneate, the apex rounded to acute; inflorescence 25– 80 cm long. Costa Rica, Lesser Antilles, northern South America; 2 subspecies, both in the flora area.

Fig. 171. Norantea guianensis subsp. japurensis

256

M ARCGRAVIACEAE

Key to the Subspecies of N. guianensis 1. Nectaries with an apiculate lip, orifice wide, protruding; petiole mostly short; nectaries often basally bifid, rarely roughish when dry; Caribbean drainage ............................... subsp. guianensis 1. Nectaries without apiculate lip, orifice elongate or ovate; petiole mostly long; nectaries often papillate-roughish when dry; Amazonian drainage ........................ ................................ subsp. japurensis N. guianensis subsp. guianensis. —Bejuco rabo de guacamayo, Guacamaya, Peine de morocoto, Rabo de guaca, Usibo akuanetete. Moist humid forests, annually burned gallery forests and savannas, near sea level to 600 m; widespread in Delta Amacuro, northern Bolívar, and Amazonas. Apure, Monagas, Nueva Esparta, Sucre; Costa Rica, Jamaica, Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Ecuador, Brazil (Amazonas). The collections from Costa Rica and Jamaica most probably originate from cultivated or escaped ornamental plants. An infusion of the bark is used to control

hemorrhaging in women during menstruation (Clark 6988, NY; Ferrigni et al. 5E, NY) and is also used as a vermifuge (Cardona 2796, VEN). N. guianensis subsp. japurensis (Mart.) Bedell, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1256. 1993. —Norantea japurensis Mart., Nov. Gen. Sp. Pl. 3: 179. 1829 [1832]. —Cola de guaca, Guacamaya, Rabo de arara. Norantea paraensis Mart., Nov. Gen. Sp. Pl. 3: 180. 1829 [1832]. Norantea guianensis var. gracilis Wittm. in Mart., Fl. Bras. 12(1): 242. 1878. Moist humid forests, Mauritia palm swamps, 50–500 m; Bolívar (30 km below La Urbana), Amazonas (widespread). Monagas, Sucre; Colombia (Meta, Vaupés), Peru (Huanuco, Loreto), Brazil (Amapá, Amazonas, Rondônia, Roraima, Maranhão, Mato Grosso, Pará). ◆Fig. 171. In Amazonas state, collections from the headwaters of Río Orinoco often show the typical nectary of subsp. japurensis. Some collections from the upper Río Negro and estuaries have the nectary of subsp. guianensis. The geographical separation between the subspecies is not clearcut in those areas.

4. SARCOPERA Bedell, Bot. Jahrb. Syst. 119: 328. 1997. Sprawling hemiepiphytic or terrestrial shrubs or lianas, rarely small trees. Leaves spirally arranged, sessile to petiolate; blade occasionally asymmetrical, chartaceous to coriaceous, with 2 basal glands and either (1)2–7 glands per side in a row, or 15–28 glands per side in 2 fairly uniform rows, these rarely lacking, the leaf apex mucronate or retuse (due to loss of mucro); margins entire, sometimes revolute. Inflorescences spicate; rachis (6–)15–45(–100) cm long with (35–)100–250 (–450) flowers. Flowers sessile or rarely shortly pedicellate, at least the flowers of the upper 1/2–3/4 of the rachis subtended by a petiolate, cyathiform, ladle- or pouchshaped, nectariferous bract, these in various shades of brilliant red. Bracteoles deltoid and 1–3 mm distant from the calyx, or sepaloid and appressed to the calyx. Sepals 5, quincuncial, coriaceous. Petals 5, free or slightly basally connate, imbricate in bud, strongly reflexed at anthesis. Stamens (6–)8–12(–25); filaments free or variously connate or adnate to corolla, broad, flattened; anthers subsagittate to cordiform. Ovary completely 2- or 3(–5)-locular; ovules 4–8(–12) per locule; style lacking; stigma smooth, sessile. Fruit a berry-like capsule, globose. Seeds small; testa reticulate, red-black or black, shiny, embedded in brightly colored pulpy mesocarp. Honduras through the Andean Cordillera to northern Bolivia; ca. 10 species, 2 in Venezuela, both in the flora area.

Sarcopera 257

Key to the Species of Sarcopera 1.

1.

Stamens 8–12; leaves mostly < 9 cm long; blade mostly with 1(2) conspicuous hypophyllous glands per side, these of a different color than blade ................................................................................................ S. flammifera Stamens 17–25; leaves usually > 9 cm long; blade with glands inconspicuous or of same color as blade ................................................... S. tepuiensis

Sarcopera flammifera de Roon & Bedell, Novon 9: 249. 1999. Small liana or sprawling shrub; leaves 5– 9(–12) × 1.5–4.5 cm, obovate to elliptic, the apex obtuse; inflorescence 24–40 cm long; stamens 8–12; filaments free; pollen yellow. (Pre)montane forests, gallery forests, open savanna vegetation, often among rocks, (100–)300–1800 m; Bolívar (widespread), Amazonas (Cerro Yutajé, Sierra Parima). Brazil (Amazonas: Sierra Aracá; Roraima: Sierra Tepequem). There are collections that show a leaf typical for Sarcopera tepuiensis but have fewer stamens, which is typical of S. flammifera (Maguire & C. Maguire 35109, NY; K.D. Phelps & Hitchcock 3, NY, VEN; Duno de Stefano & al. 467, VEN). However, the number of stamens is the only reliable distinction between these two species, and so these specimens are here considered to be S. flammifera. Sarcopera tepuiensis (de Roon) Bedell, Novon 9: 251. 1999. —Norantea tepuiensis de Roon, Acta Bot. Venez. 2: 247. 1967. Large liana or scrambling shrub; stamens 17–25; pollen magenta. Venezuela, Guyana, Brazil (Amazonas); 2 subspecies, both in the flora area. Key to the Subspecies of S. tepuiensis 1. Leaves oblanceolate to narrow-obovate, the blade with 5–9 small hypophyllous glands per side in a row ca. 2–5 mm from margin, the leaf base cuneate ................. ................................... subsp. coccinea 1. Leaves obovate, the blade with 1(2) large hypophyllous glands per side, ca. 1 cm from margin, occasionally 2–6 additional small ones, the leaf base acute ................................. subsp. tepuiensis

Fig. 172. Sarcopera tepuiensis subsp. tepuiensis

258

M ARCGRAVIACEAE

S. tepuiensis subsp. coccinea de Roon & S. Dressler, Novon 9: 251. 1999. Leaves 8–22 × 3.5–7.5 cm, mostly longnarrow-obovate or oblanceolate, the apex obtuse to rounded; filaments usually connate 1/4–3/4 of their length. Montane rain and cloud forests, 1000–1400 m; Amazonas (Cerro Aratitiyope, Cerro Duida, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yapacana, Cerro Yaví, Sierra de la Neblina). Brazil (Amazonas). S. tepuiensis subsp. tepuiensis Leaves 7.5–13(–14.5) × 3–6(–7) cm, obovate, rarely elliptic, the apex obtuse to rounded, very rarely acute (immature leaves?); filaments usually connate at base only. Montane rain and cloud forests, (400–)

700–1800 m; southeastern Bolívar (Amaruay-tepui, Auyán-tepui, near Canaima, Carrao-tepui near Ptari-tepui, Cerro Bolívar, Cerro Sarisariñama, Gran Sabana, Ilú-tepui, between Luepa and Cerro Venamo, Macizo del Chimantá [Abacapá-tepui, Toronó-tepui], Ptarí-tepui, Río Cusimi, Uaipán-tepui). Guyana (Merume Mountains, Pakaraima Mountains). ◆Fig. 172. Collections with the typical subsp. tepuiensis leaf were made also in Amazonas state (Río Coro Coro: Holst & Liesner 3127, 3384, MO; Cerro Yaví: K.D. Phelps 114, VEN). Thus the two subspecies are not completely geographically separated. The bark of the plant is boiled and used by Amerindians for healing wounds.

5. SOUROUBEA Aubl., Hist. Pl. Guiane 244. 1775. Climbing shrubs or lianas, often (hemi-)epiphytic. Leaves shortly petiolate, with hypophyllous glands of various densities, sizes, and arrangements, often in rows parallel to the margin. Inflorescences racemose. Nectaries borne on the top of the pedicel, sessile or sometimes shortly stipitate, hollow, mostly spur-like and auriculate, or tubular. Flowers 5-merous (rarely 3-, 4-, or 6-merous), often fragrant. Bracteoles sepaloid, appressed to calyx. Sepals 5, quincuncial, coriaceous with membranaceous margin. Petals free or connate to 2/3 of their length, the lobes strongly reflexed at anthesis. Stamens 3 or 5, rarely 4 or 6–8; filaments often adnate to corolla; anthers mostly ovoid. Ovary (3–)5-locular; ovules ca. 20 per locule; stigma (sub)sessile with (3–)5 radiating lobes. Fruit (depressed-)globose, with leathery pericarp. Seeds few. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 19 species, 4 in Venezuela, 2 of these in the flora area. Key to the Species of Souroubea 1.

1.

Pedicels of lowermost flowers of the inflorescence < 15 mm long; corolla < 6 mm long; auricles of the nectaries < 1 cm long; leaves papery when dry ........................................................................................ S. dasystachya Pedicels of lowermost flowers of the inflorescence > 20 mm long; corolla > 6 mm long; auricles of the nectaries > 1 cm long; leaves coriaceous when dry ................................................................................ S. guianensis

Souroubea dasystachya Gilg & Werderm. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 102. 1925. —Jidiuey, Jödöwoi (Yekwana), Pir-boi-yek, Pirboyek (last two names Arekuna). Liana, often epiphytic; leaves obovate-oblong to obovate, papery when dry, the apex

obtuse to rounded; inflorescence densely many-flowered, 8–30 cm long; pedicels 5–12 mm long; nectaries shortly auriculate, greenish or pale yellow, cup 3-10 mm long, auricles 3–6 mm long; corolla 5–6 mm long, reddish. Forests or thickets, often on riversides, 50– 500 m; Bolívar (middle Río Caura, 30 km

Souroubea 259

west of Río Caura, tributary of Río Erebato, south of Chaco-tepui), Amazonas (upper Río Orinoco, Río Ugueto). Colombia (Amazonas), Ecuador (Napo, Sucumbios), Peru (San Martin), Brazil (Roraima). ◆Fig. 174.

Souroubea guianensis Aubl., Hist. Pl. Guiane 244, t. 97. 1775. Shrub or large liana; leaves variable in shape and size, coriaceous; inflorescence laxflowered, mostly 15–30 flowers, 10–20(–25)

Fig. 173. Souroubea guianensis subsp. cylindrica

Fig. 174. Souroubea dasystachya

260

M ARCGRAVIACEAE

cm long; pedicels 1.5–3(–4) cm long; nectaries red, 2–3 cm long, spur as long as or slightly shorter than auricles; petals 8–10 mm long, yellow to orange. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 3 subspecies, 1 in Venezuela. The typical subspecies is known from western Guyana, Suriname, French Guiana, and northern Brazil (Amapá). Because of the occurrence of some intermediate collections (e.g., Liesner 4074, MO), with a rather lepidote peduncle, it is possible that subsp. guianensis may also occur in the flora area in Amazonas state. The main macromorphological differences between the two subspecies are: subsp. cylindrica has mostly large, conspicuous hypophyllous glands, the peduncle (when dry) is smooth, dark reddish brown to blackish, the rachis is tomentellous (neither lepidote nor fissured); subsp. guianensis has small and inconspicuous to large hypophyllous glands, the peduncle is mostly lepidote (rarely smooth) and mostly grayish, the rachis is tomentellous to glabrous and mostly with fissured periderm. The third subspecies, subsp. amazonica (Mart.) de Roon, occurs in Brazil in the Ama-

zon Basin (Acre, Amazonas, Pará, Rondônia), the coastal area in Bahia, and in Pernambuco. S. guianensis subsp. cylindrica (Wittm.) de Roon, Contrib. Monogr. Marcgraviaceae 183. 1975. —Souroubea guianensis var. cylindrica Wittm. in Mart., Fl. Bras. 12(1): 253. 1878. —Bejuco pene de diablo, Cabaya-cuaja, Jidiway, Jidiwoi (Yekwana), Moy sebe (Warao), Pirboiyek, Piruói (Arekuna). Souroubea guianensis var. tomentella Steyerm., Fieldiana, Bot. 28: 378. 1952. Edges of flooded forests, riparian forests, Mauritia palm swamps, forested slopes, savannas, granitic outcrops, igneous outcrops, 50–1400 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Colombia (Amazonas), western Guyana, Brazil (northwestern Amazonas, Roraima). The exudate of the bark or the bark boiled in water of Souroubea guianensis subsp. cylindrica is used to treat cuts and wounds. Also, the strong and flexible stems are occasionally used for backpack frames. ◆ Fig. 173.

MAYACACEAE by Alicia Lourteig Moss-like herbs, aquatic, submersed in shallow water or amphibious, or terrestrial at edges of aquatic habitats. Roots long, septate, fibrous. Stems very short, erect, simple or branched, or else procumbent to very long, rooting, ± submerged, the apex and inflorescences emerging from the water. Leaves alternate, spirally arranged, sessile, linear, slightly wider at the base, apex acute or more frequently bifid, the divisions sometimes prolonged into a bristle. Flowers bisexual, actinomorphic, solitary or in few-flowered, terminal or axillary, umbelliform cymes; bracts membranous at the base of the peduncle. Sepals 3, triangular-linear, stiff, acute, light green; petals 3, delicate, largely obovate, spreading, imbricate, white, pink, or violaceous. Stamens 3, free, alternate with the petals; anthers basifixed, (2)4-locular, asymmetric, introrsely dehiscent by a short apical slit, by a pore at the end of a short to long tube, or within a cup formed by an irregular projected border (M. baumii). Ovary superior, ovoid, 3-carpellate, 1-locular with parietal placentation; ovules 6–30; style slender, terminal; stigma small, 3-lobed, papillose. Fruit capsular, thin-walled, enclosed in

Mayaca 261

the persistent calyx, dehiscent by 3 valves. Seeds small, roundish with a beak at the apex, dark, spongy, finely grainy and longitudinally striate, endosperm copious, starchy. Tropical and temperate America, from southern U.S.A. to Argentina and Uruguay, tropical western Africa; 1 genus and 4 species, 3 species in Venezuela, all in the flora area. When the fruit is mature, the accrescent pedicel deflexes abruptly towards the ground. The capsule then dehisces, ejecting the seeds and sowing them in the water or moist soil. This is particulary evident in Mayaca longipes. 1. MAYACA Aubl., Hist. Pl. Guiane 42, t. 15. 1775. Biaslia Vand., Fl. Lusit. Brasil. Spec. 4, fig. 2. 1788. Syena Schreb., Gen. Pl. 39. 1789. Coletia Vell., Fl. Flumin. t. 79. 1825 [1829]. Description, distribution, and species numbers as in family. The species of Mayaca are morphologically variable depending on fluctuating water levels. Key to the Species of Mayaca 1. 1. 2(1).

2.

Anthers dehiscent by an apical slit; filaments equal or longer than the anthers .................................................................................... M. fluviatilis Anthers dehiscent by a pore at the apex of a tubular extension of the anther; filaments shorter than the anthers .............................................. 2 Flowers white, in umbelliform cymes (rarely 1-flowered); anthers oblong or obovoid-oblong, tube short; leaves 3–30 mm long, often filiform; peduncles reflexed into soil or water when in fruit ................. M. longipes Flowers pinkish to violaceous (rarely white), solitary; anthers ovoid, tube long, border oblique or slightly sinuate; leaves smaller, 2–10 mm long, linear to linear-subulate; peduncles erect in fruit ............... M. sellowiana

Mayaca fluviatilis Aubl., Hist. Pl. Guiane 42, pl. 15. 1775. Submerged or terrestrial herb, variable in size and density of leaves. Swamps and marshes, near sea level to 1300(–1900) m; Delta Amacuro (road from Tucupita to Volcán), Bolívar (scattered), Amazonas (base of Cerro Duida, base of Cerro Huachamacari, La Esmeralda, Río Samariapo, Río Sipapo, San Carlos de Río Negro). Llanos of Venezuela; southern U.S.A., Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay, Argentina. ◆Fig. 176. Two forms occur in the flora area, f. fluviatilis, with the border of the anther slit smooth, and f. kunthii (Seub.) Lourteig, with the border of the anther slit 1- or 2-lobed and thickened.

Mayaca longipes Mart. ex Seub. in Mart., Fl. Bras. 3(1): 229, pl. 31, fig. 2. 1855, non Gand. 1920. Submerged aquatic herb. Lagoons, along rivers, 100–1100 m; Bolívar (Apacará-tepui, near Divina Pastora on Río Kukenán, near El Manteco), Amazonas (Río Cunucunuma above Culebra). Guyana, Suriname, French Guiana, Amazonian Brazil, Bolivia. ◆Fig. 175. Mayaca sellowiana Kunth, Enum. Pl. 4: 32. 1843. Submerged or emergent herb. Back waters near rivers, wet areas, 100–1300 m; Bolívar (scattered), Amazonas (Caño Yagua, Cerro Yapacana, La Esmeralda, Río Casiquiare). Llanos of Venezuela; Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 177.

262

M AYACACEAE

Fig. 175. Mayaca longipes

Fig. 176. Mayaca fluviatilis

Fig. 177. Mayaca sellowiana

M ELASTOMATACEAE

263

MELASTOMATACEAE by Paul E. Berry, Andreas Gröger, Bruce K. Holst, Thomas Morley, Fabián A. Michelangeli, Navina G. Luckana, Frank Almeda, Susanne S. Renner, Alina Freire-Fierro, Orbelia R. Robinson, and Kay Yatskievych Trees, shrubs, herbs, or lianas, terrestrial, lithophytic, or epiphytic. Leaves simple, opposite, whorled, or pseudo-alternate by abortion or early dehiscence of one member of each pair, usually with 3–several palmate or subpalmate (pliveined) primary veins (used here to include the midvein as well as the pair or pairs of subparallel veins on either side of the midvein, whether they diverge at the base of the blade or higher up, as in the case of pliveined species). Inflorescence terminal or axillary, usually cymose. Flowers (3)4 or 5(–8)-merous. Hypanthium bearing the sepals, petals, and stamens on a torus (vascular plexus) at or near the base of the calyx lobes; calyx usually open in bud, sometimes calyptrate or dehiscing irregularly; petals free, contorted to the right, usually spreading. Stamens usually twice as many as the petals, usually ± dimorphic at least in size; anthers basifixed, generally subulate, 2- or 4-celled, opening by 1, 2, or rarely 4 terminal pores, connective often prolonged below the thecae and often variously appendaged ventrally and/or dorsally. Ovary 1– 5(–15)-locular, superior or inferior, usually multiovulate; placentation usually axile; style 1; stigma punctiform to capitate. Fruit a loculicidal capsule or a berry. Seeds usually numerous. Pantropics and subtropics, 4500–5000 species and 150–165 genera, 49 genera and 427 species in the flora area. Two genera, Mouriri and Votomita, are now placed in a separate family, Memecylaceae, but we include them here in accordance with their former placement as a subfamily of Melastomataceae in the Cronquist system. Many of the generic treatments herein rely heavily on floristic treatments published by the late John J. Wurdack, notably his treatments in Flora de Venezuela (Wurdack in Lasser, Fl. Venez. 8(1): pp. 1–737. 1973) and Flora of the Guianas [Wurdack in Görts, Fl. Guianas, ser. A, fam. 99 (Melastomataceae). 1993]. We also wish to acknowledge John’s help in compiling an accurate checklist of the Melastomataceae species in the flora area in the early stages of this project. Key to the Genera of Melastomataceae by Paul E. Berry and Susanne Renner 1. 1. 2(1).

2.

Ovaries inferior to partly inferior; fruits berry-like ................................. 2 Ovaries superior to 1/2-inferior (3/4-inferior in Tateanthus); fruits dehiscent capsules or indehiscent berry-like capsules (Aciotis) ......................... 27 Leaf venation pinnate or only the midrib evident (exceptionally, in Mouriri sagotiana, with a pair of prominent loop-connected primary veins); anther connective with an elongate sunken dorsal gland or a terminal rounded gland (Memecyloideae) ............................................ 3 Leaves with 1–several pairs of strongly developed primary veins diverging from midrib at or somewhat above blade base; anther connective lacking an abaxial (dorsal) gland (occasionally with glandular hairs near or at base) ...................................................................................... 4

264

M ELASTOMATACEAE

3(2).

3.

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

6. 7(6).

7.

8(5). 8. 9(8).

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

12.

13(4). 13. 14(13).

Flowers 5-merous; anthers straight or curved, usually on elongate filaments infolded in bud; seed polished at least on the enlarged outer face of the ovule, often all over except for the hilum; petals lacking stone cells; fruits 1–12-seeded ............................................................. 30. Mouriri Flowers 4-merous; anthers straight on short nonfolded filaments; seed unpolished; petals containing numerous box-shaped stone cells unattached to veins; fruits 1-seeded ............................................... 49. Votomita Inflorescences terminal, or pseudolateral by overtopping vegetative growth, but then solitary in upper leaf axils ........................................ 5 Inflorescences lateral, paired in upper leaf axils or from branchlets below leaves .................................................................................................... 13 Petals acute to acuminate at apex ............................................................. 6 Petals rounded to retuse at apex ............................................................... 8 Flowers 4-merous; leaves pliveined (main primary veins departing 0.5–1.5 cm above the base of the blade); dried fruits 8-ribbed ........................................................................... 35. Ossaea (O. micrantha) Flowers 5-merous, or if 4-merous with primary veins departing at the base of the leaf blade and the fruits cylindrical .................................... 7 Flowers 4-merous, or if 5-merous then the inflorescence and hypanthium with simple hairs at least in part; calyx usually persistent in fruit .................................................................................................. 19. Leandra Flowers 5-merous, the inflorescence and hypanthium with stellate hairs only; calyx deciduous in fruit .................................................................. ...................................... 27. Miconia (M. eugenioides, M. tetraspermoides) Calyx thick and calyptrate, circumcissilely dehiscent at anthesis; stamens 10–20 .................................................................................... 13. Conostegia Calyx truncate or lobed, usually not circumcissilely dehiscent; stamens mostly 10 or fewer (to 12 in some Clidemia and one Tococa species) ..... 9 Inflorescences in upper leaf axils or at defoliated nodes, rarely terminal and then always becoming pseudolateral; pubescence of lower surface of leaves often gland-tipped ................................................................. 10 Inflorescences in terminal panicles ......................................................... 11 Outer calyx reduced or free, or if fused laterally, not forming a skirt around the torus (top of hypanthium) ................................... 10. Clidemia Outer calyx fused laterally forming a skirt around the torus .................... ........................................................................ 47. Tococa (T. macrosperma) Ant domatia present along the petiole or at the base of the leaf blade ..................................................................................................... 47. Tococa Ant domatia (expanded, ant-inhabited cavity) lacking .......................... 12 Lower surface of leaves with amorphous or dendritic-stellulate hairs, if glabrous, then the stamens without a dorso-basal minute spur; ovaries 2–10-locular ............................................................................... 27. Miconia Lower surface of leaves glabrous or with long glandular hairs, the stamens with a dorso-basal minute spur (except for T. tepuiensis); ovary locules always 3 .......................................................................... 47. Tococa Ant domatia present along the petiole or at the base of the leaf blade .............................................................................................................. 14 Ant domatia lacking ................................................................................. 16 Leaves strongly dimorphic in each pair, one with a blade-impressed ant

M ELASTOMATACEAE 265

domatia and the other nonvesicular .......................................... 23. Maieta 14. Leaves ± isomorphic in each pair, one or both with an ant domatia ...... 15 15(14). Each leaf of pair with an ant domatia; petals white .................. 10. Clidemia 15. One leaf of each pair with an ant domatia; petals red ............................... .......................................................................... 47. Tococa (T. rotundifolia) 16(13). Flowers 6–8(9)-merous ............................................................................. 17 16. Flowers 4- or 5(6)-merous ........................................................................ 21 17(16). Flowers subtended by 2 pairs of prominent, persistent bracts .............. 18 17. Flowers lacking prominent bracts ........................................................... 19 18(17). Anther thecae 1/2–2/3 as wide as long ................................................ 7. Blakea 18. Anther thecae 1/5–1/3 as wide as long ............................................ 48. Topobea 19(17). Calyx regularly or irregularly dehiscent at anthesis; petals (20–)22– 24 mm long; stigma fluted-capitate, 2.5–4 mm diameter ......... 5. Bellucia 19. Calyx persistent; petals shorter than above; stigma < 2 mm diameter .............................................................................................................. 20 20(19). Flowers axillary, 6-merous; ovary completely adnate to the hypanthium ............................................................................ 35. Ossaea (O. mavacana) 20. Flowers 7- or 8-merous, in an inflorescence 3–6 cm long; ovary 1/10 adnate to the hypanthium ................................................ 10. Clidemia (C. octona) 21(16). Flowers in developed inflorescences with an obvious axis ..................... 22 21. Flowers solitary but aggregated in the upper leaf axils or on branchlets below leaves, the inflorescence axis not developed ............................ 23 22(21). Small to large trees, 4–30 m tall; petals with callus ridges on inner (upper) surface ................................................................................. 20. Loreya 22. Shrubs or subshrubs; petals lacking callus ridges on upper surface ................................................................................................. 10. Clidemia 23(21). Hypanthium 1.7–3 mm long .................................................................... 24 23. Hypanthium (3–)4–15 mm long ............................................................... 25 24(23). Petals rounded at apex, plane ...................... 10. Clidemia (C. conglomerata) 24. Petals bluntly acute, cross-ridged (appearing somewhat wrinkled in the middle when fresh) ............................................................. 18. Henriettella 25(23). Calyx calyptriform and closed in bud, dehiscing irregularly at anthesis; ovary 10-locular .................................................................. 31. Myriaspora 25. Calyx open in bud or dehiscing regularly and persistent at anthesis; ovary 2–6-locular ............................................................................................ 26 26(25). Anther tips spoon-shaped; stigma not or only slightly expanded .............. ............................................................................................... 17. Henriettea 26. Anther tips rounded to truncate; stigma capitate, to 2 mm diameter ..................................................................................................... 20. Loreya 27(1). Ericoid shrubs (leaves small, tough, < 10 × 5 mm, tightly decussate on the stems); generally at high elevations (> 1000 m) ................................. 28 27. Herbs or nonericoid shrubs or trees (leaves on shrubs larger and with evident internodes); at both low and high elevations ............................. 29 28(27). Flowers 4-merous; capsule 4-valved ............................................ 25. Marcetia 28. Flowers 5-merous; capsule 3-valved .......................................... 28. Microlicia 29(27). Petals yellow to orange; plants of tepui summits and slopes, or if in the lowlands, only on granitic outcrops (lajas) ......................................... 30 29. Petals of a different color, usually white, purple, or pink; plants growing in various situations ............................................................................ 32

266

M ELASTOMATACEAE

30(29). Flowers 5-merous, the hypanthium 5-winged; fruits 5-locular .................. .............................................................................................. 45. Tateanthus 30. Flowers 4-merous, the hypanthium terete; fruits 3- or 4-locular .......... 31 31(30). Shrubs generally > 50 cm tall (usually 1–5 m); petals proximally orange, distally yellow; capsule 3-locular; seeds winged .................... 1. Acanthella 31. Subshrubs < 50 cm tall; petals uniformly yellow; capsule 4-locular; seeds not winged ............................................................................. 9. Chaetolepis 32(29). Capsules and mature hypanthia 3-winged; plants herbaceous and recumbent, some leaves alternate by abortion of one member of each pair ................................................................................................. 6. Bertolonia 32. Capsules and mature hypanthia terete, 4- or 5-winged, or with 8–10 longitudinal ribs; plants arborescent, shrubby, or herbaceous, the leaves opposite ................................................................................................ 33 33(32). Tiny herbs or miniature subshrubs generally < 0.5 m tall, or occasionally broad-leaved herbs to 1.5 m tall; flowers secund in unilateral (scorpioid) cymes or else solitary or on an erect scape; capsules and mature hypanthia with 8 or 10 conspicuous longitudinal ribs, turbinate .............................................................................................................. 34 33. Plants either small as above, or commonly larger shrubs or small trees; flowers various, but not secund in unilateral cymes; capsules and mature hypanthia terete, or if winged or ribbed, then these < 8 ............ 35 34(33). Flowers 5-merous, secund in unilateral scorpioid cymes, or rarely solitary; anther connective with an ascending dorsal spur and a basal appendage; leaves same size in each pair .............................. 42. Salpinga 34. Flowers 4- or 5-merous, solitary or on an erect scape; anther connective with a single dorsal appendage; some species with one leaf of each pair substantially smaller than the other ............................ 22. Macrocentrum 35(33). Stamen connective with only abaxial (dorsal) appendages .................... 36 35. Stamen connective with adaxial (ventral), or sometimes also abaxial (dorsal) appendages .................................................................................... 47 36(35). Fruit and ovary 2- or 3-locular ................................................................ 37 36. Fruit and ovary 4–7-locular ..................................................................... 41 37(36). Anther connective with a dorsal ascending spur and a basal appendage .............................................................................................................. 38 37. Anther connective with a single dorsal appendage ................................ 39 38(37). Hairs on the stems and hypanthia simple or T-shaped, 1–4 mm long ............................................................................................... 4. Adelobotrys 38. Hairs on the stems and hypanthia stellulate, 0.1–0.2 mm long ... 26. Meriania 39(37). Flowers predominantly 4- or 5-merous ................................ 16. Graffenrieda 39. Flowers predominantly 6-merous ............................................................ 40 40(39). Hypanthium and ovary nonglandular .................................. 16. Graffenrieda 40. Hypanthium and ovary moderately glandular-setose ...... 32. Neblinanthera 41(36). Calyx calyptrate, deciduous at anthesis .................................................. 42 41. Calyx truncate or with evident lobes that are persistent at anthesis .... 43 42(41). Anther connective with an ascending dorsal appendage and a basal spur .................................................................................................. 8. Centronia 42. Anther connective lacking an ascending dorsal appendage ....................... .................................. 16. Graffenrieda (G. tricalcarata, G. pendunculata) 43(41). Flowers 4-merous ..................................................................................... 44

M ELASTOMATACEAE 267

43. Flowers 5–7-merous ................................................................................. 45 44(43). Leaf blades 1.5–4 × 0.7–3.2 cm, 5(7)-veined, apex acute; stamens 8, with 4 larger fertile anthers and 4 smaller, sterile, filiform anthers ......................................................................................... 29. Monochaetum 44. Leaf blades 6–12 × 3–6 cm, 3-veined, rounded at the apex; all 8 anthers fertile ......................................................... 16. Graffenrieda (G. reticulata) 45(43). Climbers or hemiepiphytes ....................................................... 4. Adelobotrys 45. Shrubs or trees ......................................................................................... 46 46(45). Anther connective with a dorsal ascending spur and a basal appendage ................................................................................................. 26. Meriania 46. Anther connective with a single dorsal appendage ............. 16. Graffenrieda 47(35). Stamens with a 1–4.5 mm long abaxial (dorsal) spur (in addition to 2 adaxial appendages) ......................................................................... 48 47. Stamens lacking abaxial (dorsal) appendages or with minute (> 0.5 mm) abaxial appendages (in addition to adaxial appendages) ................... 49 48(47). Inflorescences unilateral, axillary; ovaries and capsules 3-locular; seeds pyramidal ........................................................................ 34. Opisthocentra 48. Inflorescences dichasial or paniculate (or flowers solitary), terminal; ovaries and capsules 4-locular; seeds distinctly curved (cochleate) .............................................................................................. 36. Pachyloma 49(47). Perianth 5-merous ................................................................................... 50 49. Perianth 4-merous ................................................................................... 56 50(49). Fertile stamens 5, alternating with 5 staminodes .................................. 51 50. Fertile stamens 10, the two whorls ± dimorphic, but the inner one not staminodal ............................................................................................ 52 51(50). Miniature herbs 5–10 cm tall; leaves linear, 5–9 × ca. 1 mm, the margin glandular-ciliate; petals cream to pink; ovary 3-locular .... 38. Poteranthera 51. Shrubs or subshrubs 30–200 cm tall; leaves larger than above; petals bright reddish purple; ovary 3–5-locular ..................... 41. Rhynchanthera 52(50). Connective prolongation of larger anthers 2/3–3/4 as long as thecae; littlebranched, villous herbs or subshrubs with square stems, subsessile leaves, and flowers in leafy panicles .................................. 14. Desmoscelis 52. Connective prolongation of larger anthers 1/3 or less as long as thecae; some Acisanthera similar to above in habit and leaves, otherwise shrubs or trees, or if herbs or subshrubs, then without the above combination of characters ............................................................................. 53 53(52). Hypanthia and capsules with 4 or 5 stiffly ciliate longitudinal wings at least 1 mm wide .................................................................. 39. Pterogastra 53. Hypanthia and capsules lacking ciliate wings ........................................ 54 54(53). Ovaries and capsules 2- or 3-locular; small herbs 5–60 cm tall ............................................................................................... 3. Acisanthera 54. Ovaries and capsules 4- or 5-locular; mostly (sub)shrubs and trees ...... 55 55(54). Low-growing (to 20 cm high), semiprostrate subshrub, ovaries and capsules (3)4-locular; high elevations (ca. 2500 m) on Sierra de la Neblina ............................................................................................ 12. Comoliopsis 55. Generally larger herbs, erect shrubs, or trees, ovaries and capsules 5locular; widely distributed from lowlands to highlands ..... 46. Tibouchina 56(49). Ovary and capsule (or berry-like capsule) 2-locular ............................... 57 56. Ovaries and capsule 3- or 4-locular ......................................................... 62

268

M ELASTOMATACEAE

57(56). Anther connective simply articulate with filament, not appendaged .... 58 57. Anther connective basally prolonged and adaxially (ventrally) bluntly bilobed, at least in the larger stamens ................................................... 59 58(57). Herbs; petals white to pink; fruits thin-walled indehiscent berry-like capsules or capsules ........................................................................... 2. Aciotis 58. Shrubs; petals pinkish purple; fruits capsules ..................... 43. Sandemania 59(57). Herbs ........................................................................................................ 60 59. Shrubs or subshrubs ................................................................................ 61 60(59). Ventral appendices of stamen connectives subulate ................ 3. Acisanthera 60. Ventral appendices of stamen connectives obtuse ................ 44. Siphanthera 61(59). Ovary apex pubescent with glandular hairs; lower surface of leaves with globose sessile glands ............................................................. 21. Macairea 61. Ovary apex glabrous; lower surface of leaves glabrous or with a few minute glandular or eglandular hairs ..................................... 11. Comolia 62(56). Shrubs with rigid, revolute leaves densely white-woolly on the lower surface ..................................................................................... 24. Mallophyton 62. Herb, shrubs, or lianas, the leaves not rigid, revolute or woolly on the lower surface ........................................................................................ 63 63(62). Calyx hyaline and calyptrate, dehiscing at anthesis; lianas with some leaves alternate by abortion of one member of each pair or else small and opposite, or, if herbaceous, with flowers in a terminal umbellate scape .................................................................................... 37. Phainantha 63. Calyx regularly lobed, persistent; subshrubs or shrubs ......................... 64 64(63). Hypanthia with stalked stellate trichomes between the calyx lobes ................................................................................................ 40. Pterolepis 64. Hypanthia lacking stalked stellate trichomes between the calyx lobes .............................................................................................................. 65 65(64). Ovary apex pubescent with glandular hairs; lower surface of leaves densely glandular-pubescent or with globose sessile glands .............. 21. Macairea 65. Ovary apex glabrous; lower surface of leaves glabrous or with various kinds of pubescence ............................................................................. 66 66(65). Leaves rigid, with revolute margins, glabrous and viscose, the base narrowly acute ........................................................ 11. Comolia (C. vernicosa) 66. Leaves neither rigid nor revolute or viscose, the base generally rounded or cordulate .......................................................................................... 67 67(66). Stamen connectives with yellow adaxial (ventral) appendages 2–3 mm long; capsules 3- or 4-locular; lowland to highland habitats, often on rock outcrops ............................................................................ 15. Ernestia 67. Stamen connectives with purple adaxial (ventral) appendages 0.3–0.7 mm long; capsules 3-locular; in wet lowland habitats, especially Mauritia palm swamps ............................................................................ 33. Nepsera 1. ACANTHELLA Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 748. 1867. by Paul E. Berry Shrubs to small trees. Leaves entire, opposite, usually tightly clustered at the branch tips. Flowers solitary in upper leaf axils, 4-merous. Hypanthium terete, widening broadly in fruit; calyx lobes subulate, persistent; petals pale yellow to orange, flushed at the base with a tinge of orange or red, oblong-obovate. Stamens 8, slightly

Acanthella 269

dimorphic, glabrous; thecae linear-subulate and curved, with a ventrally inclined pore; connective not prolonged or only shortly so, basally with a small ventral bilobed appendage. Ovary free, 3- or 4-locular, glabrous; style glabrous; stigma punctiform. Capsule 3- or 4-valved. Seeds numerous, winged. Southeastern Colombia, southwestern Venezuela, northern Brazil; 2 species, both in the flora area. When the two species of Acanthella grow together, A. sprucei occupies drier microhabitats and sometimes drops its leaves in the dry season, whereas A. pulchra has thick leaves that persist through the dry season (S. Renner, personal communication). Key to the Species of Acanthella 1. 1.

Leaf blades 3–5 × 2–3 cm, obovate to widely obovate, apically retuse and not setulose; leaf nodes and hypanthium glabrous ................... A. pulchra Leaf blades 1–3 × 0.4–1 cm, narrowly rhombic, apically acuminate and setulose; leaf nodes setulose; hypanthium glandular-setose ..... A. sprucei

Acanthella pulchra Gleason, Fieldiana, Bot. 28: 430. 1952. Acanthella plicata Gleason, Mem. New York Bot. Gard. 8: 134. 1953. Shrub or small tree 1–6 m tall; leaves ± succulent, persistent and conduplicate

Fig. 178. Acanthella pulchra

through the dry season. In cracks on granitic outcrops, edges of rapids, 50–400 m; Bolívar (Río Parguaza), Amazonas (Caño Galipero, base of Cerro Sipapo, Isla Ratón, near Raudal de Maipures, Río Autana, Río Cuao basin, upper Río Gavilán in Río Cataniapo

Fig. 179. Acanthella sprucei

270

M ELASTOMATACEAE

basin, near Samariapo, near San Juan de Ucata in Río Guayapo basin). Colombia (Vichada). ◆Fig. 178. Acanthella sprucei Hook f. in Benth. & Hook. f., Gen. Pl. 1: 749. 1867. Acanthella montana Gleason, Mem. New York Bot. Gard. 8: 134. 1953. Chalepophyllum pungens Standl. & Steyerm., Fieldiana, Bot. 28: 568. 1953. Shrub 0.2–5 m tall, often with flattened crown; leaves ± chartaceous, partly deciduous? in the dry season. In cracks on both granitic and sandstone outcrops, 50–1900 m; Bolívar (Río Parguaza, Serranía Carichana), Amazonas (widespread on granite domes from north of Puerto Ayacucho to San Fernando de Atabapo, also Canaripó, Cerro Aratitiyope, summit of Cerro Autana, Cerro Guanay, Cerro Sipapo, Cerro Yutajé, Guarinuma, Isla Ratón, ridgetops near La Es-

meralda, Río Siapa, Samariapo, near San Juan de Ucata in Río Guayapo basin, Santa Bárbara del Orinoco, Salto Yureba). Apure; Colombia (Amazonas, Caquetá, Vaupés, Vichada), northwestern Brazil. ◆Fig. 179. The name Acanthella conferta (Vell.) Cogn. was previously applied to this taxon; however the Vellozo basionym, Melastoma conferta Vell., is based on a difficult-to-interpret plate of a plant from Rio de Janeiro, Brazil, and is most certainly not an Acanthella. Two specimens [Gröger 747 (US), 650 m elevation on granite; and Huber 13609 (US), 720 m elevation, on sandstone] from middle elevations on the Cuao massif have unusually glutinous-pubescent leaves and fruits and may represent an undescribed taxon. Both were collected in fruit in the month of February, and flowering material is needed for further evaluation.

2. ACIOTIS D. Don, Mem. Wern. Nat. Hist. Soc. 4: 283. 1823. Spennera Mart. ex DC., Prodr. 3: 115. 1828. by Alina Freire-Fierro Annual or perennial herbs, stems sometimes winged, glabrous or with simple or glandular trichomes. Leaves ovate to elliptic or oblong, thin and purplish when living, membranous or papyraceous when dried, the base attenuate to cordate, the apex acuminate to obtuse. Inflorescence a terminal cyme, panicle, or thyrse. Flowers 4-merous, usually < 15 mm long. Hypanthium terete or slightly winged; calyx lobes triangular, persistent; petals white to pink or purplish, ovate to lanceolate. Stamens 8, usually ± dimorphic, glabrous; anthers oblong to orbicular and 1-pored; connective not or somewhat prolonged, not appendaged. Ovary 1/2 or more superior, 2- or rarely 3-locular, the apex glabrous or with glandular trichomes. Fruit a 2-valved capsule, sometimes berry-like. Seeds numerous, reniform, foveolate. Southern Mexico, Central America, Lesser Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 13 species, 9 in Venezuela, all in the flora area. Key to the Species of Aciotis 1. 1. 2(1). 2.

Inflorescence with (3–)5–8 flowers per cm of terminal portion of inflorescence; bracts 2–5 mm long ............................................................ A. annua Inflorescence with 1–4 flowers per cm of terminal portion of inflorescence; bracts 0.8–1.5 mm long .......................................................................... 2 Plant prostrate; young stem with woolly, reddish, eglandular trichomes; leaf apex typically obtuse ............................................................. A. ornata Plant erect; young stem glabrous or with pinkish to whitish glandular or eglandular trichomes; leaf apex acute to acuminate ............................ 3

Aciotis 271

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

Lower surface of leaf with web-like striations; parenchyma of hypanthium with spiral tracheids ................................................. A. polystachya Lower surface of leaf without web-like striations; parenchyma of hypanthium without spiral tracheids ............................................................. 4 Inflorescence typically a cyme; fruit a fleshy capsule ........... A. acuminifolia Inflorescence typically a thryse or a panicle (a double biparous cyme in A. sp. A); fruit capsular or berry-like .................................................... 5 Fruit capsular ............................................................................................. 6 Fruit berry-like .......................................................................................... 7 Leaves papyraceous; bracts and petals dorsally glandular-pubescent ...................................................................................................... A. viscida Leaves membranous; bracts and petals dorsally glabrous ................. A. sp. A Leaf base cordate; ovary apex glandular ...................................... A. indecora Leaf base typically cuneate to attenuate; ovary apex glabrous ............... 8 Plants densely eglandular-pubescent ......................................... A. caulialata Plants totally glabrous or strigose ......................................... A. purpurascens

Aciotis acuminifolia (Mart. ex DC.) Triana, Trans. Linn. Soc. London 28: 51, pl. 3, fig. 36c. 1871. —Spennera acuminifolia Mart. ex DC., Prodr. 3: 116. 1828. Spennera dichotoma Benth., J. Bot. (Hooker) 2: 295. 1840. —Aciotis dichotoma (Benth.) Cogn. in Mart., Fl. Bras. 14(3): 460, pl. 104, fig. 1. 1885. Spennera ?anomala Miq., Linnaea 18: 274. 1844. —Aciotis dichotoma var. anomala (Miq.) Cogn. in Mart., Fl. Bras. 14(3): 461. 1885. Spennera dichotoma var. lanceolata Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 142. 1850. Spennera sphaeranthera Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 142, pl. 5, fig. 2. 1850. —Aciotis sphaeranthera (Naudin) Cogn. in Mart., Fl. Bras. 14(3): 458. 1885. Aciotis aequatorialis Cogn. in Mart., Fl. Bras. 14(3): 464, pl. 105, fig. 2. 1885. Aciotis amazonica Cogn. in Mart., Fl. Bras. 14(3): 461, pl. 105, fig. 1. 1885. Aciotis amazonica var. radicans Cogn. in Mart., Fl. Bras. 14(3): 462. 1885. Aciotis dichotoma var. longifolia S. Moore, Trans. Linn. Soc. London, Bot. 4: 361. 1895. Pubescent herb 10–50 cm tall with whitish to pinkish eglandular trichomes; leaves membranous, with base truncate to attenuate-decurrent, lower surface glabrous or with

whitish eglandular trichomes, without weblike striations; inflorescence ca. 6 cm × 3.5–6 cm; hypanthium without spiral tracheids in parenchyma, externally with glandular trichomes on the costae; ovary without spiral tracheids in parenchyma, the apex glabrous; fruit a fleshy capsule; ovary wall membranous when dry. Disturbed areas, near sea level to 700 m; Delta Amacuro (Sacupana), Bolívar (Caicara, Canaima, Isla Anacoco, La Paragua, Río Caura, Río Paramichí), Amazonas (Puerto Ayacucho, Río Atures, Río Casiquiare, Samariapo, Sierra de la Neblina). Apure, Táchira; Lesser Antilles, Colombia, Guyana, French Guiana, Suriname, Peru, Brazil. ◆Fig. 181. Aciotis annua (Mart. ex DC.) Triana, Trans. Linn. Soc. London 28: 52. 1871. —Spennera annua Mart. ex DC., Prodr. 3: 115. 1828. —Boyuyo, Garrapata, Nombró-poh (Pemón). Spennera dysophylla Benth., J. Bot. (Hooker) 2: 296. 1840. —Aciotis dysophylla (Benth.) Triana, Trans. Linn. Soc. London 28: 52. 1871. Spennera tetraptera Miq., Comm. Phytogr. 2: 76, t. 10, fig. E. 1840. Herb (2–)15–20 cm tall; stems quadrangular with conspicuous wings; leaves with lower surface whitish green to light green, pubescent, with web-like striations; inflorescence 5–7.5(–13) × 2–4.5(–10.5) cm; hypanthium with spiral tracheids in paren-

272

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chyma, externally sericeous and glandularpubescent, ovary with spiral tracheids in parenchyma, the apex glandular-pubescent; fruit a capsule. Disturbed habitats, in seasonally flooded areas as well as drier habitats or in sandy soils, 50–900 m; Bolívar (Gran Sabana, Icabarú), Amazonas (Caño Iguapo, Cerro Duida, Maroa, Río Casiquiare, Río Cunucunuma, Río Guainía, San Carlos de Río Negro, Sierra de la Neblina). Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 184. Aciotis caulialata (Ruiz & Pav.) Triana, Trans. Linn. Soc. London 28: 52, pl. 3, fig. 36b. 1871. —Arthrostemma caulialatum Ruiz & Pav., Fl. Peruv. 4: pl. 327. 1802. —Spennera caulialata (Ruiz & Pav.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 149. 1850. Spennera rubricaulis Schrank & Mart. ex DC., Prodr. 3: 115. 1828. —Aciotis rubricaulis (Schrank & Mart. ex DC.) Triana, Trans. Linn. Soc. London 28: 52. 1871. Spennera alata Beurl., Kongl. Vetensk. Acad. Handl. 40: 125. 1854. —Aciotis purpurascens var. alata (Beurl.) Cogn. in Mart., Fl. Bras. 14(3): 474. 1885, as to type, Bilberg s.n. —Aciotis alata (Beurl.) Almeda, Novon 7: 333. 1997. Aciotis indecora var. macrophylla Cogn. in Mart., Fl. Bras. 14(3): 470. 1885. Aciotis levyana Cogn. in Mart., Fl. Bras. 14(3): 460. 1885. Aciotis aristata Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 352. 1915. Aciotis purpurascens var. martinensis J.F. Macbr., Field. Mus. Nat. Hist., Bot. Ser. 13(4): 299. 1941. Aciotis asplundii Wurdack, Phytologia 35: 2. 1976. Herb 0.3–1.5 m tall, erect to prostrate; stolons present; stems quadrangular with conspicuous wings, pubescent with long (ca. 1.5 mm), white, eglandular trichomes; leaves membranous, with base cuneate, lower surface green to purplish, pubescent with long, eglandular, whitish trichomes, without weblike striations; inflorescence a thyrse; hypanthium without spiral tracheids in parenchyma, externally hirsute with glandular or eglandular trichomes; ovary without spiral tracheids in parenchyma; fruit berry-like, ovary wall membranous when dry. Dis-

turbed, especially swampy areas, 800–1200 m; Bolívar (El Dorado to Santa Elena de Uairén road, Macizo del Chimantá). Táchira; southern Mexico, Central America, Colombia, French Guiana, Suriname, Ecuador, Peru, Brazil. Aciotis indecora (Bonpl.) Triana, Trans. Linn. Soc. London 28: 52. 1871. —Rhexia indecora Bonpl. in Humb. & Bonpl., Monogr. Melast. 2: 131, pl. 49. 1823 [1821]. —Spennera indecora (Bonpl.) DC., Prodr. 3: 116. 1828. —Boyoyo blanco, Pio-yek (Arekuna). Spennera laxa DC., Prodr. 3: 116. 1828. —Aciotis laxa (DC.) Cogn. in Mart., Fl. Bras. 14(3): 476, pl. 108. 1885. Spennera acutiflora Mart., Nov. Gen. Sp. Pl. 3: 114. 1829 [1831]. —Aciotis acutiflora (Mart.) Triana, Trans. Linn. Soc. London 28: 52. 1871. Spennera kappleriana Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 146. 1850. —Aciotis laxa var. kappleriana Cogn. in Mart., Fl. Bras. 14(3): 477. 1885. Spennera rostellata Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 143. 1850. —Aciotis rostellata (Naudin) Triana, Trans. Linn. Soc. London 28: 51. 1871. Spennera viscosa Bonpl. ex Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 146. 1850. —Aciotis viscosa (Bonpl. ex Naudin) Triana, Trans. Linn. Soc. London 28: 52. 1871. Aciotis acutiflora var. parvifolia Cogn. in Mart., Fl. Bras. 14(3): 476. 1885. Aciotis indecora var. glabrescens Cogn. in Mart., Fl. Bras. 14(3): 470. 1885. Aciotis indecora var. macrophylla Cogn. in Mart., Fl. Bras. 14(3): 470. 1885. Aciotis indecora var. sagotiana Cogn. in Mart., Fl. Bras. 14(3): 470. 1885. Aciotis laxa var. robusta Cogn. in Mart., Fl. Bras. 14(3): 477, pl. 108. 1885. Aciotis anomala Brade, Publ. Inst. Nac. Pesq. Amazônica Bot. 8: 8. 1958. Herb to subshrub 0.5–2 m tall, erect; stems quadrangular with inconspicuous wings, glandular-pubescent; leaves with apex acute to acuminate, lower surface light green, with eglandular, long-strigose hairs, without web-like striations; inflorescence a thyrse or panicle, (3.5–)4.5–8 × 6–18 cm; hypanthium without spiral tracheids in paren-

Aciotis 273

chyma, externally densely glandular-pubescent; petals dorsally glandular-pubescent (rarely glabrous); ovary without spiral tracheids in parenchyma, the apex glandular-pubescent; fruit berry-like; ovary wall membranous when dry. Lowland to highland forests and openings, 50–1700 m; Bolívar (Cerro Ichún, Gran Sabana, Icabarú, Los Testigos, Macizo del Chimantá, Río Tonoro, Río Venamo, Sierra de Maigualida, Uaipán-tepui), Amazonas (Cerro Impacto, Río Atabapo, Río Atacavi, Río Casiquiare, San Carlos de Río Negro, Yavita). Trujillo; Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 185.

glandular-pubescent with dark reddish or whitish trichomes, with web-like striations; inflorescence a double biparous cyme, 4–8 × 3–6 cm; bracts dorsally glandular-pubescent; hypanthium with spiral tracheids in parenchyma, externally densely glandular-pubescent; petals dorsally with glandular trichomes; ovary with spiral tracheids in parenchyma, the apex glandular-pubescent; fruit a capsule. Open, disturbed habitats, 50–1200 m; Bolívar (Río Canaracuni, Río Orinoco), Amazonas (Cerro Duida, Puerto Ayacucho to Samariapo, Río Cunucunuma, San Carlos de Río Negro). Mérida; Colombia, Ecuador, Peru, Amazonian Brazil. ◆Fig. 182.

Aciotis ornata (Miq.) Gleason, Brittonia 1: 149. 1932. —Spennera ornata Miq., Stirp. Surinam. Select. 56, t. 13. 1850 [1851]. Aciotis herbacea Steud. ex Cogn. in Mart., Fl. Bras. 14(3): 468. 1885. Herb 5–25 cm long, prostrate; stolons present; stems quadrangular with inconspicuous wings, pubescent with woolly, reddish, eglandular trichomes < 1 mm long; leaves with the apex obtuse (rarely acute), the lower surface whitish green or purplish, pubescent with eglandular trichomes, with web-like striations; inflorescence long-pedunculate and rather few-flowered; hypanthium with spiral tracheids in parenchyma, externally densely glandular-pubescent, internally glabrous; ovary with spiral tracheids in parenchyma, the apex glandular-pubescent; fruit a capsule. Secondary areas and shaded forest understory, 100–800 m; Delta Amacuro (Serranía de Imataca), Bolívar (Maijía, upper Río Paragua, Serranía Marutaní, Sierra de Maigualida), Amazonas (Río Yatúa). Mérida; Colombia, Guyana, Suriname, French Guiana, Ecuador, Amazonian Brazil. ◆Fig. 183.

Aciotis purpurascens (Aubl.) Triana, Trans. Linn. Soc. London 28: 52, pl. 3, fig. 36a. 1871. —Melastoma purpurascens Aubl., Hist. Pl. Guiane 402, pl. 154. 1775. —Melastoma purpurea (Aubl.) Willd., Sp. Pl. 2(1): 590. 1798 [1799]. —Miconia purpurascens (Aubl.) DC., Prodr. 3: 179. 1828. —Boyuyo morado, Küdadei (Yekwana). Rhexia sileniflora Bonpl. in Humb. & Bonpl., Monogr. Melast. 2: 129, pl 48. 1823 [1821]. —Spennera sileniflora (Bonpl.) DC., Prodr. 3: 116. 1828. Aciotis sileniflora (Bonpl.) Triana, Trans. Linn. Soc. London 28: 52. 1871. Aciotis discolor D. Don, Mem. Wern. Nat. Hist. Soc. 4: 301. 1823. Spennera fragilis Rich. ex DC., Prodr. 3: 115. 1828. —Aciotis fragilis (Rich. ex DC.) Cogn. in Mart., Fl. Bras. 14(3): 475. 1885. Spennera pellucida Rich. ex DC., Prodr. 3: 115. 1828. —Aciotis purpurascens var. pellucida (Rich. ex DC.) Cogn. in Mart., Fl. Bras. 14(3): 474. 1885. —Aciotis pellucida (Rich. ex DC.) Pittier et al., Cat. Fl. Venez. 2: 244. 1947. Spennera grandifolia Miq., Linnaea 18: 273. 1844. Spennera sieberi Steud., Flora 27: 720. 1844. —Aciotis sieberi (Steud.) Triana, Trans. Linn. Soc. London 28: 52. 1871. Spennera martinicensis Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 147. 1850. —Aciotis martinicensis (Naudin) Urb., Symb. Antill. 5: 446. 1908. Spennera uliginosa Bonpl. ex Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 144. 1850. —Aciotis uliginosa (Bonpl. ex Naudin)

Aciotis polystachya (Bonpl.) Triana, Trans. Linn. Soc. London 28: 52. 1871. —Rhexia polystachya Bonpl. in Humb. & Bonpl., Monogr. Melast. 2: 98, pl. 38. 1818. —Spennera polystachya (Bonpl.) Mart. ex DC., Prodr. 3: 116. 1828. —Sirichá (Yekwana). Herb 15–50 cm tall, erect; stems quadrangular with inconspicuous wings; leaves cordiform to ovate, lower surface whitish green,

274

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Fig. 180. Aciotis purpurascens

Fig. 181. Aciotis acuminifolia

"dichotoma" form "aequatorialis" form

Aciotis 275

Fig. 182. Aciotis polystachya

Fig. 183. Aciotis ornata

Fig. 184. Aciotis annua Fig. 185. Aciotis indecora

276

M ELASTOMATACEAE

Triana, Trans. Linn. Soc. London 28: 51. 1871. Aciotis circaeifolia var. major Cogn. in Mart., Fl. Bras. 14(3): 473. 1885. Aciotis purpurascens var. alata Cogn. in Mart., Fl. Bras. 14(3): 474. 1885, as to type, Fendler 149. Spennera lancifolia Steud. ex Cogn. in Mart., Fl. Bras. 14(3): 475. 1885. —Aciotis fragilis var. lancifolia (Steud. ex Cogn.) Cogn. in Mart., Fl. Bras. 14(3): 475. 1885. Aciotis purpurascens var. longifolia Cogn. in Mart., Fl. Bras. 14(3): 474. 1885. Aciotis longifolia var. glabra Huber, Bol. Mus. Paraense Hist. Nat. 4: 595. 1906. Aciotis trinitensis Cogn. in Urb., Symb. Antill. 7: 310. 1912. Herb to subshrub 0.5–1.5(–2) m tall, erect; stems quadrangular with conspicuous wings, glabrous or strigose; leaves membranous with base attenuate to cuneate and apex acuminate, lower surface green to purplish, glabrous or strigose, without web-like striations; inflorescence a thyrse 10–18 × 3.5–6(–10) cm, bracts dorsally glabrous; hypanthium without spiral tracheids in parenchyma, externally glabrous; ovary without spiral tracheids in parenchyma, the apex glabrous; fruit berry-like; ovary wall membranous when dry. Open, disturbed areas or forest understory, in seasonally flooded or nonflooded areas, near sea level to 1000 m; Delta Amacuro (El Palmar, Sacupana, Serranía de Imataca, Tucupita), Bolívar (Anacoco, Caicara to Puerto Ayacucho road above Río Maniapure, south-southeast of Entrerios, Gran Sabana, Los Testigos, Río Icabarú, Río Nichare, Salto Pará, Serranía de Imataca, Urimán), Amazonas (Capibara, Maroa, Piedra Cocuy, Puerto Ayacucho to Samariapo, Río Atabapo, Río Guayapo, San Carlos de Río Negro, Sierra de la Neblina). Apure, Barinas, Carabobo, Mérida, Monagas, Sucre, Táchira, Zulia; southern Mexico, Central America, Lesser Antilles, Colombia, Guyana,

Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 180. Aciotis viscida (Benth.) A. Freire-Fierro, comb. nov. —Spennera viscida Benth., J. Bot. (Hooker) 2: 297. 1840. —Aciotis indecora var. glabrescens Cogn. in Mart., Fl. Bras. 14(3): 470. 1885, as to holotype R.H. Schomburgk I 999. Herb 0.4–1 m tall, erect; stems slightly quadrangular with inconspicuous wings; leaves with base obtuse to cordate, 5–7veined; lower surface green, pubescent with glandular and eglandular trichomes, without web-like striations; inflorescence a thryse 5.5–9 × 2–4(–5) cm, bracts dorsally glandular-pubescent; hypanthium without spiral tracheids in parenchyma, externally densely glandular-pubescent; petals dorsally with glandular trichomes; ovary without spiral tracheids in parenchyma, the apex glandular-pubescent; fruit a capsule. In flooded or nonflooded open areas, 100–200 m; Amazonas (La Esmeralda, Río Cuao, San Carlos de Río Negro). Guyana, Brazil. Aciotis sp. A Herb 14–30 cm tall; stems quadrangular with inconspicuous wings; leaves membranous, cordiform, 7-veined; lower surface light green, pubescent, with eglandular appressed trichomes, without web-like striations; inflorescence a double biparous cyme (rarely a thyrse), 6–14 × (3–)4–5.5(–7) cm; hypanthium ca. 1.5 mm long, 1–1.8 mm diameter, without spiral tracheids in parenchyma, externally with few glandular trichomes; ovary without spiral tracheids in parenchyma, the apex glandular-pubescent; fruit a capsule; ovary wall coriaceous. Riverbanks and evergreen lowland and lower montane forests, 100–400 m; Bolívar (Salto Pará), Amazonas (upper Río Orinoco, San Carlos de Río Negro). Distrito Federal; Colombia (Meta, Putumayo), Ecuador, Peru, Brazil (Amazonas, Pará).

3. ACISANTHERA P. Browne, Civ. Nat. Hist. Jamaica 217. 1756. by Navina G. Luckana and Paul E. Berry Herbs 5–60 cm tall, rarely lignified and often glandular-puberulous. Stem quadrangular, mostly winged, sometimes inflated at the base. Leaves isomorphic, opposite and mostly ovate, fleshy or membranous; margin obscurely serrulate or denticu-

Acisanthera 277

late. Flowers 4- or 5-merous, solitary or in terminal few-flowered panicles. Hypanthium terete; calyx lobes persistent; petals white, pink, or purple, obovate to suborbiculate, usually eciliate. Stamens 8 or 10, sometimes only one whorl fertile, with varying degrees of abortion in the other whorl, glabrous; anthers subulate to truncate-ovate, minutely to broadly pored; connective ± prolonged, ventrally bilobed, dorsally usually not appendaged. Ovary free, usually 2- or 3-locular, usually glabrous; style usually glabrous; stigma not expanded. Fruit a capsule. Seeds numerous, subcochleate, foveolate. Southern Mexico, Central America, West Indies, from northern South America to Bolivia and Argentina; ca. 20 species, 7 in Venezuela, all in the flora area. Key to the Species of Acisanthera 1. 1. 2(1). 2.

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

Leaves obviously petiolate ......................................................................... 2 Leaves sessile ............................................................................................. 3 Upper leaf surface glabrous; ventral appendages of the larger anthers not much inflated, 4–5 times longer than wide ............................. A. quadrata Upper leaf surface sparsely to moderately glandular-puberulent; ventral appendages of the larger anthers inflated, 2–3 times longer than wide .................................................................................................... A. uniflora Bases of leaf blades acute to attenuate; stems not inflated; ventral anther connective appendages subulate and acute .......................... A. hedyotidea Bases of leaf blades rounded or cordate; stems usually inflated basally; ventral anther connective appendages blunt ....................................... 4 Small and large anther thecae oblong, tapering to a narrow pore .......... 5 Small (and usually large) anther thecae broadly oval and truncate, the pore equaling the anther width ............................................................. 6 Plants glabrous; calyx lobes longer than the hypanthium ............ A. bivalvis Plants glandular-puberulous; calyx lobes usually shorter than the hypanthium .................................................................................. A. limnobios Large stamen connective prolongation distinctly shorter than the thecae, the ventral lobes and thecae erect; calyx lobes lanceolate ...... A. crassipes Large stamen connective prolongation longer than or about equaling the thecae, the ventral lobes and thecae horizontal; calyx lobes lanceolateoblong .............................................................................................. A. nana

Acisanthera bivalvis (Aubl.) Cogn. in Mart., Fl. Bras. 14(3): 216. 1885. —Melastoma bivalve Aubl., Hist. Pl. Guiane 404, t. 155. 1775, “bivalvis.” —Rhexia bivalvis (Aubl.) Vahl, Eclog. Amer. 1: 38. 1796 [1797]. —Microlicia bivalvis (Aubl.) DC., Prodr. 3: 117. 1828. —Noterophila beccabunga Mart. ex Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: 280. 1849. —Acisanthera beccabunga (Mart. ex Naudin) Triana, Trans. Linn. Soc. London 28: 33, t. 2, fig. 18d. 1871. Melastoma trivalvis Aubl., Hist. Pl. Guiane 406, t. 155. 1775. —Rhexia

trivalvis (Aubl.) Vahl, Eclog. Amer. 1: 39. 1796 [1797]. —Microlicia brevifolia DC., Prodr. 3: 117. 1828. —Noterophila brevifolia (DC.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: 280. 1849. —Acisanthera brevifolia (DC.) Griseb., Fl. Brit. W. I. 269. 1860. —Acisanthera trivalvis (Aubl.) Cogn. in Mart., Fl. Bras. 14(3): 217. 1883. Glabrous herb 20–60 cm tall, with winged, quadrangular stems; leaves sessile, fleshy, oblong-elliptic to lanceolate, acute to obtuse at the apex, rounded to cordate at the base, and entire to obscurely serrulate, 0.5–2

278

M ELASTOMATACEAE

cm × 0.2–0.5 cm, 3–5-veined; flowers terminal, predominantly 5-merous; hypanthium 3.3 mm wide, calyx lobes lanceolate-subulate, 4–5 mm long; petals pink, 6–12 mm × 4–9 mm; ovary 2- or 3-locular. Mauritia palm swamps, ca. 900 m; Bolívar (Divina Pastora, Santa Elena de Uairén). Monagas; Central America, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 186.

Acisanthera boissieriana Cogn. in Mart., Fl. Bras. 14(3): 209, t. 49, fig. 1. 1885. Herb 10–30 cm tall; stems quadrangular, winged and rarely to moderately glandularsetulose; leaves sessile, membranous, elliptic to ovate-oblong, largely to abruptly acute at the apex, broadly acute at the base, nonciliate, denticulate, 1–2 × 0.5–1 cm, 3veined; flowers solitary in upper leaf axils, 4merous; hypanthium 2 × 1 mm, glabrous to rarely glandular-puberulous; calyx lobes broadly triangular, ca. 1.5 mm long and ending in a glandular hair; petals pink, 3–4 × 2.5 mm; ovary 2-locular. Mauritia palm swamps, 200–500 m; Bolívar (Altiplanicie de Nuria, Cerro Altamira, Ciudad Piar to Cerro Toribio). Carabobo; Brazil. ◆Fig. 190.

Acisanthera crassipes (Naudin) Wurdack, Fieldiana, Bot. 29: 539. 1963. —Onoctonia crassipes Naudin, Ann. Sci. Bot. Ser. 3, 12: 277, t. 12, fig. 4. 1849. —Poteranthera crassipes (Naudin) Triana, Trans. Linn. Soc. London 28: 33, fig. 17c. 1871. Onoctonia calcarata Naudin, Ann. Sci. Bot. Ser. 3, 12: 277, t. 12, fig. 2. 1849. —Poteranthera calcarata (Naudin) Triana, Trans. Linn. Soc. London 28: 33, fig. 17a. 1871. Onoctonia pauciflora Naudin, Ann. Sci. Bot. Ser. 3, 12: 277, t. 12, fig. 3. 1849. —Poteranthera pauciflora (Naudin) Triana, Trans. Linn. Soc. London 28: 33, fig. 17b. 1871. Acisanthera gracilis Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 349. 1915. Herb 0.5–20 cm tall; stem quadrate and ± inflated, mostly glabrous; leaves sessile, fleshy, entire or moderately serrulate, ovate to oblong-ovate, abruptly acute to rounded at the apex, rounded to truncate at the base, 2– 4 mm × 1–3 mm, 1-veined; flowers terminal or solitary in upper leaf axils, 5-merous; hypanthium 1.2–2 mm wide, rarely to moderately glandular-puberulous; calyx lobes ovateoblong to obtuse, 1.4–2 mm long; petals pink, purple or white, 2.5–3.5 mm × 1–3 mm; ovary 2-locular. Wet savannas near Mauritia palm swamps, 50–600 m; Bolívar (Caicara, Canaima, La Vergareña, Uaipán-tepui, Urimán), Amazonas (Puerto Ayacucho, Río Orinoco, Sabana Huachapana). Guárico; Belize, Nicaragua, Jamaica, Colombia. ◆Fig. 189.

Acisanthera limnobios (DC.) Triana, Trans. Linn. Soc. London 28: 33. 1871. —Microlicia limnobios DC., Prodr. 3: 117. 1828. —Noterophila limnobios (DC.) Mart., Nov. Gen. Sp. Pl. 3: 112, t. 254, fig. 3. 1831. Miocarpus paludosus Naudin, Ann, Sci. Nat. Bot. Ser. 3, 12: 281, t. 13, fig. 1. 1849. Acisanthera pellucida Wright ex Griseb., Cat. Pl. Cub. 104. 1866. Herb 0.5–25 cm tall; stem inflated at the base; stem, branchlets, young leaves, and hypanthium rarely to moderately glandularpuberulous; leaves sessile, subfleshy, broadly ovate to elliptic-ovate, abruptly acute to obtuse at the apex, rounded to cordate at the base, broadly serrulate, 5–10 × 3–8 mm, slightly 3–7-veined; flowers solitary in upper leaf axils, 4- or 5- merous; hypanthium 1.2–2 mm wide, rarely to moderately glandularpuberulous; calyx lobes oblong-lanceolate and broadly acute to obtuse; petals ranging from pink to white, ending with a caducous, glandular setula; ovary 2-locular. Lowland savannas, 50–300 m; Bolívar (Ciudad Piar, La Vergareña, Serranía Carichana). Apure, Guárico; Mexico, Central America, West Indies, Brazil. ◆Fig. 187.

Acisanthera hedyotidea (C. Presl) Triana, Trans. Linn. Soc. London 28: 33. 1871. —Dicrananthera hedyotidea C. Presl, Symb. Bot. 1: 76, fig. 50. 1832. Dicrananthera salzmannii Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: 282, t. 13, fig. 1. 1849. —Acisanthera salzmannii (Naudin) Cogn. in Mart., Fl. Bras. 14(3): 209. 1885.

Acisanthera nana Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 349. 1915. Herb 0.5–15 cm tall; stems quadrangular, winged and thickened at the base; young shoots, young leaves, and hypanthium rarely to moderately glandular-puberulous; leaves sessile, fleshy, entire to obscurely ciliate-serrulate, largely ovate to suborbiculate,

Acisanthera 279

broadly acute to rounded at the base, 2–5 × 2–4 mm, inconspicuously 3–5-veined; flowers either on terminal branches or solitary in upper leaf axils, 5-merous; hypanthium 1.2– 2 mm wide, rarely to moderately glandularpuberulous; calyx lobes narrowly ovate-oblong and obtuse, 1.3–2 mm wide; petals lilac, pink, or white, 2.2–3.6 × 1.2–2.7 mm; ovary 2-locular. Moist savannas on granitic outcrops or rocky savanna slopes, 50–800 m; Bolivar (Caicara, Cerro El Médano, Santa Elena de Uairén), Amazonas (Puerto Ayacucho, Río Orinoco, Santa Bárbara). Guárico; Honduras, Colombia, Guyana, Brazil. Acisanthera quadrata Pers., Syn. Pl. 1: 477. 1805. Rhexia acisanthera L., Syst. Nat. ed. 10, 2: 998. 1759. Acisanthera quadrata Juss. ex Poir. in Lam., Encycl. suppl. 1: 111. 1810. Uranthera dicranophora Naudin, Ann. Sci. Nat. Ser. 3, 12: 283. 1849. Acisanthera adscendens C. Wright in Sauvalle, Anales Acad. Ci. Méd. Habana 52. 1868. —Tibouchina adscendens (C. Wright) M. Gómez, Anal. Hist. Nat. Madrid 23: 67. 1894. Herb; stem quadrangular and winged; stems, lower surface of leaves, and hypanthium moderately glandular-puberulous; leaves pedunculate and membranous, ovate to elliptic, acute at the apex, acute to rounded at the base, denticulate, 0.5–1 × 0.15–0.2 mm, 3-veined; flowers solitary in the upper leaf axils, 5-merous; hypanthium 2–2.5 × 1.5 mm; calyx lobes 2–2.5 mm long, oblong-lanceolate; petals pink or purple; ovary 3-locular. Savannas near granitic outcrops, 50–300 m; Amazonas (km 154 on road between Caicara and Puerto Ayacucho). Apure, Cojedes, Guárico, Táchira, Zulia; Mexico, Belize, Honduras, Nicaragua, Costa Rica, Panama, French Caribbean islands, Colombia. Acisanthera uniflora (Vahl) Gleason, Phytologia 3: 346. 1950. —Rhexia uniflora Vahl, Symb. Bot. 2: 48. 1791. Rhexia recurva Rich., Actes Soc. Hist. Nat. Paris 1: 108. 1792. —Microlicia recurva (Rich.) DC., Prodr. 3: 118. 1828. —Acisanthera recurva (Rich.) Griseb., Fl. Brit. W. I. 269. 1860. Comolia anomala Pitt., Bol. Soc. Venez.

Ci. Nat. 6: 196. 1940. Herb 15–50 cm tall; stems quadrangular, slightly winged; hypanthium and lower surface of leaf moderately puberulous; leaf blades broadly ovate to elliptic, broadly acute at the apex, acute to rounded at the base, ciliolate-denticulate, 5–15 × 3–10 mm, 3veined; flowers mostly solitary in the upper leaf axils, 5-merous; hypanthium 2–2.5 mm long; calyx lobes oblong-lanceolate, 2–2.5 mm long; petals pink, 5–6 × 3–4 mm, nonciliate; ovary 3-locular. Moist savannas, 50–500 m; Bolívar (Altiplanicie de Nuria, Camarata, Ciudad Piar, La Vergareña, Urimán), Amazonas (San Fernando de Atabapo). Anzoátegui, Guárico, Monagas, Táchira; Costa Rica, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 188.

Fig. 186. Acisanthera bivalvis

Fig. 187. Acisanthera limnobios

280

M ELASTOMATACEAE

Fig. 188. Acisanthera uniflora

Fig. 189. Acisanthera crassipes

Fig. 190. Acisanthera hedyotidea

Adelobotrys 281

4. ADELOBOTRYS DC., Prodr. 3: 127. 1828. Davya DC., Prodr. 3: 105. 1828. Xeracina Raf., Sylva Tellur. 98. 1838. Sarmentaria Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 140. 1852. Marshallfieldia J.F. Macbr., Publ. Field Columbian Mus., Bot. Ser. 4: 176. 1929. by Navina G. Luckana and Paul E. Berry Shrubs, vines, or hemiepiphytes adhering to tree trunks by adventitious roots. Leaves opposite, isomorphic. Inflorescences terminal or axillary panicles or umbels. Flowers 5-merous. Hypanthium usually terete; calyx limb expanded, usually with indistinct oblate lobes, externally with ± developed short teeth; petals usually pink, obovate and rounded at the apex, eciliate. Stamens 10, glabrous, isomorphic to anisomorphic; anthers linear-subulate, often arcuate, with a small usually dorsally inclined pore; connective not prolonged beyond the thecae but variously modified dorsally into simple and/or bifid spurs and appendages. Ovary superior, cylindric to ovoid, 5-locular; style glabrous and declinate; stigma truncate or punctiform. Capsule 2–5-locular. Seeds numerous, narrowly linear-cuneate, sometimes winged. Southern Mexico to Panama, Jamaica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 25 species, 12 in Venezuela, all in the flora area. Key to the Species of Adelobotrys 1. 1. 2(1).

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

5. 6(5). 6.

7(3).

Leaves 3-veined, 0.6–1 cm wide ................................................................ 2 Leaves 5–7-veined, 2–9 cm wide ................................................................ 3 Leaves linear-lanceolate, 4–6.5 cm long, slightly pubescent, acuminate at the apex, rounded to subcordate at the base; petioles 1–2 mm long ............................................................................................... A. linearifolia Leaves narrowly oblong-elliptic, 2.5–5 cm long, densely ferruginous, acute at both ends; petioles 3–6 mm long ............................ A. stenophylla Shrubs or small trees; ovary 3-locular ...................................................... 4 Climbers or hemiepiphytes; ovary 5-locular ............................................. 7 Leaves scabrous-strigose on upper surface, 3–6 × 2–4 cm, petioles 3–9 mm long; inflorescence subsessile, umbellate .................................... A. duidae Leaves glabrous or soon glabrescent on the upper surface, larger than above; inflorescence short to elongate ................................................... 5 Leaves narrowly elliptic, 10–28 × 4–9 cm, densely ferruginous when young, malpighiaceous trichomes persisting on lower leaf veins ..................................................................................................... A. barbata Leaves elliptic to ovate, smaller than above, glabrous on lower surface of leaf or with simple trichomes ................................................................ 6 Lower surface of leaf persistently finely strigose; peduncle < 1 cm long; calyx lobes 2.5–3 mm long ........................................................ A. fruticosa Lower surface of leaf glabrescent; inflorescence rachis 7–15 cm long with several clusters of subsessile flowers; calyx lobes 1–1.5 mm long ....................................................................................................... A. saxosa Leaf blades 10–20 cm long, densely fine-ciliolate on the margin ................................................................................................... A. permixta

282

M ELASTOMATACEAE

7.

Leaf blades < 10 cm long; margins variable, from almost lacking in hairs to prominently and persistently hairy .................................................. 8 8(7). External calyx teeth projecting 1–2 mm .................................... A. spruceana 8. External calyx teeth projecting < 1 mm .................................................... 9 9(8). Lower surface of leaves prominently ferruginous-strigulose, tardily becoming glabrescent .............................................................. A. rotundifolia 9. Lower surface of leaves initially sparsely ferruginous-strigulose, but soon glabrescent ........................................................................................... 10 10(9). Ascending anther connective appendage blunt or slightly emarginate .................................................................................................. A. monticola 10. Ascending anther connective appendage 1/3–1/2 caudate-bifid ................ 11 11(10). Leaf blades ciliate with robust, simple hairs as well as brownish bifid hairs; teeth of the calyx projecting 0.5–0.9 mm beyond the limb .......... A. ciliata 11. Leaf blades moderately and deciduously fine-ciliolate; teeth of the calyx obsolete or smaller than above ............................................. A. adscendens Adelobotrys adscendens (Sw.) Triana, Trans. Linn. Soc. London 28: t. 5, fig. 56. 1871. —Melastoma adscendens Sw., Fl. Ind. Occid. 2: 772. 1798. —Davya adscendens (Sw.) Griseb., Fl. Brit. W. I. 265. 1860. Davya guianensis DC., Prodr. 3: 105. 1828. —Adelobotrys guianensis (DC.) Gleason, Brittonia 1: 141. 1932. Sarmentaria decora Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 141. 1852. Woody vine to 8 m tall; leaves 8–13 × 5–9 cm, ovate to oblong-ovate, apex acute to short-acuminate, base rounded to cordate; petiole 2–4 cm long. Lower montane forests, 500–1500 m; Bolívar (Amaruay-tepui, Gran Sabana, Río Aparamán). Apure, Mérida, Zulia; Mexico, Central America, Jamaica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 193. Adelobotrys barbata Triana, Trans. Linn. Soc. London 28: 68. 1871. —Boyuyo. Meriania ferruginea Suess., Feddes Repert. Spec. Nov. Regni Veg. 42: 46. 1937. Shrub 1.5–5 m tall; leaves narrowly elliptic, acute at the apex, broadly acute to obtuse at the base, coriaceous, 10–26 × 4–9 cm, young growth densely ferruginous, persistent on undersides along the veins; petioles 1–4 cm long; inflorescence with peduncles 1–2 cm long, pedicels 2–5 mm long; petals white or pink. Margins of seasonally flooded forests, openings along disturbed forests, 50–200 m; Amazonas (Cano Yagua, Maroa, Río Autana, Río Baría, Río Casiquiare, Río Guainía, San Carlos de Río Negro). Colombia, Brazil. ◆Fig. 196.

Adelobotrys ciliata (Naudin) Triana, Trans. Linn. Soc. London 28: 68. 1871. —Davya ciliata Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 137. 1852. Climbing vine; leaves oblong-ovate, subcoriaceous, entire to hardly undulate-serrulate, gradually acuminate at the apex, obtuse at the base, 6–15 × 4–8 cm, ciliolate with both bifid and robust simple hairs; petioles 2–5.5 cm long. Lower montane forests, 500–1500 m; Bolívar (slopes of Ptari-tepui, headwaters of Río Chicanán, Río Tírica). Guyana?, Suriname, French Guiana. ◆Fig. 194. Adelobotrys duidae (Gleason) Wurdack, Mem. New York Bot. Gard. 10(1): 108. 1958. —Meriania duidae Gleason, Bull. Torrey Bot. Club 58: 419. 1931. Shrub 0.2–1.2 m tall, mostly densely appressed-villose with fine rust-colored trichomes; leaves ovate-elliptic to oblong-elliptic, broadly acute at the apex, rounded at the base, stiffly coriaceous, appressed-ciliate, 3– 6 × 2–4 cm; petiole 2–9 mm long; inflorescence umbellate, 4–15-flowered, peduncle 1.5–4 cm long, pedicels 4–6 mm long; petals pink. Tepui meadows, edge of gallery forests, 1700–2600 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 191. Adelobotrys fruticosa Wurdack, Mem. New York Bot. Gard. 10(1): 108. 1958. Shrub 0.3–1.5 m tall; leaves elliptic to oblong-elliptic, ± acute at the apex, obtuse at the base, coriaceous, densely fine-ciliate with appressed trichomes, 4–11 × 2–4.5 cm; peti-

Adelobotrys 283

oles 3–10 mm long; inflorescence sessile to pedunculate. Seasonally flooded sandy river margins, 100–200 m; Amazonas (Caño Pimichín, Caño San Miguel, Río Atacavi, Río Pasimoni, Yapacana Sabana at the base of Cerro Yapacana). Endemic. ◆Fig. 192.

Amazonas (Cerro Aracamuni, Maroa, Río Casiquiare, San Carlos de Río Negro). Colombia, Peru, Brazil, Bolivia.

Adelobotrys linearifolia L. Uribe, Caldasia 9: 295. 1966. Vine ca. 1 m high; leaves subsessile, linear-lanceolate, 40–65 × 7–9 mm; flowers white. Seasonally flooded evergreen lowland forests, lower montane forests, 100–600 m; Bolívar (Cerro Camarón), Amazonas (Río Baría, Río Yatúa, Serranía Batata in CuaoSipapo Massif). Colombia (Amazonas, Caquetá, Vaupés), Brazil (Amazonas). Adelobotrys monticola Gleason, Recueil Trav. Bot. Néerl. 32: 207. 1935. Adelobotrys monticola subsp. occidentalis Wurdack, Mem. New York Bot. Gard. 18(2): 287. 1969. Vine or hemiepiphyte on tree trunks to 5 m tall; leaves broadly elliptic, shortly acuminate to acute at the apex, obtuse to rounded at the base, subcoriaceous, the margins appressed-ciliate and inconspicously denticulate, 3–7 × 2.5–5 cm; inflorescence 3–7 cm long, few-flowered. Montane forests, 1000– 2100 m; Bolívar (Cerro Jaua, Cerro Venamo, Gran Sabana, Macizo del Chimantá, Ptaritepui), Amazonas (slopes of Cerro Duida, Cerro Marahuaka, Cerro Sipapo, Sierra Parima). Suriname, Brazil. ◆Fig. 195. Adelobotrys permixta Wurdack, Phytologia 20: 372. 1970. Liana; leaves coriaceous, broadly ovateelliptic to elliptic, shortly-acuminate at the apex, obtuse to truncate at the base, entire to obscurely undulate-serrulate, margin densely fine-ciliate, 10–20 × 6–13 cm. Lower montane forests, 100–500 m; Bolivar (La Escalera). Guyana. Adelobotrys rotundifolia Triana, Trans. Linn. Soc. London 28: 68. 1871. Liana; leaves broadly elliptic, abruptly shortly-acuminate at the apex, rounded at the base, thinly coriaceous densely fine-ciliate on the margin, the upper surface glabrous, the lower surface moderately finestrigulose. Evergreen lowland to lower montane forests, 100–900 m; Bolívar (El Paují),

Fig. 191. Adelobotrys duidae

284

M ELASTOMATACEAE

Fig. 192. Adelobotrys fruticosa

Fig. 193. Adelobotrys adscendens

Adelobotrys 285

Fig. 194. Adelobotrys ciliata

Fig. 195. Adelobotrys monticola

286

M ELASTOMATACEAE

Fig. 196. Adelobotrys barbata

Adelobotrys saxosa Wurdack, Mem. New York Bot. Gard.10(5): 146. 1964. Shrub or small tree 1–5 m tall; leaves elliptic, obtuse and mucronate at the apex, obtuse at the base, coriaceous, 5–15 × 3–7 cm, becoming glabrescent except for a dense line of rusty trichomes along the margin; petioles 1.5–3 cm long; inflorescence with peduncles 7–14 cm long, flowers in groups of 3–8, alone at the tip of the rachis or in several groups, pedicels 2–5 mm long. Low open forest on granitic or quartzitic outcrops, 100–600 m; Amazonas (slopes of Cerro Aracamuni along Río Siapa, Piedra Arauicaua). Endemic. Adelobotrys spruceana Cogn. in Mart., Fl. Bras. 14(4): 18, t. 5, fig. 1. 1888. Climber on tree trunks to 5–6 m high; leaves elliptic-ovate to elliptic, acute to subacuminate at the apex, obtuse to rounded-truncate at the base, the upper leaf

surface lacking simple hairs (but the margins ciliate with simple hairs), ca 3.5 × 2–4.5 cm; inflorescence usually 1–2 cm long, compact and subumbellate. Riparian forests, near sea level to 500 m; Delta Amacuro (San Victor on Serranía de Imataca), Bolívar (Cerro Ichún, upper Río Caura), Amazonas (La Esmeralda, Río Baría, Río Pasimoni, Río Yatúa). Colombia, French Guiana, Ecuador, Peru, Brazil. Adelobotrys stenophylla Wurdack, Phytologia 19: 192. 1969. Small shrub, most of the plant densely covered with rusty-colored, appressed trichomes; leaves narrowly oblong-elliptic, acute at both ends, 2.5–5 × 0.5–1 cm, 3veined, ciliate-margined; petioles 3–6 mm long; inflorecence 3–5-flowered. Tepui meadows, ca. 1000 m; Amazonas (Cerro Duida). Endemic.

Bellucia 287

5. BELLUCIA Raf., Sylva Tellur. 92. 1838, nom. cons. Apatitia Desv. ex Ham., Prodr. Pl. Ind. Occid. 42. 1825. Axinanthera H. Karst., Linnaea 30: 157. 1859. by Paul E. Berry and Susanne S. Renner Small or medium-sized trees to 25 m tall, or shrubs. Leaves opposite, simple, petiolate, large, coriaceous and glossy green when fresh, the lower surface glaucous, 3- or 5-pliveined; margin entire or serrulate. Inflorescences 2- or 3-branched cymes or congested clusters of few flowers, rarely a single flower in the leaf axils, on branches below the leaves, or on the trunk. Flowers bisexual, 5–8-merous. Hypanthium subglobose; calyx a calyptra, splitting open in an irregular semicircle and drying as a hyaline membrane, often persistent on the limb after anthesis, or else dehiscing regularly into triangular lobes, the lobes valvate in bud, erect and persistent or truncate; petals obovate, clawed, apically truncate or emarginate, fleshy, with 2 longitudinal and 1 tranverse callus ridge on adaxial surface, spreading at anthesis, white or sometimes pink flushed outside. Stamens twice as many as the petals, isomorphic, glabrous, laterally compressed; anther and filament ± of equal length, anthers thick, with 2 slightly introrse pores. Ovary completely inferior, 10– 14(15)-locular; style glabrous, thick, terete, longer than the stamens; stigma capitate and 10–16-lobed. Fruit a semiglobose, greenish yellow berry, many-seeded. Seeds minute, 0.5–1 mm long, ± ovoid. Southern Mexico, Central America, tropical South America to northern Bolivia; 7 species, 3 in Venezuela, all in the flora area. Species of Bellucia have edible fruits. This treatment was extracted in large part from the generic revision of S. S. Renner (Mem. New York Bot. Gard. 50: 1–112. 1989). Key to the Species of Bellucia 1.

1. 2(1). 2.

Calyx a calyptra dehiscing in an irregular semicircle, drying as a hyaline membrane, usually persistent on the limb; flowers mostly 8-merous ......................................................................................... B. grossularioides Calyx dehiscing regularly into persistent triangular lobes, or calyx truncate; flowers 5- or 6-merous ................................................................... 2 Calyx truncate, ca. 2 mm long, buds completely smooth; petals ca. 12 mm long ................................................................................................ B. huberi Calyx dehiscing regularly into persistent, thick triangular lobes 5–8 mm long, lines of dehiscence visible in buds; petals 15–20 mm long ................................................................................................ B. pentamera

Bellucia grossularioides (L.) Triana, Trans. Linn. Soc. London 28: 141. 1871. —Melastoma grossularioides L., Sp. Pl. 390. 1753. —Cara-re-yek (Pemón), Guayabo de danta, Pomarrosa. Blakea quinquenervia Aubl., Hist. Pl. Guiane 525, pl. 210. 1775. —Bellucia quinquenervia (Aubl.) H. Karst., Linnaea 30: 159. 1859. Apatitia blakeoides Desv. ex Ham., Prodr. Pl. Ind. Occid. 42. 1825.

Bellucia brasiliensis Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 104. 1851. Bellucia hostmannii Naudin, Ann. Sci. Nat. Bot. sér 3, 16: 104. 1851. Bellucia circumscissa Spruce ex Cogn. in Mart., Fl. Bras. 14(4): 515, pl. 108. 1888. Tree 3–25(–30) m tall. Nonflooded forests, river banks, open, marshy areas, 50–800 m; Bolívar (Gran Sabana, lower Rio Caura, Rio Parguaza), Amazonas (widespread along Rio Negro and Río Orinoco, base of Sierra de la

288

M ELASTOMATACEAE

Neblina). Apure, Barinas, Táchira, Trujillo, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 197. The red inner bark is rubbed on paddles, wooden tools, ropes, and drinking vessels to give a waterproof surface and a varnished appearance.

Bellucia huberi (Wurdack) S.S. Renner, Mem. New York Bot. Gard. 50: 28. 1989. —Loreya huberi Wurdack, Phytologia 50: 306. 1982. Shrub or treelet 1–5 m tall. White-sand savannas, dwarf forests on sandstone, 100– 1000 m; Amazonas (Maroa, Rio Coro Coro near Yutajé, west of San Juan de Manapiare). Endemic. ◆Fig. 198.

Fig. 197. Bellucia grossularioides

Bellucia 289

Fig. 198. Bellucia huberi

290

M ELASTOMATACEAE

Bellucia pentamera Naudin, Ann. Sci. Nat. Bot. sér 3, 16: 105. 1851. Axinanthera macrophylla H. Karst., Linnaea 30: 157. 1859. Bellucia aricuaizensium Pittier, Explor. Cuenca de Maracaibo [reprinted from Bol. Comerc. Industr. Venez. Año IV. 1923]: 40. 1923. Shrub or tree 3–15 m tall; twigs with pronounced nodes and rows of pustule-like lenticels. Nonflooded forests, riparian forests,

near sea level to 300 m; Delta Amacuro (Río Amacuro), Bolívar (Las Trincheras), Amazonas (Mavaca, Río Casiquiare, Río Cunucunuma). Apure, Barinas, Carabobo, Cojedes, Mérida, Monagas, Trujillo, Yaracuy, Zulia; Mexico, Central America, Guadeloupe, Colombia, Ecuador, Peru, Brazil, Bolivia, naturalized in Africa. Several collections have been made that appear to be natural hybrids between Bellucia pentamera and B. grossularioides.

6. BERTOLONIA Raddi, Atti Soc. Ital. Sci. 18: 384, fig. 3. 1820, nom. cons. by Paul E. Berry Erect or prostrate (rarely creeping) herbs or subshrubs, terrestrial, rupicolous, or epiphytic. Leaves opposite and decussate, alternate by suppression of one member of each pair in B. venezuelensis, sometimes appearing rosulate, with 3, 5, 7, or 9 principal veins, petiolate. Inflorescences terminal, of dichasia, thyrsoid dichasia, scorpioid cymes, or umbels of scorpioid cymes. Flowers 5-merous. Hypanthium cupuliform or 10-costate; sepals persistent; petals white to pink or lilac, elliptic, obovate, or suborbicular. Stamens 10, (sub)isomorphic, glabrous; thecae oblong-lanceolate, nonrostrate, the connective slightly prolonged and nonappendaged or with a dorsal appendage at the base. Ovary 3-locular; style glabrous or with glandular trichomes at the base; stigma punctiform. Fruit a triquetrous capsule with 3 small wings. Seeds numerous, obovoid to clavoid, rarely oblongoid. Southern Venezuela, southeastern Brazil; 18 species, 2 in Venezuela, both in the flora area. Key to the Species of Bertolonia 1. 1.

Leaves ovate to oblong-elliptic, apex acuminate, base acute to truncate ............................................................................................ B. venezuelensis Leaves suborbicular or broadly ovate-elliptic, apex rouded, base slightly cordate .............................................................................................. B. sp. A

Bertolonia venezuelensis Wurdack, Mem. New York Bot. Gard. 10(1): 111, fig. 12b– f. 1958. Prostrate to creeping herb; stems and petioles with shaggy rufous pubescence; one of each leaf pair suppressed, blade ovate to oblong-elliptic, 6–18 × 3–6 cm; peduncle 10– 15 cm long, the umbellate branches 2–3 cm long; petals white-pink; stamens yellow. Rocky escarpments, moist mossy ground in cloud forests, 1500–1700 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 200.

Bertolonia sp. A Creeping herb; stems and petioles densely pubescent; leaves alternate by suppression of one of the leaf pairs in the lower portion of the stem, opposite in the distal (aerial) part of stem; leaf blade suborbicular or broadly ovate-elliptic, 4–8 × 2.5–6 cm; peduncle 7–15 cm long; buds white (young petals pink), nodding. Mossy, shady ground along stream in cloud forests, 1400–1500 m; Amazonas (Sierra de la Neblina ca. 5 km north-northeast of Pico Phelps). Endemic. ◆Fig. 199. This species is in press by F. Michelangeli under the name of Bertolonia repens Michelangeli.

Bertolonia 291

Fig. 199. Bertolonia sp. A

Fig. 200. Bertolonia venezuelensis

292

M ELASTOMATACEAE

7. BLAKEA P. Browne, Civ. Nat. Hist. Jamaica 323. 1756. by Paul E. Berry Trees, shrubs, lianas, or epiphytes. Leaves opposite, isomorphic, or rarely pseudoalternate, entire, and petiolate (in Venezuelan species). Flowers 6-merous, axillary and pedicellate in the upper leaf axils, 4-bracteate (the bracts free or ± united in pairs). Hypanthium terete; calyx 6-lobed to truncate, persistent; petals white to pink, usually diminutively ciliolate, glabrous on both surfaces but often verruculose when dry. Stamens 12, glabrous; anthers 1/2–2/3 as wide as long, laterally compressed and coherent in a ring, very minutely biporate at the apex; connective basally enlarged, ventrally nonappendaged, dorsally with a divergent or descending tooth. Ovary (4)6-locular, completely inferior, usually with a stylar cone; style glabrous; stigma truncate (in Venezuelan species). Fruit a berry. Seeds numerous, ovoid to pyramidal. Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia; 85 species, 7 in Venezuela, 1 of these in the flora area.

Fig. 201. Blakea rosea

Blakea rosea (Ruiz & Pav.) D. Don, Mem. Wern. Nat. Hist. Soc. 4: 325. 1823. —Valdesia rosea Ruiz & Pav., Fl. Peruv. 4: pl. 408. 1802. Blakea caudata Triana, Trans. Linn. Soc. London 28: 148. 1871. Blakea calycanthus Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 13: 463. 1937. Liana or epiphytic shrub to 8 m tall. Leaf blades oblong-elliptic to obovate-elliptic, abruptly acuminate, broadly acute at the base, thin-coriaceous, 6–13 × 3–7 cm, shortly 3-pliveined; petioles 1–2 cm long; petals white, 10–14 × 6–10 mm; ovary 6-locular. Evergreen lowland and lower montane forests, 100–700 m; Bolívar (Río Caura), Amazonas (Río Yatúa, slopes of Sierra de la Neblina). Apure, Táchira, Zulia; Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 201.

Centronia 293

8. CENTRONIA D. Don, Mem. Wern. Nat. Hist. Soc. 4: 314. 1823. by Paul E. Berry Trees and shrubs. Leaves opposite, isomorphic. Flowers 5–7-merous, in terminal panicles. Calyx calyptrate, dehiscent at anthesis; petals white, pink, purplish, or orange in some non-Venezuelan species, usually glabrous and eciliate. Stamens subisomorphic or dimorphic, glabrous; anthers subulate, 1-porate, connective not or only slightly prolonged, with a prominent basal spur and often with a short, ascending dorsal appendage. Ovary free, 5–7-locular, the apex truncate or prolonged. Capsule 5–7-valved. Seeds numerous, narrowly oblong-pyramidal. Costa Rica, Colombia, Ecuador, Peru, Bolivia; 18 species, 3 in Venezuela, 1 of these in the flora area. Centronia is doubtfully distinct from Meriania.

Fig. 202. Centronia neblinae

294

M ELASTOMATACEAE

Centronia neblinae Wurdack, Mem. New York Bot. Gard. 10(5): 154. 1964. Tree 6–12 m tall; stems, leaves, and young inflorescences densely appressed-scurfy with minute reddish hairs, soon glabrescent; leaf blades obovate-oblong to oblong, shortly and abruptly acuminate at the apex, broadly acute at the base, coriaceous and entire, 15– 40 × 6–16 cm, 3-veined, with the primary

veins submarginal, the secondary veins 1–2 cm apart; petioles 3.5–8 cm long; panicle of dichasial cymes 10–15 cm long, lateral branches 0.5–2 cm long; petals white, 17–20 × 11–12 mm, obovate and obtuse at the apex; ovary 5-locular. Montane forests, 1100–1500 m; Amazonas (Sierra de la Neblina along Cañon Grande). Endemic. ◆Fig. 202.

9. CHAETOLEPIS (DC.) Miq., Comm. Phytogr. 2: 72. 1840. —Osbeckia sect. Chaetolepis DC., Prodr. 3: 140. 1828. Haplodesmium Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 150. 1850. Trimeranthus H. Karst., Linnaea 30: 159. 1859. Chaetogastra sect. Adesmogastra Griseb., Cat. Pl. Cub. 103. 1866. by Frank Almeda Shrubs or subshrubs. Leaves small, isomorphic. Flowers 4-merous, mostly solitary or in simple cymes at the branchlet ends. Hypanthium terete; calyx lobes triangular to lanceolate, persistent; petals ovate to elliptic, or obovate, acute, usually eciliate, often with a terminal setula. Stamens 8, essentially isomorphic or nearly so, glabrous; thecae linear-oblong, minutely 1-pored; connective prolonged below the thecae to the indistinct articulation with the filament but not appendaged. Ovary 4locular, free, setulose at the apex; style glabrous; stigma not expanded. Capsule 4valved. Seeds numerous, ovoid-cochleate, the surface appearing smooth or finely and diminutively muriculate. Costa Rica, Cuba, Colombia, Venezuela, Guyana; 12 or 13 species, 6 in Venezuela, 2 of these in the flora area. If additional study corroborates the recent transfer of Nerophila gentianoides Naudin of lowland tropical west Africa to Chaetolepis, this will be the only genus in the family with a trans-Atlantic distribution. Key to the Species of Chaetolepis 1. 1.

Upper internodes moderately to copiously beset with spreading, prevailingly eglandular trichomes to 0.25 mm long ......................... C. anisandra Upper internodes glabrous or very sparsely and caducously beset with spreading (sometimes gland-tipped) trichomes 0.5–1 mm long .................................................................................................. C. phelpsiae

Chaetolepis anisandra Naudin, Ann. Sci. Nat. Bot. sér 3, 14: 141. 1850. Chaetolepis citrina Gleason, Brittonia 3: 174. 1939. Diffuse shrub 20–40 cm tall with prostrate branches; lower surface of primary leaves moderately beset with smooth (rarely glandular) trichomes; flowers solitary or ternate at branchlet ends, petals and anthers yellow. Moist upper-escarpment faces of tepuis, 2100–2600 m; Bolívar (Auyán-tepui,

Roraima-tepui), Amazonas (Cerro Yutajé). Colombia, Guyana. ◆Fig. 203.

Chaetolepis phelpsiae Gleason, Brittonia 7: 82. 1950. Subshrub 10–30 cm tall, the cauline internodes quadrate; flowers solitary on the branchlet ends; petals and anthers yellow. Venezuela; 2 subspecies, both endemic in the flora area.

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Fig. 203. Chaetolepis anisandra

Fig. 204. Chaetolepis phelpsiae subsp. chimantensis

Key to the Subspecies of C. phelpsiae 1. Leaf blades 5-veined, the abaxial midvein glabrous or very sparingly beset with glandular trichomes < 0.5 mm long; hypanthium basally glabrous, upper part strigillose ............... subsp. chimantensis 1. Leaf blades 3-veined, the abaxial midvein sparsely but consistently beset with eglandular trichomes 0.75–1 mm long; hypanthium moderately strigillose ........ .................................. subsp. phelpsiae

C. phelpsiae subsp. chimantensis Wurdack, Mem. New York Bot. Gard. 10(5): 138. 1964. Slopes and talus forests, vicinity of waterfalls, 1900–2000 m; Bolívar (Macizo del Chimantá [Sarvén-tepui]). Endemic. ◆Fig. 204. C. phelpsiae subsp. phelpsiae On rocks by waterfalls, vertical sandstone cliffs, 1400–1900 m; Amazonas (Cerro Yaví). Endemic.

10. CLIDEMIA D. Don, Mem. Wern. Nat. Hist. Soc. 4: 306. 1823. by Bruce K. Holst Erect or rarely scandent or radicant shrubs. Leaves opposite, rarely pseudoalternate (by abortion of one member of the pair), rarely with ant domatia. Inflorescences lateral or pseudolateral cymes in the upper leaf axils or at branchlet nodes below the leaves. Flowers 4- or 5(–8)-merous, diplostemonous (pleiostemonous in C. bullosa). Hypanthium terete; calyx usually lobed and persistent, the external teeth developed and often projecting; petals white to pink, usually glabrous, rounded at the apex. Stamens glabrous, isomorphic or slightly dimorphic in size; anthers usu-

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ally slender and 1-pored; connective not or slightly prolonged, usually exappendiculate or with a dorso-basal tooth. Ovary (2)3–5(–10)-locular, partly to completely inferior; style usually glabrous; stigma not or slightly expanded. Fruit a manyseeded berry. Seeds ovoid to pyramidal, smooth to granulate. Neotropics; ca. 175 species, about 60 in Venezuela, 47 of these in the flora area. Key to the Species of Clidemia 1. 1. 2(1). 2. 3(2).

Flowers predominantly 5–8-merous .......................................................... 2 Flowers predominantly 4-merous ............................................................ 24 Ant domatia present at petiole apices or at stem nodes ........................... 3 Ant domatia absent .................................................................................... 5 Ant domatia geminate on the stem nodes; inflorescence many-flowered, ca. 10 cm long ............................................................................. C. neblinae 3. Ant domatia solitary at petiole apex; inflorescence few-flowered, 0.5–2 cm long ......................................................................................................... 4 4(3). Leaf pairs strongly dimorphic, the smallest usually without ant domatia; flowers (6)7-merous; anthers 2.7 mm long; ovary 3/4-inferior ................ ................................................................................................ C. heptamera 4. Leaf pairs isomorphic, both with ant domatia; flowers 5-merous; anthers 5.6 mm long; ovary 1/10-inferior ................................................ C. juruensis 5(2). Ovary 3-locular ........................................................................................... 6 5. Ovary (4)5(–10)-locular ............................................................................ 13 6(5). External calyx teeth extending 1.5–1.8 mm beyond the internal lobes ................................................................................................................ 7 6. External calyx teeth extending < 0.8 mm beyond the internal lobes ...... 9 7(6). Indument of branchlets and inflorescences of all simple hairs; flowers sessile; petals 3.8–4.2 mm long ........................................... C. morichensis 7. Indument of branchlets and inflorescences in part of stellate hairs; flowers short-pedicellate; petals 2.2–2.7 mm long ...................................... 8 8(7). Leaf margins entire; vestiture moderately short-setose with gland-tipped hairs underlain with stellate hairs; hypanthium 3–3.5 mm long .................................................................................................. C. simpsonii 8. Leaf margins ciliolate-serrulate; vestiture of dense stellate hairs with sparse gland-tipped hairs; hypanthium 1.6–2 mm long .......... C. stellipilis 9(6). Leaves subalternately 5- or 7-pliveined .................................................. 10 9. Leaves basally veined .............................................................................. 11 10(9). Flowers pedicellate, the pedicels 0.5–2 mm long; inflorescences paniculate; leaf veins densely reticulate on lower surface ............... C. japurensis 10. Flowers sessile; inflorescences pedunculate, bracteate heads; leaf veins laxly reticulate on lower surface .......................................... C. involucrata 11(9). Stems, leaves, hypanthia, and ovary apices moderately stellatepuberulous, without simple hairs ........................................... C. capillipes 11. Stems, leaves, hypanthia, and ovary apices with solely simple hairs ... 12 12(11). Flowers nearly sessile, the pedicels < 0.3 mm long; cymes one per node; bracteoles 4.5–5 mm long; petals glabrous or with a simple subapical seta ........................................................................................... C. buntingii 12. Flowers pedicellate, the pedicels 1–2 mm long; cymes in opposite leaf ax-

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13(5). 13. 14(13). 14. 15(14). 15. 16(15). 16. 17(16). 17. 18(17). 18. 19(16). 19. 20(19). 20. 21(20). 21. 22(21). 22. 23(22). 23. 24(1). 24. 25(24). 25. 26(25). 26. 27(26). 27.

ils of upper nodes; bracteoles 0.3 mm long; petals with an externally setulose tooth 0.7–1 mm long .............................................. C. epibaterium Hairs on branchlets flattened, with erose to fimbriate borders; flowers in axillary, sessile heads......................................................... C. conglomerata Hairs on branchlets terete or stellate; flowers in cymes or panicles, usually pedicellate ..................................................................................... 14 Flowers with 3 times or more stamens as petals (5-merous with 15–21 stamens) ............................................................................................ C. bullosa Flowers with 2 times as many stamens as petals ................................... 15 Flowers predominantly 7- or 8-merous; ovary 7–10-locular............ C. octona Flowers predominantly 5(6)-merous; ovary 5(6)-locular ........................ 16 Anther pores dorsally inclined ................................................................ 17 Anther pores ventrally inclined ............................................................... 19 Upper surface of leaf blades glabrous; torus with fine hairs ...... C. acurensis Upper surface of leaf blades pubescent; torus with 10 fimbriate or ciliate scales .................................................................................................... 18 Leaf blades subalternately pliveined; stamen connective unappendaged ..................................................................................................... C. dentata Leaf blades basally veined; stamen connective with a dorso-basal tooth ......................................................................................................... C. hirta Anther connective basally prolonged 0.7–1.2 mm ................. C. novemnervia Anther connective not prolonged or at most prolonged 0.1–0.2 mm ...... 20 Pubescence of upper surface of leaf blades nearly exclusively of sessilestellate hairs; ovary apex without gland-tipped hairs ............ C. siapensis Pubescence of upper surface of leaf blades of soft setae; ovary apex glandular-setulose ....................................................................................... 21 Inflorescence usually spicate, occasionally with two suprabasal branches; floral bracts conspicuous, 1–2 mm wide ............................... C. capitellata Inflorescence paniculate, with secondary branches evident above the basal pair; floral bracts inconspicuous, < 1 mm wide ......................... 22 Branchlet hairs 2–3 mm long, gland-tipped, numerous ............. C. urceolata Branchlet hairs 0.4–1 mm long, usually gland-tipped ........................... 23 External calyx teeth extending 0.5–0.8 mm beyond the internal lobes; stellate pubescence on ovary sparse ....................................... C. pustulata External calyx teeth extending 1.7–2.2 mm beyond the internal lobes; stellate pubescence on ovary dense ........................................ C. strigillosa Inflorescences dense, prominently 2-bracteate heads; peduncle 1.5–4 cm long ............................................................................................. C. capitata Inflorescences developed panicles, few-flowered cymes, or glomerate and essentially sessile in the leaf axils; bracts inconspicuous .................. 25 Ovary 4-locular ......................................................................................... 26 Ovary (2)3-locular .................................................................................... 34 Leaves of a pair strongly dimorphic, one strongly reduced (1 mm long) and early deciduous, leaf phyllotaxy appearing alternate ......... C. alternifolia Leaves of a pair similar in size and shape .............................................. 27 Flowers solitary or glomerate in the leaf axils, the inflorescence axis not developed .............................................................................................. 28 Flowers in evident cymes or racemes ...................................................... 30

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28(27). Anthers 2–3.5 mm long; leaf blades basally veined or weakly (< 1 cm) pliveined ......................................................................................... C. rubra 28. Anthers 0.9–1.1 mm long; leaves strongly pliveined, the internal pair of primary veins diverging 2–4 cm above the blade base ....................... 29 29(28). Leaves and stems sparsely strigulose with simple hairs; bracteoles broadly ovate, 1 mm wide ........................................................ C. bernardii 29. Leaves and stems lacking simple hairs; bracteoles subulate, 0.2 mm wide .............................................................................................. C. heteroneura 30(27). Leaf blades pliveined ............................................................... C. aphanantha 30. Leaf blades basally veined ....................................................................... 31 31(30). Ant domatia present on branchlets; petals ca. 1.5 mm long ..... C. tococoidea 31. Ant domatia absent on branchlets; petals 2.1–2.5 mm long ................... 32 32(31). Leaf blades obtuse to truncate at base; inflorescences ca. 1 cm long .............................................................................................. C. microthyrsa 32. Leaf blades emarginate to cordate at base .............................................. 33 33(32). Leaf blades 0.5–1.5 cm cordate at base; inflorescences 3–5 cm long .................................................................................................... C. silvicola 33. Leaf blades emarginate at base; inflorescences to 2.5 cm long ............ C. ulei 34(25). Leaves evidently pliveined ...................................................................... 35 34. Leaves basally veined (rarely pliveined 1 cm or less above blade base) 40 35(34). Hypanthium moderately lax-strigulose .................................................. 36 35. Hypanthium granulose, but not setulose ................................................ 38 36(35). Upper surface of leaf blades with persistent setae ca. 1 mm long .................................................................................................. C. attenuata 36. Upper surface of leaf blades with deciduous setulae ca. 0.2 mm long ... 37 37(36). Leaves (2.5–)5–8(–9) × 1.5–3 cm ......................................... C. marahuacensis 37. Leaves 10–16 × 4–8 cm ............................................................... C. piperifolia 38(35). Young branchlets and primary veins on lower surfaces of leaf blades setulose; petals 1.5 × 0.4 mm .................................................. C. trinitensis 38. Young branchlets and primary veins on lower surfaces of leaf blades not setulose; petals 2–2.7 × 0.8–1 mm ....................................................... 39 39(38). External calyx teeth not extending beyond the internal ones; calyx tube 0.1–0.2 mm long ..................................................................... C. andersonii 39. External calyx teeth extending 0.3–0.7 mm beyond the internal ones; calyx tube ca. 0.4 mm long .............................................................. C. sessiliflora 40(34). Flowers solitary and multifasciculate in the leaf axils ........................... 41 40. Flowers in developed cymes or panicles .................................................. 44 41(40). Leaf blades lacking setae; petals 1–1.2 mm long ..................... C. minutiflora 41. Leaf blades setulose, at least on lower surface; petals 2–2.5 mm long .............................................................................................................. 42 42(41). Petioles usually 0.2–0.4 cm long ....................................................... C. sericea 42. Petioles usually 2–5 cm long .................................................................... 43 43(42). Hypanthium hairs not gland-tipped ................................................. C. debilis 43. Hypanthium hairs gland-tipped, at least in part ..................... C. micrantha 44(40). Leaf blades persistently stellate-puberulent on either upper and/or lower surface .................................................................................................. 45 44. Leaf blades not persistently stellate-puberulent on upper surface ....... 46 45(44). Leaf blades persistently stellate-puberulent on both surfaces; petals 2.2–2.4 mm long; hypanthium ca. 2.5 mm long ...................... C. capillipes

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45.

Leaf blades persistently stellate-puberulent only on lower surface; petals 3.6–4.2 mm long; hypanthium 3–3.5 mm long ....................... C. pycnaster 46(44). Leaf blades narrowly lanceolate-oblong, 0.5–0.9 cm wide, 3-veined ..................................................................................................... C. linearis 46. Leaf blades ovate to oblong-ovate, 4–9 cm wide, 5(7)-veined ................. 47 47(46). Hypanthium moderately stellate-puberulent and sparsely glandularsetulose ......................................................................................... C. duidae 47. Hypanthium glabrous ................................................................. C. tepuiensis Clidemia acurensis Wurdack, Acta Bot. Venez. 2: 376, fig. 5. 1967. Shrub to 2 m tall. Lowland evergreen forests, ca. 100 m; Delta Amacuro (Río Acure). Endemic.

Shrub 1–2.5 m tall. Lowland to upland evergreen forests, 100–1200 m; Bolívar (Aprada-tepui, Gran Sabana, Río Erebato), Amazonas (Cerro Duida). Brazil (Amazonas: base of Serra da Neblina, Roraima).

Clidemia alternifolia Wurdack, Mem. New York Bot. Gard. 10(5): 184. 1964. Shrub to 1.5 m tall. Lowland evergreen forests, 100–200 m; Amazonas (near Maroa, San Carlos de Río Negro, Yavita). Brazil (Amazonas). ◆Fig. 205.

Clidemia bullosa DC., Prodr. 3: 158. 1828. —Clidemia candolleana Cogn. in Mart., Fl. Bras. 14(4): 490. 1888, nom. illeg. Clidemia rariflora Benth., J. Bot. (Hooker) 2: 308. 1840. Clidemia manoacensis Kraenzl., Vierteljahrsschr. Naturf Ges. Zürich 76: 157. 1931. Clidemia novemnervia var. depauperata Pittier, Bol. Soc. Venez. Ci. Nat. 11: 23. 1947. Shrub 1–2(–3) m tall. Lowland evergreen forests, along streams, near sea level to 400 m; Delta Amacuro (Manoa), Bolívar (Icabarú, La Unión, Macarapa, Río Caura), Amazonas (widespread). Portuguesa, Zulia; Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, and Bolivia. ◆Fig. 206.

Clidemia andersonii Wurdack, Mem. New York Bot. Gard. 51: 104. 1989. Shrub ca. 2.5 m tall. Montane forests, 1800–1900 m; Amazonas (Sierra de la Neblina). Endemic. Clidemia aphanantha (Naudin) Sagot, Ann. Sci. Nat. Bot. sér. 6, 15: 327. 1883. —Staphidiastrum aphananthum Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 333. 1852. Shrub to 2 m tall. Lowland evergreen to semideciduous forests, 200–400 m; Amazonas (Río Cunucunuma, Río Sipapo), Bolívar (Sierra de Imataca). Suriname, French Guiana, Brazil (Amazonas). ◆Fig. 209. The specific distinctions between Clidemia aphanantha, C. micrantha, C. microthyrsa, and C. rubra are tenuous.

Clidemia buntingii Wurdack, Phytologia 19: 195. 1969. Scandent shrub to 4 m tall. Montane evergreen forests, 1000–1200 m; Bolívar (La Escalera). Guyana. ◆Fig. 212.

Clidemia attenuata (Naudin) Cogn. in A. DC. & C. DC., Monogr. Phan. 7: 1006. 1891. —Staphidiastrum attenuatum Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 332. 1852. Shrub to 2 m tall. Evergreen lowland to upland shrub forests, 200–800 m; Bolívar (Río Túriba headwaters), Amazonas (Río Parú). Trinidad, Guyana, French Guiana.

Clidemia capillipes (Triana) Cogn. in Mart., Fl. Bras. 14(4): 509, pl. 107, fig. 2. 1888. —Sagraea capillipes Triana, Trans. Linn. Soc. London 28: 138. 1871. Shrub to 1.5 m tall. Lowland evergreen forests, ca. 100 m; Amazonas (Río Yatúa, San Carlos de Río Negro, Santa Lucía, Tamatama). Amazonian Colombia and Brazil. ◆Fig. 216.

Clidemia bernardii Wurdack, Phytologia 19: 196. 1969.

Clidemia capitata Benth., J. Bot. (Hooker) 2: 307. 1840.

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Shrub 1–2(–3) m tall. Disturbed areas in premontane to montane slope and gallery forests, scrub forests, riverbanks, 300–1800 m; Bolívar (Cerro Bolívar, Cerro Guaiquinima, Cerro Jaua, Gran Sabana), Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Sierra de la Neblina). Adjacent Guyana and Brazil (Amazonas, Roraima). ◆Fig. 211. Clidemia capitellata (Bonpl.) D. Don, Mem. Wern. Nat. Hist. Soc. 4: 310. 1823. —Melastoma capitellata Bonpl. in Humb. & Bonpl., Monogr. Melast. 1: 5, t. 3. 1806. Shrub to 2 m tall. Evergreen lowland to montane forests, 100–1200 m. Mexico and the Antilles to Amazonian Brazil and Bolivia; 4 varieties, 3 in Venezuela, all in the flora area. Key to the Varieties of C. capitellata 1. Inflorescences 1–1.5 cm long ..................... ............................................ var. levelii 1. Inflorescences 5–10 cm long ................... 2 2. Hairs of branchlets and inflorescence axes 1–1.5 mm long ........... var. capitellata 2. Hairs of branchlets inflorescence axes 2– 5(–8) mm long ............ var. dependens C. capitellata var. capitellata 100–600 m; Bolívar (locality unknown), Amazonas (Río Baría, Río Cuao, Río Manapiare, San Carlos de Río Negro). Anzoátegui, Barinas, Mérida, Monagas, Portuguesa, Sucre, Táchira, Zulia; southern Mexico and the Guianas to Amazonian Bolivia. C. capitellata var. dependens (D. Don) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4): 484. 1941. —Clidemia dependens D. Don, Mem. Wern. Nat. Hist. Soc. 4: 307. 1863. 100–1200 m; Bolívar (Caicara, Corozal, El Paují, Río Reforma), Amazonas (La Esmeralda, Puerto Ayacucho, Río Casiquiare, Río Matacuni). Apure, Barinas, Carabobo, Guárico, Mérida, Portuguesa, Táchira; southern Mexico and the Guianas to southeastern Brazil. C. capitellata var. levelii Wurdack, Mem. New York Bot. Gard. 10(5): 181. 1964. 100–200 m; Amazonas (Río Ventuari). Endemic.

Clidemia conglomerata DC., Prodr. 3: 156. 1828. —Staphidium conglomeratum (DC.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 321. 1852. Shrub 0.5–2(–4) m tall. Evergreen lowland to lower montane forests, 50–500 (–1000) m; Delta Amacuro (Río Amacuro), Bolívar (La Escalera, Río Aparurén, Río Tírica, Urimán). Trinidad to northeastern Brazil (Amapá). ◆Fig. 218. Clidemia debilis Crueg., Linnaea 20: 104. 1847. Shrub 1–1.5 m tall. Semideciduous forests, 200–800 m; Bolívar (Cerro Bolívar, Lago Guri), Amazonas (Sierra Parima). Distrito Federal, Miranda, Sucre, Táchira; Trinidad, eastern Brazil. Clidemia dentata D. Don, Mem. Wern. Nat. Hist. Soc. 4: 306. 1823. Shrub 1–3(–6) m tall. Evergreen lowland and lower montane forests, 100–800 m; Bolívar (95 km south of El Dorado, Kilómetro 88, Río Samay). Apure, Barinas, Mérida, Monagas, Portuguesa, Táchira, Zulia; Mexico, Central America, Trinidad, tropical South America. Clidemia duidae Gleason, Bull. Torrey Bot. Club 58: 432. 1931. Shrub. Tepui summit, 2100–2200 m; Amazonas (Cerro Duida). Endemic. Clidemia epibaterium DC., Prodr. 3: 157. 1828. Terrestrial or epiphytic woody vine with adventitious roots. Evergreen lowland and lower montane forests, 100–1100 m; Amazonas (base of Cerro Duida, upper Río Caura, San Carlos de Río Negro, base of Sierra de la Neblina, Yavita), Bolívar (Cerro Ichún, Macizo del Chimantá [Toronó-tepui]). Amazonian Colombia, Guyana, Peru, northern Brazil. ◆Fig. 213. Clidemia heptamera Wurdack, Mem. New York Bot. Gard. 10(5): 179. 1964. Shrub 0.5–2 m tall. Evergreen tepui-slope forests, (500–)1000–1500 m; Bolívar (Amaruay-tepui, Macizo del Chimantá [Abacapátepui], Sierra de Lema). Adjacent Guyana. ◆Fig. 220. Clidemia heteroneura (DC.) Cogn. in Mart., Fl. Bras. 14(4): 506. 1888.

Clidemia 301

—Melastoma heteroneura DC., Prodr. 3: 201. 1828. Shrub 0.5–1.5 m tall. Evergreen lowland to montane forests, 100–1200 m; Amazonas (Cerro Aracamuni, Cerro Marahuaka, Maroa, San Carlos de Río Negro, Sierra de la Neblina). Upper Río Negro basin in Colombia and Brazil.

Shrub 0.5–3(–4) m tall. Evergreen lowland to montane forests, riverbanks, 300– 1600 m; Bolívar (Gran Sabana, Ptari-tepui, upper Río Caura, Sierra de Lema, Sierra de Maigualida, Sororopán-tepui), Amazonas (Cerro Huachamacari, Cerro Yutajé). Sucre, Yaracuy; Belize, Nicaragua, disjunct to Trinidad, Guyana, Suriname, French Guiana, Brazil (Roraima). ◆Fig. 222.

Clidemia hirta (L.) D. Don, Mem. Wern. Nat. Hist. Soc. 4: 309. 1823. —Melastoma hirta L., Sp. Pl. 7: 35. 1880. Shrub to 2(–3) m tall. Evergreen lowland and lower montane forests, riverbanks, open areas, 100–800 m. Widespread in the Neotropics; 3 varieties, all in the flora area.

Clidemia japurensis DC., Prodr. 3: 159. 1828. Shrub 1–4 m tall. Evergreen lowland and lower montane forests, 100–700 m. Monagas, Táchira; Nicaragua, Costa Rica, tropical South America; 2 varieties, both in the flora area.

Key to the Varieties of C. hirta

Key to the Varieties of C. japurensis

1. Leaf margins clearly crenulate; leaf blades cordate at base .... var. elegans 1. Leaf margins entire to obscurely crenulate, rounded to cordulate at the base ........................................................... 2 2. Branchlet hairs 0.5–1 mm long ................ ...................................... var. tiliaefolia 2. Branchlet hairs 2–3 mm long ................... ............................................. var. hirta

1. Branchlets and petioles densely glandular-setose with hairs 0.5–1 mm long eglandular hairs ....... var. heterobasis 1. Branchlets and petioles moderately finesetose with mostly eglandular hairs 1.5– 2 mm long .................. var. japurensis

C. hirta var. elegans (Aubl.) Griseb., Fl. Brit. W. I. 247. 1860. —Melastoma elegans Aubl., Hist. Pl. Guiane 427, t. 167. 1775. 200–800 m; Delta Amacuro (Caño Mánamo, Sacupana), Bolívar (Altiplanicie de Nuria, Las Patos south of El Manteco, Río Paramichi). Other distribution as in the species. ◆Fig. 217. C. hirta var. hirta 200–800 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela and the Neotropics, introduced into Africa and the Pacific and invasive there. C. hirta var. tiliaefolia (DC.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4.1): 488. 1941. —Clidemia tiliaefolia DC., Prodr. 3: 158. 1828. 100–400 m; Bolívar (Río Caura), Amazonas (widespread). Amazonian Colombia, Ecuador, Peru, and Brazil. Clidemia involucrata DC., Prodr. 3: 163. 1828.

C. japurensis var. heterobasis (DC.) Wurdack, Phytologia 21: 125. 1971. —Clidemia heterobasis DC., Prodr. 3: 164. 1828. 100–700 m; Amazonas (Río Casiquiare, Río Pasimoni, Río Siapa, Río Yatúa, San Carlos de Río Negro, Simarawochi, Tamatama). Táchira; Costa Rica, Nicaragua, Amazonian Colombia to Bolivia. ◆Fig. 223. C. japurensis var. japurensis 100–300 m; Delta Amacuro (Caño Jotajana), northern Bolívar. Monagas; Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, and Brazil. Clidemia juruensis (Pilg.) Gleason, Bull. Torrey Bot. Club 58: 84. 1931. —Maieta juruensis Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 178. 1905. Shrub 1–2 m tall. Lowland forests along blackwater rivers, ca. 100 m; Amazonas (Río Negro between San Carlos and Piedra Cocuy). Amazonian Colombia, Ecuador, Peru, Brazil, and Bolivia. Clidemia linearis (Gleason) Wurdack, Phytologia 19: 197. 1969. —Leandra linearis Gleason, Bull. Torrey Bot. Club 15: 425. 1931.

302

M ELASTOMATACEAE

Shrub 30–70 cm tall. Tepui slopes and summits, 1100–1700 m; Amazonas (Cerro Duida, Cerro Marahuaka). Endemic. ◆Fig. 215. Clidemia marahuacensis Wurdack, Mem. New York Bot. Gard. 51: 104. 1989. Shrub 1–3 m tall. Evergreen montane forests, 1000–1600 m; Amazonas (Caño Iguapo, Cerro Marahuaka). Endemic. Clidemia micrantha Sagot, Ann. Sci. Nat. Bot. sér. 6, 15: 327. 1883. Shrub 1–2 m tall. Lowland evergreen forests; Bolívar (elevation and distribution unknown). Guyana, French Guiana, Brazil (Pará). Clidemia microthyrsa R.O. Williams, Fl. Trinidad 1: 348. 1934. Shrub to 2 m tall. Lowland semideciduous forests, 300–500 m; Bolívar (El Manteco, Río Botanamo). Trinidad, Guyana, Suriname, French Guiana. Clidemia minutiflora (Triana) Cogn. in Mart., Fl. Bras. 14(4): 507. 1888. —Sagraea minutiflora Triana, Trans. Linn Soc. London 28: 137. 1871. Shrub 0.5–2 m tall. Lowland to montane evergreen forests, 100–1400 m; Bolívar (Cerro Ichún, Macizo del Chimantá, Río Canarakuni, Río Chicanán, Río Hacha in Río Icabarú basin, Río Paragua, Sierra de Lema), Amazonas (Cerro Duida, Cerro Huachamacari, Río Casiquiare, Río Mawarinuma, San Carlos de Río Negro). Guyana, Suriname, French Guiana, Brazil (Amazonas: upper Rio Negro). ◆Fig. 221. Clidemia morichensis Wurdack, Mem. New York Bot. Gard. 10(5): 183. 1964. Shrub ca. 1 m tall. Evergreen lowland to lower montane forests, 200–900 m; Bolívar (Cerro Guaiquinima, Río Paragua), Amazonas (Cerro Moriche). Endemic. Clidemia neblinae Wurdack, Mem. New York Bot. Gard. 10(4): 45. 1961. Small tree to 6 m tall. Montane evergreen forests, 1400–1600 m; Amazonas (Sierra de la Neblina). Endemic. Clidemia novemnervia (DC.) Triana, Trans. Linn. Soc. London 28: 137. 1871.

—Heterotrichum novemnervium DC., Prodr. 3: 173. 1828. Shrub to 3 m tall. Lowland evergreen or semideciduous forests, 100–600 m; widespread in Bolívar and Amazonas. Apure, Zulia; Belize, Nicaragua, Costa Rica, Panama, Amazonian Colombia, Peru, Brazil, and Bolivia. ◆Fig. 208. Clidemia octona (Bonpl.) L.O. Williams, Fieldiana, Bot. 29: 558. 1963. —Melastoma octona Bonpl. in Humb. & Bonpl., Monogr. Melast. 1: 7, pl. 4. 1816 [1806]. Shrub 1–3 m tall. Lowland to montane evergreen or semideciduous forests, 100–300. Widespread in the Neotropics; 2 varieties, both in the flora area. Key to the Subspecies of C. octona 1. Hairs on stems 1–2 mm long .................... ............................. subsp. guayanensis 1. Hairs on stems 5–10 mm long .................. ...................................... subsp. octona C. octona subsp. guayanensis Wurdack, Phytologia 19: 193. 1969. 100–300 m; Bolívar (Los Patos south of El Manteco, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil (Goiás). C. octona subsp. octona 100–200 m; Bolívar (Los Patos south of El Manteco, Tumeremo), Amazonas (Río Mawarinuma). Aragua, Carabobo, Falcón, Lara, Miranda, Mérida, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Neotropics. Clidemia piperifolia Gleason, Bull. Torrey Bot. Club 58: 433. 1931. Shrub 1–4 m tall. Montane evergreen forests, 1000–1500 m; Amazonas (Cerro Cuao, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Peru, Brazil (Acre). Clidemia pustulata DC., Prodr. 3: 159. 1828. Shrub 1–3 m tall. Lowland to montane evergreen forests, savanna edges, (200–)500– 1300 m; Bolívar (Gran Sabana, Río Chicanán, Río Paragua), Amazonas (Isla Ratón). Falcón; Belize, Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, northern Brazil. ◆Fig. 210.

Clidemia 303

Clidemia pycnaster Tutin, J. Bot. 72: 306. 1934. Shrub to 1 m tall. Venezuela, Guyana, Suriname; 2 varieties, 1 in Venezuela. Clidemia pycnaster subsp. robusta Wurdack is known from the interior savannas of Guyana and Suriname. C. pycnaster subsp. pycnaster Sandy savannas, 400–1300 m; Bolívar (Cerro Venado, El Paují, Kavanayén, Río Carrao). Guyana, Suriname. ◆Fig. 207. Clidemia rubra (Aubl.) Mart., Nov. Gen. Sp. Pl. 3: 152. 1832. —Melastoma rubra Aubl., Hist. Pl. Guiane 416. 1775. Shrub to 3 m tall. Evergreen lowland forests, savannas, 100–300 m; Bolívar (Río Cuyuní), Amazonas (Río Casiquiare, Río Coromoto, Río Guayapo, Río Mawarinuma, San Carlos de Río Negro). Guárico, Zulia; Colombia and Guyana to Amazonian Peru, Brazil, and Bolivia. Clidemia sericea D. Don, Mem. Wern. Nat. Hist. Soc. 4: 310. 1823. Shrub to 1 m tall. Savannas, commonly in disturbed areas, 100–900(–1200) m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Barinas, Cojedes, Distrito Federal, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; Neotropics. Six varieties with varying amounts of foliar indument have been described. The Venezuelan populations fall under var. microphylla Naudin. Clidemia sessiliflora (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 505. 1888. —Staphidium sessiliflorum Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 311. 1852. Shrub 1–5 m tall. Evergreen montane forests, 900–1000 m; Amazonas (Sierra de la Neblina). Costa Rica, Colombia (Boyacá), Ecuador, Peru, Bolivia. Clidemia siapensis Wurdack, Mem. New York Bot. Gard. 10(5): 180. 1964. Shrub 1–2 m tall. Forest on igneous outcrops, 100–1800 m; Amazonas (Río Siapa, Serranía Uasadi). Brazil (Amazonas). Clidemia silvicola Gleason, Lloydia 2: 202. 1939.

Shrub 1–3 m tall. Evergreen montane forests, rocky slopes, 800–900 m; Bolívar (San Ignacio de Yuruaní). Guyana, Suriname, French Guiana. Clidemia simpsonii Wurdack, Phytologia 24: 205. 1972. Shrub ca. 1 m tall. Lowland evergreen forests, ca. 100 m; Amazonas (Río Baría). Amazonian Colombia, Peru, and Bolivia. Clidemia stellipilis (Gleason) Wurdack, Phytologia 19: 194. 1969. —Leandra stellipilis Gleason, Fieldiana, Bot. 28: 434. 1952. Shrub 1.5–3 m tall. Evergreen montane forests, 900–2100 m; Bolívar (Cerro Venamo, Gran Sabana, Sororopán-tepui). Guyana. Clidemia strigillosa (Sw.) DC., Prodr. 3: 159. 1828. —Melastoma strigillosa Sw., Prodr. 71. 1788. Shrub 0.5–2 m tall. Lowland to montane forests and savannas, 100–1000 m; Bolívar (Canaima, Cuchivero, Gran Sabana), Amazonas (La Esmeralda, Río Asisa, Río Parhueña). Anzoátegui, Aragua; Central America, Antilles, Colombia, Guyana, Suriname, Amazonian Peru and Brazil. Clidemia tepuiensis Wurdack, Phytologia 19: 197. 1969. —Staphidiastrum coriaceum Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 329. 1852. —Clidemia coriacea (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 510. 1888, non Naudin 1852. Shrub 0.3–1.5 m tall. Scrub forests, 900– 2400 m; Bolívar (Amaruay-tepui, Auyántepui, Cerro Guaiquinima, Cerro Jaua, Cerro Venamo, La Danta, Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Aracamuni, Cerro Marahuaka, Cerro Sipapo, Cerro Uasadi, Cerro Uei, Sierra de Maigualida). Adjacent Guyana, Brazil (Amazonas). ◆Fig. 219. Clidemia tococoidea (DC.) Gleason, Bull. Torrey Bot. Club 58: 81. 1931. —Calophysa tococoidea DC., Prodr. 3: 166. 1828. Shrub 1–3 m tall. Savannas, lowland evergreen forests, 100–400 m; Amazonas (widespread). Amazonian Colombia, Brazil (Amazonas: Rio Negro basin). ◆Fig. 214.

304

M ELASTOMATACEAE

Fig. 205. Clidemia alternifolia

Fig. 206. Clidemia bullosa

Fig. 207. Clidemia pycnaster subsp. pycnaster

Clidemia 305

Fig. 208. Clidemia novemnervia

Fig. 210. Clidemia pustulata

Fig. 209. Clidemia aphanantha

306

M ELASTOMATACEAE

Fig. 211. Clidemia capitata

Fig. 213. Clidemia epibaterium

Fig. 212. Clidemia buntingii

Clidemia 307

Fig. 214. Clidemia tococoidea

Fig. 215. Clidemia linearis

Fig. 216. Clidemia capillipes

308

M ELASTOMATACEAE

Fig. 217. Clidemia hirta var. elegans

Fig. 218. Clidemia conglomerata

Clidemia 309

Fig. 219. Clidemia tepuiensis

Fig. 220. Clidemia heptamera

310

M ELASTOMATACEAE

Fig. 221. Clidemia minutiflora

Fig. 222. Clidemia involucrata

Fig. 223. Clidemia japurensis var. heterobasis

Comolia 311

Clidemia trinitensis (Crueg.) Griseb., Fl. Brit. W. I. 249. 1860. —Ossaea trinitensis Crueg., Linnaea 20: 105. 1847. Shrub 1–2.5 m tall. Evergreen montane forests, 2100–2300 m; Bolívar (Sororopántepui). Trinidad, Tobago. Clidemia ulei Pilg, Verh. Bot. Vereins Prov. Brandenburg 47: 180. 1905. Shrub 1–2 m tall. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro, Sierra de la Neblina base).

Colombia, Peru. Clidemia urceolata DC., Prodr. 3: 158. 1828. Shrub 1–2 m tall. Lowland to montane evergreen forests, 400–1500 m; Bolívar (Auyán-tepui, Gran Sabana, Macizo del Chimantá, Cerro Jaua base), Amazonas (Cerro Yutajé, Sierra Parima). Anzoátegui, Carabobo, Miranda, Nueva Esparta, Sucre, Táchira; Central America, Colombia, Trinidad, French Guiana, Brazil.

11. COMOLIA DC., Prodr. 3: 123. 1828. Tricentrum DC., Prodr. 3: 123. 1828. Leiostegia Benth., J. Bot. (Hooker) 2: 294. 1840. Tetrameris Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 120. 1850. by Navina G. Luckana and Paul E. Berry Shrubs or subshrubs. Leaves opposite, isomorphic, linear to obovate, 1–5veined, margins serrulate, denticulate, or ciliate with glandular or eglandular hairs. Flowers solitary on terminal or lateral branches, occasionally glomerate in upper leaf axils, 4-merous. Hypanthium usually terete; calyx lobes persistent; petals pink to violet or purple, obovate to oblong, glabrous or ciliate, sometimes ending in a gland-tipped terminal setula. Stamens somewhat dimorphic, glabrous; anthers subulate to linear-subulate, with a small ventrally inclined pore; connective somewhat prolonged, bluntly bilobed ventrally, dorsally exappendiculate or with a blunt spur. Ovary 2–4-locular, glabrous; style glabrous; stigma not expanded. Fruit a capsule. Seeds numerous, cochleate, foveolate. Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Brazil; 25–30 species, 10 in Venezuela, all restricted to the flora area. This genus may eventually need to be subsumed under Acisanthera. Key to the Species of Comolia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5. 5(3).

Leaf blades 3–5 cm long; ovary 4-locular .................................... C. vernicosa Leaf blades < 3 cm long; ovary 2-locular ................................................... 2 Leaf blades linear to oblanceolate, 1–3 mm wide, 10–25 mm long ............................................................................................... C. leptophylla Leaf blades wider than above .................................................................... 3 Leaf blades thick-rigid, the margin entire; tepui summits in Bolívar state ................................................................................................................ 4 Leaf blades chartaceous to thin-rigid, the margin denticulate at least apically; more widespread ..................................................................... 5 Leaves and hypanthium glabrous ................................................ C. montana Leaves and hypanthium glandular-pubescent .............................. C. coriacea Hypanthium glabrous or finely setulose with nonglandular trichomes .. 6 Hypanthium fine-setulose, with gland-tipped trichomes ......................... 8

312

M ELASTOMATACEAE

Hypanthium slightly 8-ribbed; leaf blades 2–3.5 × 1.5–2.5 cm ...... C. smithii Hypanthium terete; leaf blades generally smaller than above ................ 7 Leaf blade suborbicular to broadly elliptic-ovate, 1–1.7 × 0.8–1.5 cm; petals not ending with a gland-tipped setula .................. C. nummularioides 7. Leaf blade elliptic to obovate-elliptic, 1–2.5 × 0.4–1.4 cm; petals ending with a gland-tipped setula ............................................................ C. villosa 8(5). Leaves with evident primary and secondary venation; erect (sub)shrubs ................................................................................................................ 9 8. Leaves without evident primary and secondary venation; usually prostrate subshrubs or climbers ................................................................. 10 9(8). Hypanthium 2–2.5 mm long; petals 5–7 mm long .................. C. microphylla 9. Hypanthium 3–5 mm long; petals 9–12 long ................................... C. villosa 10(8). Internodes glabrous ...................................................................... C. prostrata 10. Internodes laxly setulose .......................................................... C. serpyllacea 6(5). 6. 7(6).

Comolia coriacea Gleason, Fieldiana, Bot. 28: 429. 1952. Shrub 0.5–4 m tall, the whole plant glandular-puberulent, otherwise similar to C. montana. Rocky tepui meadows, 1800–2500 m; Bolívar (Aprada-tepui, Auyán-tepui, Macizo del Chimantá [Acapará-tepui, Chimantátepui, Churí-tepui, Murey-tepui], Ptaritepui). Endemic. ◆Fig. 226. Comolia leptophylla (Bonpl.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 27. 1850. —Rhexia leptophylla Bonpl. in Humb. & Bonpl., Monogr. Melast. 2: 64, pl. 24. 1823 [1812]. —Tricentrum leptophyllum (Bonpl.) DC., Prodr. 3: 123. 1828. Subshrub 6–30 cm tall; leaves 1–2.5 × 0.1–0.3 cm, linear to oblanceolate, 1-veined, margins ciliate-denticulate; petioles lacking or to 0.5 mm long; petals 9–12 × 5–9 mm, pink. Savannas, mostly on white sand, 50– 200 m; Bolívar (southwest of Caicara, Río Horeda near Ciudad Bolívar, savannas near Serranía Carichana), Amazonas (Canaripó, Caño Camani, Parú savannas, Puerto Ayacucho, Raudal de Atures, Raudal de Maipures, Río Ventuari, near San Juan de Ucata, Santa Bárbara, Yapacana savannas). Apure; Colombia (Vichada). ◆Fig. 229. Comolia microphylla Benth., J. Bot. (Hooker) 2: 295. 1840. Shrub or subshrub 0.3–2 m tall, often cespitose; leaf blades 0.7–2 × 0.6–1.2 cm, elliptic to ovate-elliptic, 3–5 veined; petioles 2– 6 mm long; petals 5–7 × 3.5–4.5 mm, pale pink to purple and violet. Open granitic outcrops and edges of savannas, 50–1300 m;

widespread in Bolívar and Amazonas. Guyana, Suriname, French Guiana, Brazil. ◆Fig. 224. Comolia montana Gleason, Brittonia 3: 174. 1939. Shrub 0.1–1.5 m tall; leaves 7–13 × 4–10 mm, obovate to elliptic-obovate, 3-veined; petioles 1–2.5 mm long; petals 16–23 × 10– 21 mm, pink. Rocky tepui meadows, 1900– 2500 m; Bolivar (Auyán-tepui, Macizo del Chimantá [Amurí-tepui]). Endemic. ◆Fig. 227. Comolia nummularioides (Bonpl.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 27. 1850. —Rhexia nummularioides Bonpl., Rhex. 62, pl. 23. 1812. —Arthrostemma nummularioides (Bonpl.) DC., Prodr. 3: 137. 1828, “Arthrostema .” Shrub 0.5–1.5 m tall; leaves 1–1.7 × 0.8– 1.5 cm, suborbicular to widely elliptic-ovate, 5-veined; petioles 3–5 mm long; petals 6.5–8 × 5–6 mm, reddish purple. Open granitic outcrops and savannas, 50–200 m; Bolívar (Caicara, Río Suapure), Amazonas (La Esmeralda, Puerto Ayacucho). Colombia (Guainía, Vichada). ◆Fig. 231. Comolia prostrata Wurdack, Mem. New York Bot. Gard. 10(5): 143. 1964. Densely branched subshrub, reddishtinged; branches prostrate and glabrous, to 20 cm long; leaves 3–5 × 2–4 mm, obovate-elliptic, 3-veined, blades and margins with glandular hairs; petals 7–8 × 4.5 mm, pink. White-sand savannas, ca. 100 m; Amazonas (Río Atabapo). Colombia (Guainía).

Comolia 313

Comolia serpyllacea Wurdack, Mem. New York Bot. Gard. 10(1): 102, fig. 11a–g. 1958. Subwoody, prostrate climber; leaves 2.5–4 × 1–2 mm, elliptic to elliptic-ovate, 1-veined; petioles 0.3–0.8 mm long; petals 13–15 × 10– 11 mm. Moist rock faces of tepui slopes, 1300–2500 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Endemic. ◆Fig. 230.

Comolia vernicosa (Benth.) Triana, Trans. Linn. Soc. London 28: 37. 1871. —Leiostegia vernicosa Benth., J. Bot. (Hooker) 2: 294. 1840. Shrub 0.5–2 m tall, glabrous and viscose; leaf blades 3–5 × 0.8–1.5 cm, oblong-elliptic, 3-veined; petioles 1–4 mm long; petals 5–7 × 3.5–4 mm, glabrous, rose to violet. Lowland savannas, 50–100 m; Bolívar (Río Asa). Guyana, Suriname, Brazil (Pará). ◆Fig. 228.

Comolia smithii Wurdack in Görts, Fl. Guianas, ser. A, fam. 99 (Melastomataceae) 85. 1993. Shrub 0.5–1 m tall, entire plant covered with gland-tipped trichomes; leaves 20–35 × 15–25 mm, ovate to elliptic-ovate, 5-veined; petiole 6–10 mm long; petals 12 × 8–9 mm, pale pink. Savannas, granitic outcrops, 50– 300 m; Bolívar (Corozal, Serranía Cerbatana), Amazonas (Piedra Nunca north of Piedra Cocuy, Puerto Ayacucho). Guyana, Brazil (Amazonas).

Comolia villosa (Aubl.) Triana, Trans. Linn. Soc. London 28: 37. 1871. —Rhexia villosa Aubl., Hist. Pl. Guiane 334, t. 129, fig. 1. 1775. —Arthrostemma villosum (Aubl.) DC., Prodr. 3: 137. 1828. —Tetrameris villosa (Aubl.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 14: 127. 1850. Comolia purpurea Miq., Linnaea 18: 617. 1840. Comolia veronicaefolia Benth., J. Bot. (Hooker) 2: 295. 1840. Comolia hirtella Naudin, Ann. Sci. Nat.

Fig. 224. Comolia microphylla

Fig. 225. Comolia villosa

Fig. 226. Comolia coriacea

314

M ELASTOMATACEAE

Fig. 227. Comolia montana

Fig. 228. Comolia vernicosa

Fig. 230. Comolia serpyllacea

Fig. 229. Comolia leptophylla

Fig. 231. Comolia nummularioides

Comoliopsis 315

Bot. sér. 3, 13: 26. 1850. Comolia lythrarioides Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 27. 1850. Comolia surinamensis Miq., Stirp. Surinam. Select. 55. 1850 [1851]. Comolia neglecta Cogn. in Mart., Fl. Bras. 14(3): 423. 1883. Comolia affinis Hoehne, Anexos Mem. Inst. Butantan, Secc. Bot. 1(5): 91, t. 13, fig. 1. 1922. Comolia kuhlmannii Hoehne, Anexos

Mem. Inst. Butantan, Secc. Bot. 1(5): 92, t. 13, fig. 2. 1922. Comolia angustifolia Gleason, Brittonia 1: 151. 1932. Shrub 20–70 cm tall; leaves 10–25 × 4–14 mm, elliptic to obovate-elliptic, 3- or 5veined; petals 8–12 × 5–7 mm, pink-lavender. Savannas, rock outcrops, forest edges, 100–1000 m; widespread in Bolívar and Amazonas. Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 225.

12. COMOLIOPSIS Wurdack, Acta Bot. Venez. 14(3): 23. 1984. by Paul E. Berry Prostrate subshrubs. Flowers 5-merous, mostly solitary or ternate in upper leaf axils. Hypanthium terete; calyx lobes subulate, persistent. Stamens slightly dimorphic; anthers narrowly lanceolate, pore ventrally inclined; connective prolonged 1 mm or 0.6–0.7 mm, ventrally with slightly bilobed appendage, dorsally non-

Fig. 232. Comoliopsis neblinae

316

M ELASTOMATACEAE

appendaged. Ovary superior, (3)4-locular, the apex with glanduliferous hairs; style glabrous; stigma not expanded. Seeds cochleate, minutely papillate. Endemic to the Sierra de la Neblina in Venezuela and Brazil; 1 species. Comoliopsis neblinae Wurdack, Acta Bot. Venez. 14(3): 23. 1984. Prostrate shrub 10–30 cm tall; leaves coriaceous, rigid, 1–1.7 × 0.8–1.2 cm, 3-veined, apex broadly acute or obtuse and shortly mucronulate, base obtuse or rounded-trun-

cate, petiole 1–3 mm long; pedicels 1–1.5 mm long; petals dark pink, obovate, apex broadly acute, 13–17 × 8–9 mm; filaments reddish pink. Moist tepui meadows and streamsides, 2200–2300 m; Amazonas (Sierra de la Neblina). Brazil (Serra da Neblina). ◆Fig. 232.

Fig. 233. Conostegia superba

Desmoscelis 317

13. CONOSTEGIA D. Don, Mem. Wern. Nat. Hist. Soc.4: 316. 1823. by Paul E. Berry and Kay Yatskievych Shrubs or trees to 20 m tall. Leaves often moderately anisophyllous, narrowly ovate to elliptic or obovate; margins entire to dentate; tertiary venation never prominent. Inflorescences terminal or subterminal panicles. Calyx calyptrate, circumscissile at anthesis; petals (4)5–12, obovate and rounded at the apex, white to pink, glabrous and minutely granulose. Stamens 10–30(–96), glabrous, the thecae narrowly oblong, with a single pore; connective simple, not prolonged. Ovary 5–17(– 20)-locular, completely inferior, apex glabrous; style glabrous; stigma usually wider than style, subcapitate or capitate to truncate or punctiform. Fruit berry-like. Seeds many, often irregularly pyramidal, angular. Mexico, Central America, West Indies, Colombia, Venezuela, Tobago, Ecuador, Peru, Brazil; 42 species, 7 in Venezuela, 1 of these in the flora area. Conostegia superba Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 108. 1851. Small tree 3–10 m tall; branchlets strongly tetragonal with ridged corners; leaf blades 9–36 × 4–25 cm; petioles 1–8 cm long; panicle terminal, 8–25 cm long; petals 4–6,

white or pink; stamens 10–17, yellow. Montane tepui-slope forests, 1200–1800 m; Amazonas (Sierra de la Neblina). Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil. ◆Fig. 233.

14. DESMOSCELIS Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: 29. 1849. by Paul E. Berry Little-branched herbs or subshrubs. Flowers 5-merous, in leafy panicles. Hypanthium terete; calyx lobes lanceolate, persistent; petals pink to purple-pink, obovate and rounded to emarginate, ciliate. Stamens 10, dimorphic, glabrous; thecae oblong, with a single small pore; large stamens with connective well prolonged and ventrally with a pair of filiform lobes; small stamens with connective not prolonged and with short, thick ventral lobes. Ovary superior, the apex densely fine-setose with deciduously gland-tipped hairs; style glabrous; stigma not expanded. Fruit a capsule, 5-locular. Seeds numerous, cochleate, tuberculate. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 species, 1 in Venezuela. Desmoscelis villosa (Aubl.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 30. 1850. —Melastoma villosa Aubl., Hist. Pl. Guiane 428, t. 168. 1775. Desmoscelis mollis Pittier, J. Wash. Acad. Sci. 13: 387. 1923. Herb or subshrub 0.5–1 m tall; stems quadrangular, moderately to densely fine-setose with obscurely scabrous hairs 2–7(–10) mm long; leaf blades (1.5–3–5(–6) × 1–2.5 cm, ovate to oblong-elliptic, apex acute, base rounded to cordulate, firm-chartaceous and

entire, 5–7-veined; petiole 2–5 mm long; flowers usually solitary in upper bract axils of short lateral branches; pedicels 1.5–3 mm long; petals pink, 6–9 × 4–7 mm. Lowland and upland savannas, Mauritia palm swamps, white-sand scrub (bana), weedy open areas along evergreen lowland forests, 50–1200 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Carabobo, Guárico, Monagas, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 234.

318

M ELASTOMATACEAE

Fig. 234. Desmoscelis villosa

Ernestia 319

15. ERNESTIA DC., Prodr. 3: 121. 1828. by Navina G. Luckana and Paul E. Berry Shrubs or subshrubs. Leaves opposite, isomorphic. Flowers 4- or 5-merous, usually in terminal panicles (rarely solitary or terminal-glomerulous in some nonVenezuelan species). Hypanthium terete; calyx lobes persistent; petals pink or white, obovate and obtuse. Stamens 8 or 10, dimorphic, glabrous; anthers subulate and arched, minutely 1-pored; connective evidently or slightly prolonged toward the insertion of the filament, ventrally bilobed with aristate appendices, dorsally ± tuberculate. Ovary superior, 3- or 4- locular, glabrous or glandular-puberulous; stigma not expanded. Fruit a capsule, 3- or 4-locular. Seeds numerous, cochleate, muriculate. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 12–14 species, 5 in Venezuela, all in the flora area. This genus is very closely related to Comolia. Key to the Species of Ernestia 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Leaf blade 3–10 × 1–6 cm; flowers 4- or 5-merous .................................... 2 Leaf blade 1.5–3(–4.5) × 1–2.5(–3) cm; flowers 4-merous ......................... 3 Leaf blade 7- or 9-veined; flowers 5-merous, petal margins glandularciliolate ..................................................................................... E. cordifolia Leaf blade 5- or 7-veined; flowers 4-merous, petal margins glabrous ....................................................................................................... E. tenella Ovary 3-locular ................................................................................... E. pullei Ovary 4-locular ........................................................................................... 4 Apex of petals ending in 1–3 glandular hairs; style glabrous; leaf blade 5- or 7-veined .......................................................................... E. cataractae Apex of petals ending in 1 glandular hair; style glandular-puberulent in the middle; leaf blade 5-veined ................................................ E. maguirei

Ernestia cataractae Tutin, J. Bot. 72: 308. 1934. Subshrub 10–50 cm tall; leaf blades ovate-elliptic, apex acute, base rounded to cordate, 1.5–3 × 1–2.5 cm, 5- or 7-veined; petioles 0.3–1.5 cm long; flowers 4-merous, petals pink, each one ending in 1–3 glandular hairs; style glabrous, ovary 4-locular, glabrous. Rocky, open tepui summits, 1000– 1800 m; Bolívar (Macizo del Chimantá [Toronó-tepui]), Amazonas (Cerro Guanay, Cerro Parú, Cerro Ualipano, Cerro Yaví, Serranía Uasadi, Sierra de Maigualida). Guyana, Brazil (Amazonas). ◆Fig. 237.

This species may be conspecific with Comolia ayangannae Wurdack. Ernestia cordifolia Berg. ex Triana, Trans. Linn. Soc. London 28: 36, pl. 2, fig. 23b. 1871.

Subshrub; leaf blades ovate, apex acute or acuminate, base cordate, 6–10 × 4–6 cm, 7or 9-veined, weakly puberulent; petioles 1.5– 3 cm long; flowers 5-merous, petals pink, glandular-ciliolate; style glabrous; ovary 3locular, glabrous. Along exposed granitic rocks along rivers, 50–100 m; Amazonas (Raudal de Atures, Raudal de Maipures). Adjacent Colombia. Ernestia maguirei Wurdack, Mem. New York Bot. Gard. 10(5): 136, fig. 63k–q. 1964. Shrub 0.5–1.5 m tall; leaf blades oblongovate, apex narrowly acute, base rounded or cordate, 2–3 × 1–1.5 cm, 5-veined; petioles 4– 10 mm long; petals pink, each ending in a glandular hair; style centrally glandularpuberulous; ovary 4-locular, glabrous. Upper tepui slopes and summits, 1800–1900 m;

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Fig. 235. Ernestia tenella var. sprucei

Fig. 236. Ernestia maguirei

Fig. 237. Ernestia cataractae

Graffenrieda 321

Bolívar (Auyán-tepui, Ptari-tepui, Sororopán-tepui). Endemic. ◆Fig. 236.

nian Colombia, Venezuela, Amazonian Brazil; 2 varieties, both in the flora area.

Ernestia pullei Gleason, Recueil Trav. Bot. Néerl. 32: 203. 1935. Subshrub 0.5–2 m tall; leaf blades ovateelliptic to ovate, apex acute to obtuse, base rounded to cordate; 1.5–3(–4.5) × 1–2(–3) cm, 5-veined; petioles 5–20 mm long; flowers 4merous, petals pink and ending in 1 or 3 glandular hairs; ovary 3-locular, apex of ovary and style glabrous. Open slopes of iron-rich mountains, 600–800 m; Bolívar (Cerro Altamira east of Ciudad Piar, Cerro Bolívar). Guyana, Suriname, French Guiana, Brazil.

Key to the Varieties of E. tenella

Ernestia tenella (Bonpl.) DC., Prodr. 3: 121. 1828. —Rhexia tenella Bonpl., Rhex. 79, pl. 30. 1813. Subshrub 0.5–2 m tall; branchlets 4angled, moderately to densely finely-setulose with glandular hairs; leaf blades ellipticovate to broadly lanceolate, apex acute to acuminate, base rounded to cordate, 3–7 × 1– 4 cm, 5- or 7- veined; petioles 4–15 mm long; flowers 4-merous, petals pink or white; style glabrous; ovary 3- or 4-locular, apex rarely to moderately glandular-puberulous. Amazo-

1. Lower surface of leaf reddish brown when dry; trichomes on stem ± 0.3 mm long; apex of ovary moderately glandular-setose ................................... var. sprucei 1. Lower surface of leaf pale when dry; trichomes on stem 0.8–1 mm long; apex of ovary very sparsely glandular-puberulent ................................... var. tenella E. tenella var. sprucei Cogn. in Mart., Fl. Bras. 14(3): 227. 1885. Ernestia lata Gleason, Bull. Torrey Bot. Club 52: 328. 1925. Openings of lowland or lower montane forests, 100–500(–1500) m; Amazonas (slopes of Cerro Duida, slopes of Cerro Moriche, ridge above La Esmeralda). Colombia, Brazil. ◆Fig. 235. E. tenella var. tenella Disturbed areas along lowland forest edges, 50–200 m; Amazonas (Pimichín, Río Temi, San Carlos de Rio Negro, San Fernando de Atabapo, Yavita). Colombia, Brazil.

16. GRAFFENRIEDA DC., Prodr. 3: 105. 1828. Calyptrella Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 115. 1852. Ptilanthus Gleason, Bull. Torrey Bot. Club 72: 472. 1945. by Andreas Gröger Deciduous or evergreen shrubs or small trees to 15 m tall (seldom vines). Leaves entire, coriaceous or rigid, 1–7-veined (occasionally pliveined). Flowers in terminal panicles, 4–6(–8)-merous. Hypanthium terete or costate; calyx open or closed in bud, dehiscing regularly or irregularly into persistent lobes at anthesis (seldom transversally dehiscent); petals white to pink, glabrous or inconspicuously pulverulent, obovate to lanceolate, rounded to acuminate at the apex. Stamens isomorphic or nearly so; anthers linear-subulate and arcuate, with an apical, ventrally inclined pore; connective not or slightly dorso-basally prolonged into a sharp spur. Ovary 2–5(–7)-locular, glabrous or granulose; style glabrous; stigma not expanded. Fruit a capsule. Seeds numerous, linear-pyramidal. Mexico, Central America, Greater and Lesser Antilles, Colombia, Venezuela, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia; ca. 45 species, 30 in Venezuela; 24 of these in the flora area. The generic limits of Graffenrieda are difficult to define. They are mainly based on pubescence, leaf venation, calyx structure, ovary surface, anther spurs, and seed morphology. In some characters, Graffenrieda resembles Meriania (differs in struc-

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ture of the anther connective), Centronia (differs in structure and indumentum of the calyx), and Miconia (differs in indumentum and fruit). Key to the Species of Graffenrieda 1. 1. 2(1). 2. 3(2). 3. 4(2). 4. 5(4). 5. 6(5). 6. 7(6). 7. 8(6). 8. 9(4). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(1). 13. 14(13).

Flowers predominantly 4-merous .............................................................. 2 Flowers predominantly 5- or 6-merous ................................................... 13 Ovary 2-locular ........................................................................................... 3 Ovary 3(4)-locular ...................................................................................... 4 Leaf with 1 primary vein; apex obtuse to acute; blade 4–7 times as long as wide; petals 8–13 mm long ................................................. G. pedunculata Leaf 5-pliveined; apex acuminate; blade 2–3 times as long as wide; petals 2.5 mm long ................................................................................. G. obliqua Leaves with 1–3 primary veins (seldom 5, with the outer laterals close to the margin and weakly developed) ....................................................... 5 Leaves with 5–7 primary veins ................................................................. 9 Leaves with 1 (seldom 3) primary veins; apex of ovary with a dentate tube (1–1.5 mm long) around the style base ................................... G. versicolor Leaves with 3 primary veins (seldom 5); apex of ovary without dentate tube ......................................................................................................... 6 Leaves 1.3 cm or less wide ......................................................................... 7 Leaves 2.5–21 cm wide ............................................................................... 8 Leaf blade 1–2 cm long, 2–3 times as long as wide; apex of ovary granulose ............................................................................................. G. steyermarkii Leaf blade 4–8 cm long, 5–7 times as long as wide; ovary glabrous ................................................................................................. G. lanceolata Leaf apex rounded or retuse; pedicels thick (0.6–0.9 mm diameter) .................................................................................................. G. reticulata Leaf apex acute or acuminate; pedicels thin (0.2–0.3 mm diameter) .............................................................................................. G. miconioides Leaves pliveined ....................................................................................... 10 Leaves not pliveined ................................................................................ 11 Axils of primary veins on the lower surface of leaf with simple hairs 2–5 mm long (seldom completely glabrous) .................................. G. kralii Primary veins of the lower surface of leaves with barbed hairs < 1 mm long ........................................................................................... G. cinnoides Lower surface of leaf with a dense, ferruginous cover and with a prominent secondary venation ........................................................ G. hitchcockii Leaf surface of leaf glabrous to nearly glabrous (hairs only in the primary vein axils); secondary venation not prominent ................................... 12 Tertiary leaf venation reticulate; lower surface of leaves totally glabrous .................................................................................................. G. reticulata Tertiary leaf venation not obvious; lower surface of leaves frequently with long hairs in the axils of primary veins ................................. G. fantastica Ovary 5-locular; leaf blade usually 20–40 cm long ...................... G. rupestris Ovary 2–4-locular; leaf blade usually < 20 cm long ................................ 14 Leaves sessile; leaf blade > 15 times as long as petiole; petiole 1–7 mm long; leaf base cordate ........................................................... G. sessilifolia

Graffenrieda 323

14.

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

18. 19(18). 19. 20(15).

20.

21(20). 21. 22(21). 22. 23(22). 23.

24(23). 24. 25(24). 25. 26(25). 26. 27(24). 27. 28(27). 28.

Leaves obviously petiolate; leaf blade at most 10 times longer than petiole; petiole 4–70 mm long; leaf base acute to obtuse or cordate .............................................................................................................. 15 Leaves with 3 primary veins ................................................................... 16 Leaves with 5–9 primary veins ............................................................... 20 Leaves 5–6.5 times as long as wide ............................................ G. lanceolata Leaves < 3 times as long as wide ............................................................. 17 Leaf blade 1–4 cm long; calyx lobes with external teeth ............. G. fruticosa Leaf blade 7–21 cm long; calyx lobes without teeth ............................... 18 Leaf blade suborbicular to elliptic-ovate (< 1.5 times as long as wide); plants deciduous, with flowers appearing before or together with the foliage ................................................................................... G. rotundifolia Leaf blade elliptic (at least 1.5 times as long as wide); plants evergreen .............................................................................................................. 19 Calyx margin entire in bud, splitting irregularly into 2–5 lobes at anthesis ......................................................................................... G. caryophyllea Calyx margin entire or equally oblately lobed ............................ G. polymera Leaves obviously 7- or 9-pliveined, the inner pair of primary veins diverging 3–4 cm from the base of the leaf blade; petiole with 2 thick protuberances .................................................................................. G. sipapoana Primary veins diverging at the base of the leaf blade or shortly pliveined, the inner pair of primary veins diverging < 1 cm from the base of the blade; petiole without protuberances .................................................. 21 Liana; calyx lobes with small thickened external teeth .................. G. patens Shrub or tree; calyx lobes with acute or no teeth ................................... 22 Calyx hyaline and calyptrate, at anthesis transversally dehiscent, only external teeth of calyx lobes remaining .............................. G. tricalcarata Calyx firm, open or closed in bud, at anthesis longitudinally dehiscent into persistent lobes ............................................................................. 23 Primary veins of the lower surface of leaves with barbed hairs to 0.8 mm long ........................................................................................... G. cinnoides Lower surface of leaves with primary veins glabrous (occasionally simple hairs only in the axils of the veins), pulverulent, or with small simple hairs covering the whole lower surface ............................................... 24 Lower surface of leaves with long hairs in the primary vein axils ........ 25 Lower surface of leaves without long hairs in the primary vein axils .............................................................................................................. 27 Each calyx lobe with a projecting external tooth ............................ G. jauana Calyx lobes without teeth ........................................................................ 26 Petals 4–6 mm long; apex of ovary rounded ............................. G. intermedia Petals 7.5–14 mm long; apex of ovary with a collar around the style base .................................................................................................. G. weddellii Petals lanceolate (4–5 times as long as wide) ........................... G. intermedia Petals obovate (1–2 times as long as wide) ............................................. 28 Pedicels 2–3 mm long; petals white; leaf blade 7–25 cm long ................................................................................................ G. hitchcockii Pedicels 0.5–1.7 mm long; petals pink; leaf blade 1.5–7 cm long ........................................................................................................... G. rufa

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Graffenrieda caryophyllea Triana, Trans. Linn. Soc. London 28: 69. 1871. Shrub or much-branched tree 3–12 m tall; young leaves and inflorescences densely appressed-squamate; petiole 1.5–2.5 cm long; leaf blade 7–17 × 2–10 cm, oblong-elliptic to ovate-elliptic, apex broadly acute to obtuse, base obtuse to rounded, thin-rigid, lower surface of leaves sometimes with an amorphous, light brownish indumentum, 3-veined or shortly pliveined with the primary laterals near the margin; panicle 10–20 cm long, many-flowered; flowers 5-merous; pedicels 1–1.7 mm long, thick; hypanthium 4–8 mm long; calyx margin dehiscing at anthesis irregularly into 2–5 ovate persistent lobes (0.7–2.7 mm long); petals 7–10 × 4–6.5 mm, obovate, apex rounded, white (rarely pinkish); anther thecae 6.5–9.5 mm long, dorsal spur 1.2–2 mm long; ovary 3-locular, glabrous. Scrub forests on granitic outcrops (lajas), low forests on sandstone, 100–1900 m; Bolívar (Amaruay-tepui, Auyán-tepuí, Cerro Guaiquinima, Cerro Pitón, Cerro Venado, Gran Sabana, Macizo del Chimantá, upper Río Cuyuní, Río Parguaza, Río Urimán, Salto Pará, Sierra de Maigualida), Amazonas (base of Cerro Aracamuni, Cerro Duida, Cerro Yutajé, Laja Catipán, Piedra Arauicaua, Piedra Cocuy, Río Cataniapo, Río Coro Coro, Río Cuao, upper Río Guayapo, Ucata southeast of Síquita). Colombia (Vaupés), Guyana (Pakaraima Mountains). ◆Fig. 253. Graffenrieda caryophyllea is a highly variable taxon with a large altitudinal amplitude. Most obvious variability appears in inflorescence structure. Lowland specimens show a lax, many-branched inflorescence. In species from above 1000 m, flowers are grouped in nearly spherical tufts in intervals along an unbranched or few-branched axis. Graffenrieda caryophyllea is closely related to G. polymera. Both taxa form a species complex that will require a more detailed revision. Graffenrieda cinnoides Gleason, Mem. New York Bot. Gard. 8: 135. 1953. Shrub or tree to 14 m tall; petiole 1–3 cm long; leaf blade 7–12 × 2.5–5.5 cm, ovate-lanceolate to elliptic-ovate, apex subacuminate, base broadly acute to obtuse, membranous, 5-veined or shortly (to 0.8 cm) 5-pliveined; primary veins on the lower surface of leaf

densely covered with barbed hairs (0.1–0.8 mm long); panicle 5 cm long, few-flowered; pedicels 1 mm long; hypanthium 2–3.5 mm long; calyx lobes 1.5 mm long, lanceolate; petals unknown; anther thecae 3.5 mm long, dorsal spur 0.1–0.2 mm long; ovary 3- or 4locular, densely granulose. Forests along streamlets, 1500–2000 m; Amazonas (Cerro Sipapo). Endemic. Graffenrieda fantastica R.E. Schult. & L.B. Sm., Bot. Mus. Leafl. 13: 306, pls. 35, 36. 1949. Tree 4–10 m tall; young branchlets with a succulent cortex parenchyma; petiole 2–4.5 cm long; leaf blade 8–17 × 6–10 cm, narrowly elliptic to ovate-elliptic, apex rounded, obtuse or acute, base obtuse; panicle manyflowered; flowers sessile, 4-merous; hypanthium 3–4 mm long; calyx margin dehiscing at anthesis into 3 or 4 ovate or rounded lobes; petals 3–4.5 × 2–3 mm, ovate, apex narrowly acute, white; anther thecae 3.5 mm long, curved, dorsal spur 0.4–0.5 mm long; ovary 3-locular, glabrous. Low forests on sandstone, 300–1800 m; Bolívar (Cerro Guanay), Amazonas (Cerro Camani, Cerro Cuao, Cerro Moriche, Cerro Sipapo, Cerro Yutajé). Colombia. Graffenrieda fruticosa Wurdack, Mem. New York Bot. Gard. 10(1): 109. 1958. Shrub 0.2–2 m tall; branchlets, lower surface of leaves, and inflorescences completely covered with an appressed indumentum; petiole 0.4–0.9 cm long; leaf blade 1–4 × 0.6– 2 cm, elliptic to ovate-elliptic, apex acute, base broadly acute to obtuse, thin-coriaceous, 3-veined; inflorescences terminal dichasia or flowers solitary; flowers sessile to subsessile, (4)5- or 6-merous; hypanthium 3– 5 mm long; calyx lobes rounded, with external teeth projecting 1–2.5 mm; petals 8–14 × 5–9 mm, obovate, apex obtuse, pink; anther thecae 4–5 mm long, dorsal spur 0.8–1 mm long; ovary 3-locular, apex densely granulose, forming a collar (1 mm high) around the style base. Bonnetia tepui forests, 1700–2300 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 248. Graffenrieda hitchcockii Gleason, Brittonia 7: 83. 1950.

Graffenrieda 325

Graffenrieda hitchcockii var. parvifolia Wurdack, Mem. New York Bot. Gard. 10(1): 108. 1958. Tree 4–15 m tall; branchlets, lower surface of leaves, and inflorescences completely covered with a dense, ferruginous indumentum; branchlets obtusely rectangular; petiole 0.5–8 cm long; leaf blade 7–25 × 3–13 cm, elliptic to ovate-elliptic, apex obtuse to broadly acute, base rounded to cordate, coriaceous, 5–7(–9)-veined; panicle many-flowered, 7–21 cm long; flowers 4- or 5-merous, clustered subumbellately at the end of the branchlets; pedicels 2–3 mm long, thick; hypanthium 4–4.5 mm long; calyx lobes 0.7–1.3 mm long, ovately rounded, with external teeth hardly exceeding the lobe apices; petals 5.5 × 7 mm, obovate, apex rounded, white; anther thecae 4.2–5 mm long, dorsal spur 0.5–0.7 mm long; ovary 3(4)-locular, granulose at apex, forming a regularly lobed collar (0.6 mm high) around the style base. Tepui forests, scrub forests, 1200–2200 m; Bolívar (Cerro Guanay), Amazonas (Cerro Coro Coro, Cerro Moriche, Cerro Yavi, Cerro Yutajé). Endemic. ◆Fig. 244. Wurdack differentiated two varieties (var. hitchcockii and var. parvifolia) by size of leaves, size of inflorescences, and by the intensity of pubescence of the lower surface of leaves. Geographically, these two taxa are not separated, and the mentioned characters are within the continuous range of variability of the species. Graffenrieda intermedia Triana, Trans. Linn. Soc. London 28: 71. 1871. Graffenrieda stenopetala Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 353. 1915. Tree 3–12 m tall; petiole 3–7 cm long; leaf blade 10–22 × 5–13 cm, oblong-ovate, apex shortly acuminate, base acute to rounded, thin-coriaceous, glabrous (rarely with hairs in the primary vein axils on the lower surface), 5–7(–9)-veined or shortly pliveined; panicle many-flowered, 10–20 cm long; flowers (4)5-merous, clustered terminally; pedicels 1 mm long, thick; hypanthium 2 mm long; calyx lobes 1–2 mm long, obtuse-triangular; petals 4–6 × 1–1.8 mm, lanceolate-oblong, acute at apex, white; anther thecae 2– 2.5 mm long, arched, dorsal spur 0.3–0.4 mm long; ovary 3-locular, granulose at apex. Tepui-slope forests, 700–2200 m; Bolívar

(Auyán-tepuí, Cerro Guanacoco, Kukenántepui, Macizo del Chimantá, Ptari-tepui, Roraima-tepui), Amazonas (Cerro Marahuaka, Sierra Parima, Sierra de la Neblina). Táchira; Guyana, Ecuador, Peru, northern Brazil. ◆Fig. 251. Graffenrieda jauana Wurdack, Mem. New York Bot. Gard. 23: 874. 1972. Shrub or small tree 1.5–5 m tall; petiole 1.5–2 cm long; leaf blade 5.5–9 × 2–4 cm, elliptic to ovate-elliptic, apex abruptly acute to acuminate, base broadly acute to obtuse, thin-coriaceous, shortly 5-pliveined; panicle many-flowered, 6–9 cm long; flowers 5merous; pedicels 2–5 mm long; hypanthium 2.5–4 mm long; calyx lobes 0.7–0.8 mm long, with external teeth projecting 0.4–0.6 mm; petals 5.3–7.5 × 2.2–3 mm, elliptic-lanceolate, apex acute, white; anther thecae 4 mm long, curved, dorsal spur 0.2–0.25 mm long; ovary 3-locular, granulose at the apex. Low forests along streamlets, 1700–2200 m; Bolívar (Cerro Jaua). Endemic. ◆Fig. 250. Graffenrieda kralii Wurdack, Mem. New York Bot. Gard. 51: 102. 1989. Shrub or small tree 1-6 m tall; petiole 0.7– 2.5 cm long; leaf blade 6–13 × 2–7 cm, elliptic to ovate-elliptic, apex gradually shortly acuminate, base broadly acute, firmly papery, 5pliveined; panicle 4–6 cm long; flowers 4merous; pedicels 1–1.5 mm long; hypanthium 1.3–2.5 mm long; calyx lobes 1–1.3 mm long, triangular, apex acute to subacuminate; petals 2.5–4 × 1–1.4 mm, lanceolate to oblong-lanceolate, apex acute, white; anther thecae 2.2–2.3 mm long, subulate, dorsal spur 0.3 mm long; ovary 3-locular, granulose at the apex, forming a lobed collar (0.2–0.3 mm high) around the style base. Low forests along streamlets, on granite or sandstone, 1400–2100 m; Bolívar (Cerro Jaua), Amazonas (Serranía Uasadi?, Serranía Yutajé?, Sierra de la Neblina). Endemic. Graffenrieda lanceolata Gleason, Fieldiana, Bot. 28: 432. 1952. Shrub 0.3–1 m tall; petiole 0.3–0.7 cm long; leaf blade 4–8 × 0.5–1.3 cm, narrowly lanceolate, apex narrowly acute, base obtuse to rounded, rigid, 3-veined; few-flowered panicle; flowers subsessile, 4-merous, verticillately clustered in intervals along the in-

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florescence axis or terminally on the short side branches; hypanthium 4 mm long; calyx lobes 3 or 4, rounded, 1 mm long; petals and anthers unknown; ovary 3-locular, glabrous. Along rocky banks on tepui summits, 1000– 1600 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 242. Graffenrieda miconioides Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 118. 1852. Graffenrieda floribunda Triana, Trans. Linn. Soc. London 28: 71, pl. 5, fig. 58a. 1871. Tree 3–15 m tall; petiole 2–5 cm long; leaf blade 14–24 × 6–15 cm, elliptic to oblong-elliptic, apex acute or shortly acuminate, base obtuse, thin-coriaceous, 3-veined with the primary laterals very close to the margin, short hairs (1–2 mm long) in the primary vein axils on the lower surface; panicle many-flowered, 20–35 cm long; flowers 4merous, clustered in intervals along the branchlet; pedicels 1–1.5 mm long; hypanthium 2.5 mm long; calyx margin dehiscing at anthesis into (3)4 lobes 1 mm long, broadly ovate; petals 5–6 × 2–4 mm, obovate to oblong-lanceolate, apex abruptly acute to obtuse, white to pinkish; anther thecae 3–3.5 mm long, arched, dorsal spur 0.6–0.7 mm long; ovary 3-locular, the apex scurfy. Lowland forests, 100–400 m; Bolívar (Río Apacará), Amazonas (Río Negro). Colombia (Vaupés), Ecuador, Peru, Amazonian Brazil. Graffenrieda obliqua Triana, Trans. Linn. Soc. London 28: 71. 1871. Shrub or small tree 2–6 m tall; petiole 0.7–2.5 cm long; leaf blade 3.5–10 × 2–4.5 cm, elliptic-lanceolate, the apex gradually shortly acuminate, base broad and frequently unequal-sided, thin-coriaceous, 5pliveined, secondary veins inserting subalternately, frequently with hairs to 6 mm long in the vein axils on the lower surface; panicle few-flowered, 2–4 cm long; flowers 4-merous; pedicels 1–1.5 mm long; hypanthium 1.5–2.2 mm long; calyx lobes 1 mm long, lanceolate; petals 2.5 × 0.8 mm, oblong-lanceolate, apex acuminate, white; anther thecae 2 mm long, dorsal spur 0.2–0.3 mm long; ovary 2-locular, apex granulose, slightly 4-lobed. Low forests of tepui slopes, 400–2500 m; Bolívar (Cerro Jaua, Gran Sabana, Macizo del Chimantá, Macizo Ilú-Tramen, Ptari-tepui, Roraimatepui, Uaipán-tepui). Guyana. ◆Fig. 240.

Graffenrieda patens Triana, Trans. Linn. Soc. London 28: 70. 1871. Woody liana to 15 m tall; petiole 1–3 cm long; leaf blade 7–18 × 3.5–9 cm, narrowly ovate to oblong-ovate, apex shortly acuminate, base rounded to cordate, thin-coriaceous, 5-veined; inflorescence many-flowered; flowers 5-merous, clustered terminally on divaricate long branches; pedicels 0.5–1.5 mm long; hypanthium 2–2.5 mm long; calyx in bud hyaline, dehiscing at anthesis irregularly, with small, thickend external teeth; petals 3.5–4.5 × 3–4 mm, broadly ovate, apex rounded, white; anther thecae 3.2–3.5 mm long, dorsal spur 0.3–0.4 mm long; ovary 3locular, glabrous. Evergreen lowland forests, 100–200 m; Amazonas (Yavita). Colombia (Amazonas, Vaupés, Vichada). Graffenrieda pedunculata Gleason, Mem. New York Bot. Gard. 8: 135. 1953. Shrub 0.2–1 m tall; young branchlets with a succulent cortex parenchyma; lower surface of leaves with a densely appressed, amorphous, silvery-grayish cover; petiole 0.5–1 cm long; leaf blade 4–6.5 × 0.8–2 cm, narrowly oblong to lanceolate-oblong, apex shortly acute to obtuse, base narrowly acute, thin-rigid, 1-veined; flowers 4-merous, verticillately clustered in intervals along the unbranched inflorescence axis; pedicels 1–2 mm long, thick; hypanthium 5–8 mm long; calyx calyptrate, dehiscing at anthesis circumscissilely; petals 8–13 × 7–12 mm, broadly elliptic, apex apiculate, white flushed with pink; anther thecae 7 mm long, slightly arched, dorsal spur 0.5–0.7 mm long; ovary 2-locular, apex slightly scurfy, with a tube-like extension (0.5 mm long) around the style base. Herb meadows with shrubs, on granite or sandstone, 1300–2000 m; Amazonas (Cerro Cuao, Cerro Sipapo). Endemic. ◆Fig. 239. Technically Graffenrieda pedunculata should be placed within Centronia because of its calyptrate calyx completely dehiscing at anthesis. But its similarity with G. versicolor justifies a provisional position within Graffenrieda. Graffenrieda polymera Gleason, Bull. Torrey Bot. Club 58: 420. 1931. Small tree 2–8 m tall; branchlets, lower surface of leaves, and inflorescences densly covered with fine, scurfy, ferruginous (later

Graffenrieda 327

grayish brown) indumentum; petiole 1–2.5 cm long; leaf blade 8–20 × 5–9 cm, oblong-elliptic to elliptic, apex shortly blunt-acuminate, base broadly acute to rounded, rigid-coriaceous, 3-veined with the primary laterals near the margin; panicle scarcely branched; flowers (4)5- or 6(8)-merous; pedicels 3–5 mm long, sometimes not well differentiated from the attenuated hypanthium base; margin of calyx 2–3 mm, entire or oblately lobed; petals obovate, apex rounded; anther thecae 6–8.5 mm long, dorsal spur 2–3 mm long; ovary 3- or 4-locular, glabrous. Venezuela, Brazil; 2 subspecies, both in the flora area. Graffenrieda polymera is a variable taxon, which will deserve at least a further subspecific differentiation. Collections from Cerro Sipapo [Maguire & Politi 27978 (F, NY, US)] and Cerro Duida [Steyermark 58307 (F)] with very large leaves, a ferruginous indumentum on the lower surface, and relatively small capsules certainly represent a separate taxon, but presently only fruiting material is known. Other collections from Cerro Aracamuni [Liesner & Carnevali 22423 (MO, US), 22699 (MO,US), 22730 (MO, US)] resemble in habit G. pedunculata by their crassulent branchlets and the oblong-lanceolate leaf shape, but have to be placed within the G. polymera-complex because of their floral characters. Graffenrieda polymera is closely related to G. caryophyllea, and both taxa need a more detailed revision. Key to the Subspecies of G. polymera 1. Pedicels 1 mm diameter; hypanthium and calyx 6–7 mm long; petals 9–11 × 7–9 mm, white ............. subsp. neblinensis 1. Pedicels 1.5–2 mm diameter; hypanthium and calyx 9–11 mm long; petals 14–17 × 10–12 mm, white flushed with pink .................................. subsp. polymera G. polymera subsp. neblinensis Wurdack, Mem. New York Bot. Gard. 10(5): 151. 1964. Scrub forests on sandstone, 1300–1900 m; Amazonas (Sierra de la Neblina). Brazil (Serra Pirapucú). G. polymera subsp. polymera Scrub forests on sandstone, 1200–2300 m; Bolívar (Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Duida, Cerro Huacha-

macari, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina?). Endemic. ◆Fig. 249. Graffenrieda reticulata Wurdack, Mem. New York Bot. Gard. 10(1): 110. 1958. Tree 4–8 m tall; young branchlets slightly quadrangular, with a succulent cortex parenchyma; petiole 1–2 cm long; leaf blade 6–12 × 3–6 cm, elliptic, apex rounded or slightly retuse, base broadly acute, rigid, 3-veined (seldom 5 primary veins, with the outer laterals close to the margin and very weak); flowers 4-merous; pedicels 2 mm long, thick; hypanthium 4–6 mm long; calyx margin dehiscing at anthesis into 3 or 4 ovate, persistent lobes; petals 5.5–8 × 3.5–5.5 mm, obovate, apex rounded, white; anther thecae 5.5 mm long, dorsal spur 0.5 mm long; ovary 3- or 4locular, glabrous. Low forests on tepui slopes and summits, 1600–2100 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 238. Graffenrieda rotundifolia (Bonpl.) DC., Prodr. 3: 106. 1828. —Rhexia rotundifolia Bonpl., Rhex. 66, pl. 25. 1812. Deciduous shrub or small tree 2–8 m tall; young leaves and branchlets densely covered with a brownish indumentum, but soon glabrescent; petiole 2–3.5 cm long; leaf blade 8– 23 × 7–18 cm, suborbicular to elliptic-ovate, apex and base obtuse to rounded, thin-coriaceous, very shortly 3-pliveined; flowers 5merous; pedicels 3–8 mm long; hypanthium 5–7 mm long; margin of calyx 2–2.5 mm, entire or weakly 5-lobed; petals 12–15 × 11.5– 13 mm, broadly obovate, apex rounded, pink; anther thecae 10–11 mm long, dorsal spur 2– 2.5 mm long; ovary 3- or 4(5)-locular, glabrous. Cracks and scrub forests on granitic outcrops (lajas), 50–300 m; Bolívar (Río Maniapure, Río Parguaza, Río Suapure), Amazonas (Nericagua, Puerto Ayacucho, Río Asisa, Río Parhueña, Río Yureba). Colombia (Vichada). ◆Fig. 252. Graffenrieda rufa Wurdack, Mem. New York Bot. Gard. 10(1): 109. 1958. Shrub 2–5 m tall; branchlets, lower surface of leaves, and inflorescences with a reddish brown (later ash gray), amorphous cover; petiole 0.8–1.5 cm long; leaf blade 1.5– 7 × 2–4 cm, narrowly ovate, apex gradually blunt-acuminate, base weakly cordate, thincoriaceous, 5-veined; panicle few-flowered; flowers 5-merous; pedicels 0.5–1.7 mm long;

328

M ELASTOMATACEAE

hypanthium 4–5 mm long; calyx lobes 3–4 mm long, acute; petals 8–8.5 × 6 mm, obovate, apex rounded, pink; anther thecae 4– 4.5 mm long, dorsal spur 1 mm long; ovary 3locular, densely granulose at the apex, forming a short collar around the style base. Exposed sandstone bluffs, 1200–1900 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. Graffenrieda rupestris Ducke, Arq. Inst. Biol. Veg. 2: 66. 1935. Tree 5–15 m tall; petiole 3–9 cm long; leaf blade 17–41 × 8.5–22 cm, oblong-elliptic to elliptic-ovate, apex obtuse and very shortly acuminate, base rounded to cordate, coriaceous, 3-veined or pliveined with the primary laterals near the margin; panicle many-flowered, 25–40 cm long; flowers 5-merous; pedicels 1–2 mm long, thick; hypanthium 4– 9 mm long; calyx margin dehiscing at anthesis irregularly into 2 or 3 ovate persistent lobes (3–5 mm long); petals 11–17 × 7–9.5 mm, obovate, apex rounded, white; anther thecae 6–7 mm long, dorsal spur 1.5–2 mm long; ovary (4)5-locular, glabrous. Forest edges along rocky river banks, mainly lowlands, ascending to 900 m; Bolívar (Cerro Guaiquinima, El Paují, Icabarú, Río Apacará, Río Caroní, Río Ichún, Río Tírica, Río Tonoro), Amazonas (Caño Canarabén, Cerro Aratitiyope, bases of Cerro Sipapo and Cerro Cuao, Río Atabapo, Río Autana, Río Casiquiare, Río Matacuni, Río Yureba). Colombia (Vaupés), Brazil (Amazonas). ◆Fig. 254. Graffenrieda sessilifolia Triana, Trans. Linn. Soc. London 28: 70. 1871. Shrub or small tree 0.3–5 m tall; branchlets of small plants with a succulent cortex parenchyma; petiole 0.1–0.7 cm long; leaf blade 5–14 × 3.5–9 cm, oblong-ovate to ovate, apex broadly acute to rounded, base cordate, coriaceous, 5(7)-veined or pliveined with the primary laterals near the margin; flowers 5-merous; pedicels 1–3 mm long, thick; hypanthium 3.5–7 mm long; calyx margin at anthesis dehiscing irregularly into 2 or 3 ovate persistent lobes; petals 9.5–12 × 7–9 mm, obovate, apex rounded; anther thecae 6–6.5 mm long, dorsal spur 0.4–1 mm long; ovary 2- or 3-locular, glabrous. Venezuela; 4 subspecies, all in the flora area.

Key to the Subspecies of G. sessilifolia 1. Ovary apex with a dentate tube (1 mm long) around the style base ..................... .............................. subsp. occidentalis 1. Ovary apex truncate ........................... 2 2. Anther connective conspicuously covered with hairs between thecae ... subsp. A 2. Anther connective with few hairs or glabrous .................................................. 3 3. Lower surface of leaf persistently scurfy; petals pink ............... subsp. cardonae 3. Lower surface of leaf caducously scurfy; petals white ........... subsp. sessilifolia G. sessilifolia subsp. cardonae Wurdack, Mem. New York Bot. Gard. 10(5): 151. 1964. Lower surface of leaves densely and persistently scurfy, chocolate brown. Scrub forests, herb meadows, 1600–2400 m; Bolívar (Auyán-tepui). Endemic. ◆Fig. 245. G. sessilifolia subsp. occidentalis Wurdack, Mem. New York Bot. Gard. 10(1): 110. 1958. Lower surface of leaves glabrous; petals pink; anther connective glabrous. Scrub forests, among boulders, 1100–2200 m; Amazonas (Cerro Coro Coro, Cerro Yaví, Cerro Yutajé). Endemic. G. sessilifolia subsp. sessilifolia Lower surface of the younger leaves sparsely scurfy, soon glabrescent. Scrub forests, herb meadows, forest edges, 1700–2200 m; Bolívar (Los Testigos, Macizo del Chimantá, Macizo Ilú-Tramen, Ptari-tepui, Roraima-tepui). Endemic. G. sessilifolia subsp. A Lower surface of leaves glabrous; petals white or tinted pink. Scrub forests, forest edges, sandstone cliffs, 300–1200 m; Bolívar (Arekuna, Cerro Guaiquinima, Cerro Sarisariñama, Chiguao, upper Río Caura, middle Río Paragua, San Salvador de Paúl). Endemic. Graffenrieda sipapoana Wurdack, Phytologia 14: 266. 1967. Small tree to 6 m tall; branchlets quadrangular; petiole 4–7 cm long, with a pair of thick, flattened protuberances (2–5 mm di-

Graffenrieda 329

ameter) next to the base of the blade; leaf blade 12–18 × 8–14 cm, elliptic-ovate to oblong-elliptic, apex rounded or broadly bluntobtuse, base cordate, coriaceous, 7- or 9pliveined; primary veins on the lower surface of leaves covered with reddish brown hairs; panicle many-flowered; flowers 5-merous; pedicels 2.5–3 mm long; hypanthium (in fruit) 6.5–7.5 mm long; calyx lobes 2 or 3, persistent; petals and anthers unknown; ovary 3-locular, apex with a dentate tube (2– 2.5 mm long) around the style base. Forest edges along rocky river banks, 200–1500 m; Amazonas (Cerro Sipapo). Endemic. Graffenrieda steyermarkii Wurdack, Mem. New York Bot. Gard. 10(5): 152. 1964. Shrub 0.3–1.6 m tall; branchlets, lower surface of leaves and inflorescences with a brownish, scurfy cover; petiole 0.5–2 cm long; leaf blade 1–2 × 0.4–0.7 cm, oblong-lanceolate, apex abruptly acute, base acute, rigid, 3-veined (inconspicuously pliveined); panicle 3–7-flowered; flowers 4-merous; pedicels 2.5–4 mm long; hypanthium 2.5 mm long; calyx lobes 0.6 mm long, oblately rounded; petals 9 × 4–5 mm, obovate-elliptic, apex abrupty acute, white; anther thecae 3– 3.2 mm long, dorsal spur 0.1 mm long; ovary 3-locular, densely granulose at the apex, forming a collar (0.5–1 mm high) around the style base. Bonnetia tepui forests, 1800– 2300 m; Bolívar (Los Testigos, Macizo del Chimantá). Endemic. ◆Fig. 243. Graffenrieda tricalcarata Gleason, Bull. Torrey Bot. Club 58: 421. 1931. Small tree 3–8 m tall; petiole 2–3 cm long; leaf blade 5–10.5 × 2.5–5 cm, narrowly ovate, apex gradually long-acuminate, base obtuse to rounded, thick-membranous, 5-veined, inconspicuously fine-setulose along the primary vein axils beneath; flowers 5(6)merous; pedicels 11.5 mm long; hypanthium 2–2.5 mm long; calyx hyaline, dehiscing at anthesis, only triangular external teeth (0.4 mm long) remaining; petals 5 × 2 mm, oblong-lanceolate, apex long-acuminate, white; anther thecae 2 mm long, dorsal spur 0.7 mm long; ovary 3(4)-locular, pulverulent at the apex. Tepui-slope forests, 1100–1800 m; Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Parú). Endemic. ◆Fig. 246.

Graffenrieda versicolor Gleason, Mem. New York Bot. Gard. 8: 136. 1953. Shrub or small tree 1–8 m tall; young branchlets with a succulent cortex parenchyma; petiole 0.5–1.5 cm long; leaf blade 4– 10 × 1.5–3.5 cm, elliptic-oblong to lanceolateelliptic, apex rounded, base obtuse to rounded, rigid-coriaceous, 1-veined; panicle scarcely branched; flowers 4(5)-merous; pedicels 1–3 mm long; hypanthium 5 mm long; calyx margin dehiscing at anthesis irregularly into 2 or 3 ovate persistent lobes (2–3 mm long); petals 8–9 × 8–9 mm, broadly obovate, apex rounded, pink; anther thecae 5.5–6.5 mm long, dorsal spur 1 mm long; ovary 3-locular, glabrous, apex with a dentate tube (1–1.5 mm long) around the style base. Low forests and scrub forests, on granite or sandstone, 1100–2200 m; Bolívar (Cerro Guanay), Amazonas (Cerro Coro Coro, Cerro Sipapo, Cerro Yutajé). Endemic. ◆Fig. 241. Graffenrieda weddellii Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 117. 1852. Graffenrieda ovalifolia Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 117, t. 5, fig. 4. 1852. Shrub or small tree 1–3(–10) m tall; petiole (0.7-)1.5–3(–4) cm long; leaf blade 8–15 × 3–9 cm, ovate to oblong-ovate, apex bluntly acute to acuminate, base rounded to cordate, coriaceous, 5- or 7-veined, the lower surface usually inconspicuously setulose in the primary vein axils; panicle multiflorous, 6–10 cm long; flowers 5-merous; pedicels 1.5–3.5 mm long; hypanthium 2.5–4.5 mm long; calyx margin dehiscing at anthesis into several lanceolate to ovate lobes; petals 7.5–14 × 2.5–5.5 mm, narrowly obovate, apex acute, white; anther thecae 3.5–5.5 mm long, dorsal spur 1–2 mm long; ovary 3-locular, apex granulose, with a collar of 6–8 lobes (0.5–1 mm long) around the style base. Rocky savannas, 100–1400 m; Bolívar (Gran Sabana, Karaurín-tepui, Los Testigos, Macizo de Chimantá, Río Canaracuni, Roraima-tepui, San Ignacio de Yuruaní, Urimán), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Parú, Río Casiquiare). Distrito Federal; Colombia (Vaupés, Amazonas), Guyana (Pakaraima Mountains), northern and central Brazil, Bolivia. ◆Fig. 247.

330

M ELASTOMATACEAE

Fig. 238. Graffenrieda reticulata

Fig. 239. Graffenrieda pedunculata

Fig. 240. Graffenrieda obliqua

Graffenrieda 331

Fig. 241. Graffenrieda versicolor

Fig. 242. Graffenrieda lanceolata

Fig. 243. Graffenrieda steyermarkii

332

M ELASTOMATACEAE

Fig. 244. Graffenrieda hitchcockii

Graffenrieda 333

Fig. 245. Graffenrieda sessilifolia subsp. cardonae

Fig. 246. Graffenrieda tricalcarata

Fig. 247. Graffenrieda weddellii

334

M ELASTOMATACEAE

Fig. 248. Graffenrieda fruticosa

Fig. 249. Graffenrieda polymera subsp. polymera

Graffenrieda 335

Fig. 250. Graffenrieda jauana

Fig. 251. Graffenrieda intermedia

336

M ELASTOMATACEAE

Fig. 252. Graffenrieda rotundifolia

Graffenrieda 337

Fig. 253. Graffenrieda caryophyllea

338

M ELASTOMATACEAE

Fig. 254. Graffenrieda rupestris

17. HENRIETTEA DC., Prodr. 3: 178. 1828. Phyllopus DC., Prodr. 3: 177. 1828. by Navina G. Luckana and Paul E. Berry Shrubs to small trees. Leaves usually large, entire and ciliate. Flowers 5- or 6merous, either few-fasciculate at the branchlet nodes below the leaves or in the axils of mature leaves; bracteoles basal, small. Hypanthium terete, externally pubescent, glabrous or pubescent with retrorse hairs within; calyx limb entire to lobed and per-

Henriettea 339

sistent, the external teeth usually inframarginal; petals white or rarely pink, usually puberulous, clawed, obtuse to acute at the apex, with an external subapical tooth. Stamens isomorphic, glabrous; anthers usually attenuate-rostrate and with a small dorsally inclined pore; connective usually slightly bilobed at the base but completely adnate to the thecae. Ovary 5-locular and completely inferior, truncate or with a short column; style glabrous or puberulous; stigma not or slightly clavate-expanded. Fruit a berry. Seeds numerous, ovoid to oblong-pyramidal, papillate. Mexico, Central America, Jamaica, Colombia, Venezuela, Trinidad, Guyana, Suriname, Ecuador, Brazil, ca. 15 species, 11 in Venezuela, all in the flora area. Key to the Species of Henriettea 1. 1. 2(1). 2. 3(1). 3. 4(3).

4.

5(4).

5.

6(5).

6.

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

Flowers 6-merous ....................................................................................... 2 Flowers 5-merous ....................................................................................... 3 Leaves lanceolate to oblong-lanceolate, apex acuminate; petioles 1–1.5 cm long ........................................................................................... H. martiusii Leaves elliptic-ovate, apex obtuse; petioles 2.5–3 cm long ...... H. mucronata Hypanthium within (below the torus) densely retrorse-strigulose .......... 4 Hypanthium within (below the torus) glabrous or very sparsely retrorsestrigulose ................................................................................................ 7 Upper surface of leaf blades moderately to densely bullate-setulose, the bases of the trichomes expanded and 0.5–0.7 mm wide; anther thecae 3–4 mm long ............................................................................ H. granulata Upper surface of leaf blades glabrous or sparsely strigulose or appressedsetulose, the bases of the trichomes 0.1–0.3 mm wide; anthers thecae 5–8 mm long ........................................................................................... 5 Upper surface of leaf blades sparsely setulose with trichomes 0.5–1.2 mm long; pubescence of the hypanthium loose, mostly 1.5–2 mm long ................................................................................................. H. horridula Upper surface of leaf blades glabrous or sparsely strigulose with trichomes 0.1–0.25 mm long; pubescence of the hypanthium appressed, mostly 0.2–1 mm long ............................................................................ 6 Upper surface of leaf blades sparsely and persistently strigulose with trichomes mostly 0.2–0.25 mm long; calyx lobes usually 2–3 mm long .................................................................................................... H. stellaris Upper surface of leaf blades glabrous or with very sparse, quickly deciduous trichomes ca. 0.1 mm long; calyx lobes mostly 4–5 mm long ................................................................................................ H. spruceana Calyx lobes 4–6 mm long; style moderately to densely fine-setulose ...... 8 Calyx lobes 0.3–0.6(–2) mm long; style glabrous or very sparsely finesetulose ................................................................................................... 9 Upper surface of leaves persistently short-setose; hypanthial trichomes 2–3 mm long ........................................................................ H. maroniensis Upper surface of leaves caducously setulose; hypanthial trichomes 1– 1.5 mm long ................................................................................. H. succosa Pedicels ca. 1 mm long at anthesis; corolla 4.5–5 mm long; anthers ca. 4 mm long ............................................................................... H. patrisiana Pedicels 2–5 mm long at anthesis; corolla 7–10 mm long; anthers 5–8 mm long ....................................................................................................... 10

340

10(9). 10.

M ELASTOMATACEAE

Lower surface of leaves with stellate trichomes lacking a terminal seta; petals white ............................................................................. H. multiflora Lower surface of leaves with trichomes at least in part with a terminal seta; petals pink .......................................................................H. ramiflora

Henriettea granulata O. Berg ex Triana, Trans. Linn. Soc. London 28: 144. 1871. Shrub or small tree 3–8(–20) m tall; flowers nearly sessile; petals white, style glabrous. Along forest edges, along streams in white-sand savannas, shrublands, 100–900 m; Bolívar (Canaima, base of Cerro Guaiquinima, Parupa, Río Asa, Río Cuyuní, San Ignacio de Yuruaní), Amazonas (Caño Caname, Caño Cupueni near San Fernando de Atabapo, base of Cerro Yapacana, Río Negro, Río Pasimoni, Río Yatúa). Colombia, Guyana, Brazil. ◆Fig. 255. Henriettea horridula Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 182. 1905. Shrub or small tree 4–8 m tall; petals white, stamens purple. Riparian forests, edges of granitic outcrops, 100–200 m; Amazonas (mouth of Río Casiquiare, Pimichín, Yavita). Colombia (Guainía), Brazil (Amazonas). Henriettea maroniensis Sagot, Ann. Sci. Nat. Bot. sér. 6, 15: 330. 1883. Shrub or small tree 3–12 m tall; petals white. Rocky montane savannas, riparian forests, 500–1400 m; Bolívar (Altiplanicie de Nuria, Amaruay-tepui, Cerro Abismo, El Zamuro, Ilú-tepui, Macizo del Chimantá [Toronó-tepui], Ptari-tepui, Río Samay, tributaries of Río Karaurín and Río Asadón). Anzoátegui; Colombia (Vaupés), Guyana, Suriname, French Guiana, Brazil. ◆Fig. 257. Henriettea martiusii (DC.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 105. 1852. —Phyllopus martiusii DC., Prodr. 3: 177. 1828. Henriettea oblongifolia Naudin, Ann. Sci. Nat. Bot. sér. 3, 4: 56. 1845. Henriettea orinocensis Naudin, Ann. Sci. Nat. Bot. sér. 3, 4: 57, pl. 3, fig. 6. 1845. Shrub or small tree 1.5–4(–6) m tall; flowers solitary or in groups of 3, petals white. Riparian forests, 100–200 m; Amazonas (Caño Caname, Caño Yapacana, Río Atabapo, Río Casiquaire, Río Pasiba, Río

Pasimoni, Río Samariapo, mouth of Río Ventuari, San Antonio del Sipapo, San Carlos de Río Negro, San Juan de Ucata, Yavita). Colombia (Vichada), Brazil (Amazonas). ◆Fig. 260. Henriettea mucronata (Gleason) Renner, Mem. New York Bot. Gard. 50: 52. 1989. —Loreya mucronata Gleason, Mem. New York Bot. Gard. 8: 143. 1953. —Boyo boyo, Guayaba de danto rebalsera. Small tree 2–12 m tall; young stems 4angled, densely strigillose; petals reddish. Evergreen lowland forests, 100–200 m; Amazonas (base of Cerro Sipapo, El Pozo southeast of San Fernando de Atabapo, Río Cuchakén in Río Atabapo basin, Río Yatúa). Colombia (Vichada). ◆Fig. 258. Henriettea multiflora Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 105. 1852. Shrub or small tree 3–10 m tall; petals white. Seasonally flooded riparian forests, wet savannas, near sea level to 400 m; Delta Amacuro (Jotajana, Río Amacuro), Bolívar (Río Cuyuní). Trinidad, Guyana, Suriname, French Guiana. Henriettea patrisiana DC., Prodr. 3: 178. 1828. Shrub or small tree 1.5–7 m tall; petals white. 200–1000 m; Bolívar (Represa Guri, Río Caura), Amazonas (Cerro Yutajé). Monagas; Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 259. Henriettea ramiflora (Sw.) DC., Prodr. 3: 178. 1828. —Melastoma ramiflora Sw., Fl. Ind. Occid. 69. 1788. — Henriettella ramiflora (Sw.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 106. 1852. Henriettea surinamensis Miq., Stirp. Surinam. Select. 45. 1850 [1851]. Henriettea trinervia Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 105. 1852. Henriettea succosa var. guianensis Gleason, Bull. Misc. Inform. Kew 1939: 551. 1939.

Henriettea 341

Fig. 255. Henriettea granulata

Fig. 256. Henriettea stellaris

342

M ELASTOMATACEAE

Fig. 257. Henriettea maroniensis

Henriettea 343

Fig. 258. Henriettea mucronata

344

M ELASTOMATACEAE

Fig. 259. Henriettea patrisiana

Fig. 260. Henriettea martiusii

Henriettea 345

Fig. 261. Henriettea spruceana

Fig. 262. Henriettea succosa

346

M ELASTOMATACEAE

Shrub to small tree 4–15(–18) tall; petals pink to light purple; filaments, stigma, and style white. Evergreen lowland to montane forests, 50–1300 m; Bolívar (base of Auyántepui, Kavanayén, near Luepa, Río Paramichí, Santa Elena de Uairén), Amazonas (lower slopes of Cerro Huachamacari, Río Baría, Río Guayapo, San Carlos de Río Negro). Barinas, Mérida, Táchira; Guatemala, Honduras, Nicaragua, Jamaica, Trinidad, Guyana, Suriname, French Guiana, Brazil. Henriettea spruceana Cogn. in Mart., Fl. Bras. 14(4): 530. 1888. Tree 3–15 m tall, ants sometimes present in hollow stems; petals white. Seasonally flooded riparian forests, 50–300 m; Bolívar (Caño Pablo on Río Caura, Río Tabaro on Río Nichare), Amazonas (near La Esmeralda, near Puerto Ayacucho, Río Atabapo, Río Baría, Río Casiquiare, Río Cataniapo, Río Emoni, Río Negro, Río Yatúa, San Carlos de Río Negro). Colombia (Vaupés), Brazil (Amazonas: Rio Negro basin). ◆Fig. 261. Henriettea stellaris O. Berg ex Triana, Trans. Linn. Soc. London 28: 145. 1871.

Tree 3–10 m tall; petals white. Riparian forests and savannas, 100–500 m; Bolívar (Río Acanán, Río Caroni near Urimán, Río Caura, Río Hacha, Río Icabarú, Río Paragua above Salto Pará, Río Trueno at base of Cerro Guaiquinima), Amazonas (Río Atacavi, Río Baría, Río Cunucunuma, Río Mawarinuma). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 256. Henriettea succosa (Aubl.) DC., Prodr. 3: 178. 1828. —Melastoma succosa Aubl., Hist. Pl. Guiane 418, t. 162. 1775. —Lala-ak (Arekuna). Tree 3–12 m tall; petals white to pink. Riparian and nonflooded lowland forests, 100–1200 m; Bolívar (Arabupu, Río Uaiparú at junction with Río Icabarú), Amazonas (road between San Carlos de Río Negro and Solano). Zulia; southern Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 262. Henriettea succosa is similar to H. maroniensis, but has shorter and caducous setae on the upper leaves and hypanthial hairs only 1–1.5 mm long.

18. HENRIETTELLA Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 107. 1852. by Paul E. Berry Shrubs or small trees. Leaves opposite, isomorphic. Flowers 4- or 5-merous, multifasciculate at the branchlet nodes below the leaves; bracteoles basal, mostly minute. Hypanthium terete, glabrous within; calyx truncate or shortly lobed, glabrous within, with minute external teeth; petals white, ovate to lanceolate and bluntly acute, usually with an external mucro, glabrous or externally puberulous, usually cross-ridged or appendaged within. Stamens isomorphic, glabrous; anthers with 1 or 2 pores; connective not or slightly prolonged, usually not appendaged. Ovary (3-)4- or 5-locular, completely inferior, mostly glabrous at the apex; style glabrous; stigma not or somewhat expanded. Fruit a berry. Seeds numerous, obovateangulate. Guatemala to Panama, West Indies, Colombia, Venezuela, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia; 35–40 species, 10 in Venezuela, 6 of these in the flora area. Key to the Species of Henriettella 1. 1.

Leaves basally veined or shortly (< 1 cm) pliveined ......................... H. ovata Leaves 5-pliveined, the lowermost pair of primary veins diverging 1–4 cm above the base of the blade .................................................................... 2

Henriettella 347

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

Flowers 4-merous ....................................................................................... 3 Flowers 5-merous ....................................................................................... 4 Flowers sessile; leaf blades 2.5–5 cm wide, petioles 1–2.5 cm long .............................................................................................. H. heteroneura Flowers with pedicels 2.5–3.6 mm long; leaf blades 6–8 cm wide, petioles 3–4 cm long ................................................................................. H. prancei Stems and main veins on lower surface of leaves densely fine-strigulose ............................................................................................. H. steyermarkii Stems and main veins on lower surface of leaves glabrous or sparsely and deciduously strigulose ........................................................................... 5 Flowers with pedicels 4–6 mm long .............................................. H. caudata Flowers sessile .............................................................................. H. duckeana

Henriettella caudata Gleason, Recueil Trav. Bot. Néerl. 32: 210. 1935. Tree or shrub 1.5–8 m tall; leaf blade oblong-elliptic to oblong-lanceolate, apex subabruptly long-acuminate, base acute, 15–25 × 5–9 cm, chartaceous and apically distantly undulate-serrulate, 5-pliveined with laxly reticulate venules; petiole 2–5 cm long; flowers 5-merous; pedicels 4–6 mm long; hypanthium minutely glandular; petals 2.6–3 × 1.5–1.8 mm. Lowland and lower montane forests, 200–700 m; Bolívar (Altiplanicie de Nuria, Serranía de Imataca), Amazonas (Sierra Parima). Guyana, Suriname, French Guiana, Ecuador, Brazil (Amapá, Amazonas). Henriettella duckeana Hoehne, Anexos Mem. Inst. Butantan, Secc. Bot. 1(5): 170. 1992. Shrub or small tree 1–3(–10) m tall, vegetatively similar to Henriettella caudata, the leaf venules somewhat smaller (0.5–1 mm); flowers 5-merous, sessile; petals ca. 3.5 × 1.2 mm. Lower montane forests, ca. 300 m; Bolívar (Río Oris in upper Río Paragua basin). French Guiana, Brazil (Pará, Roraima). Henriettella heteroneura Gleason, Mem. New York Bot. Gard. 8: 143. 1953. Shrub, the young growth soon glabrescent; leaf blades oblong-elliptic to oblanceolate-elliptic, subabruptly obtuse-acuminate for 0.7–2 cm at the apex, base narrowly acute to long-attenuate, 7–15 × 2.5–5 cm, entire and firmly membranous, 5-pliveined; petioles 1–2.5 cm long; flowers 4-merous, sessile and densely fasciculate; petals white, ca. 2 × 1.2 mm; ovary 3-locular, scurfy at the

apex. Tepui scrub, 1500–2000 m; Amazonas (Caño Profundo on Cerro Sipapo). Endemic. Henriettella ovata Cogn. in Mart., Fl. Bras. 14(4): 540, t. 115. 1888. Henriettella longistyla Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 366. 1915. Henriettella micrantha Gleason, Bull. Torrey Bot. Club 58: 414. 1931. Shrub or small tree 2–8 m tall; leaf blades oblong-ovate, apex acute to subacuminate, base rounded to cordulate, 7–19 × 3.5–9.5 cm, subcoriaceous, 5-veined or shortly 5pliveined; petioles 0.6–2 cm long; flowers 5merous; pedicels ca. 1.5 mm long; petals 2.1– 3 × 1.4–1.9 mm. Riparian and lower montane forests, 50–1100 m; Bolívar (Caicara, between El Callao and Supamo, Icabarú, Río Aparamán, Río Asa, Río Caura, Río Canarakuni, Río Paragua, Río Trueno), Amazonas (Caño Yutajé, Cerro Duida, Cerro Yutajé, Puerto Ayacucho, Río Coromoto, Santa Bárbara del Orinoco, Simarawochi). Apure, Monagas; Colombia, Guyana?, Brazil. ◆Fig. 265. Henriettella prancei Wurdack, Phytologia 24: 206. 1972. Small tree; leaf blades elliptic, apex narrowly acute, base broadly acute or obtuse, 11–18 × 6–8 cm, entire and coriaceous, 5pliveined; petioles 3–4 cm long; flowers 4merous, few-fasciculate; petals white, 4.7– 5.2 × 2.5–3 mm. Evergreen lowland, riparian, and caatinga forests, 100–200 m; Amazonas (Pimichín, Río Yatúa, Yavita). Brazil (Amazonas: upper Rio Negro). ◆Fig. 264.

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Henriettella steyermarkii Wurdack, Bol. Soc. Venez. Ci. Nat. 25: 54. 1963. Tree to 10 m tall; young growth densely appressed-bristly with hairs 1–2 mm long; leaf blades elliptic to ovate-elliptic, apex gradually acuminate, base broadly acute, 11–18 × 4–7 cm, membranous and slightly

ciliate-crenulate, 5-pliveined; petioles 1–1.5 cm long; flowers 5-merous; pedicels 1.5–3 mm long; petals white, ca. 2.8 × 1.5 mm. Evergreen lowland and lower montane forests, 100–800 m; Bolívar (northeast of Cerro Marutaní), Amazonas (Boca Mavaca). Endemic. ◆Fig. 263.

Fig. 263. Henriettella steyermarkii

Henriettella 349

Fig. 264. Henriettella prancei

Fig. 265. Henriettella ovata

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19. LEANDRA Raddi, Mem. Mat. Fis. Soc. Ital. Modena, Pt. Mem. Fis. 18: 385. 1820. Oxymeris DC., Prodr. 3: 190. 1828. Tschudya DC., Prodr. 3: 155. 1828. Clidemiastrum Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 87. 1852. Platycentrum Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 114. 1852. by Andreas Gröger Shrubs, rarely woody lianas, with isomorphic (anisomorphic in L. aristigera) leaves. Leaves membranous to chartaceous, seldom coriaceous, with margin mostly ciliate, 1–7-veined (occasionally weakly pliveined). Inconspicuous flowers in terminal (sometimes pseudoterminal, seldom lateral) panicles, 4- or 5(–7)-merous; inflorescence branches frequently pink to reddish violet, especially when fruiting. Hypanthium terete; interior calyx lobes usually inconspicuous, the persistent external teeth often well developed; petals white to pink, usually glabrous, linear to ovate, acute. Stamens essentially isomorphic, glabrous; anthers usually oblong; connective not or barely prolonged, usually not appendaged. Ovary 2–6-locular, mostly 1/2-inferior to completely inferior, glabrous or pubescent at the apex; style usually glabrous; stigma usually not expanded. Fruit baccate, when mature blue-violet to purpleblack (rarely white). Seeds numerous, small, ovoid-cochleate or oblong-pyramidal. Southern Mexico to northern Argentina, especially numerous in southeastern Brazil, but absent from the Antilles; more than 200 species, 26 in Venezuela, 22 of these in the flora area. Leandra belongs to the tribe Miconieae. It differs from Miconia by its narrow, acute to acuminate petals, and by the frequently dense pubescence with simple, barbellate, and/or gland-tipped hairs, mostly lacking a stellulate indumentum. Very closely related to Leandra is Ossaea, which differs in its petals with an exterior tooth. According to Triana (Trans. Linn. Soc. London 28: 90–96. 1871), several sections can be distinguished within Leandra. The most distinctive is the section Secundiflorae (see key below). Key to the Species of Leandra 1.

1.

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

Flowers borne secundly, i.e., arranged in unilateral rows on the inflorescence branchlets (most obvious in fruit); anthers oblong (with the exception of L. maguirei), rounded at the apex, and with a ventrally inclined pore; seeds ovoid-cochleate with a frequently tuberculate surface (Sect. Secundiflorae) ...................................................................... 2 Flowers not borne secundly; anthers subulate, tapering to the apex, and with a frequently dorsally inclined pore; seeds oblong-pyramidal with a smooth surface ....................................................................................... 9 Flowers 6(7)-merous ..................................................................... L. solenifera Flowers 5-merous ....................................................................................... 3 Ovary 5-locular ........................................................................................... 4 Ovary 3-locular ........................................................................................... 5 Hypanthia covered with gland-tipped hairs; calyx teeth 0.1–0.8 mm long ..................................................................................................... L. secunda Hypanthia covered with simple eglandular hairs; calyx teeth 1–2 mm long ..................................................................................... L. francavillana

Leandra 351

5(3). 5. 6(5). 6. 7(6). 7. 8(7). 8. 9(1). 9. 10(9). 10. 11(10). 11. 12(9). 12. 13(12). 13. 14(12). 14. 15(14). 15. 16(15). 16. 17(16). 17. 18(17).

Base of the leaf blade cordulate, leaf blade above rugose; anthers subulate ................................................................................................... L. maguirei Base of the leaf blade rounded to acute, leaf blade above plane; anthers oblong ..................................................................................................... 6 Branchlets and petioles with simple, eglandular hairs to 5 mm long; calyx teeth 0.2–0.5 mm long ........................................................... L. neblinensis Indumentum with hairs < 3 mm; calyx teeth 0.5–3 mm long .................. 7 Petals 3.2–3.8 mm long; calyx teeth 2.2–3 mm long; ovary apex with a bowl-shaped corona ............................................................... L. longisepala Petals 1.5–3 mm long; calyx teeth 0.5–2.1 mm long; ovary apex knobby or lobed ....................................................................................................... 8 Leaf blade 5–11 cm long; leaf margin nearly entire; inflorescence rather few-flowered, 3–9 cm long ...................................................... L. divaricata Leaf blade 13–30 cm long; leaf margin serrulate; inflorescence many-flowered, 10–25 cm long ................................................................ L. sanguinea Woody liana or creeping shrub, with internodal roots; flowers 4- or 5-merous ............................................................................................... 10 Erect shrub; flowers 5-merous ................................................................. 12 Creeping or climbing shrub in tepui vegetation; flowers 4-merous .............................................................................................. L. procumbens Liana in lowland forests; flowers 5-merous ............................................ 11 Leaves coriaceous, basally 3-veined with the primary laterals near the margin; panicle many-flowered, 8–13 cm long .....................L. candelabra Leaves membranous, 5-veined or shortly pliveined; panicle few-flowered, 2.5–4 cm long ...................................................................... L. steyermarkii Ovary 5-locular ......................................................................................... 13 Ovary 3-locular (seldom 2- or 4-locular) .................................................. 14 Petals 0.4–0.7 mm long, fringed with gland-tipped hairs; leaf apex sharply acuminate .................................................................... L. purpurea Petals 0.9–1.3 mm long, externally covered with close-pressed, eglandular hairs; leaf apex gradually acuminate ....................................... L. rufescens Lower surface of leaf blade totally glabrous; inflorescences small, lateral ................................................................................................. L. chaetodon Lower surface of leaf blade pubescent, at least on primary veins; inflorescences terminal or pseudoterminal ..................................................... 15 Indumentum of inflorescence axes and hypanthia partially gland-tipped .............................................................................................................. 16 Indumentum of inflorescence axes and hypanthia totally eglandular .............................................................................................................. 19 Petals 2–2.4 mm long, externally sparsely covered with gland-tipped and with minute stellulate hairs; mature berry white .................. L. aristigera Petals 0.3–0.6 mm long, glabrous; mature berry purple-black .............. 17 Base of the leaf blade acute to obtuse, shortly 5-pliveined; anthers 2.1– 2.4 mm long ......................................................................... L. glandulifera Base of the leaf blade cordulate, basally 7–9-veined; anthers 3–4 mm long .............................................................................................................. 18 Petioles, inflorescence axes, and hypanthia with gland-tipped hairs 0.2– 0.6 mm long (glands 0.1 mm long); upper surface of leaf blade plane, with hairs only on the veins .................................................... L. phelpsiae

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18.

Petioles, inflorescence axes, and hypanthia with gland-tipped hairs 0.3– 3 mm long (glands 0.2–0.3 mm long); upper surface of leaf blade rugose, with hairs not restricted to the veins ..................................... L. polyadena 19(15). Petals 3–5 mm long; indumentum on branchlets, petioles, inflorescence axes, and hypanthia with an underlayer of minute barbellate hairs ................................................................................................ L. lindeniana 19. Petals 0.8–2.6 mm long; indumentum without an underlayer of minute barbellate hairs .................................................................................... 20 20(19). Anther connective elevated at its dorsal base into a conspicuous, apically ascending protuberance; calyx teeth 0.2–0.5 mm long ...... L. clidemioides 20. Anther connective without this kind of appendage; calyx teeth 0.5– 1.5 mm long .......................................................................................... 21 21(20). Petals linear, 1–1.2 × 0.15–0.25 mm; petioles 1–2.5 cm long ....... L. edentula 21. Petals lanceolate, 1.5–2.6 × 0.5–1.1 mm; petioles 0.2–0.5 cm long .................................................................................................... L. gorzulae Leandra aristigera (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 185. 1886. —Clidemia aristigera Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 366. 1851. —Oxymeris aristigera (Naudin) Triana, Trans. Linn. Soc. London 28: 94. 1871. Shrub 1–2.2 m tall; branchlets, petioles, and primary veins on lower surface of leaves moderately covered with simple hairs 0.9– 3.5 mm long; inflorescence axes moderately covered with gland-tipped hairs 0.5–2 mm long; hypanthia moderately covered with gland-tipped and very sparsely with minute stellulate hairs; petiole 1–6 cm long; leaf blade often differently sized in each pair, 10– 25 × 5–10 cm, elliptic to ovate-elliptic, apex caudate-acuminate, base broadly acute to obtuse, thin, 5(7)-veined or pliveined with the inner pair of primary veins diverging to 1.5 cm above the blade base; panicle few-flowered, 5–11 cm long; pedicels 0.2–0.6 mm long; hypanthium 2–2.5 mm long; calyx teeth 0.5–1.3 mm long; petals 2–2.4 × 0.6–0.8 mm, oblong-lanceolate, externally sparsely covered with gland-tipped and with minute stellulate hairs, white; anthers 1.8–2.2 mm long, with a slightly dorsally inclined pore; ovary 3(4)-locular, sparsely puberulous and with a short fringed tube at the apex; mature berry white. Forest understory, forest edges, 100–1100 m; Bolívar (Auyán-tepui, El Paují, base of Perai-tepui, Río Erebato), Amazonas (Boca Mavaca, Río Baría, Río Cataniapo, Río Coro Coro, Río Negro, Serranía del Santo). Apure, Táchira; Amazonian region of Colombia, Ecuador, Peru, and Brazil. ◆Fig. 267.

Leandra candelabrum (J.F. Macbr.) Wurdack, Phytologia 52: 61. 1982. —Graffenrieda candelabrum J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4.1): 321. 1941. Woody liana to 10 m tall, with internodal roots; branchlets, petioles, primary veins on lower surface of leaves, inflorescence axes, and hypanthia glabrous or very sparsely covered with minute stellulate hairs; petiole 1– 2.2 cm long; leaf blade 14–18 × 4.5–6.5 cm, ovate, apex acuminate, base rounded to subacute, papery-coriaceous, 3-veined with the primary laterals near the margin; leaf margin not ciliate; panicle many-flowered, spreading, 8–13 cm long; flowers sessile; hypanthium 2.2–2.5 mm long; calyx teeth 0.4– 0.8 mm long; petals 1.1–1.6 × 0.8 mm, oblong-lanceolate, apex acuminate, whitish to reddish; anthers 2–2.2 mm long, with a slightly dorsally inclined pore; ovary 3-locular, with a densely puberulous tube at the apex. Nonflooded lowland forests, 100–300 m; Bolívar (upper Río Caura), Amazonas (San Carlos de Río Negro, base of Sierra de la Neblina). Amazon basin periphery in Colombia, Peru, and northwestern Brazil. ◆Fig. 268. Leandra chaetodon (DC.) Cogn. in Mart., Fl. Bras. 14(4): 178. 1886. —Spennera chaetodon DC., Prodr. 3: 117. 1828. —Clidemia chaetodon (DC.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 360. 1852. —Oxymeris chaetodon (DC.) Triana, Trans. Linn. Soc. London 28: 94. 1871.

Leandra 353

Ossaea trichopoda Gleason, Brittonia 2: 324. 1937. Shrub 1–3 m tall; branchlets and leaf blades glabrous; adaxial side of petioles moderately covered with gland-tipped hairs 0.5– 1 mm long and simple hairs 0.5–2 mm long; inflorescence axes and hypanthia sparsely covered with caducous gland-tipped hairs or glabrous; petiole 0.5–2 cm long; leaf blade 15–21 × 3–6 cm, elliptic to lanceolate-elliptic, apex gradually long-acuminate, base acute, membranous, 5-pliveined, the inner pair of primary veins faint, diverging 0.5–2.5 cm above the blade base; panicle few-flowered, 3–6 cm long; pedicels 0.5–1 mm long; hypanthium 2–4 mm long; calyx teeth 0.7–2 mm long; petals 1–1.5 × 0.3–0.8 mm, lanceolate to oblong-lanceolate, with an external subapical tooth projecting 0.5–1.5 mm, white; anthers 2–2.7 mm long, with a terminal to slightly dorsally inclined pore; ovary (2)3-locular, glabrous. Forest understory, 200–1100 m; Bolívar (Amaruay-tepui, slopes of Auyán-tepui, base of Cerro Guaiquinima, base of Chaco-tepui, El Paují, upper Río Caura, Río Erebato, Río Icabarú, Río Kukenán, Río Mereware, Río Nichare, upper Río Paragua, Río Suapure, Río Tírica), Amazonas (Cerro Huachamacari, base of Cerro Yutajé, Río Autana, Río Mavaca, Río Mawarinuma). Apure, Táchira; Panama, Colombia, Guyana, Amazon basin periphery in Ecuador, Peru, and northern Brazil. ◆Fig. 266. Leandra clidemioides (Naudin) Wurdack, Phytologia 55: 145. 1984. —Platycentrum clidemioides Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 114. 1852. Tschudya strigillosa Griseb., Fl. Brit. W. I. 250. 1860. Leandra hylophila Gleason, Bull. Torrey Bot. Club 68: 246. 1941. Shrub or small tree 2–5 m tall; branchlets, petioles, primary leaf veins, lower surface of leaves, inflorescence axes, and hypanthia moderately to densely covered with simple and barbellate hairs (both 0.1–0.8 mm long); petiole 1–4.5 cm long; leaf blade 7–30 × 4–12 cm, oblong-elliptic to lanceolate-elliptic, apex acute to gradually acuminate, base acute, chartaceous, 5pliveined with the inner pair of primary

veins diverging 0.5–1.5 cm above the blade base; panicle many-flowered, 4–12 cm long; pedicels 0.5–0.8 mm long; hypanthium 1.5–2 mm long; calyx teeth 0.2–0.5 mm long; petals 0.8–2 × 0.2–0.5 mm, narrowly oblong, white; anthers 1.5–2.5 mm long, with a ventrally inclined pore, base of connective with a conspicuous dorsal protuberance ascending apically; ovary 3-locular, glabrous or very sparsely puberulous, with a small collar at the apex. Understory of montane forests, 1000–1800 m; Bolívar (Sierra de la Neblina). Sucre; Colombia, Trinidad, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Leandra divaricata (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 196. 1886. —Clidemia divaricata Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 373. 1852. Clidemia rhamnifolia var. macrodon Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 373. 1852. —Leandra rhamnifolia var. macrodon (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 195. 1886. Subshrub 0.3–1.3 m tall, branches often decumbent and rooting at the nodes; branchlets, petioles, primary veins on lower surface of leaves, inflorescence axes, and hypanthia moderately covered with simple hairs 0.3–1.3 mm long; petiole 1–4 cm long; leaf blade 5–11 × 2.5–6 cm, elliptic to ovateelliptic, apex short-acuminate, base broadly acute to obtuse, chartaceous, shortly 5(7)pliveined with the inner pair of primary veins diverging up to 1 cm above the blade base; panicle rather few-flowered, 3–9 cm long; flowers aggregated on the lateral inflorescence branches; secund arrangement more obvious in fruit; pedicels 0.1–0.5 mm long; hypanthium 1.6–1.8 mm long; calyx teeth 0.5–1.4 mm long; petals 1.7–3 × 0.5–0.8 mm, oblong-lanceolate, white; anthers 1–1.5 mm long; ovary 3-locular, lobed at the apex. Understory of dense forests, forest edges, 50–1000 m; Delta Amacuro (Manoa, Río Amacuro), Bolívar (Aparurén, base of Chacotepui, El Paují, base of Perai-tepui, Río Paragua, Urimán). Guyana, Suriname, French Guiana, northeastern Brazil (Amapá, Pará). ◆Fig. 269. Putative hybrids with Leandra sanguinea are reported from the flora area (Fernández 3243, PORT, US, VEN), and with L. solen-

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ifera from adjacent Guyana (McDowell & Gopaul 2702, F). Leandra francavillana is closely related to L. divaricata, but is geographically separate. Leandra edentula Gleason, Fieldiana, Bot. 28: 434. 1952. Shrub 1.5–3 m tall; branchlets, petioles, and primary veins on lower surface of leaves densely covered with simple, or occasionally barbellate hairs 0.1–1.1 mm long; inflorescence axes and hypanthia with simple hairs; petiole 1–2.5 cm long; leaf blade 9–15 × 3–5 cm, oblong-lanceolate, apex gradually acuminate, base broadly acute to obtuse, the upper surface rugose, the lower surface foveolate, firm-chartaceous, 5(7)-veined or shortly pliveined with the inner pair of primary veins diverging 0.5 cm above the blade base; panicle rather few-flowered, 8–14 cm long; pedicels 0.2–0.4 mm long; hypanthium 2–2.3 mm long; calyx teeth 0.5–0.8 mm long; petals 1–1.2 × 0.15–0.25 mm, narrowly oblong, white; anthers 2.5 mm long; connective with a blunt ascending protuberance dorsally; ovary (2)3-locular, with a very sparsely puberulous collar at the apex. Forest understory of tepui slopes and ridges, 1100–2200 m; Bolívar (Sororopán-tepui). Guyana. Leandra francavillana Cogn. in Mart., Fl. Bras. 14(4): 197, pl. 43, fig. 1. 1886. Shrub 1–5 m tall; branchlets, petioles, primary veins on lower surface of leaves, inflorescence axes, and hypanthia densely to moderately covered with simple hairs 0.8– 1.4 mm long, on lower surface of leaf blade frequently as an appressed silky indumentum; petiole 1–4 cm long; leaf blade 8–18 × 3–8 cm, narrowly ovate to oblong-ovate, apex gradually acuminate, base obtuse to rounded-truncate, membranous, 7-veined or pliveined with the inner pair of primary veins diverging to 1.5 cm above the blade base; panicle many-flowered, elongate, 8–30 cm long; flowers aggregated on the lateral inflorescence branches; secund arrangement more obvious in fruit; pedicels 0.1–0.2 mm long; hypanthium 1.2–2.5 mm long; calyx teeth 1–2 mm long; petals 1.5–2.4 × 0.4–0.7 mm, lanceolate, pure white; anthers 0.8–1.6 mm long; ovary 5-locular, sparsely puberulous at the apex. Understory of lowland forests, 50–600 m; Bolívar (upper Río Caura,

Río Erebato), Amazonas (Maroa, Piedra Nunca north of Piedra Cocuy, Río Baría, Río Cunucunuma, Río Matacuni, Río Mawarinuma, Río Siapa, Río Yatúa, San Carlos de Río Negro). Colombia (Vaupés), Brazil (Amazonas). ◆Fig. 273. For comments, see Leandra divaricata. Leandra glandulifera (Triana) Cogn. in Mart., Fl. Bras. 14(4): 207, pl. 44, fig. 2. 1886. —Oxymeris glandulifera Triana, Trans. Linn. Soc. London 28: 95. 1871. Shrub 1–2 m tall; branchlets, petioles, primary veins on lower surface of leaves, inflorescence axes, and hypanthia densely covered with simple hairs 0.3–0.8 mm long, and moderately covered with gland-tipped hairs 0.8–1.6 mm long; petiole 1–4 cm long; leaf blade 6–19 × 3–6 cm, oblong-laneolate to elliptic, apex acuminate, base acute to obtuse, membranous, margin serrate, 5-pliveined with the inner pair of primary veins diverging 0.5–1 cm above the blade base; panicle few-flowered, 4–8 cm long; pedicels 0.2–0.3 mm long; hypanthium 1.5–2 mm long; calyx teeth 0.3–0.6 mm long; petals 0.3–0.5 × 0.3– 0.4 mm, triangular, whitish to pink; anthers 2.1–2.4 mm long, with a slightly dorsally inclined pore; ovary 3-locular, densely puberulous and with a short fringed tube at the apex. Understory and edges of lowland forests, 50–200 m; Amazonas (Maroa, Río Baría, Río Casiquiare, Río Guainía, Río Yatúa, San Carlos de Río Negro, Yavita). Colombia, Peru, Brazil (Amazonas). ◆Fig. 272. Leandra gorzulae Wurdack, Phytologia 69: 325. 1990. Small shrub 0.3–1 m tall; branchlets, petioles, and inflorescence axes densely covered with simple and barbellate hairs 0.1– 0.9 mm long; hypanthia only with simple hairs; upper surface of leaf blade warty or with short hairs with swollen base (hairs 0.1–0.2 mm long); petiole 0.2–0.5 cm long; leaf blade 3.5–5 × 1.5–2.5 cm, elliptic, apex broadly acute, base obtuse, coriaceous, margin rolled inward, 5-veined; panicle few-flowered, 2–4.5 cm long; pedicels 0.5–1.1 mm long; hypanthium 2.1–3.7 mm long; calyx teeth 1–1.5 mm long; petals 1.5–2.6 × 0.5–1.1 mm, lanceolate, pink; anthers 1.6–1.7 mm long, with a slightly ventrally inclined pore; ovary 3-locular, with a minute glabrous col-

Leandra 355

lar at the apex. Tepui meadows and scrub, 2100–2200 m; Bolívar (Sierra de Maigualida). Endemic. Leandra lindeniana (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 136. 1886. —Clidemia lutescens var. lindeniana Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 348. 1852. —Oxymeris lindeniana (Naudin) Triana, Trans. Linn. Soc. London 28: 90. 1871. Leandra fendleri Cogn. in A. DC. & C. DC., Monogr. Phan. 7: 653. 1891. Shrub 0.5–3 m tall; branchlets, petioles, primary veins on lower surface of leaves, inflorescence axes, and hypanthia densely to moderately covered with simple hairs 0.5– 2.4 mm long and an underlayer of minute barbellate hairs 0.1–0.6 mm long; petiole 0.5–1.7 cm long; leaf blade 5–11 × 2.5–5 cm, ovate-oblong, apex blunt-acute, base rounded to cordulate, rigid, 5–7-veined; panicle 4–12 cm long; flowers sessile, clustered in intervals along the unbranched inflorescence axis or on short lateral branches; hypanthium 3–5 mm long; calyx teeth 1.3– 2.5 mm long; petals 3.5–5 × 1–2.5 mm, lanceolate-oblong, white; anthers 2.7–3 mm long, with a slightly dorsally inclined pore; ovary 3-locular, with a thickened, sparsely puberulous collar at the apex. Montane forest understory, 700–1200 m; Bolívar (Río Kukenán, Río Yuruaní). Aragua, Distrito Federal, Mérida, Miranda, Táchira; Colombia, Guyana. ◆Fig. 270. Leandra longisepala Wurdack, Mem. New York Bot. Gard. 10(4): 32. 1961. Shrub 1.5–2.5 m tall; branchlets, petioles, primary leaf veins, inflorescence axes, and hypanthia densely to moderately covered with simple hairs 0.5–1.7 mm long; petiole 3–7 cm long; leaf blade 9–17 × 4–9 cm, ovate, apex gradually acuminate, base roundedtruncate, membranous, 7-pliveined with the inner pair of primary veins diverging 0.6–1.1 cm above the blade base; panicle many-flowered, 12–17 cm long; pedicels 0.3–0.5 mm long; hypanthium 2.3–2.5 mm long; calyx teeth 2.2–3 mm long; petals 3.2–3.8 × 1.3–1.6 mm, acuminate-lanceolate, pale pink; anthers 2.4–2.6 mm long; ovary 3-locular, the apex with a bowl-shaped corona, ca. 1.5 mm diameter. Understory of tepui-slope forests,

1600–1700 m; Bolívar (Sierra de la Neblina). Endemic. Leandra maguirei Wurdack, Mem. New York Bot. Gard. 10(5): 162. 1964. Shrub 1–2 m tall; branchlets, petioles, and inflorescence axes densely covered with simple and with gland-tipped hairs (both 0.2–0.9 mm long), gland-tipped hairs most evident on inflorescence axes; primary veins on lower surface of leaves with only simple hairs; petiole 1.5–3 cm long; leaf blade 5–11 × 3.5–7.5 cm, ovate, apex narrowly acute to gradually acuminate, base cordulate, the upper surface slightly rugose (hairs projecting from elevated bases), the lower surface foveolate, rigid, (5)7-veined; panicle manyflowered, 7–12 cm long; inflorescence branches pale pink; pedicels 0.2–1 mm long; hypanthium 2.1–3 mm long; calyx teeth 1.5– 2.4 mm long; petals 2–3.5 × 0.6–0.9 mm, oblong-lanceolate, white to pink; anthers 1.7– 1.9 mm long; ovary 3-locular, lobed and puberulous at the apex. Tepui meadows and forests, 1700–2500 m; Amazonas (Sierra de la Neblina). Brazil (Serra da Neblina). ◆Fig. 276. Leandra neblinensis Wurdack, Mem. New York Bot. Gard. 10(5): 163. 1964. Shrub 1–1.5 m tall; branchlets, petioles, and primary veins on lower surface of leaves densely covered with simple hairs 0.7–5 mm long; inflorescence axes and hypanthia mainly with gland-tipped hairs 0.8–1.5 mm long; petiole 1–3.5 cm long; leaf blade 9–20 × 5–13.5 cm, broadly elliptic, apex shortacuminate, base rounded, membranous, 5–7veined; panicle many-flowered, 13–15 cm long; pedicels 0.2–0.3 mm long; hypanthium 1.8–2.2 mm long; calyx teeth 0.2–0.5 mm long; petals 3.5–4 × 1.2–1.8 mm, oblongovate, pale pink; anthers 2 mm long; ovary 3locular, lobed and sparsely puberulous at the apex. Understory of scrub forests, ca. 700 m; Amazonas (headwaters of Río Yatúa in Sierra de la Neblina). Endemic. Leandra phelpsiae Gleason, Phytologia 3: 351. 1950. Shrub 1.5–3 m tall; branchlets, petioles, inflorescence axes, and hypanthia densely covered with simple or barbellate hairs 0.1– 0.4 mm long, and moderately covered with

356

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gland-tipped hairs 0.2–0.6 mm long; lower surface of leaf blade and primary veins on the upper surface with simple and barbellate hairs; petiole 2–4 cm long; leaf blade 7–12 × 4–6.5 cm, ovate-lanceolate to ovate, apex acute to inconspicuously acuminate, base cordulate, subcoriaceous, 7-veined; panicle rather many-flowered, 10–17 cm long; pedicels < 0.2 mm long; hypanthium 1.5–3 mm long; calyx teeth 0.5–0.8 mm long; petals 0.4–0.6 × 0.4–0.6 mm, ovate, white to reddish; anthers 2.7–3.4 mm long, with a dorsally inclined pore; ovary 3-locular, with a small fringed collar at the apex. Forest understory of tepui slopes and ridges, 1200– 2000 m; Amazonas (slope of Cerro Marahuaka, Cerro Parú, Sierra de la Neblina). Endemic. ◆Fig. 277. Leandra phelpsiae represents the southern vicariant relative of L. polyadena. Leandra polyadena Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 354. 1915. Shrub 1.5–2.5 m tall; branchlets, petioles, primary and secondary veins on the lower surface of the leaves, inflorescence axes, and hypanthia moderately to densely covered with simple hairs 0.3–1 mm long and different-sized gland-tipped hairs 0.3–3 mm long, the glands dark purple; petiole 3–9 cm long; leaf blade 7–18 × 4–10 cm, ovate, chartaceous, 7(9)-veined, apex gradually acuminate, base cordulate, the upper surface slightly rugose, with hairs projecting from elevated bases, the lower surface foveolate; panicle rather few-flowered, 10–18 cm long; petioles, veins on the lower surface of leaves, and young branches purple-red; pedicels 0.2– 0.5 mm long; hypanthium 1.9–2.6 mm long; calyx teeth 0.5–0.8 mm long; petals 0.5 × 0.4 mm, ovate-oblong, whitish to pale pink; anthers 3–4 mm long, with a slightly ventrally inclined pore; ovary 3-locular, with a small fringed collar at the apex. Understory of tepui-slope forests, 1300–2200 m; Bolívar (Auyán-tepui, Los Testigos, Luepa, Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Sipapo). Guyana. ◆Fig. 275. For comments, see Leandra phelpsiae. Leandra procumbens Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 355. 1915. Leandra chimantensis Wurdack, Mem. New York Bot. Gard. 10(5): 161. 1964.

Creeping or climbing shrub with internodal roots; branchlets, petioles (especially on adaxial side), inflorescence axes, and hypanthia moderately to densely covered with simple, sometimes barbellate, hairs 0.1–1.2 mm long, and occasionally with gland-tipped hairs 0.5–3 mm long; petiole 0.5–1.5 cm long; leaf blade 2–7.5 × 1.2–3.8 cm, oblong-ovate to oblong-elliptic, apex acute to short-acuminate, base rounded, thin-rigid, 5-veined; panicle rather few-flowered, 2–10 cm long; flowers 4-merous; pedicels 0.2–0.4 mm long; hypanthium 1.6–2.7 mm long; calyx teeth 0.5–1.2 mm long; petals 1.7–3.5 × 0.7–1.5 mm, oblong-lanceolate, whitish to pink; anthers 2.2–2.5 mm long, with a terminal pore; ovary 3-locular, glabrous, with a short sulcate tube at the apex. Understory of scrub forests, shaded canyons on tepuis, 1900– 2500 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Roraima-tepui). Guyana (Mount Roraima). ◆Fig. 271. J. J. Wurdack described Leandra chimantensis, differentiating it from L. procumbens mainly by the occurrence of gland-tipped and barbellate hairs on inflorescence branches and hypanthia. These taxa are neither geographically nor ecologically separated. Duplicates of one and the same collection series may contain characters of both taxa (e.g., Steyermark & Wurdack 1051, F, NY, US, Steyermark et al. 128740, F, MO, NY, US, VEN). For those reasons, L. chimantensis should be designated as a synonym of L. procumbens. Leandra purpurea Gleason, Bull. Torrey Bot. Club 52: 376. 1925. Shrub 2.5–5 m tall; branchlets, petioles, lower surface of leaves, inflorescence axes, and hypanthia covered with simple hairs 0.2–1.5 mm long and gland-tipped hairs 0.3– 1.5 mm long; petiole 2–4 cm long; leaf blade 13–20 × 5.5–8.5 cm, ovate-elliptic, apex sharply acuminate, base rounded, thin-membranous, 5-veined; panicle many-flowered, 12–15 cm long; pedicels 0.1–0.7 mm long; hypanthium 1.2–2 mm long; calyx teeth 0.3–0.4 mm long, with terminal hairs 0.3–0.6 mm long; petals white, 0.4–0.7 × 0.3–0.5 mm, fringed with gland-tipped hairs; anthers 1.9– 2.2 mm long, with a ventrally inclined pore; ovary 5-locular, densely puberulous at the apex. Forest understory, 400–900 m; Bolívar

Leandra 357

(Cerro Abismo, El Paují, Los Testigos, base of Perai-tepui). Guyana, French Guiana. Leandra rufescens (DC.) Cogn. in Mart., Fl. Bras. 14(4): 204. 1886. —Tschudya rufescens DC., Prodr. 3: 155. 1828. —Oxymeris rufescens (DC.) Triana, Trans. Linn. Soc. London 28: 95. 1871. Tschudya asperiuscula DC., Prodr. 3: 155. 1828. —Oxymeris asperiuscula (DC.) Triana, Trans. Linn. Soc. London 28: 95. 1871. —Leandra rufescens var. asperiuscula (DC.) Cogn. in Mart., Fl. Bras. 14(4): 205. 1886. Leandra rufescens var. glandulosa Cogn. in Mart., Fl. Bras. 14(4): 205. 1886. Shrub 1–4.5 m tall; branchlets, petioles, and primary veins on lower surface of leaves densely covered with simple, or occasionally barbellate, hairs 0.1–1.7 mm long; inflorescence axes and hypanthia densely covered with simple and sparsely covered with glandtipped hairs 0.6–0.8 mm long; petiole 1–4.5 cm long; leaf blade 10–31 × 4–12 cm, oblonglanceolate to ovate-elliptic, apex gradually acuminate, base broadly acute to obtuse, firm-chartaceous, 5(7)-veined; panicle manyflowered, 10–20 cm long; flowers sessile; hypanthium 1.5–2.5 mm long; calyx teeth 0.3– 0.7 mm long; petals 0.9–1.3 × 0.3–0.5 mm, obovate-oblong, externally covered with close-pressed short hairs; anthers 1.5–1.7 mm long, with a terminal pore; ovary 5-locular, densely puberulous at the apex. Forest understory, 100–1200 m; Delta Amacuro (lower Río Orinoco), Bolívar (near Kilómetro 88, Los Testigos, Serranía de Imataca). Trinidad, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil (Amapá, Amazonas, Bahia, Pará). Leandra sanguinea Gleason, Bull. Misc. Inform. Kew 1939: 550. 1939. Shrub 1–4 m tall, with angled branches; petiole 2.5–11 cm long; leaf blade 13–30 × 6– 16 cm, elliptic-ovate to ovate, apex caudateacuminate, base obtuse, chartaceous, shortly 7-pliveined with the inner pair of primary veins diverging 0.5–2 cm above the blade base; panicle many-flowered, 10–25 cm long; flowers sessile, aggregated on the lateral inflorescence branches; secund arrangement more obvious in fruit; hypanthium 1.5–1.9 mm long; petals 1.5–3 × 0.4–0.8 mm, lan-

ceolate-oblong, white; anthers 1–1.5 mm long; ovary 3-locular. Venezuela, Guyana, Suriname, Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of G. sanguinea 1. Indumentum consisting of simple and gland-tipped hairs (best recognizable in inflorescences); calyx teeth subulate; ovary apex with hairs .............................. ................................ subsp. sanguinea 1. Indumentum eglandular; calyx teeth ovate-oblong; ovary apex usually hair less ........................... subsp. tepuiensis L. sanguinea subsp. sanguinea Branchlets, petioles, primary veins on lower surface of leaves, inflorescence axes, and hypanthia densely covered with simple hairs 0.2–2 mm long and sparsely covered with gland-tipped hairs 0.3–0.9 mm long, the glands very tiny; calyx teeth subulate, 1.5– 2.1 mm long; ovary apex knobby, with hairs 0.3–0.5 mm long. Understory of sparse, lower montane forests, 300–900 m; Bolívar (La Escalera). Guyana. ◆Fig. 274. L. sanguinea subsp. tepuiensis Wurdack, Mem. New York Bot. Gard. 10(5): 163. 1964. Indumentum consisting of simple hairs 0.2–2 mm long; calyx teeth ovate-oblong, 0.6–1.5 mm long; ovary apex knobby, hairless or with few hairs 0.1 mm long. Understory and edges of tepui-slope and summit forests, 800–2200 m; Bolívar (Cerro Guaiquinima, Gran Sabana, La Escalera, Macizo del Chimantá, Perai-tepui, Sororopán-tepui), Amazonas (base of Cerro Cuao, slope of Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Guyana, Suriname, Brazil (Amazonas, Roraima). Leandra secunda (D. Don) Cogn. in Mart., Fl. Bras. 14(4): 199. 1886. —Clidemia secunda D. Don, Mem. Wern. Nat. Hist. Soc. 4: 308. 1823. —Oxymeris secunda (D. Don) Triana, Trans. Linn. Soc. London 28: 95. 1871. Shrub 0.3–1 m tall; branchlets, petioles, primary veins on lower surface of leaves, and inflorescence axes moderately to densely covered with appressed simple hairs 0.3–1.5

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mm long and sparsely with caducous glandtipped hairs 0.2–0.7 mm long; hypanthia densely covered with gland-tipped hairs; petiole 1.5–5 cm long; leaf blade 10–15 × 5–8 cm, ovate to elliptic-ovate, apex short-acumi-

nate, base obtuse, chartaceous, (5)7-veined or shortly pliveined with the inner pair of primary veins diverging 0.7 cm above the blade base; panicle rather few-flowered, 4– 10 cm long; pedicels 0.1–0.4 mm long or ab-

Fig. 266. Leandra chaetodon

Fig. 267. Leandra aristigera

Leandra 359

Fig. 268. Leandra candelabrum

Fig. 270. Leandra lindeniana

Fig. 269. Leandra divaricata

Fig. 271. Leandra procumbens

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Fig. 272. Leandra glandulifera

Fig. 273. Leandra francavillana

Fig. 274. Leandra sanguinea subsp. sanguinea

Leandra 361

Fig. 275. Leandra polyadena

Fig. 276. Leandra maguirei

Fig. 277. Leandra phelpsiae

362

M ELASTOMATACEAE

sent; hypanthium 1.8–2.3 mm long; calyx teeth 0.1–0.8 mm long; petals 1.2–2.7 × 0.5– 0.8 mm, oblong-lanceolate, white; anthers 0.9–1.3 mm long; ovary 5-locular, sparsely puberulous at the apex. Understory of lowland forests, 100–200 m; Amazonas (Río Mavaca, Río Mawarinuma). Colombia (Amazonas, Putomayo, Vaupés), Ecuador, Peru, Brazil (Acre, Amazonas, Rondônia). Leandra solenifera Cogn. in Mart., Fl. Bras. 14(4): 192, t. 42, fig. 1. 1886. Shrub 0.5–3 m tall; branchlets, petioles, primary veins on lower surface of leaves, and inflorescence axes moderately to densely covered with simple and gland-tipped hairs (both 0.2–0.7 mm long); petiole 2–6 cm long; leaf blade 8–16 × 5–9 cm, ovate, apex gradually acuminate, base weakly cordulate to cordate, chartaceous, 7–9-veined or shortly pliveined with the inner pair of primary veins diverging to 1 cm above the blade base; panicle many-flowered, 10–18 cm long; flowers sessile, 6(7)-merous; hypanthium 1.7–2 mm long; calyx teeth 0.3–0.7 mm long; petals 2–2.8 × 1.1–1.6 mm, elliptic-ovate, white flushed with pink; anthers 1.2–2 mm long; ovary 4–6-locular, with a glabrous or sparsely puberulous collar at the apex. Un-

derstory and edges of forests, 200–900 m; Bolívar (Perai-tepui, Río Maniapure), Amazonas (Coromoto, Río Cunucunuma, headwaters of Río Orinoco). Cojedes, Mérida, Táchira; Colombia, Guyana, Suriname, French Guiana, Amazon basin periphery in Ecuador, Peru, and western Brazil, to Bolivia. Leandra steyermarkii Wurdack, Phytologia 21: 355. 1971. Woody liana, with internodal roots; branchlets, primary veins on lower surface of leaves, inflorescence axes, and hypanthia densely covered with gland-tipped hairs 0.6– 1 mm long and a moderate underlayer of minute stellulate hairs; petiole 0.6–1.2 cm long; leaf blades slightly dissimilar in size in each pair, 3–7.5 × 1.5–4 cm, elliptic-oblong, apex acuminate, base cordulate, membranous, 5-veined; panicle few-flowered, 2.5–4 cm long; pedicels 0.3–0.5 mm long; hypanthium 2 mm long; calyx teeth 0.6–0.7 mm long; petals 2.8 × 1 mm, lanceolate-oblong, white, externally covered with gland-tipped hairs; anthers 2.2–2.3 mm long, with a dorsally inclined pore; ovary 4-locular, granulose and sparsely puberulous at the apex. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro). Endemic.

20. LOREYA DC., Prodr. 3: 178. 1828. Heteroneuron Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 768. 1867. by Paul E. Berry and Susanne S. Renner Small to large trees 3–38 m tall. Leaves opposite, simple, petiolate, usually large and membranous, drying dark brown; venation with a midvein and 2 or 3 pairs of suprabasal primary veins, rarely pinnate (L. nigricans); margin entire or serrulate. Inflorescences borne on the old wood, few-branched cymes on a well-developed peduncle, or contracted cymes with the peduncle short or lacking, or clusters of few flowers. Flowers 5(6)-merous. Hypanthium hemispherical; calyx limb truncate, lobes vestigial; petals ovate, elliptic, or broadly triangular, apex obtuse, irregular, or subacute, fleshy, often with adaxial callus ridges and some development of lateral flaps, pink outside, pink and white inside, or else completely pink or white. Stamens twice as many as the petals, isomorphic, glabrous, laterally compressed, the anthers and filaments of ± equal length; anthers thick, 1- or 2-pored, the connective not prolonged but fleshy. Ovary completely inferior, 5(6)- or 10-locular, apically with 10 radial ridges; style glabrous, thick, longer than the stamens; stigma capitate, 5(6)lobed. Fruit a hemispherical berry, many-seeded, yellow or red to blue. Seeds minute, 0.5–1.4 mm long, ovoid or wedge-shaped. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 13 species, 5 in Venezuela, all in the flora area.

Loreya 363

This treatment is extracted in large part from the monograph of Loreya by S. S. Renner (Mem. New York Bot. Gard. 50: 1–112. 1989). Key to the Species of Loreya 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Anthers 1-pored; leaf blades pubescent; fruits > 1 cm diameter, yellow ................................................................................................................ 2 Anthers 2-pored; leaf blades completely glabrous or sericeous; fruits < 1 cm diameter, red, blue, or black ......................................................... 3 Lower surface of leaves sericeous; ovary 5-locular .................. L. mespiloides Lower surface of leaves puberulous to laxly strigulose; ovary 10-locular ................................................................................................. L. spruceana Flowers in cymes, the peduncle 5–30 mm long ........................ L. arborescens Flowers in sessile or subsessile clusters, the peduncle short, < 5 mm long or lacking ................................................................................................ 4 Leaf blades glabrous; cymes once-branched, the peduncles 1–5 mm long ......................................................................................................... L. ovata Leaf blades sericeous on both surfaces and on the veins; flowers in sessile clusters ........................................................................................ L. strigosa

Loreya arborescens (Aubl.) DC., Prodr. 3: 179. 1828. —Melastoma arborescens Aubl., Hist. Pl. Guiane 420, pl. 163. 1775. —Bellucia arborescens (Aubl.) Baill., Hist. Pl. 7: 34. 1877. —Trompo de lamorado. Loreya acutifolia O. Berg ex Triana, Trans. Linn. Soc. London 28: 142. 1871. Loreya maguirei Wurdack, Phytologia 18: 160. 1969. Tree 10–25(–38) m tall. Evergreen lowland forests, 100–300 m; Bolívar (lower part of La Escalera), Amazonas (around Maroa, Río Coro Coro, Río Cuao, San Carlos de Rio Negro). Colombia (Vaupés), Guyana, French Guiana, Peru, Brazil. Loreya mespiloides Miq., Linnaea 18: 619. 1844. —Bellucia mespiloides (Miq.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(4.1): 497. 1941. Henriettea brunnescens Standl., Field Mus. Nat. Hist., Bot. Ser. 4: 247. 1929. —Loreya brunnescens (Standl.) Gleason, Phytologia 3: 346. 1950. Tree (2–)4–10(–18) m tall; branchlets with conspicuous, brown, velvety hairs. Nonflooded lowland to upland forests, 100–900 m; Bolívar (near El Manteco, southern Gran Sabana), Amazonas (Puerto Ayacucho to Gavilán). Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil. ◆Fig. 278.

Loreya ovata O. Berg ex Triana, Trans. Linn. Soc. London 28: 142. 1871. —Palo de escoba. Loreya minor Cogn. in Mart., Fl. Bras. 14(4): 522. 1888. Tree 5–10 m tall; leaves with a drip tip 1– 2 cm long. Nonflooded evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Rio Negro, Yavita). Colombia (Vaupés), Brazil (Amazonas). ◆Fig. 280. Loreya spruceana Benth. ex Triana, Trans. Linn. Soc. London 28: 143. 1871. Loreya collatata Wurdack, Phytologia 18: 162. 1969. Small tree 3–10 m tall. Nonflooded forests, open forest on white sand, often in waterlogged soil, 100–800 m; Bolívar (30 km south of El Dorado, Cerro Ichún, northeast of Cerro Marutaní, Rio Chicanán), Amazonas (near San Carlos de Rio Negro, base of Sierra de la Neblina). Barinas, Táchira, Zulia; Colombia (Guajira), Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 279. Loreya strigosa Gleason, Bull. Torrey Bot. Club 60: 386. 1933. Loreya quadrifolia Gleason, Bull. Torrey Bot. Club 60: 386. 1933. Small tree 4–10 m tall. Lower montane and riparian forests, 300–400 m; Amazonas (upper Río Matacuni). Colombia (Caquetá), Brazil.

364

M ELASTOMATACEAE

Fig. 278. Loreya mespiloides

Loreya 365

Fig. 279. Loreya spruceana

Fig. 280. Loreya ovata

366

M ELASTOMATACEAE

21. MACAIREA DC., Prodr. 3: 109. 1828. Siphantheropsis Brade, Publ. Inst. Nac. Pesq. Amazônia Bot. 8: 4. 1958. by Paul E. Berry and Susanne S. Renner Shrubs, subshrubs, small trees, or rarely herbs; indument on young branchlets densely hirsute, sericeous, or lepidote, rarely glabrous. Leaves opposite, petiolate, 3–9-veined, subcoriaceous or thin and brittle. Inflorescence a terminal, many-flowered, elongate or congested paniculate cyme, or groups of 3–5 subsessile flowers in the axils of the uppermost leaves; pedicels with bracts and bracteoles. Flowers bisexual, 4-merous. Hypanthium campanulate; calyx lobes narrowly to broadly triangular, spreading, persistent on the limb after anthesis; petals thin, obovate, apically ± emarginate, minutely clawed at the base, basally white, turning red with age, apically pink or magenta, or petals white or purple, margins with short-stalked globose glands. Stamens 8, dimorphic or subisomorphic, ± same length as petals; filaments ventrally with globose, short-stalked glands or glabrous; anthers linear, 1pored, rostrate, rarely truncate; connective dorso-basally prolonged and expanded into a cordate appendage often surrounding the insertion of the filament. Ovary free, perigynous, 2–4-locular, apically with short glandular hairs; style thin, terete or subclavate, sigmoid, longer than the stamens and exserted from the corolla; stigma punctiform. Fruit a loculicidal, many-seeded, dark brown capsule. Seeds 0.5– 1.4 mm long, cochleate or polyhedric. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, most species restricted to the Guayana Highlands; 22 species, 17 in Venezuela, all in the flora area. This treatment was extracted in large part from the generic revision by S. S. Renner (Mem. New York Bot. Gard. 50: 1–112. 1989). Key to the Species of Macairea 1. 1. 2(1). 2. 3(2). 3. 4(2). 4. 5(4). 5. 6(5). 6. 7(6).

Flowers subsessile in the axils of the uppermost leaves; petals 3–4 m long ................................................................................................. M. axilliflora Flowers pedicellate, borne in a pedunculate inflorescence; petals > 4 mm long ......................................................................................................... 2 Lower surface of leaves glabrous or with minute, papillate-like hairs, without sessile glands ............................................................................ 3 Lower surface of leaves with either long, glandular hairs or globose, sessile glands ......................................................................................... 4 Leaf blades narrowly elliptic, 3–12 × 1–3 cm .................................. M. stylosa Leaf blades linear, 1.2–1.8 × ca. 0.2 cm ......................................... M. linearis Lower surface of leaves with long, eglandular hairs, with sessile glands ................................................................................................................ 5 Lower surface of leaves with long, glandular hairs, lacking sessile glands .............................................................................................................. 10 Inflorescence with long, glandular hairs ..................................... M. maroana Inflorescence without glandular hairs ...................................................... 6 Ovary apically sericeous with long, eglandular hairs .............. M. spruceana Ovary apically with few, short, glandular hairs ....................................... 7 Leaf blades pseudopinnately veined, the single pair of lateral primary veins inconspicuous, upper surface glabrous and smooth, lower surface

Macairea 367

7.

8(7).

8.

9(8). 9. 10(4). 10. 11(10). 11. 12(11).

12. 13(11). 13. 14(13). 14. 15(14). 15. 16(14). 16.

sparsely hirsute with long, prostrate, awl-shaped hairs which are apically frayed .......................................................................... M. rufescens Leaf blades with 1 or 2 pairs of conspicuous lateral primary veins, upper surface verrucose or scabrous, with stout, conical hairs, lower surface with erect, basally broad and roughened hairs .................................... 8 Leaf blades narrowly elliptic, 3.5–5.5 × 1.8–2.5 cm; calyx lobes 1–2 mm long; filaments 2.8–3.2 (longer whorl) or 1.8–2 mm long, thecae 1.5– 2 mm long .......................................................................... M. chimantensis Leaf blades broadly elliptic or elliptic or obovate, 3.5–13 × (2–)2.5–4.5 cm long: calyx lobes to 5 mm long; filaments 4–9 or 3–8 mm long, thecae 3– 5 mm long ............................................................................................... 9 Leaf blades 3.5–6 × 2.5–4 cm; calyx lobes 1–2 mm long; filaments 3.8– 4.2 (longer whorl) or 2.8–3.3 mm long ........................................ M. duidae Leaf blades 4.5–13 × (2–)3–4.5 cm; calyx lobes 2–5 mm long; filaments 8– 9 or 5–8 mm long ................................................................... M. thyrsiflora Cymes in the axils of the uppermost foliage leaves ........................ M. lanata Cymes terminal ........................................................................................ 11 Leaf blades with 3 or 4 pairs of lateral primary veins; petals 14–15 × 10– 15 mm ................................................................................................... 12 Leaf blades with 1 or 2 pairs of lateral primary veins; petals ≤ 9 mm long, ≤ 6(–8) mm broad ................................................................................. 13 Petioles 0.5–1.5 cm long; inflorescence congested; filaments 5–6 mm (longer whorl) or 4–5 mm long, with short-stalked glands; ovary 4-locular ................................................................................... M. cardonae Petioles 3–8 mm long; inflorescence elongate; filaments 8–10 or 7–8 mm long; ovary 3(4)-locular ........................................................ M. multinervia Petioles 3–5 mm long; filaments glabrous ............................... M. lasiophylla Petioles > 5 mm long; filaments with short-stalked glands ................... 14 Ovary (2)3-locular .................................................................................... 15 Ovary 4-locular ......................................................................................... 16 Leaf blades elliptic, 2–4 × 0.8–1.8 cm ......................................... M. parvifolia Leaf blades obovate, 4–7 × 2–4 cm ................................................... M. rigida Leaf blades narrowly elliptic, (3.5–)6–8 × 2–3.5 cm .................... M. neblinae Leaf blades elliptic or obovate, (4.5–)7–15 × (1.8–)3–9 cm .... M. pachyphylla

Macairea axilliflora Wurdack, Mem. New York Bot. Gard. 10(1): 99. 1958. Shrub 0.3–2 m tall; leaves often sticky. Seasonally flooded white-sand savannas, black-water river banks, 100–200 m; southwestern Amazonas. Colombia (Guianía). ◆Fig. 283. Macairea cardonae Wurdack, Mem. New York Bot. Gard. 10(5): 139, fig. 64a–d. 1964. Shrub 0.5–2 m tall. Exposed areas of tepui summits, 1700–2500 m; Bolívar (Auyán-tepui, Macizo del Chimanta). Endemic. ◆Fig. 282.

Macairea chimantensis Wurdack, Mem. New York Bot. Gard. 10(5): 138. 1964. Shrub or treelet 0.3–7 m tall. Exposed areas of tepui summits, 1600–2500 m; Bolívar (tepuis of northern and central Gran Sabana area). Endemic. ◆Fig. 286. Macairea duidae Gleason, Bull. Torrey Bot. Club 58: 416. 1931. Shrub 1.5–2 m tall. Tepui ridge tops, Heliamphora swamp savannas, 1200–2100 m; Amazonas (Cerro Duida). Endemic. Macairea lanata Gleason, Bull. Torrey Bot. Club 58: 416. 1931.

368

M ELASTOMATACEAE

Macairea lanata subsp. eglandulosa Wurdack, Mem. New York Bot. Gard. 10(1): 99. 1958. Shrub 0.2–1 m tall. Moist to swampy savannas or tepui meadows, 100–2100 m; Amazonas (scattered in central and southern parts). Colombia (Guianía). ◆Fig. 285. Macairea lasiophylla (Benth.) Wurdack, Phytologia 13: 65. 1966. —Chaetogastra? lasiophylla Benth., J. Bot. (Hooker) 2: 291. 1840. —Pterolepis lasiophylla (Benth.) Triana, Trans. Linn. Soc. London 28: 40. 1871. —Tibouchina lasiophylla (Benth.) Cogn. in Mart., Fl. Bras. 14(3): 297. 1885. —Acisanthera lasiophylla (Benth.) Gleason, Phytologia 3: 243. 1950. Acisanthera erecta Gleason, Bull. Torrey Bot. Club 58: 412. 1931. Herb or subshrub 10–50 cm tall; leaves narrowly elliptic, 0.6–1.6 × 0.3–0.8 cm; anther connective appendages purple, ventrally 2-lobed. Lowland to upland white-sand savannas, 100–1200 m; Bolívar (Gran Sabana), Amazonas (Canaripó, Cerro Vinilla, La Esmeralda, near Santa Bárbara del Orinoco). Guyana, Brazil (Roraima). ◆Fig. 288. Macairea linearis Gleason, Bull. Torrey Bot. Club 58: 417, pl. 35, fig. 1. 1931. Shrub ca. 1 m tall. Along stream beds on tepui summit, 1600–2100 m; Amazonas (Cerro Duida). Endemic. Macairea maroana Wurdack, Mem. New York Bot. Gard. 10(5): 141. 1964. Shrub 1.2–2 m tall. Seasonally flooded white-sand savannas, forest-savanna borders, 100–200 m; Amazonas (around Maroa). Brazil (northern Amazonas). ◆Fig. 284. Macairea multinervia Benth., J. Bot. (Hooker) 2: 219. 1840. —Tibouchina multinervia (Benth.) Baill., Adansonia 12: 75. 1876. Subshrub or shrub 0.2–1 m tall; leaves thin, basally cordate, 7–9-veined. Lowland and upland white-sand savannas, 200–1300 m; Bolívar (Gran Sabana). Brazil (Amazonas). ◆Fig. 281. Macairea neblinae Wurdack, Mem. New York Bot. Gard. 10(1): 100. 1958.

Shrub 1–2 m tall. Rock outcrops on tepui slopes, 1700–2200 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 290. Macairea pachyphylla Benth., J. Bot. (Hooker) 2: 292. 1840. Macairea calvescens Naudin, Ann. Sci. Nat. Bot. sér 3, 13: 35. 1850. Shrub or treelet 1.5–4(–12) m tall, with long, patent, glandular hairs. Montane and lower montane forests and gallery forests, moist savannas, 400–1500 m; Bolívar (Gran Sabana and included tepuis), Amazonas (Cerro Duida, base of Cerro Huachamacari, Cerro Parú). Guyana, Suriname, Brazil. ◆Fig. 291. Macairea parvifolia Benth., J. Bot. (Hooker) 2: 293. 1840. Shrub 0.8–2 m tall. White-sand savannas, tepui-summit shrublands, 800–1800; Bolívar (Cerro Guaiquinima, Gran Sabana, Río Karún). Adjacent Guyana, Brazil (Amazonas: Serra Aracá). ◆Fig. 289. Macairea rigida Benth., J. Bot. (Hooker) 2: 292. 1840. Shrub or treelet 1–5 m tall. Tepui summit shrublands or shrub islands, 700–2000 m; Amazonas (Cerro Duida, Cerro Guanay, Cerro Marahuaka, Cerro Parú, Cerro Yutajé). Endemic. ◆Fig. 292. Macairea rufescens DC., Prodr. 3: 109. 1828. Shrub 1–2 m tall; leaves 7–15 × 4–7 cm, coriaceous. Granitic outcrops, white-sand savannas, 100–200 m; Amazonas (Maroa, near San Carlos de Rio Negro). Southwestern Colombia, Amazonian Peru. ◆Fig. 287. Macairea spruceana O. Berg ex Triana, Trans. Linn. Soc. London 28: 38. 1871. Shrub or treelet 0.5–2 m tall. White-sand savannas, black-water river banks, 100–200 m; Amazonas (uppermost Río Negro and tributaries, Río Temi). Brazil (northern Amazonas). ◆Fig. 295. Macairea stylosa Triana, Trans. Linn. Soc. London 28: 38. 1871. Shrub or treelet 1–3(–5) m tall. Seasonally flooded savannas and shrublands along black-water rivers, 50–200 m; central and

Macairea 369

Fig. 281. Macairea multinervia

Fig. 282. Macairea cardonae

Fig. 283. Macairea axilliflora

370

M ELASTOMATACEAE

Fig. 285. Macairea lanata

Fig. 284. Macairea maroana

Fig. 286. Macairea chimantensis

Macairea 371

Fig. 287. Macairea rufescens

Fig. 288. Macairea lasiophylla

372

M ELASTOMATACEAE

Fig. 290. Macairea neblinae Fig. 289. Macairea parvifolia

Fig. 291. Macairea pachyphylla

Macairea 373

Fig. 292. Macairea rigida

Fig. 293. Macairea thyrsiflora

374

M ELASTOMATACEAE

Fig. 294. Macairea stylosa

Fig. 295. Macairea spruceana

Macrocentrum 375

southwestern Amazonas. Brazil (Amazonas). ◆Fig. 294. In this species, the hypanthium is often greatly enlarged due to insect galls. Macairea thyrsiflora DC., Prodr. 3: 109, 484. 1828. Macairea albiflora Cogn. in Mart., Fl. Bras. 14(3): 246. 1885. Macairea aspera N.E. Br., Trans. Linn. Soc. London 6: 27, pl. 2. 1901.

Macairea schultesii Wurdack, Bot. Mus. Leafl. 18: 164. 1958. Macairea maguirei Wurdack, Phytologia 20: 36. 1970. Shrub or treelet 0.5–2(–8) m tall. Humid savannas, open areas, montane and tepui scrub, 100–2000 m; Bolívar (Cerro Pitón, Gran Sabana, Ilú-tepui, Ptari-tepui, Uaipántepui), Amazonas (widespread). Southeastern Colombia, Guyana, Peru, Brazil, Bolivia. ◆Fig. 293.

22. MACROCENTRUM Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 756. 1867. by Paul E. Berry, Navina Luckana, and Kay Yatskievych Terrestrial herbs or subshrubs. Leaves opposite, isomorphic to strongly dimorphic. Flowers solitary at the ends of leafy branchets or secund in terminal few-flowered cymes, 4- or 5-merous. Hypanthium 8- or 10-costate; calyx lobes oblate to lanceolate or aristate, persistent; petals white or pinkish white, oblong-lanceolate to oblong, cuspidate-acute to rounded at the apex, glabrous or tipped with a single setula. Stamens essentially isomorphic, glabrous; anthers linear-subulate, minutely 1-pored; connective slightly prolonged, ventrally not appendaged, dorsally with an acute, descending tooth. Ovary usually 3-locular, free, with a truncate or collared apex; style glabrous; stigma punctiform. Fruit a capsule, 8- or 10-ridged. Seeds numerous, ovoid-clavate, minutely roughened. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil; ca. 22 species, 15 in Venezuela, 14 of these in the flora area. Members of the genus Macrocentrum are particularly fond of waterfalls, mostly occurring in moist, mossy, shady habitats. Key to the Species of Macrocentrum 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(4). 6.

Acaulescent, rosette-forming herb with 1–4-flowered scapes ... M. droseroides Caulescent, nonrosette-forming herbs or subshrubs; flowers solitary or in secund cymes ......................................................................................... 2 Inflorescence a secund, multi-flowered cyme; petioles 10–15 mm long ................................................................................................. M. cristatum Flowers solitary; petioles < 5 mm long ...................................................... 3 Flowers 5-merous ....................................................................................... 4 Flowers 4-merous ....................................................................................... 8 Leaves isomorphic in each pair ................................................................. 5 Leaves strongly dimorphic in each pair .................................................... 6 Leaf blade sharply serrate in the distal half ....................... M. angustifolium Leaf entire except for a single small serration on both sides near the tip ............................................................................................ M. chimantense Longer leaf blade 2–5 cm long, narrowly elliptic; calyx lobes aristate, the arista 3–5 mm long ................................................................... M. neblinae Longer leaf blade 1–2.5 cm long, elliptic to narrowly elliptic; calyx lobes acute, not aristate .................................................................................. 7

376

M ELASTOMATACEAE

7(6). 7. 8(3). 8. 9(8). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(12). 13.

Longer leaf blades narrowly elliptic, < 12 mm long ........................ M. repens Longer leaf blades elliptic, 12–27 mm long ................................ M. longidens Leaf blades uniformly pubescent on both surfaces ....... M. brevipedicellatum Leaf blades glabrous or pubescent only near the margins ....................... 9 Internodes bearing minute, incurved trichomes .................................... 10 Internodes glabrous ................................................................................. 11 Pedicels 1–3 mm long; petals esetulose ............................................ M. minus Pedicels 7–10 mm long; petals setulose-glandular ............... M. steyermarkii Pedicels 12–16 mm long; leaves not ciliate ..................................... M. huberi Pedicels < 6 mm long; at least the young leaves leaves ciliate .............. 12 Hypanthium with fine trichomes over the entire surface ......... M. rubescens Hypanthium with fine trichomes only on the torus ............................... 13 Petals 12–14 mm long; anthers with filaments ca. 6 mm long and thecae 3.5 mm long; leaf length:width ratio 2–2.3 ......................... M. anychioides Petals ca. 5.5 mm long; anthers with filaments ca. 3 mm long and thecae 1.1 mm long; leaf length:width ratio 3–4.5 ............................. M. maguirei

Macrocentrum angustifolium Gleason, Mem. New York Bot. Gard. 8: 137. 1953. Densely branched subshrub 20–40 cm tall; petals pink; stamens yellow. Moist shady streambanks, 1500–2000 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 296. Macrocentrum anychioides Gleason, Mem. New York Bot. Gard. 8: 138. 1953. Terrestrial or epiphytic herb 5–15 cm tall; leaves dimorphic in each pair, 1-veined; petals pink to purple; stamens white. On wet rocks along streams in montane forests, 1200–2000 m; Amazonas (Cerro Cuao, slope of Cerro Marahuaka, Cerro Sipapo). Endemic. ◆Fig. 300. Macrocentrum brevipedicellatum Wurdack, Mem. New York Bot. Gard. 10(5): 156. 1964. Terrestrial or epiphytic herb 5–12 cm tall; leaves entire or indistinctly crenulate-serrulate, strongly dimorphic in each pair, the small leaves to 5 mm long, the large leaves to 30 mm long; petals white; calyx red. Mossy areas on rocks and tree trunks in montane forests, 500–1200 m; Bolívar (summit of Amaruay-tepui, La Escalera, Macizo del Chimantá [slopes of Tirepón-tepui], Sierra de Lema). Guyana. ◆Fig. 302. Macrocentrum chimantense Wurdack, Mem. New York Bot. Gard. 50: 103. 1989. Herb to subshrub 10–20 cm tall; leaves isomorphic or slightly dimorphic in each pair; petals pink; anthers yellow. Moist rocks at base of canyon along streamlets, 1800–

1900 m; Bolívar (Macizo del Chimantá [Amurí-tepui]). Endemic. Macrocentrum cristatum (DC.) Triana, Trans. Linn. Soc. London 28: 79. 1871. —Salpinga cristata DC., Prodr. 3: 113. 1828. Herb or subshrub 10–30 cm long; leaves isomorphic in each pair. Venezuela, Guyana, Suriname, French Guiana, Brazil; 3 varieties, 1 in Venezuela. M. cristatum var. parviflorum (DC.) Cogn. in Mart., Fl. Bras. 14(4): 59. 1886. —Salpinga parviflora DC., Prodr. 3: 113. 1828. Moist shady boulders near waterfall in lower montane forests, ca. 500 m; Bolívar (Sierra de Lema). Guyana, French Guiana. Macrocentrum droseroides Triana, Trans. Linn. Soc. London 28: 80. 1871. Acaulescent terrestrial to epiphytic herb 6–50 cm tall; leaves rosulate, spatulate, dimorphic in each pair; flowers 1–4 per peduncle; peduncle scapose, 3–5 cm long; petals white to light pink or white with pink blush on back; fruit erect. Cliff overhangs, shady streambanks, on mossy boulders in lowland to montane forests, 100–1800 m; Bolívar (Amaruay-tepui, Carrao-tepui, La Escalera, Macizo del Chimantá [Abacapátepui, Tirepón-tepui], Río Tírica, Sierra de Lema), Amazonas (Río Mawarinuma, Sierra de la Neblina). Guyana. ◆Fig. 303. The collection from Amazonas are much larger plants than those from Bolívar.

Macrocentrum 377

Fig. 296. Macrocentrum angustifolium

Fig. 297. Macrocentrum maguirei

Fig. 298. Macrocentrum minus

Fig. 299. Macrocentrum repens

Fig. 300. Macrocentrum anychioides

378

M ELASTOMATACEAE

Fig. 301. Macrocentrum neblinae

Fig. 302. Macrocentrum brevipedicellatum

Fig. 303. Macrocentrum droseroides

Fig. 304. Macrocentrum rubescens

Fig. 305. Macrocentrum steyermarkii

Maieta 379

Macrocentrum huberi Wurdack, Phytologia 69: 317. 1990. Subshrub 20–40 cm tall; leaves fleshy; petals lilac, stamens yellow. Meadows over granitic base, ca. 2100 m; Bolivar (Sierra de Maigualida). Endemic. Macrocentrum longidens (Gleason) Wurdack, Mem. New York Bot. Gard. 10(1): 114. 1958. —Diolena longidens Gleason, Mem. New York Bot. Gard. 8: 140. 1953. Herb to subshrub 10–20 cm tall; leaves dimorphic in each pair; petals white. Wet cliffs, 1000–1300 m; Amazonas (Cerro Sipapo). Endemic. Macrocentrum maguirei Wurdack, Mem. New York Bot. Gard. 10(1): 115. 1958. Herb to subshrub 5–20 cm tall; petals pink. Moist, mossy, shady areas near waterfalls, 1900–2200 m; Amazonas (Cerro Coro Coro, Cerro Yutajé). Endemic. ◆Fig. 297. Macrocentrum minus Gleason, Fieldiana, Bot. 28: 432, t. 92. 1952. Terrestrial or epiphytic herb 6–15 cm tall; branchlets moderately setulose with deciduously gland-tipped hairs; leaves dimorphic in each pair, subcoriaceous; petals white. Mossy sites in moist montane forests, 500–2600 m; Bolívar (Carrao-tepui, Ilú-tepui, Kamarkawarai-tepui, Sierra de Lema). Guyana. ◆Fig. 298. Macrocentrum neblinae Wurdack, Mem. New York Bot. Gard. 10(1): 114. 1958. Small shrub 10–60 cm tall; leaves strongly

dimorphic in each pair, the smaller ones ovate and sessile, < 2 mm long, thin and distantly ciliolate-serrulate, the large ones lanceolate and narrowly acute at both ends, 1.8– 5 cm long; calyx lobes aristate; petals white. Mossy rocks along streamsides in montane and lower montane forests, 300–1400 m; Amazonas (Cerro Cuao, Río Coro Coro, Sierra de la Neblina). Guyana, Brazil. ◆Fig. 301. Macrocentrum repens (Gleason) Wurdack, Mem. New York Bot. Gard. 10(1): 114. 1958. —Diolena repens Gleason, Bull. Torrey Bot. Club 75: 541. 1948. Herb to subshrub 7–20 cm tall; leaves dimorphic in each pair, the large ones lance-elliptic, acute at broad ends, the small ones broadly lanceolate and sessile. Base of cliffs with water seepage, streamsides, 500–1500 m; Bolívar (La Escalera, Ptari-tepui, Sierra de Lema). Guyana. ◆Fig. 299. Macrocentrum rubescens Gleason, Mem. New York Bot. Gard. 8: 138. 1953. Herb to subshrub 5–20 cm tall; petals white, pink-tipped. Mossy montane forests, wet cliffs, 900–1200 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 304. Macrocentrum steyermarkii Wurdack, Phytologia 14: 267. 1967. Herb 7–15 cm tall; leaves strongly dimorphic in each pair; flowers white. Rocky ledges by waterfalls, moist bluffs, 1000–1700 m; Bolívar (Amaruay-tepui, Sierra de Lema). Endemic. ◆Fig. 305.

23. MAIETA Aubl., Hist. Pl. Guiane 443. 1775. by Navina G. Luckana Glandular-setose shrubs or subshrubs. Leaves strongly dimorphic in each pair, one with a blade-impressed ant domatium and the other nonvesicular. Flowers solitary or in short cymes in upper leaf axis, 5-merous. Hypanthium terete or winged; external calyx teeth subulate or shortly projecting; petals white to pink, rounded or retuse at the apex. Stamens isomorphic, glabrous; anthers yellow, subulate, emarginate at the apex and with a dorsally inclined pore, prolonged ventrally below the filament insertion and bilobed, not appendaged. Ovary 5-locular, 1/2–2/3-inferior, with a conic truncate apex; style glabrous; stigma capitellate. Fruit a many-seeded berry. Seeds 0.5–0.8 mm long, pyramidal, minutely pebbled, with lateral raphe. Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, all in the flora area.

380

M ELASTOMATACEAE

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

Hypanthium 10-winged, the wings glandular-setose but the rest glabrous ............................................................................................... M. neblinensis Hypanthium terete, sparsely to moderately stellate-scurfy as well as glandular-setose ..................................................................................... 2 Flowers solitary or in cymes of 2 or 3 flowers; bracteoles in 2 or 3 pairs, 4–7 mm long, the outer pair basally united ......................... M. guianensis Flowers in short cymes; bracteoles 1 pair, ca. 1.5 mm long, free .................................................................................................. M. poeppigii

Maieta guianensis Aubl., Hist. Pl. Guiane 443, t. 176. 1775. —Melastoma maieta Desr. in Lam., Encycl. 4: 34. 1797. —Tococa maieta D. Don, Mem. Wern. Nat. Hist. Soc. 4: 305. 1823. Shrub (0.5–)1–2 m tall; petals white to pink; fruits sessile, red or orange-red. Evergreen lowland to lower montane forests, 100–1100 m; Bolívar (Amaruay-tepui, Auyántepui, Cerro Camarón, Gran Sabana, Kavanayén, Macizo del Chimantá, Rio Cusimi, Rio Erebato basin, Río Canarakuni, near Urimán), Amazonas (Cerro Aratitiyope, slopes of Cerro Duida, Cerro Huachamacari, Platanal, Rio Caura, Río Cunucunuma, Rio Mawarinuma, Rio Siapa, San Carlos de Rio Negro). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 308.

Maieta neblinensis Wurdack, Mem. New York Bot. Gard. 10(5): 178. 1964. Shrub 1–6 m tall; petals cream-colored to pink; fruit yellow or pale orange turning dark purple when ripe. Moist montane and lower montane forests, 600–1600 m; Amazonas (Cerro Aracamuni, slopes of Sierra de la Neblina). Brazil (Serra da Neblina). ◆Fig. 306. Maieta poeppigii Mart. ex Cogn. in Mart., Fl. Bras. 14(4): 464, t. 99. 1888.

Fig. 306. Maieta neblinensis

Maieta 381

Fig. 307. Maieta poeppigii

Fig. 308. Maieta guianensis

382

M ELASTOMATACEAE

Shrub 0.5–2 m tall; petals white; fruits yellow to orange. Evergreen lowland to lower montane forests, secondary vegetation, 50– 900 m; Bolivar (Cerro Abismo, La Escalera, Río Caura, Rio Erebato, Río Tabaro, 53 km south of Urimán), Amazonas (Cerro Aracamu-

ni, Cerro Duida, Cerro Huachamacari, Puerto Ayacucho, Rio Cataniapo, Rio Cucucunuma, Rio Mawarinuma, Río Siapa, San Carlos de Rio Negro, Victorino). Costa Rica (Cocos Island), Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 307.

24. MALLOPHYTON Wurdack, Mem. New York Bot. Gard. 10(5): 145. 1964. by Paul E. Berry Small shrubs. Leaves opposite, stiffly coriaceous. Flowers 4-merous, in sessile, terminal dichasia. Hypanthium terete; calyx lobes subulate and persistent; petals pink, obovate. Stamens 8, slightly dimorphic, glabrous; anthers linear-subulate, minutely 1-porate; connective slightly prolonged to the insertion of the filament, basally expanded into an obovate or oblong-obovate appendage, dorsally nonappendaged. Ovary superior, 3-locular, glabrous. Fruit a 3-valved capsule. Seeds numerous, cochleate, muriculate. Endemic to the summit of the Macizo del Chimantá, Venezuela; 1 species. Mallophyton chimantense Wurdack, Mem. New York Bot. Gard. 10(5): 145, fig. 63a–j. 1964. Shrub 0.2–1 m tall; stems, lower surface of the leaves, and hypanthia woolly-pubescent; leaf blades broadly ovate to broadly elliptic-ovate, obtuse to rounded at the apex, slightly cordate at the base, rigid and with entire, recurved margins, 2–4 × 2–3 cm, 5- or 7-veined; petioles 2–4 mm long; petals 16–17 × 15–16 mm. Tepui shrublands, 2000–2500 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 309.

Fig. 309. Mallophyton chimantense

Marcetia 383

25. MARCETIA DC., Prodr. 3: 124. 1828. by Paul E. Berry and Navina G. Luckana Generally ericoid shrubs with small, decussate, needle-like leaves tightly clustered on ascending branches. Flowers 4-merous, usually solitary in upper leaf axils. Hypanthium terete; calyx lobes persistent; petals pink or white, obovate, eciliate (in the flora area). Stamens 8, basically isomorphic, glabrous; anthers oblong-linear, 1pored; connective not prolonged, thickened ventrally at the base and joined with the thecae, not appendaged. Ovary free, 4-locular (in the flora area), glabrous; style glabrous; stigma not expanded. Capsule 4-valved (in the flora area). Seeds numerous, cochleate, impressed-punctate. Colombia, Venezuela, Guyana, Brazil, Uruguay (most diverse in Brazil and Uruguay); ca. 20 species, 1 in Venezuela. Marcetia taxifolia (A. St.-Hil.) DC., Prodr. 3: 124. 1828. —Rhexia taxifolia A. St.Hil. in Humb. & Bonpl., Monogr. Melast. 2: 150. t. 57. 1823. Rhexia juniperina Humb. in Spreng., Syst. Veg. 2: 310. 1825. —Marcetia juniperina (Humb.) DC., Prodr. 3: 125. 1828. Marcetia andicola Naudin, Ann. Sci. Nat. Bot. sér. 3, 15: 44. 1851. —Marcetia cordigera var. andicola (Naudin) Cogn. in Mart., Fl. Bras. 14(3): 446. 1885. Shrub 0.5–3 m tall; young branchlets, leaves, pedicels, and hypanthia sparsely to moderately glandular-puberulous; leaf blade 4–8 × 2–6 mm, broadly lanceolate to ovate, acute and mucronulate at the apex, cordulate at the base, subrigid and entire with margins usually strongly recurved, moderately glandular-punctate, 3-veined; pedicels 1 mm long; hypanthium 2.5 mm long; calyx lobes 2–2.5 mm long, lanceolate to oblong-lanceolate; petals pink (occasionally white), 5–7 × 3–4 mm, acute to shortly acuminate. Rocky outcrops (mostly Roraima sandstone), 800–2800 m; Bolívar (most tepuis, Gran Sabana, Santa Elena de Uairén to El Paují), Amazonas (most tepuis). Venezuelan Andes and Coastal Cordillera; Colombia (Huila, Santander), Guyana, southeastern Brazil. ◆Fig. 310.

Fig. 310. Marcetia taxifolia

384

M ELASTOMATACEAE

26. MERIANIA Sw., Fl. Ind. Occid. 2: 823. 1798, nom. cons. Adelbertia Meisn., Pl. Vasc. Gen. 1: 114. 1838. Pachymeria Benth., Pl. Hartw. 2: 130. 1844. Schwerinia H. Karst., Auswahl Gew. Venez. 12. 1848. Notocentrum Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 131. 1852. by Navina G. Luckana and Paul E. Berry Trees or shrubs. Leaves opposite, isomorphic, mostly coriaceous to rigid. Inflorescences usually terminal panicles, occasionally flowers solitary or ternate. Flowers generally 5-merous. Hypanthium usually terete; calyx truncate or with distinct persistent lobes, occasionally closed in bud and dehiscing irregularly at anthesis; petals usually purple-pink, obovate, rounded to truncate at the apex. Stamens isomorphic or dimorphic, glabrous; thecae subulate, 1-pored, connective not or slightly prolonged, dorsally with a basal spur and often with a blunt ascending appendage. Ovary 3- or 5-locular; style glabrous; stigma not expanded. Fruit a capsule. Seeds numerous, narrowly oblong-pyramidal. Central America, Antilles, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; ca. 50 species, 11 in Venezuela, 6 of these in the flora area. Meriania is not clearly distinguished from Graffenrieda, Centronia, Axinaea, or Adelobotrys. Many of its species have large, showy petals. Key to the Species of Meriania 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4).

5.

Leaves nearly sessile (petiole 2–5 mm long), cordate at the base ............................................................................................. M. sclerophylla Leaves with petioles ≥ 5 mm long, obtuse to rounded at the base ........... 2 Petioles 5–10 mm long; leaf blades 4–8 × 2–4.5 cm ...................... M. broccha Petioles > 15 mm long; leaf blades 7–23 × 3–13 cm .................................. 3 Petioles 7–11 cm long; petals 31–35 × 21–35 mm; lowland forests ...................................................................................................... M. ornata Petioles 1.5–6 cm long; petals 11–28 × 7–20 mm; lowland to mostly upland forests ..................................................................................................... 4 Shrubs < 1 m tall; leaf blades 7–8.5 × 3–4 cm, 3-veined; petioles 2–4 cm long; petals ca. 28 × 16–17 mm ............................................ M. neblinensis Shrubs or trees 2–15 m tall; leaf blades 7–16 × 4–9 cm, 3- or 5-veined; petioles 1.5–6 cm; petals 11–25 × 7–20 mm .......................................... 5 Leaf apex obtuse to rounded, base without auricles at base; petioles 1.5– 2.5 cm long; calyx dehiscing into irregularly deciduous lobes; petals 20– 25 × 18–20 mm ..................................................................... M. crassiramis Leaf apex shortly acuminate, base with inconspicuous, deflexed auricles on lower surface; petioles 2–6 cm long; calyx truncate; petals 11–17 × 7–10 mm ................................................................................... M. urceolata

Meriania broccha Wurdack, Phytologia 69: 316. 1990. Small shrub 30–100 cm tall; leaf blades (4–)6–8 × (2–)2.5–4.5 cm, 3(5)-veined; petioles 5–10 mm long; inflorescence 5–7-flowered; calyx lobes with thick teeth 5–9 mm long; petals 21–2 × 15–17 mm, pink-purple. Among rocks on tepui slopes, 1600–1900 m;

Amazonas (Sierra de la Neblina). Endemic. Meriania crassiramis (Naudin) Wurdack, Brittonia 39: 159. 1987. —Davya crassiramis Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 138. 1852. —Centronia crassiramis (Naudin) Triana, Trans. Linn. Soc. London 28: 72. 1871.

Meriania 385

Tree (3–)9–15 m tall; leaf blades 7–15 × 4–7 cm, 3(5)-veined; petioles 1.5–2.5 cm long; calyx 10–13 m long and externally 5-carinate, dehiscing 3–4 mm above the torus into irregular deciduous lobes; petals 20–25 × 18– 20 mm, initially pink, then white. Montane and upper montane forests, 1000–2100 m; Bolívar (Aprada-tepui, Ilú-tepui, Karauríntepui, Ptari-tepui, Roraima-tepui). Guyana. ◆Fig. 313. Meriania neblinensis Wurdack, Mem. New York Bot. Gard. 18(2): 288. 1969. Shrub ca. 80 cm tall; leaf blades 7–8.5 × 3–4 cm, 3-veined; petioles 2–4 cm long; inflorescence ca. 5-flowered; petals ca. 28 × 16–17 mm, pink. Montane scrub forests, 1300–1700 m; Amazonas (expected on Sierra de la Neblina). Brazil (Serra Pirapucú). Meriania ornata Wurdack, Acta Bot. Venez. 13: 135. 1978. Tree ca. 10 m tall; indumentum rust-colored; leaves coriaceous, 13–23 × 6–13 cm, 5veined, the lower surface silvery green, the upper surface deep green; petioles 7–11 cm long; petals 31–35 × 21–35 mm, pink. Evergreen lowland forests, 100–200 m; Amazonas

(10–30 km from Puerto Ayacucho toward Gavilán). Endemic. Meriania sclerophylla (Naudin) Triana, Trans. Linn. Soc. London 28: 67. 1871. —Davya sclerophylla Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 138. 1852. Shrub 0.3–3 m tall; leaf blades 6–10 × 4–7 cm, 3- or 5-veined, bases cordate; petioles 2– 5 mm long; calyx lobes glandular-setose; petals 15–18 × 10–12 mm. Upland savannas, 700–1300 m; Bolívar (Gran Sabana, base of Roraima-tepui). Guyana. ◆Fig. 312. Meriania urceolata Triana, Trans. Linn. Soc. London 28: 67. 1871. Meriania paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 224. 1922. Shrub or tree 2–12 m tall; leaf blades 10– 16 × 5–9 cm, base obtuse to rounded with inconspicuous deflexed auricles, 5-veined; petioles 2–6 cm long; petals 11–17 × 7–10 mm, purple-red. Mauritia palm swamps, whitesand savannas, evergreen lowland to montane forests, especially along streams, 100– 1700 m; widespread in Bolívar and Amazonas. Colombia, Guyana, Peru, Brazil, Bolivia. ◆Fig. 311.

Fig. 311. Meriania urceolata

386

M ELASTOMATACEAE

Fig. 312. Meriania sclerophylla

Fig. 313. Meriania crassiramis

Miconia 387

27. MICONIA Ruiz & Pav., Fl. Peruv. Prodr. 60. 1794, nom. cons. by Paul E. Berry Shrubs or trees (rarely woody vines). Leaves mostly opposite and isomorphic. Flowers 4, 5(–10)-merous, mostly in terminal, many-flowered panicles. Hypanthium usually terete; calyx usually regularly and weakly lobed (rarely calyptrate in bud, then hyaline and dehiscing irregularly into lobes at anthesis), usually persistent in fruit, generally with inconspicuous external teeth. Petals generally white (rarely pink), usually small, rounded to retuse or occasionally acute at the apex, usually eciliate, glabrous or granulose (rarely stellulate-puberulous). Stamens anisomorphic or isomorphic; filaments usually glabrous, rarely strigulose or glandular-setulose; anthers of various forms, usually l-pored (rarely 2- or 4-pored or with several fissures), the connective simple or basally appendaged and sometimes prolonged. Ovary barely to completely inferior, (2–)5(–10)-locular; style usually glabrous, occasionally setulose or glandular-puberulous; stigma not or somewhat expanded. Fruit a berry. Seeds pyramidate to ovoid, usually smooth. Widespread throughout the Neotropics, most speciose in the tropical Andes; ca. 1000 species, ca. 180 species in Venezuela, 120 of these in the flora area. The following key is based initially on readily seen leaf pubescence characters. Since certain species vary or are intermediate in this respect, they may be keyed out in more than one group. Within each of the four main groupings, the emphasis remains on vegetative characters, but the keys also use inflorescence types or other floral characters when they are diagnostic. To assist with the terminology of primary leaf venation types and the main types of inflorescences, see Figure 314. Some pubescence terms that appear frequently in the keys and descriptions are defined below: cobwebby (synonym, arachnoid): dense, matted indument with individual hairs generally not evident dendritic: hairs branched in a more or less tree-like fashion granulose: surface covered by a grainy or knobby texture, mainly applied to ovaries and fruits lepidote: with rounded, shield-shaped scales (often overlapping to make a lightercolored covering on the lower surface of the leaf) scurfy (synonym, furfuraceous): covered with bran-like scales or powder stellate (diminutive stellulate): hairs with narrow divisions radiating from a center like the rays of a star There are several main differences in inflorescence morphology that are highly diagnostic of individual species. The first is the case of species with secund flowers, meaning having all the flowers to the same side, and which is often associated with scorpioid side branches, or ones that are segmented-curved like a scorpion’s tail. Among the other more regularly arranged inflorescence types, there are some that are spicate or spike-like in appearance, and some species have in addition short or long basal branches. Other species are paniculately and dichotomously or polytomously branched, with the flowers clustered in different ways towards the tips of the side branches, and the flowers pedicellate or not. Sometimes the flowers form umbellate-like clusters at the ends of the side branches.

388

M ELASTOMATACEAE

SECONDARY VENATION TYPES (lower order veins not shown)

basally veined Miconia superba

pliveined Miconia splendens

strongly pliveined Miconia tomentosa

MAIN INFLORESCENCE TYPES

spicate-verticillate (type I) Miconia aplostachya

secund-scorpioid (type IV) Miconia trinervia

spicate-verticillate with basal branches (type II) Miconia radulaefolia

paniculate-umbellate (type V) Miconia mirabilis

Fig. 314. Miconia venation and inflorescence types.

paniculate-spicate (type III) Miconia fragrans

open-paniculate (type VI) Miconia splendens

Miconia 389

Key to the Four Subkeys of Miconia 1.

1.

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

Lower surface of the mature leaf blades glabrous between the primary and secondary veins (note that the veins themselves may be pubescent) ................................................................................... GROUP D (page 396) Lower surface of the mature leaf blades persistently pubescent between the veins (note that when the lower surface is strongly discolored, that is, usually pale or tan contrasting with the green upper surface, the indument is sometimes so fine as to appear glabrous without high magnification) ............................................................................................... 2 Pubescence on the lower surface of leaves of discrete, smooth, unbranched hairs ............................................................................ GROUP A (page 389) Pubescence on the lower surface of leaves of matted (non-discrete) hairs, or of dendritic, stellate, or lepidote hairs .............................................. 3 Pubescence on the lower surface of leaves completely covering the surface between the veins ..................................................... GROUP C (page 392) Pubescence on the lower surface of leaves only partially covering the leaf surface between the veins (the actual surface can be seen between or through the hairs) ..................................................... GROUP B (page 390) Key to the Species of Miconia in GROUP A (Lower surface of leaves between the veins with discrete, smooth, unbranched hairs) 10 species

1. 1. 2(1).

2.

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

6(5).

Leaf blades glabrous on the upper surface ............................................... 2 Leaf blades variously pubescent on the upper surface, at least on the younger leaves ....................................................................................... 5 Inflorescence broadly paniculate, the flowers shortly pedicellate (2– 4 mm); leaves 5-veined, somewhat subcordate at the base, short-petiolate (5–10 mm) ............................................................... M. acinodendron Inflorescence spicate or paniculate with short side branches, the flowers sessile; leaves 3-veined or 3- or 5-pliveined, the base attenuate, cuneate, or subcordate, usually with longer petioles (> 10 mm long) ..... 3 Leaf base subcordate, the blade 6–11(–15) × 2–4 cm, 3-pliveined; inflorescence interrupted-spicate .................................................. M. abysmophila Leaf base acute, attenuate, or cuneate, the blade (8–)10–30 × (2.5–)4– 13 cm, 5-pliveined, 3-veined, or shortly 3-pliveined ............................. 4 Opposite members of leaf pairs ± unequal in size, the blade shortly 5pliveined and 8–13 cm wide ............................................. M. brachybotrya Opposite members of leaf pairs ± equal in size, the blade 3-veined or often shortly 3-pliveined and 4–7 cm wide ....................................... M. matthaei Leaf blades strongly 5- or 7-pliveined, densely reticulate; inflorescence spicate ........................................................................................ M. nervosa Leaf blades basally 3- or 5-veined or shortly 5- or 7-pliveined, not densely reticulate; inflorescence branched, densely paniculate to subspicate, or with secund side branches ..................................................................... 6 Young stems, leaf nodes, and petioles with long hispid hairs 2–6 mm long; leaf margins long-ciliate; inflorescence narrowly paniculate with short side branches bearing secund flowers ......................................... M. pileata

390

6.

7(6).

7.

8(7).

8.

9(8). 9.

M ELASTOMATACEAE

Young stems, leaf nodes, and petioles with shorter pubescence; leaf margins nonciliate; inflorescence spicate-verticillate with basal branches (flowers not secund), or else the inflorescence paniculately and regularly branched ........................................................................................ 7 Inflorescence spicate-verticillate with basal branches (type II on Fig. 314), the rachis red-purple when fresh; leaf blades with fuzzy hairs 1– 2 mm long on the upper surface ......................................... M. radulaefolia Inflorescence paniculately and regularly branched, the rachis not markedly red-purple when fresh; leaf blades densely ferruginous-pubescent (hairs to 2 mm long) or with shorter pubescence than above on the upper surface .............................................................................................. 8 Leaf blades 7–13 × 2.5–5 cm, short-pubescent on the upper surface; petioles 5–10 mm long; plants typical of savannas or forest margins ................................................................................................ M. ibaguensis Leaf blades 8–20 × 4–12 cm, densely ferruginous-pubescent on the upper surface; petioles 15–50 mm long; plants typical of forest understories or riparian forests ...................................................................................... 9 Leaf blades 5-veined; teeth of calyx lobes < 2 mm long .............. M. bracteata Leaf blades shortly 5- or 7-pliveined; teeth of calyx lobes linear-bristly, 6– 8 mm long ................................................................................... M. strigosa Key to the Species of Miconia in GROUP B (Lower surface of leaves with dendritic, stellate, or lepidote hairs, the pubescence only partially covering the leaf surface between the veins [i.e., the actual surface can be seen between or through the hairs]) 21 species

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

7.

Leaves sessile or very short-petiolate (< 1 cm long and usually on largeleaved species) ........................................................................................ 2 Leaves distinctly petiolate ......................................................................... 9 Stems winged, the wings 2–3 mm wide ............................................ M. alata Stems terete to quadrangular, but not winged ......................................... 3 Bases of most leaf blades ± amplexicaul (clasping the stem) ................... 4 Bases of leaf blades not amplexicaul ......................................................... 6 Leaf blades elliptic-pandurate (much narrowed basally), strongly pliveined ................................................................................. M. tomentosa Leaf blades basally veined, not strongly narrowed towards the base ...... 5 Opposite leaves unequal in size (in a ratio of up to 1:2.5); leaves membranous, 7–15 cm long; fruit 10-ribbed ................................. M. inaequalifolia Opposite leaves equal in size; leaves coriaceous, 20–35 cm long; fruit terete ................................................................................... M. impetiolaris Leaves basally 5- or 7-veined, the blade ovate and the base cordulate; lower surface often rugose-bullate .......................................... M. rufescens Leaves 3(5)-pliveined, the blade not ovate and cordulate together; lower surface smooth to rugose ....................................................................... 7 Inflorescence spicate-verticillate; pubescence on stems and on main veins on lower surface of leaves of flexuous-erect hairs 2–7 mm long (the surface in between the veins with stellate or dendritic hairs) ........ M. rugosa Inflorescence narrowly to broadly paniculate; pubescence on leaves of

Miconia 391

8(7). 8. 9(1). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(12). 13. 14(13). 14.

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

stellate or dendritic hairs, lacking the long erect hairs on the stems and veins of the lower surface of leaves ....................................................... 8 Inflorescence rachis quadrangular, with brownish pubescence, the flowers ± umbellately clustered at the branch tips ................ M. biglandulosa Inflorescence rachis ± terete, with tan pubescence, the flowers not umbellately clustered at branch tips .................................... M. tomentosa Flowers secund on the ultimate inflorescence branches (type IV on Fig. 314) ....................................................................................................... 10 Flowers not secund .................................................................................. 11 Leaves basally 3-veined, 2.5–5 cm wide, the pubescence stellulate-scurfy; stems terete .............................................................................. M. ruficalyx Leaves shortly 3-pliveined, 5–10 cm wide, the pubescence lepidotestellulate; young stems compressed-flattened ......................... M. trinervia Inflorescence spicate or with some basal verticillate branches (types I and II on Fig. 314) ....................................................................................... 12 Inflorescence paniculate to pyramidal .................................................... 15 Leaf margin subentire, ciliate; fruits strongly 10-ribbed ....M. rhytidophylla Leaf margin denticulate-undulate, at least apically, not ciliate; fruits not ribbed.................................................................................................... 13 Leaf margin conspicuously bluntly toothed (teeth ca. 5 mm apart and 1.5– 2 mm deep at the sinuses); inflorescence 3–6 cm long .... M. grossidentata Leaf margin less conspicuously denticulate, the teeth smaller and sharper; inflorescence 5–14 cm long ................................................... 14 Mature leaves 2.5–4 cm wide; inflorescence glomerulate-verticillate, without short side branches, 8–14 cm long; flowers 5-merous ........ M. aristata Mature leaves 5–8 cm wide; inflorescence subracemose, 5–8 cm long, the lower part with very short 3-flowered branches; flowers mostly 6merous .................................................................................... M. carassana Mature leaf blades 3–5(–8) × 1.5–3 cm .......................................... M. tinifolia Mature leaf blades larger......................................................................... 16 Leaf blades 7-pliveined, 10–17 cm wide; petioles 4–13 cm long ............................................................................................... M. perobscura Leaf blades basally veined, narrower and with shorter petioles ........... 17 Pedicels 6–20 mm long; petals 8.5–9 × 4.5–5 mm; inflorescence paniculate-umbellate (type V on Fig. 314) ........................................... M. iluensis Pedicels shorter or flowers sessile; petals smaller than above; inflorescences of various types ........................................................................ 18 Inflorescence ± paniculate-umbellate; flowers with pedicels 0.7–1.3 mm long ...................................................................................... M. phaeophylla Inflorescence broad to narrow panicles, not noticeably umbellate; flowers sessile or sometimes with pedicels to 1 mm long ............................... 19 Inflorescence narrowly paniculate, at least twice as long as wide; petals 2.3–3 mm long ....................................................................... M. rubiginosa Inflorescence broader, nearly as wide as long; petals 1–2 × 1–1.5 mm .............................................................................................................. 20 Petals 1.7–2 × 1–1.5 mm; leaf blades 6–13 × 2–5(–6) cm with petioles 0.6– 1 cm long; savanna shrubs ........................................................ M. brevipes Petals 1–1.5 × ca. 1 mm; leaf blades 12–35 × 3–12 cm with petioles 1– 2.5 cm long; forest trees ....................................................................... 21

392

M ELASTOMATACEAE

21(20. Leaf blades 15–35 × 5–12 cm, with hairs 0.4–0.5 mm diameter; petioles 1.5–2.5 cm long ........................................................................... M. solmsii 21. Leaf blades 5–20 × 3–5(–8) cm, with hairs 0.1–0.3 mm diameter; petioles 1–2 cm long .......................................................................................... 22 22(21). Leaf blades 5—14 x 3—6.5 cm; anther connective with a dorso-basal spur .................................................................................................. M. buntingii 22. Leaf blades 12—20 x 3—5(—8) cm; anther connective with a simple base ................................................................................................ M. pilgeriana Key to the Species of Miconia in GROUP C (Lower surface of leaves with matted, dendritic, stellate, or lepidote hairs that completely cover the surface between the veins) 45 species 1. 1. 2(1). 2. 3(2).

3.

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

6. 7(6).

7. 8(7).

8. 9(8).

Leaves pliveined ......................................................................................... 2 Leaves basally veined .............................................................................. 12 Leaves sessile (or very shortly petiolate in relation to the size of the leaf blade) ...................................................................................................... 3 Leaves clearly petiolate ............................................................................. 4 Leaf blades 10–30 × 5–10 cm, pandurate (narrowed at the base), the main primary veins diverging 1/3–1/2 the length of the blade above the base; panicle 8–12 cm long ................................................................ M. silicicola Leaf blades 30–60 × 10–25 cm, the main primary veins diverging subalternately 1/4–1/3 the length of the blade above the base; panicle 15–35 cm long ......................................................................... M. plukenetii Inflorescence with flowers secund on the side branches .......................... 5 Inflorescence spicate or paniculate with regular branching (flowers not secund) ................................................................................................... 6 Leaf blades equal in size in each pair, 11–28 × 4–13 cm, petioles 2–3 cm long ..................................................................................... M. argyrophylla Leaf blades generally unequal in each pair, 6–20 × 2–7 cm, petioles 0.2– 0.5 cm long .................................................................................. M. serialis Leaf blades narrowly lance-oblong, 6–15 × 1.5–3.5 cm, entire, shortly (0.3–1 cm) 3-pliveined; petiole 0.3–1 cm long; inflorescence 10–15 cm long and spicate (interrupted-verticillate) ......................... M. aplostachya Leaf blades wider and larger; petioles usually longer; inflorescence paniculate ..................................................................................................... 7 Leaf blades elliptic to oblong-elliptic, apex abruptly short-acuminate, base obtuse to rounded, 20–45 × 8–25 cm, 5, 7(–11)-pliveined diverging 3–8 cm above the base; petiole 0.1–0.5(–1) cm long; panicle 20–40 cm long and nearly as wide ............................................................... M. ampla Leaves smaller or proportionately narrower and longer-petiolate; panicles smaller ..................................................................................... 8 Leaves mostly 3- or 4-whorled (some can be opposite), the blades obscurely (0.2–0.5 cm) 3-pliveined, narrowly oblong-elliptic, 10–20 × 2– 4(–6) cm; petioles 0.5–1 cm long ........................................ M. chrysophylla Leaves all opposite; leaves proportionately wider than above; petioles 0.7–3 cm long ......................................................................................... 9 Leaf blades elliptic-ovate (wider at base than at middle) ...................... 10

Miconia 393

9. Leaf blades widest close to the middle .................................................... 11 10(9). Apex of leaf blade caudate-acuminate (1.5–2 cm), base acute, blade 11–19 × 4.5–8.5 cm, 5(7)-pliveined, with the primary lateral veins emerging from different points on the midvein; petioles winged to the base due to the decurrent leaf base; panicle 6–18 cm long, many-flowered; flowers 5-merous, sessile ............................................... M. decurrens 10. Apex of leaf blade slightly acuminate, base obtuse, blade 8–18 × 3.5–7 cm, very slightly (to 0.7 cm) 5-pliveined; petioles unwinged; panicles 3– 10 cm long, few-flowered; flowers 6-merous, pedicels 2–5 cm long ................................................................................................... M. truncata 11(9). Leaf blades lance-elliptic, 10–19 × 4–7 cm, the pairs of internal primary veins diverging 0.8–2.5(–3.5) cm above the base; petioles 0.7–1.5 cm long; flowers 5-merous .................................................................... M. aguitensis 11. Leaf blades ovate-elliptic to oblong-elliptic, 12–25 × 6–12 cm, the pairs of internal primary veins diverging < 1 cm above the base; petiole 2–4 cm long; flowers 6-merous .......................................................... M. holosericea 12(1). Leaves strongly cordate-auriculate at the base, the auricles 3–5 cm wide, the blades 35–75 × 12–25 cm, 7- or 9-veined; petioles robust, 1–3 cm long; inflorescence a panicle 10–25 cm long with 6-merous flowers ................................................................................................... M. macrotis 12. Leaves acute, obtuse, or cordulate at the base, but not strongly auriculate as above, the blades generally smaller, but if as large as above, then with fewer veins; inflorescence of various shapes and sizes, if as large as above, then with 5-merous flowers ................................................. 13 13(12). Leaves cordulate (with two short rounded lobes at the base) ................ 14 13. Leaves acute to obtuse or rounded at the base ....................................... 19 14(13). Leaves subsessile (petioles to 0.5 cm long) ...................................... M. fallax 14. Leaves with petioles 0.5–9 cm long ......................................................... 15 15(14). Petioles 2–9 cm long; leaf blades 15–35 × 7–20; inflorescence 15–40 cm long ....................................................................................................... 16 15. Petioles 0.5–2 cm long; leaf blades entire to obscurely crenulate or minutely serrulate, 6–15 × 3.5–10 cm; inflorescence 5–20 cm long ....... 17 16(15). Petioles 2–4 cm long; leaf blades entire or obscurely undulatesubdenticulate, 15–30 × 7–18 cm; inflorescence verticillate-spicate, 15–40 cm long ..................................................................... M. longispicata 16. Petioles 3–9 cm long; leaf blades distinctly coarsely serrulate, 20–35 × 10– 20 cm; inflorescence branched-paniculate, 20–40 cm long ..... M. serrulata 17(15). Panicle 5–8 cm long, flowers not secund, glomerulate at branchlet ends ........................................................................................... M. alborufescens 17. Panicle 6–20 cm long, flowers secund on lateral branches ..................... 18 18(17). Panicle 6–15 cm long, the side branches divaricately branched; leaf blades elliptic-oblong, apex bluntly acute to obtuse, 6–10 × 2.5–5 cm, 5-veined ................................................................................................... M. albicans 18. Panicle 10–20 cm long, the side branches not branched; leaf blades broadly ovate-elliptic to elliptic, apex acute to rounded, 6–14 × 3.5– 10 cm, 5- or 7-veined .......................................................... M. macrothyrsa 19(13). Inflorescences spicate, spicate-verticillate, spicate-verticillate with basal branches, or spicate-paniculate (types I, II , or III on Fig. 314) ........ 20 19. Inflorescences well branched, either paniculate or secund-scorpioid (types

394

M ELASTOMATACEAE

IV, V, or VI on Fig. 314) ........................................................................ 24 20(19). Leaf blades 3–6(–10) × 1–2.5 cm, subrigid; subshrubs 0.3–1 m tall in lowland white-sand savannas .................................................. M. biglomerata 20. Leaf blades mostly larger than above, membranous to subcoriaceous; shrubs or small trees mostly of forest or forest-edge habitats ........... 21 21(20). Leaf blades 14–20 × 5–9 cm, stem and leaf with fine-stellate orangish pubescence; petioles 2–3 cm long; inflorescence 8–15 cm long ..... M. maroana 21. Leaf blades 5–17 × 2–7 cm, stem and leaf pubescence stellate-lepidote; petioles 0.5–3 cm; inflorescence 1.5–10 cm long ................................. 22 22(21). Leaf blades elliptic to obovate-elliptic, apex abruptly acuminate, 5–11 × 2–5 cm, 3-veined; petioles 0.5–1.2 cm long; inflorescence 1.5–4 cm long ............................................................................................... M. abbreviata 22. Leaf blades ovate-elliptic to oblong-lanceolate, apex gradually acuminate, 8–17 × 2.5–7 cm, 3- or 5-veined; petioles 1–4 cm long; inflorescence 3– 15 cm long ............................................................................................ 23 23(22). Leaf blades narrowly ovate to ovate-elliptic, apex gradually acuminate, base broadly acute to obtuse, 8–17 × 2.5–7 cm, 3- or 5-veined; petioles 1.5–3 cm long; inflorescence spicate with basal branches (type II on Fig. 314), 3–10 cm long ............................................................ M. borjensis 23. Leaf blades oblong-lanceolate, apex gradually long-acuminate, base obtuse to rounded, 12–18 × 3–5 cm, 3-veined; petioles 1–2(–4) cm long; inflorescence spicate (type I on Fig. 314), 10–15 cm long .................................................................................. M. pseudoaplostachya 24(19). Inflorescences narrowly paniculate with flowers secund on scorpioid side branches ............................................................................................... 25 24. Inflorescences ± dichotomously branched or umbellate panicles ........... 33 25(24). Lower surface of leaves with amorphous or cobwebby pubescence ....... 26 25. Lower surface of leaves with pubescence of distinct stellate, lepidote, or dendritic hairs ...................................................................................... 29 26(25). Branchlets distinctly quadrangular ........................................................ 27 26. Branchlets terete or flattened, but not quadrangular ............................ 28 27(26). Indument reddish, forming fuzzy bands along the primary veins on the lower surface of leaves; petals 2–2.5 mm long .................. M. argyrophylla 27. Indument whitish; petals 3.5–4 mm long ............................. M. stenostachya 28(26). Leaf blades with acuminate apex and acute to obtuse base ..... M. lourteigiana 28. Leaf blades with obtusely acute apex and rounded base ............ M. albicans 29(25). Leaves 3- or 4-whorled ........................................................... M. chrysophylla 29. Leaves opposite ........................................................................................ 30 30(29). Branchlets and lower surface of leaves with stellate-dendritic hairs; veinlets on lower surface of leaves raised-reticulate ................................. 31 30. Branchlets and lower surface of leaves with lepidote hairs; veinlets on lower surface of leaves plane ............................................................... 32 31(30). Leaf blade 10–20(–30) × 4–10(–13) cm, oblong-ovate, apex gradually acuminate, base obtuse, entire to inconspicuously undulate-denticulate; petioles 1.5–4 cm long; panicle 9–17 cm long ............ M. crassinervia 31. Leaf blade 20–35 × 7–18 cm, elliptic-oblong, apex rather abruptly shortacuminate, base broadly acute to obtuse, obscurely undulate-serrulate; petioles 2.5–5 cm long; panicle 20–30 cm long, narrowly oblong ...................................................................................................... M. dispar

Miconia 395

32(30). Branchlets alternately bluntly compressed; pubescence light brown .................................................................................................... M. lepidota 32. Branchlets sharply compressed-quadrangular; pubescence reddish brown .................................................................................................. M. punctata 33(24). Inflorescences panicles with branches of umbel-like clusters of flowers, the branches often 4-verticillate ......................................................... 34 33. Inflorescences panicles with racemose or dichasially branched side branches ............................................................................................... 40 34(33). Branchlets tetragonal, with thick wings 1 mm wide ........... M. kavanayensis 34. Branchlets terete to tetragonal, without thick wings ............................. 35 35(34). Leaf blades 6–15 × 2–5 cm; petioles 1–2(–3.5) cm long; flowers sessile or on pedicels < 1 mm long ...................................................................... 36 35. Leaf blades 10–25 × 4–13 cm; petioles 2–5 cm long; flowers on pedicels 2– 5 mm long ............................................................................................. 37 36(35). Leaf blades 5- or 7-veined, the upper surface rugulose and bullate-setulose and stellulate; hypanthium glandular-setulose ........... M. campestris 36. Leaf blades 3-veined, the upper surface glabrous with age and smooth; hypanthium not glandular-setulose ................................ M. guaiquinimae 37(35). Leaf blades 15–25 × 8–13 cm; petals externally densely scurfy ............................................................................................... M. pubipetala 37. Leaf blades 10–17 × 4–8(–11) cm; petals glabrous or pruinose-granulose .............................................................................................................. 38 38(37). Leaves 7-veined; petioles 3–6.5 cm long; flowers 6-merous, pedicels 8– 10 mm long at anthesis ............................................................. M. mituana 38. Leaves 5- or 7-veined; petioles 2–5 cm long; flowers 5- or less often 6merous; pedicels 2–5 mm long at anthesis ......................................... 39 39(38). Flowers subtended by a pair of deciduous, broadly elliptic bracts ca. 6 mm long; hypanthium puberulous with stellulate hairs ........... M. dodecandra 39. Flowers subtended by a pair of deciduous, broadly elliptic to obovate-elliptic bracts 5–8(–15) mm long; hypanthium glabrous ............... M. mirabilis 40(33). Branchlets and petioles thick-robust (to ca. 1 cm diameter), with smooth, bristly, reddish brown hairs 5–12 mm long; leaf blades 20–35 × 10– 25 cm; petioles 3–9 cm long; panicle 20–30 cm long ................. M. superba 40. Branchlets and petioles less robust, never with hairs as long as above; leaf blades and petioles smaller; panicle generally smaller as well ......... 41 41(40). Lower surface of leaves with an appressed cobwebby indument ........... 42 41. Lower surface of leaves with pubescence of distinct stellate, lepidote, or dendritic hairs ...................................................................................... 44 42(41). Petioles 4–7 cm long; leaf blades 15–25 × 8–17 cm; panicles 12–20 cm long and nearly as broad ........................................................................ M. elata 42. Petioles 0.5–5 cm long; leaf blades 10–30 × 4–12 cm; panicles 8–18 cm long, narrow to pyramidal (not as broad as long) ............................... 43 43(42). Petioles 0.5–2(–3) cm long; leaf blades elliptic to lance-elliptic; flowers sessile to pedicellate (to 3 mm long); hypanthium 7–8 mm long; ovary glandular-puberulous ........................................................... M. gratissima 43. Petioles 2–5 cm long; leaf blades elliptic to ovate-oblong; flowers sessile; hypanthium 1.2–1.4 mm long; ovary glabrous ...................... M. hypoleuca 44(41). Flowers 6-merous; petals 7–9 × 3–6 mm ................................................. 45 44. Flowers 5-merous; petals 2.7–7 × 1.5–4 mm ........................................... 46

396

M ELASTOMATACEAE

Petioles 1–3 cm long; panicle 3–8 cm long ............................... M. holosericea Petioles 3–6 cm long; panicle 10–20 cm long ................................... M. trailii Margins of leaf blades crenulate-serrulate ................................. M. bubalina Margins of leaf blades entire ................................................................... 47 Petioles 0.3–1 cm long; pubescence on lower surface of leaves reddish, with short-stalked stellate hairs .......................................... M. rubiginosa 47. Petioles 0.7–7 cm long; pubescence on lower surface of leaves either light brown-dendritic or else lepidote .......................................................... 48 48(47). Leaf blades rigid-coriaceous, the base acute; pubescence on petioles, inflorescence, and veins on lower surface of leaves dendritic, light tan; petioles 2–7 cm long ................................................................ M. acutifolia 48. Leaf blades subcoriaceous, the base obtuse to rounded; pubescence on petioles, inflorescence, and veins on lower surface of leaves lepidote; petioles 0.7–3 cm long .................................................................... M. wittii 45(44). 45. 46(44). 46. 47(46).

Key to the Species of Miconia in GROUP D (Lower surface of the mature leaf blades glabrous on the surface between the secondary and lower-order veins — the secondary veins may themselves be pubescent) 52 species 1.

1.

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

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

7(6). 7.

Mature leaf blades 2–10(–12) × 1–4(–6) cm, the leaf apex never longacuminate; plants restricted to tepui-summit vegetation (over 1500 m elevation) ................................................................................................ 2 Mature leaf blades either larger than above, or if similarly small, then not occurring on tepui summits and sometimes with the leaf apex longacuminate ............................................................................................. 11 Leaves subsessile (petioles 2–5 mm long), the base cordate-auriculate ............................................................................................ M. steyermarkii Leaves petiolate (petioles 4–25 mm long), the base acute, obtuse, rounded, or cordulate ............................................................................................ 3 Panicle 8–10 cm long; flowers 4-merous, pedicels 2–5 mm long; hypanthium ca. 3.5 mm long; petals 7–7.3 × 4.3–4.5 mm .............. M. huberi Panicle generally < 8 cm long; flowers 5-merous, sessile or with pedicels less than or equal to 3 mm long; hypanthia and petals smaller than above ....................................................................................................... 4 Leaf blades broadly elliptic to ovate-elliptic, 5.5–8.5 × 3.5–6 cm, basally 5-veined; petioles 2–2.5 cm long ............................................. M. cacumina Leaf blades narrower, 2–12 × 1–5 cm, basally veined or pliveined; petioles 0.4–2.5 cm long ...................................................................................... 5 Leaf blades 2–3 × 1.3–2 cm, basally 3-veined .......................... M. neblinensis Leaf blades 3–12 × 1–5 cm, basally veined or shortly pliveined .............. 6 Leaf blades 3–4 × 1–1.7 cm, shortly pliveined, apex acute to obtuse; petioles 0.7–1 cm long; stigma expanded, 0.9–1 mm diameter .... M. elaeoides Leaf blades 3–12 × 1.5–5 cm, basally veined or pliveined, apex acute to shortly acuminate; petioles 0.5–2.5 cm long; stigma not or slightly expanded, smaller than above .................................................................. 7 Anthers 4-pored; leaf blades 5–10 × 1.5–4 cm .......................................... 8 Anthers 1- or 2-pored; leaf blades 3–12 × 1.5–5 cm .................................. 9

Miconia 397

8(7). 8. 9(7). 9. 10(9).

10.

11(1). 11. 12(11). 12. 13(12). 13. 14(13). 14. 15(14). 15. 16(15). 16. 17(16). 17. 18(14).

18. 19(18). 19.

Calyx obviously 5-lobed (lobes 0.5–0.7 mm long); petals usually 1.9–2.1 mm long; young stems scaly-scurfy ................................................ M. rupestris Calyx barely lobed (lobes 0.1–0.3 mm long), 5-undulate; petals 0.8– 1.3 mm long; young stems glabrous or deciduously scaly ..... M. theaezans Flowers unisexual (plants dioecious); pedicels ca. 1 mm long; petals 0.7– 0.8 × 0.7–0.9 mm ........................................................................... M. dioica Flowers bisexual; pedicels 1–3 mm long; petals 1.5–2.3 × 1.5–2.2 mm .............................................................................................................. 10 Leaf blades ovate-elliptic to elliptic, apex subabruptly acuminate, base obtuse to rounded, 6–10 × 2.5–5 cm; anthers 1-pored; pedicels 2–3 mm long ...................................................................................... M. roraimensis Leaf blades elliptic to oblong-elliptic, apex shortly blunt-acuminate, base acute to obtuse, 3–8 × 1.5–3 cm; anthers 2-pored; pedicels 1–2 mm long .................................................................................................... M. tinifolia Leaves basally veined .............................................................................. 12 Leaves pliveined ....................................................................................... 30 Leaves clearly unequal in size in each pair, subsessile (petioles 2–5 mm long), the base cordate-auriculate ................................... M. inaequalifolia Leaves ± equal in size in each pair, distinctly petiolate, the base not cordate-auriculate ..................................................................................... 13 Flowers 6-merous, with pedicels 2–5 mm long; fruit 12-ribbed when dry ............................................................................................ M. amacurensis Flowers 4- or 5-merous; flowers sessile or pedicellate; fruit generally terete ......................................................................................................... 14 Flowers secund or on scorpioid side branches of the inflorescence ........ 15 Flowers not secund or on scorpioid side branches, the inflorescence regularly paniculate .................................................................................... 18 Trees 8–20 m tall; young growth densely stellulate-scurfy with appressed reddish hairs ............................................................................ M. ruficalyx Shrubs < 5 m tall; young growth fine-setulose or sparsely to moderately and deciduously stellate-scurfy ........................................................... 16 Leaf margins serrulate and densely ciliolate; blades basally 5-veined; petioles 2–6 cm long ................................................................. M. racemosa Leaf margins entire to inconspicuously serrulate, appressed-ciliate or deciduously ciliolate; petioles 0.4–3 cm long .......................................... 17 Leaf blades basally 3-veined, the margins persistently appressed-ciliate and inconspicuously serrulate ..................................................... M. ciliata Leaf blades basally 5-veined, the margins deciduously ciliolate and entire or inconspicuously serrulate .................................................... M. laevigata Leaf blades lance-oblong with a long-acuminate apex, 6–13 × 1.5–5 cm, petioles 0.5–1.8 cm long; inflorescences much-branched, pyramidal, small-flowered (petals 2–3.2 × 1–1.4 mm), with pedicels 1–2 mm long .............................................................................................................. 19 Leaf blades either larger than above, or else lacking a long-acuminate apex... ................................................................................................... 21 Petals lance-oblong with an acute apex, 2.8–3.2 × 1–1.4 mm; leaf blades basally 3-veined ................................................................... M. eugenioides Petals ovate-elliptic, 2–2.3 × 1.2 mm, the apex rounded-obtuse; leaf blades basally 3- or 5-veined ........................................................................... 20

398

M ELASTOMATACEAE

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

22.

23(21). 23. 24(23). 24. 25(24). 25. 26(25). 26. 27(26). 27. 28(27).

28.

29(28). 29. 30(11).

30.

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

Lower surface of leaf blades not noticeably glaucous ............. M. minutiflora Lower surface of leaf blades noticeably glaucous ..................... M. myriantha Flowers 4-merous ..................................................................................... 22 Flowers 5-merous ..................................................................................... 23 Trees 5–8 m tall; leaf margins undulate-serrulate, blades (3)5- or 7veined; petioles mostly 4–11 cm long; panicle 10–12 long and wide, many-flowered, the calyx conical and apiculate, dehiscing irregularly at anthesis .............................................................................. M. rupticalyx Shrubs 1–3 m tall; leaf margins entire or inconspicuously undulate-serrulate, blades 3(5)-veined; petioles 0.8–3.5 cm long; panicle shorter than above; flowers few and in groups of 3-flowered dichasia, the calyx lobes free in bud and with acute external teeth projecting 0.3–1.2 mm ................................................................................................ M. lateriflora Inflorescence branches, pedicels, and hypanthia glandular-setulose with slender hairs ................................................................................. M. polita Inflorescence branches, pedicels, and hypanthia glabrous or without glandular hairs ............................................................................................ 24 Leaves with strongly ciliate margins and rounded to cordulate base .......................................................................................... M. acinodendron Leaves with nonciliate margins, the base acute to rounded .................. 25 Calyx apiculate-conical in bud, opening irregularly at anthesis ... M. mariae Calyx open and lobed in bud .................................................................... 26 Leaf margins distantly but distinctly undulate-serrulate; pedicels 1.5– 3.5 mm long ................................................................... M. tetraspermoides Leaf margins entire to obscurely undulate- or crenulate-serrulate; pedicels 0.5–2 mm long ........................................................................ 27 Lower surface of leaves glaucous ........................................... M. phaeophylla Lower surface of leaves not glaucous ...................................................... 28 Young stems obtusely 4-angled; leaf blades 5-veined, 8–14 × 5–7 cm, the margin entire; inflorescence branches usually 4(–6)verticillate ............................................................................................... M. perturbata Young stems ± rounded; leaf blades 3- or 5-veined, 10–25 × 4–10 cm, the margin entire to obscurely and distantly undulate- or crenulate-serrulate ........................................................................................................ 29 Pedicels 0.5–1 mm long; leaf blades 10–20 × 4–8 cm, 3- or 5-veined ...................................................................................................... M. affinis Pedicels ca. 0.5 mm long; leaf blades mostly 16–25 × 7–10 cm, 5-veined ..................................................................................................... M. egensis Leaves 17–45 × 8–25 cm, petioles 0.1–1 cm long, 5, 7(–11)-pliveined, with the two lowermost pairs of primary veins diverging 3–8 cm above the base; panicles 20–40 cm long; hypanthium 6–7 mm long, calyx 3–4 mm long ............................................................................................... M. ampla Leaves smaller than above, with fewer primary veins diverging closer to the base; inflorescences and floral dimensions smaller than above .............................................................................................................. 31 Flowers 6-merous ............................................................................. M. lasseri Flowers 4- or 5-merous ............................................................................ 32 Flowers secund on somewhat curved or scorpioid side branches of the inflorescence ............................................................................................ 33

Miconia 399

32. Flowers not secund, in spicate or regularly branched panicles ............. 34 33(32). Panicle 15–25 cm long, many-flowered, the side branches with secund flowers, but not strongly scorpioid; leaf blades 13–30 × 5–10 cm, 3pliveined, the margins entire; petioles 0.5–1.5 cm long .......... M. trinervia 33. Panicle considerably shorter than above, sparsely flowered, the side branches strongly scorpioid; leaf blades 12–23 × 5.5–12 cm, 5-pliveined, the margins often crenulate; petioles 1–7 cm long ....M. pseudocapsularis 34(32). Leaves mostly 3- or 4-verticillate ............................................... M. longifolia 34. Leaves mostly opposite ............................................................................ 35 35(34). Inflorescence spicate-verticillate with basal branches or paniculate-spicate (types II and III on Fig. 314) ....................................................... 36 35. Inflorescence paniculate, more heavily branched ................................... 37 36(35). Leaves 5-pliveined, 11–30 × 6–14 cm, the base broadly acute to obtuse; leaf surface clearly showing the secondary venation .............. M. fragrans 36. Leaves 3-pliveined, 9–30 × 3–9 cm, the base narrowly attenuate; leaf surface smooth-plane, the secondary venation not evident ......M. marginata 37(35). Panicle 2–3 cm long, the short branches divaricate, subfiliform, and 1–3flowered; leaf blade with apex abruptly long-acuminate, base long-attenuate and decurrent on the petiole, 10–15 × 4–6 cm, typically undulate-margined .............................................................................. M. sprucei 37. Panicle longer than above, generally with more flowers; leaf blades of various types, but not with the above combination of characters ...... 38 38(37). Shrub 1–2 m tall, the branchlets, petioles, and inflorescences densely scurfy with dendritic hairs; leaves sometimes very unequal in size in each pair or even pseudoalternate, the blades elliptic, apex subabruptly short-acuminate to caudate, base cuneate, margins clearly undulate-crenate, 15–23 × 6–9 cm, moderately scurfy along the central vein on the lower surface, the rest of the surface glabrous; petioles 4–11 cm long; inflorescence ca. 7 cm long, the branches densely flowered; flowers 4-merous, sessile ......................................................... M. yatuensis 38. Subshrubs to trees, young growth glabrous to variously pubescent; leaves generally equal in size in each pair, the blades without the above combination of characters; inflorescences usually longer than above ...... 39 39(38). Subshrubs 0.3–1 m tall, the branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia sparsely to moderately covered with fine, smooth hairs to 1(–1.5) mm long; leaf blades inconspicuously ciliolate-serrulate, 5(7)-pliveined, the inner pair of primary veins diverging 0.5–1 cm and the second pair 1.5–4 cm above the base; inflorescence 2–9 cm long, oblong and rather few-flowered, the rachis reddish ................................................................................ M. ceramicarpa 39. Larger shrubs or trees, usually lacking the kind of hairs described above; leaf blades mostly 3- or 5-pliveined; inflorescences generally larger than above and more heavily flowered ............................................... 40 40(39). Leaf blades 5-pliveined, the lower pair of primary veins essentially submarginal and giving a thickened appearance to the margins, the next pair diverging 1.5–3.5 cm from the base; hypanthium 2.5–3 mm long, 10-ribbed, the calyx 0.6–0.7 mm long with rounded-triangular, persistent lobes ...................................................................................... M. tillettii 40. Leaf blades variously 3- or 5-pliveined, but without the above kind of ve-

400

M ELASTOMATACEAE

41(40). 41. 42(41).

42. 43(41).

43. 44(43).

44. 45(44). 45. 46(45). 46. 47(46).

47.

48(45).

48.

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

50.

51(49). 51.

nation; hypanthium usually smaller than above and not conspicuously ridged; calyx lobes usually less evident than above or lacking .......... 41 Calyx apiculate-conical in bud, dehiscing irregularly at anthesis ......... 42 Calyx open in bud, truncate or regularly lobed ...................................... 43 Flowers 4-merous, in 3-flowered dichasia at the tips of the inflorescence branches, these mostly 2 per node; petioles 1.5–5 cm long ............................................................................................ M. centrodesma Flowers 5-merous, in many-flowered panicles with primary branches mostly 4 per node; petioles 1–2 cm long ................................... M. pyrifolia Petals 5–5.5 × 2.3–2.6 mm long, bracts 2.5–3.5 mm long; branchlets and inflorescence densely stellate-puberulent; petioles 4–13 cm long ............................................................................................... M. perobscura Petals and bracts smaller than above; branchlets and inflorescences glabrous to variously pubescent; petioles shorter than above ................ 44 Leaf blades narrowly elliptic to narrowly obovate, 15–22 × 4–6 cm, the apex caudate for 3–4 cm, the base attenuate, the upper surface nitid and with the secondary veins strongly impressed, the lower surface with the same veins prominent and raised ............................... M. undata Leaf blades proportionately wider and without the caudate tip or the strongly impressed secondary veins on the upper surface ................. 45 Flowers 4-merous ..................................................................................... 46 Flowers 5-merous ..................................................................................... 48 Petioles 2–4 cm long ........................................................................... M. curta Petioles 0.7–2 cm long .............................................................................. 47 Branchlets subnodose, with distinct narrow interpetiolar ridges; leaf blades oblong-lanceolate, apex acuminate to caudate, 6–10 × 2–3(–4) cm ................................................................................................ M. fragilis Branchlets not subnodose, lacking distinct interpetiolar ridges; leaf blades oblong-elliptic, apex short or subabruptly acuminate, 9–22 × 3.5–7 cm ........................................................................................ M. cautis Trees mostly 15–30 m tall; inflorescence branches, pedicels, and hypanthia glandular-setulose with slender hairs; petals glabrous .................................................................................................. M. poeppigii Shrubs or smaller trees, mostly 1–12 m tall; inflorescence and floral parts without glandular-setulose hairs; petals densely granulose externally .............................................................................................................. 49 Leaf blades ovate-oblong (wider below the middle than above it), apex gradually acuminate, base rounded .................................................... 50 Leaf blades elliptic to oblong or obovate-elliptic (widest at the middle or above the middle), apex acute to short-acuminate, base acute .......... 51 Leaf blades 8–17 × 4–7 cm, the base rounded to slightly decurrent, the petioles 0.5–1 cm long; veinlet areoles on lower surface of leaves 0.5– 0.7 mm wide ............................................................................ M. alternans Leaf blades 12–29 × 6–14 cm, the base decurrent on the petiole almost to the base; veinlet areoles on lower surface of leaves ca. 0.3 mm wide ....................................................................................... M. stephananthera Veinlet areoles on lower surface of leaves 1–3 mm wide .............. M. prasina Veinlet areoles on lower surface of leaves 0.2–0.8 mm wide ..... M. splendens

Miconia 401

Miconia abbreviata Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 9: 1145. 1927. Shrub to small tree 1–3(–8) m tall; leaf blades elliptic to obovate-elliptic, apex abruptly acuminate, base broadly acute, 5– 11 × 2–5 cm, basally 3-veined, densely tanlepidote on lower surface; petioles 0.5–1.2 cm long; inflorescence 1.5–4 cm long, few-flowered, a few basal branches to 2 cm long; flowers 5-merous, sessile, hypanthium and calyx tan-lepidote; petals white and densely lepidote outside, ca. 2.5 mm long. Lowland to lower montane forests and forest edges, 100– 800 m; Bolívar (Cerro Marutaní, El Abismo, Río Caura, Río Erebato, Río Paragua, Río Samay), Amazonas (Río Jénita, Río Negro). Amazonian Colombia, Peru (Loreto, San Martin), Brazil (Roraima). Miconia abysmophila Wurdack, Mem. New York Bot. Gard. 10(4): 39. 1961. Shrub 0.7–1.5 m tall; inflorescence, hypanthium, and primary veins on lower surface of leaves with curved-bristly trichomes 1.5–2.5 mm long; leaf blades narrowly elliptic-oblong, apex acute to shortly acuminate, base narrowly acute, subcoriaceous, margin entire, 6–11(–15) × 2–4 cm, inconspicuously 3-pliveined; petioles 1–3 cm long; panicle 4–6 cm long, interrupted-spicate; flowers 5merous, sessile, densely glomerulate; petals white and glabrous except for some marginal setae. Tepui-slope forests, ca. 1100 m; Amazonas (Cañon Grande of Sierra de la Neblina). Endemic. Miconia acinodendron (L.) Sweet, Hort. Brit. ed. 1, 1: 159. 1826. —Melastoma acinodendron L., Sp. Pl. 389. 1753. Shrub 1–4 m tall; leaf blades elliptic-oblong to ovate-oblong, apex acuminate, base rounded to cordulate, 8–18 × 3–8 cm, chartaceous and distinctly ciliolate-serrulate, usually sparsely pubescent on both sides with flexuous, smooth hairs 1–4 mm long, basally 5-veined, densely reticulatevenulose on lower surface; petioles 0.5–1 cm long; panicle 10–20 cm long, many-flowered, with lax side branches to 6 cm long; flowers 5-merous, sessile or nearly so, few-glomerulate at branchlet ends; petals white, 3 × 1.5– 1.7 mm, glabrous or externally very sparsely stellulate-puberulous, obscurely ciliolate. Ri-

parian forests, edges of granitic outcrops, seasonally flooded forests, 100–200 m; Amazonas (Mavaca, Río Casiquiare, Río Pamoni, Río Siapa, San Antonio, Síquita). Falcón, Mérida, Monagas, Sucre, Táchira, Yaracuy, Zulia; Lesser Antilles, Colombia, Trinidad, Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 317. Miconia acutifolia Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 356. 1915. Shrub or tree 2–12 m tall, the branchlets, inflorescences, petioles, and primary leaf veins on lower surface densely covered with dendritic hairs mostly 0.3–0.5 mm long; leaf blade elliptic to oblong-elliptic, apex acute to short-acuminate, base acute, 9–21 × 3–7 cm, coriaceous and entire, the upper surface glabrous and nitid, the lower surface completely covered with rufous stellate or dendritic-stellate hairs, basally 3- or 5-veined, tertiary venation impressed on upper surface and prominent on lower surface; petioles 2–7 cm long; panicle (4–)10–15 cm long and submultiflorous; flowers 5-merous and sessile; petals white and glabrous, 6–7 × 3–4 mm, obovate. Dwarf tepui-summit forests and cloud forests on tepui slopes, 1400–2300 m; Bolívar (Auyán-tepui, Cerro Jaua, Macizo del Chimantá, Roraima-tepui, Uaipán-tepui), Amazonas (slopes of Cerro Marahuaka, Cerro Yutajé, Sierra de la Neblina). Guyana (Mount Roraima), Brazil (Amazonas: Serra da Neblina). ◆Fig. 320. Miconia affinis DC., Prodr. 3: 187. 1828. Miconia microcarpa DC., Prodr. 3: 189. 1828. Miconia planinervia Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 160. 1850. Shrub or small tree 3–8 m tall; leaf blades 10–20 × 4–8 cm, oblong-elliptic to elliptic, apex subabruptly acuminate, base broadly acute to obtuse, chartaceous and entire to obscurely and distantly undulate-serrulate, the upper surface glabrous, the lower surface sparsely and deciduously stellulate-puberulous, basally 3- or 5-veined with lax venule areoles; petioles 1–2 cm long; panicle mostly 8–15 cm long; flowers 5-merous, pedicels 0.5–1 mm long; petals white, 1.8–3 × 0.8–1.2 mm, narrowly obovate. Understory of semideciduous and evergreen lowland forests,

402

M ELASTOMATACEAE

edges of granitic outcrops, near sea level to 400 m; Delta Amacuro (Río Toro, Sacupana), Bolívar (35 km southwest of Caicara, 30 km south of Manteco), Amazonas (Puerto Ayacucho to Samariapo, Raudal de Atures, Río Coro Coro west of Cerro Yutajé). Carabobo, Lara, Mérida, Miranda, Táchira, Zulia; southern Mexico to Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 316. Miconia aguitensis Gleason, Bull. Torrey Bot. Club 58: 426. 1931. Small tree 1–4 m tall; stems, inflorescence, lower surface of leaves, and hypanthium completely covered by appressed stellate trichomes; leaf blades lance-elliptic, apex acute, base acute to obtuse, firmly membranous and entire or inconspicuously undulate, 10–19 × 4–7 cm, clearly (3)5pliveined with the pairs of internal primary veins diverging 0.8–2.5(–3.5) cm above the base; petioles 0.7–1.5 cm long; panicle 7–10 cm, few-flowered, subspicate; flowers 5merous, the pedicels thick, 1–2 mm long; petals glabrous, ca. 5.5 mm × 2.5 mm, oblong. Elfin tepui-summit and slope forests, 1000– 2000 m; Bolívar (Cerro Guanay), Amazonas (Cerro Duida, Cerro Parú, Cerro Yutajé). Endemic. ◆Fig. 315. Miconia alata (Aubl.) DC., Prodr. 3: 184. 1828. —Melastoma alata Aubl., Hist. Pl. Guiane 410, pl. 158. 1775. Miconia pterophora Miq., Linnaea 18: 622. 1844. Shrub 1–3 m tall; stems 4-winged, the wings 2–3 mm wide; leaves (sub)sessile, the blades interconnected by nodal ridges, elliptic to pandurate-elliptic, 9–25 × 5–20 cm, 5(7)-pliveined with inner pair of primary veins diverging 1–4 cm above the blade base; panicle 8–15(–20) cm long; flowers 5-merous, sessile; petals white, 2–2.5 × ca. 1 mm. Disturbed forest edges, granitic outcrops, upland savannas and shrublands, 100–1300 m; Bolívar (widespread, throughout the northern part of the state and into the Gran Sabana), Amazonas (Cacurí, Cerro Parú, Ocamo, Puerto Ayacucho, Río Cataniapo). Anzoátegui, Falcón, Lara, Sucre, Trujillo, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 321.

Miconia albicans (Sw.) Triana, Trans. Linn. Soc. London 28: 116. 1871. —Melastoma albicans Sw., Prodr. 70. 1788. Shrub mostly 0.5–2 m tall; young growth completely covered with a cobwebby (at first rufous, later white) tomentum; leaf blades elliptic-oblong, apex bluntly acute to obtuse, base rounded to cordulate, 6–10 × 2.5–5 cm, basally 5-veined, coriaceous and entire to obscurely crenulate, glabrous and nitidulous on upper surface; petioles 0.5–2 cm long; panicle 6–15 cm long, many-flowered; flowers 5-merous, sessile, secund on the divaricate branches; petals white, 2.5–3 × 1.4– 1.6 mm, glabrous; ovary 3-locular, glabrous. Broadly distributed in open and mostly disturbed areas, especially savannas, 50–1300 m; widely scattered in Bolívar and Amazonas. Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 323. Miconia alborufescens Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 160. 1850. Miconia arirambae Huber, Bull. Soc. Bot. Genève sér. 2, 6: 192. 1915. Miconia cachimbensis Brade, Arch. Jard. Bot. Rio de Janeiro 16: 14. 1959. Shrub or small tree 0.5–5 m tall; young growth densely scurfy with short dendriticstellulate hairs; leaf blades ovate to ellipticovate, apex acute to short-acuminate, base cordulate, 6–15 × 4–8 cm, coriaceous and entire to obscurely crenulate, sparsely and deciduously stellulate-puberulent on upper surface, lower surface completely covered by a cobwebby indument, basally 5- or 7-veined; petioles 0.5–1.5 cm long; panicle 5–10 cm long, many-flowered, narrowly pyramidal; flowers 5-merous, sessile, glomerulate at branchlet ends, with subpersistent bracteoles ca. 1 mm long; petals white, 2.5–2.8 × 1.5–1.8 mm, obovate and emarginate, glabrous; ovary 3-locular, the apex moderately granulose-puberulent. Lowland to upland savannas, 100–1200 m; Bolívar (Cerro Bolívar, Cerro Guaiquinima, Ciudad Piar, El

Miconia 403

Paují, Gran Sabana, Kavanayén, Río Caruay, Río Hacha, Río Kukenán, Río Paragua near Raudal Auraima, Río Suapure, Santa Elena de Uairén, Serranía de los Pijiguaos), Amazonas (Cerro Calentura in headwaters of Río Parucito, Puerto Ayacucho, Rincones del Chacorro, Río Coro Coro west of Cerro Yutajé, Río Manapiare, Río Mawarinuma, Río Ocamo, Río Parú, Santa Bárbara, slopes of Sierra de la Neblina). Adjacent Guyana, Brazil (Pará, Roraima, and then disjunct to southern planalto). ◆Fig. 325. Miconia alternans Naudin, Melast. Monogr. Descr. 720. 1853. Shrub 1–4 m tall, the branchlets roundedquadrangular; young growth moderately but deciduously stellulate-puberulous; leaf blades ovate-oblong, apex gradually acuminate, base rounded and shortly decurrent, 8–17 × 4–7 cm, ovate-oblong, chartaceous and entire to undulate-denticulate, glabrous on upper surface, lower surface glabrous but microscopically dense-granulose, shortly (0.5–0.8 cm) 3-pliveined (excluding the tenuous marginal veins), with the veinlets rather laxly reticulate; petioles 0.5–1 cm long; panicle 10–20 cm long, many-flowered, ± pyramidal; flowers 5-merous, pedicels ca. 0.5 mm long; petals white, 2.7–3 × 1.2–1.4 mm, narrowly obovate-oblong, densely granulose; ovary 3locular, apically granulose. Edges of evergreen lowland forests, sea level to 300 m; Delta Amacuro (Caño Jotajana, Río Amacuro, Río Cuyubini), Amazonas (Río Asisa). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 324. Miconia amacurensis Wurdack, Acta Bot. Venez. 2: 373, fig. 4. 1967. Small tree 2–16 m tall; young growth sparsely and deciduously stellulate-scurfy; leaf blades elliptic, apex gradually bluntacuminate, base acute, 6–11 × 2–5 cm, chartaceous and entire, glabrous, basally 3veined; petioles 0.7–1.5 cm long; panicle 5–7 cm long, few-flowered, oblong; flowers 6merous, pedicels 3.5–4.5 mm long; petals white, 6–6.5 × 2–2.5 mm, oblong, obscurely granulose; ovary 4-locular, the apex granulose; fruit 12-ribbed when dry. Evergreen lowland and semideciduous forests, sea level to 400 m; Delta Amacuro (Río Toro, San Vic-

tor), Bolívar (Maravaca south of Los Castillos, Río Paramichí). Guyana, French Guiana, Brazil (Pernambuco). Miconia ampla Triana, Trans. Linn. Soc. London 28: 101. 1871. Miconia involucrata Donn. Sm., Bot. Gaz. (Crawfordsville) 37: 209. 1904. Miconia megaphylla Gleason, Bull. Torrey Bot. Club 59: 363. 1932. Shrub or tree (1.5–)5–15 m tall; young branchlets and lower surface of leaves with an evanescent, amorphous, appressed white indument, the nodes with an interpetiolar line; leaf blades elliptic to oblong-elliptic, apex abruptly short-acuminate, base obtuse to rounded, 20–45 × 8–25 cm, chartaceous and entire to obscurely undulate, glabrous on upper surface, 5, 7(–11)-pliveined with pairs of primary lateral veins diverging 3–8 cm above the base; petioles 0.1–0.5(–1) cm long; panicle 20–40 cm long, many-flowered, the branchlets and hypanthia deciduously stellulate-puberulous; flowers 6-merous, sessile, enveloped by a pair of caducous bracts 6–8 mm long; hypanthium 6–7 mm long, calyx 3–4 mm long; petals white, ca. 5–8 × 3– 4.5 mm, obovate-oblong, glabrous; ovary 4locular, glabrous. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (Río Paramichí, El Paují), Amazonas (Caño Yureba in lower Río Ventuari basin, La Esmeralda, Raudal Gallineta on Río Siapa, San Carlos de Río Negro). Anzoátegui, Sucre, Táchira, Zulia; Central America, Jamaica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 327. Miconia aplostachya (Bonpl.) DC., Prodr. 3: 183. 1828. —Melastoma aplostachya Bonpl. in Humb. & Bonpl., Monogr. Melast. 1, t. 1. 1806. —Boyuyo rebalsero. Miconia macrostachya DC., Prodr. 3: 190. 1828. Shrub mostly 1–3(–4) m tall; young growth completely covered by an appressed stellulate indument; leaf blades narrowly lance-oblong, apex acute and apiculate, base narrowly decurrent-acute, 6–15(–20) × 1.5– 3.5 cm, coriaceous and entire, deciduously stellulate-scurfy on upper surface, shortly (0.3–1 cm) 3-pliveined; petioles 0.3–1 cm

404

M ELASTOMATACEAE

long; inflorescence mostly 10–15 cm long, spicate; flowers 5-merous, sessile, interrupted-verticillate, with small deltoid bracteoles; petals white, 2–2.5 × 2 mm, minutely granulose; ovary 3-locular, glabrous. Savannas, edge of granitic outcrops, upper limit of seasonally flooded riparian forests, 50–900 m; widely scattered in Delta Amacuro, Bolívar, and Amazonas. Apure, Anzoátegui, Guárico; Guyana, Suriname, Amazonian Colombia and Brazil. ◆Fig. 319. Miconia argyrophylla DC., Prodr. 3. 181. 1828. Miconia longistyla Steud., Flora 27: 724. 1844. Miconia argyrophylla var. attenuata Cogn. in Mart., Fl. Bras. 14(4): 296. 1887. Shrub or small tree (1–)3–5(–10) m tall; young growth covered with a rusty, rather deciduous, stellulate pubescence underlain by a persistent, appressed cobwebby indument; branchlets quadrangular, becoming terete with age; leaf blades elliptic to oblongelliptic, apex subabruptly acuminate, base acute to subrotund, 11–28 × 4–13 cm, chartaceous and entire or obscurely undulate, glabrous on the upper surface, completely covered with a pale cobwebby tomentum and flocculose along the costa on the lower surface, basally 3-veined (excluding the tenuous marginal veins) or obscurely pliveined; petioles 2–3 cm long; panicle 5–17 cm long, many-flowered; flowers 5-merous, sessile, secund on the short branchlets, with persistent bracteoles ca. 1 mm long; petals white, 2–2.5 × 1–1.3 mm, narrowly obovate-oblong, minutely granulose and apically glandularciliolate; ovary 3-locular, the apex sparsely granulose. Evergreen lowland to montane forests, riparian forests, 100–1200 m; Bolívar (Gran Sabana, Guayaraca, Rio Caura, Rio Karaurín, Sierra Imataca), Amazonas (slopes of Cerro Duida, western base of Cerro Guaiquinima, El Pozo southeast of San Fernando de Atabapo, Ocamo, Pimichín, San Carlos de Río Negro, Yavita). Carabobo, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 326. Miconia aristata Gleason, Bull. Torrey Bot. Club 58: 427. 1931. Shrub 1–2 m tall; young growth densely covered by stipitate-stellate hairs; leaf

blades narrowly elliptic-oblong, apex shortly and gradually acuminate, base acute, 9–16 × 2.5–4 cm, membranaceous and deeply ciliolate-denticulate, glabrous and rugose on the upper surface, sparsely stellate-puberulent on the lower surface between the veins, basally 3-veined (excluding the faint marginal veins) and the veinlets on the lower surface reticulate; petioles 1–2 cm long; panicle many-flowered, with several elongate branches to 14 cm long; flowers 5-merous, sessile, and glomerulate-verticillate; petals ca. 2.2 × 1 mm, each with a minute subterminal seta; ovary 3-locular, glabrous. Tepuislope forests and base of cliffs, 1000–1300 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Adjacent Brazil. Miconia biglandulosa Gleason, Bull. Torrey Bot. Club 59: 367. 1932. Shrub or small tree 2–8 m tall; stems 4angled; stems, branches, and lower surface of leaves stellate-scurfy with brownish trichomes; leaves (sub)sessile, the blades oblong-obovate, apex sharply acuminate, cuneately narrowed from below the middle to a subcordate base, 25–35 × 10–15 cm, subentire, 5-pliveined, lower surface densely stellate, especially on the veins; panicle narrow, densely scurfy, rachis 4-angled, 10–25 cm long, freely branched; flowers 5-merous, sessile, ± umbellately clustered at the branch tips; ovary 3-locular, glabrous. Montane and lower montane forests, 200–1000 m; Bolívar (base of Cerro Camarón), Amazonas (saddle between Cerro Duida and Cerro Marahuaka). Colombia, Ecuador, Peru, Brazil (Amazonas). Miconia biglomerata (Bonpl.) DC., Prodr. 3: 183. 1828. —Melastoma biglomerata Bonpl. in Humb. & Bonpl., Monogr. Melast. 1: 34, t. 15. 1807. Clidemia campestris var. pauciflora Benth., J. Bot. (Hooker) 2: 308. 1840. Subshrub 0.3–1 m tall; young growth densely stellate-puberulent; leaf blades lanceolate to oblong-lanceolate, apex narrowly acute, base rounded, 3–6(–10) × 1–2.5 cm, subrigid and inconspicuously denticulate (but not ciliolate), deciduously stellate-puberulent on upper surface, very sparsely glandular-setulose (hairs ca. 0.2 mm long) on lower surface, basally 3- or 5-veined with the

Miconia 405

veinlets reticulate-raised on the lower surface; petioles 0.5–1 cm long; inflorescence unbranched, with 2 interrupted glomerules of flowers; flowers 5-merous, sessile; petals white, glabrous, ca. 2 × 1 mm, obovate-oblong; ovary 3-locular, sparsely stellate-puberulent at the apex. White-sand savannas, 100–200 m; Amazonas (Caño Caname, Río Atabapo, Yapacana savannas). Endemic. ◆Fig. 318.

shaggy-pubescent, with few branches near the base; flowers 5-merous, sessile in bracteolate verticels; petals white and very minutely granulose, ca. 3 × 1.7–2 mm, obovate-oblong; ovary 3-locular, the apex inconspicuously glandular-puberulent. Evergreen lowland to lower montane forests, 100–300 m; Amazonas (base of Cerro Imeri, Río Pasimoni, Sierra de la Neblina). Amazonian Ecuador, Peru, and Brazil.

Miconia borjensis Wurdack, Mem. New York Bot. Gard. 10(4): 40. 1961. Shrub 1–3 m tall; young growth densely covered with appressed, stellate-sublepidote hairs; leaf blades narrowly ovate to ovate-elliptic, apex gradually acuminate, base broadly acute to obtuse, 8–17 × 2.5–7 cm, thin-coriaceous and undulate-serrulate, glabrous on the upper surface, basally 3- or 5veined or shortly pliveined, the veinlets on the lower surface inconspicuous and laxly reticulate; petioles 1.5–3 cm long; inflorescence paniculate-spicate, 3–10 cm long, oblong and several- to many-flowered; flowers mainly 6merous, sessile or shortly (1–2 mm) pedicellate; petals white, externally moderately granulose-sublepidote in the middle, 6–6.5 × 4–4.5 mm, obovate; ovary 6-locular, apically stellate-puberulent; mature fruits ribbed. Seasonally flooded river banks, river edges, gallery forests, savannas, near sea level to 300 m; Delta Amacuro (Corisal, Sacupana), Bolívar (Cerro San Borja, Macarapa on Río Paragua, Puerto Ordaz, Río Caura), Amazonas (Isla Ratón, Río Manapiare near San Juan, San Fernando de Atabapo). Apure, Anzoátegui. ◆Fig. 328.

Miconia bracteata (DC.) Triana, Trans. Linn. Soc. London 28: 1871. —Clidemia bracteata DC., Prodr. 3: 162. 1828. Miconia demerarensis Gleason, Bull. Torrey Bot. Club 75: 551. 1948. Shrub 0.5–3 m tall, the young growth moderately to densely fine-setose with smooth or obscurely barbellate hairs mostly 1.5–2 mm long; leaf blades elliptic to oblongelliptic, apex acuminate, base broadly acute to obtuse, 8–20 × 4–12 cm, firm-chartaceous and minutely serrulate to subentire and ciliolate at the margin, sparsely appressedsetulose or setose on upper surface, sparsely fine-setose on lower surface, basally 5-veined (including the attenuated outer pair); petioles 1.5–3.5 cm long; inflorescence 5–10 cm long and glomerulate-spicate or with short branches basally; flowers 5-merous, sessile; petals white, 4.5–6 × 1.7–2.5 mm, oblong, glabrous; ovary 5-locular, the apex densely setulose. Evergreen lowland to montane forests and forest edges, near sea level to 1400 m; Delta Amacuro (Güiniquina), Bolívar (Altiplanicie de Nuria, Amaruay-tepui, El Paují, Karaurín-tepui, La Escalera, Macizo del Chimantá [slopes of Abacapá-tepui, Apacará-tepui], Río Chicanán, Río Cuyuní, Río Tírica, Sierra de Lema), Amazonas (Cerro Duida and saddle between Duida and Cerro Marahuaka). Trinidad, Guyana, Suriname, French Guiana, Brazil (Amapá). ◆Fig. 330.

Miconia brachybotrya Triana, Trans. Linn. Soc. London 28: 111. 1871. Small tree 2.5–4 m tall, the young growth densely loose-strigulose and with fine, soft hairs mostly 1–2 mm long; leaf blades somewhat different in size in each pair, the blades oblong-elliptic to elliptic, apex acute to shortly obtuse-acuminate, base broadly acute, 15–30 × 8–13 cm, membranous and entire but densely ciliolate, upper surface glabrous, lower surface sparsely fine-setulose, shortly 5-pliveined (the lower pair of primary veins diverging 0.7–2 cm above the base) with the veinlets on the lower surface rather densely reticulate (areoles 0.4–0.5 mm wide); petioles 1–5 cm long; panicle 3–5 cm long,

Miconia brevipes Benth., J. Bot. (Hooker) 2: 313. 1840. Miconia plebeia Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 170. 1850. Miconia rubiginosa var. kuhlmannli Hoehne, Anexos Mem. Inst. Butantan, Secc. Bot. 1(5): 136. 1922. Shrub or small tree 0.5–4 m tall, the young growth densely covered with reddish stipitate- and sessile-stellate hairs; leaf blades oblong-lanceolate, apex acuminate,

406

M ELASTOMATACEAE

base rounded, 5–8(–13) × 2–4(–6) cm, rather firm and entire, upper surface glabrous and nitid except along the costa, lower surface densely covered with mostly sessile-stellate hairs and eventually glabrescent, basally 3(5)-veined; petioles 0.3–1 cm long; panicle 5–15 cm long, many-flowered, ± pyramidal; flowers 4, 5(–7)-merous, usually sessile; petals white, 1.7–2 × 1–1.5 mm, obovate, granulose. Rocky savannas, open areas, edges of Mauritia palm swamps, shrublands on bauxite deposits, 100–1000 m; Bolívar (Cerro Bolívar, base of Cerro Jaua, Cerro Perro in upper Río Paragua basin, southwest of El Manteco, Guri area, La Paragua to Ciudad Piar, Maripa to Aripao, Río Caroní near Arekuna, Río Parguaza, headwaters of Río Túriba, base of Roraima-tepui, San Francisco de Yuruaní, Santa Elena de Uairén), Amazonas (La Esmeralda, Puerto Ayacucho, Río Cunucunuma near base of Cerro Duida, Samariapo, Santa Bárbara). Sucre; Guyana, Suriname, northern Brazil. ◆Fig. 333.

Colombia, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 334.

Miconia bubalina (D. Don) Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 244. 1850. —Chitonia bubalina D. Don, Mem. Wern. Nat. Hist. Soc. 4(2): 319. 1823. —Diplochita bubalina (D. Don) DC., Prodr. 3: 177. 1828. Shrub 1–5 m tall, the branchlets, primary and secondary veins on lower surface of leaves, and inflorescences densely covered with minute dendritic hairs; leaf blades oblong-elliptic, apex acuminate, base roundedobtuse or cordulate, mostly 12–20 × 5–7 cm, thin-coriaceous and crenulate-serrulate, glabrous on upper surface, lower surface completely covered with minute stellulate hairs (much lighter colored than upper surface), basally 5-veined or barely pliveined; petioles 1–5 cm long; panicle 5–20 cm long, rather few-flowered; flowers white, predominantly 5-merous and essentially sessile, subtended by deciduous bracts 3–4 × 2–3 mm; ovary 3locular, apically stellulate-puberulous. Riparian and lowland to montane gallery forests, 100–1000 m; Bolívar (Matacuchillo ca. 25 m east-southeast of Santa Elena de Uairén, Peray-tepui west of Santa Elena, Río Acanán, Río Mawela tributary of Río Erebato), Amazonas (Río Mawarinuma, above mouth of Río Ugueto). Apure, Táchira, Zulia; southern Mexico, Central America,

Miconia cacumina Wurdack, Phytologia 64: 296. 1988. Shrub 1–1.5 m tall, the sharply quadrate branchlets, inflorescences, primary leaf veins on lower surface, and hypanthia sparsely to moderately covered with deciduous glands, otherwise glabrous; leaf blades broadly elliptic to ovate-elliptic, apex shortly and abruptly blunt-acuminate, base broadly obtuse to cordulate, 5.5–8.5 × 3.5–6 cm, coriaceous and entire, basally 5-veined; petioles mostly 2–2.5 cm long; panicle 3–7.5 cm long and submultiflorous; flowers 5-merous, the thick pedicels 0.5–1 mm long, the subpersistent subulate bracteoles 0.3 mm long; petals pinkish, 3 × 2.7-2.8 mm, subrotund, granulose; ovary 3-locular, somewhat glandular on the cone. Tepui scrub forests, 2200– 2300 m; Bolívar (expected on upper slopes of Roraima-tepui). Guyana.

Miconia buntingii Wurdack, Phytologia 21: 357. 1971. Tree ca. 8 m tall, the terete branchlets, primary leaf veins on lower surface, and inflorescences densely puberulous with dendritic hairs 0.1–0.15 mm long; leaf blades elliptic, apex subabruptly narrowed and acuminate to caudate (1–1.5 cm), base broadly acute, membranous and entire, 5–14 × 3–6.5 cm, sparsely dendritic-pubescent when young on both surfaces, but becoming glabrescent on upper surface, basally 5veined; petioles 1–2 cm long; panicle ca. 10 cm long and nearly as broad, many-flowered, the primary and secondary branches usually 4 per node; flowers 5-merous, shortly (0.4– 0.6 mm) pedicellate; petals white, inconspicuously granulose, ca. 1.2 × 1 mm, suborbicular; ovary 3-locular. Evergreen lowland forests, 100–200 m; Amazonas (Yavita). Endemic.

Miconia campestris (Benth.) Triana, Trans. Linn. Soc. London 28: 113. 1871. —Clidemia campestris Benth., J. Bot. (Hooker) 2: 308. 1840. Shrub 1–2(–5) m tall, the branchlets, lower surface of leaves, inflorescences, and hypanthia densely stellate-puberulous; leaf blades oblong-lanceolate, apex narrowly acute, base rounded, 6.5–11 × 2–4 cm,

Miconia 407

subrigid and finely ciliolate-denticulate, upper surface rugulose and deciduously bullulate-setulose as well as stellate-puberulous, lower surface sparsely glandular-setulose on the primary veins, basally 5- or 7-veined, with the veinlets on lower surface prominently elevated-reticulate; petioles 1–3.5 cm long; panicle 6–10 cm long, submultiflorous, moderately glandular-setulose; flowers 5merous and usually verticillate-glomerulate, very shortly pedicellate; petals white, 3.5–4 × ca. 2 mm, obovate-oblong, slightly granulose; ovary 3- or 4-locular, the apex densely stellulate-puberulous. Riversides, savannas, Mauritia palm swamps, gallery forests, 100– 500 m; Bolívar (Río Acanán, Río Hacha, Río Ichún, Río Toronó, Río Tírica), Amazonas (Caño Caname, Caño Cotua, La Esmeralda). Guyana, Suriname, Brazil (Río Negro). ◆Fig. 332. Miconia carassana Cogn. in Mart., Fl. Bras. 14(4): 346. 1887. Miconia compacta Gleason, Bull. Torrey Bot. Club 58: 230. 1931. Miconia semota Markg., Notizbl. Bot. Gart. Berlin-Dahlem 12: 177. 1934. Shrub 1–2(–3.5) m tall, the young growth densely tomentose with sessile or shortly stipitate stellate hairs; stems terete; leaf blades elliptic to oblong-elliptic, apex acute or subabruptly short-acuminate, base obtuse to rounded, membranous and entire to inconspicuously undulate, 9–17 × 5–8 cm, upper surface initially sparsely stellate-puberulous but soon glabrescent, lower surface sparsely to moderately stellate-puberulent on the veinlets, basally 3- or sparsely 5-veined or very shortly pliveined; petioles 1–2 cm long; inflorescence 5–8 cm long, narrowly oblong and subracemose (the short basal branchlets with 3-flowered dichasia); flowers essentially sessile, (5)6-merous; petals white, glabrous, 2.5–3 × ca. 1 mm wide; ovary 3-locular, the apex glabrous but sparsely punctate. Riparian forests, 100–200 m; Amazonas (Río Guainía near Victorino). Colombia (Amazonas, Vaupés), Ecuador (Napo), Peru (Loreto), Brazil (Amazonas). ◆Fig. 331. Miconia cautis Wurdack, Mem. New York Bot. Gard. 10(5): 169. 1964. Shrub 1.5–2.5 m tall, the branchlets, inflorescence, and hypanthia densely stellatescurfy; leaves often somewhat dimorphic in

each pair, straight or slightly curved, blades oblong-elliptic, apex short or subabruptly acuminate, base acute to obtuse, 9–22 × 3.5– 7 cm, firmly membranous and entire, upper surface glabrous, lower surface glabrous and sparsely to moderately stellate-scurfy on the primary veins, barely (0.2–0.5 cm) 3pliveined with the veinlets on the lower side densely elevated-reticulate; petioles 0.7–2 cm long; panicle 7–11 cm long, relatively fewflowered, rachis reddish; flowers 4-merous, sessile or very shortly pedicellate; petals white and inconspicuously granulose, ca. 0.7 × 0.4 mm, broadly elliptic; ovary 2-locular, apex moderately stellate-scurfy. Riparian forests, evergreen lowland to lower montane forests, 100–600 m; Bolívar (Río Ichún, 53 km southwest of Urimán), Amazonas (slopes of Cerro Aracamuni, slopes of Cerro Huachamacari, slopes of Cerro Duida and Cerro Marahuaka, Piedra Tururumeri on Río Yatua, Río Mawarinuma at base of Sierra de la Neblina, San Carlos de Río Negro to Solano). Adjacent Colombia and Brazil. ◆Fig. 336. Miconia centrodesma Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 164. 1850. Clidemia martii Naudin, Ann. Sci. Nat. Bot. sér. 3, 17: 375. 1851. Miconia buchtienii Cogn., Repert. Spec. Nov. Regni Veg. 8: 2. 1910. Miconia centrandra Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 361. 1915. Miconia subtriloba Gleason, Bull. Torrey Bot. Club 58: 429. 1931. Miconia sylvestris Gleason, Brittonia 2: 321. 1937. Shrub or small tree 1–4 m tall, the terete branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia very sparsely and deciduously stellulate-scurfy; leaf blades elliptic to ovate-elliptic, apex gradually acuminate, base acute to obtuse, 10–25 × 5–11 cm, thin and entire or obscurely serrulate-undulate, deciduously ciliolate, upper surface usually glabrous and nitid, lower surface glabrous on the surface, shortly 5-pliveined with very laxly reticulate areoles; petioles 1.5–5 cm long; panicle 4–10 cm long and submultiflorous, the primary branches 2–4 per node; flowers 4-merous and mostly on pedicels 1–2 mm long; petals white, ca. 1.2 × 0.8 mm, obovate-oblong, glabrous; ovary usually 3-locular, minutely

408

M ELASTOMATACEAE

stellulate-puberulous at the apex. Montane or occasionally lowland forests, (100–)600– 1400 m; Bolívar (Majagua, Río Uei, Río Venamo, Sierra de Lema, Sierra Pakaraima), Amazonas (slopes of Cerro Duida, upper slopes of Cerro Huachamacari, Río Cataniapo, slopes of Sierra de la Neblina, Sierra Parima). Aragua, Mérida, Sucre, Táchira, Yaracuy; Guatemala to Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 329.

apex sparsely strigulose in part with glandtipped hairs; fruiting hypanthium sparsely to moderately setulose with gland-tipped hairs. Moist areas, lowland to lower montane forests, near sea level to 700 m; Delta Amacuro (Río Grande, Sacupana, San Victor), Bolívar (Altiplanicie de Nuria, El Paují, Icabarú, Kilómetro 88, Río Botanamo, Río Erebato, Río Paragua, Río Tonoro), Amazonas (mouth of Río Matacuni). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 322.

Miconia ceramicarpa (DC.) Cogn. in Mart., Fl. Bras. 14(4): 338. 1887. —Clidemia? ceramicarpa DC., Prodr. 3: 160. 1828. Clidemia coccinea DC., Prodr. 3: 162. 1828. Clidemia violacea DC., Prodr. 3: 162. 1828. —Miconia ceramicarpa var. violacea (DC.) Cogn. in Mart., Fl. Bras. 14(4): 339. 1887. Miconia erythropila Steud., Flora 27: 723. 1844. Miconia iodopila Steud., Flora 27: 723. 1844. Miconia spondylantha var. leschenaultiana Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 188. 1850. —Miconia leschenaultiana (Naudin) Sagot, Ann. Sci. Nat. Bot. sér. 6. 15: 325. 1883. Subshrub or suffrutescent (sometimes ± creeping) herb 0.3–1 m tall, the branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia sparsely to moderately covered with fine, smooth hairs to 1(– 1.5) mm long; leaf blades elliptic-ovate to elliptic, apex acute to shortly acuminate, base broadly acute to obtuse, 8–20 × 4–10 cm, often purple or reddish-tinged on lower surface, thin and inconspicuously ciliolate-serrulate, 5(7)-pliveined (the inner pair of primary veins diverging 0.5–1 cm and the second pair 1.5–4 cm above the base), sparsely strigulose on upper surface, lower surface only strigulose on the veins but mostly glabrous on the surface; petioles mostly 1–3 cm long; inflorescence 2–9 cm long, oblong and rather few-flowered, the rachis reddish; flowers 5-merous, sessile, fewaggregated at short branchlet ends; petals white, 2.5–3.5 × 1.5–2 mm, oblong to obovate-oblong, glabrous; ovary 3-locular, the

Miconia chrysophylla (Rich.) Urb., Symb. Antill. 4: 459. 1910. —Melastoma chrysophylla Rich., Actes Soc. Hist. Nat. Paris 109. 1792. Melastoma fulva Rich. in Humb. & Bonpl., Monogr. Melast. 1: 23, t. 11. 1807. —Miconia fulva (Rich.) DC., Prodr. 3: 180. 1828. Miconia longifolia var. aubletiana Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 157. 1850. —Miconia fulva var. aubletiana (Naudin) Cogn. in Mart., Fl. Bras. 14(4): 389. 1887. Miconia fulva var. angustifolia Cogn. in Mart., Fl. Bras. 14(4): 389. 1887. Shrub or tree 4–15 m tall, the young branchlets, lower leaf surfaces, inflorescence, and hypanthia completely covered with yellow-brown lepidote pubescence; young branchlets 2- or 4-angled; leaf blades mostly 3- or 4-whorled; blade narrowly oblong-elliptic, apex acuminate, base acute, 10–20 × 2–4(–6) cm, firm-chartaceous and obscurely undulate-serrulate to entire, upper surface sparsely and deciduously lepidote-puberulous, basally 3-veined or obscurely (2–5 mm) 3-pliveined; petioles 0.5–1 cm long; panicle 6–15 cm long, basal branches nearly as broad, many-flowered; flowers 5-merous, sessile; petals white, 2 × 1.2–1.3 mm, oblong-obovate, glabrous; ovary 3-locular, glabrous. Evergreen lowland to montane and gallery forests, semideciduous forests, near sea level to 1300 m; Delta Amacuro (Sacupana), Bolívar (Los Patos, base of Ptari-tepui, Río Botanamo, Río Caruay, Río Parguaza, Río Supamo, Río Toro, Santa Elena de Uairén), Amazonas (Caño Negro on Cerro Duida, Ocamo, Río Coro Coro west of Cerro Yutajé). Apure, Miranda, Monagas, Táchira, Zulia; southern Mexico,

Miconia 409

Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 338. Miconia ciliata (Rich.) DC., Prodr. 3: 179. 1828. —Melastoma ciliata Rich., Actes Soc. Hist. Nat. Paris 1: 109. 1792. —Miconia racemosa var. ciliata (Rich.) Griseb., Fl. Brit. W. I. 258. 1860. Miconia virgulata Gleason, Bull. Torrey Bot. Club 52: 386. 1925. Shrub (rarely small tree) 0.5–2.5(–4) m tall, fine-setulose at the branchlet nodes and petiole apices and appressed-ciliate at the leaf margins (sometimes sparsely appressedsetulose on the lower leaf surfaces), otherwise usually glabrous; leaf blades elliptic to oblong, apex acute, base obtuse to rounded, 4–20 × (1–)1.5–7 cm, subrigid and obscurely serrulate, basally 3-veined (excluding the tenuous marginal veins), often reddish on lower surface; petioles mostly 0.4–2.5 cm long; inflorescence 4–10 cm long, submultiflorous, rachis reddish; flowers 5-merous, sessile, crowded-secund on the branchlets; petals pinkish; ovary 3-locular, the apex inconspicuously glandular. Savannas, forest edges, gallery forests, burned-over areas on tepui summit (Auyán-tepui), 100–2200 m; Bolívar (Acopán-tepui, Altiplanicie de Nuria, Auyán-tepui, Cerro Guaiquinima, Cerro Ichún, Cerro Venado, Ciudad Bolívar, El Paují, Kavanayén, Kurún-tepui, Río Caroní near Arekuna, Río Karaurín, Río Paragua, Sororopán-tepui, Uarama), Amazonas (Cerro Aratitiyope, Cerro Autana, Cerro Yapacana, La Esmeralda, Río Cunucunuma, Río Siapa, Sierra Parima). Aragua, Anzoátegui, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 341. Miconia crassinervia Cogn. in Mart., Fl. Bras. 14(4): 391, pl. 79. 1887. Shrub or tree 1.5–12 m tall, the branchlets and primary and secondary veins on the lower leaf surface, inflorescence, and hypanthia completely covered by ashy to reddish, stellate-dendritic hairs to 0.1 mm long; branchlets initially rounded-quadrangular,

terete with age; leaf blades oblong-ovate, apex gradually acuminate, base obtuse, subcoriaceous, and inconspicuously undulatedenticulate to entire, 10–20(–30) × 4–10(–13) cm, upper surface glabrous, lower surface covered by a subamorphous, appressed indument, basally 5-veined, with the venation densely reticulate on the lower side but inconspicuous due to the pubescence; petioles 1.5–4 cm long panicle 9–17 cm long, many-flowered; flowers 5-merous, sessile, secund on the short upper side shoots; petals white and minutely granulose, 1.6–1.8 × 0.7– 1 mm, obovate-oblong; ovary (2)3-locular, apex minutely granulose. Evergreen lowland to montane forests, riverbanks, near sea level to 1200 m; Bolívar (base of Cerro Guaiquinima, base of Ptari-tepui, Río Caura, Río Hacha, Río Ikabarú, Río Karaurín, Ucaima near Canaima), Amazonas (16 km east of San Fernando de Atabapo). Brazil (Acre, Amapá, Amazonas). Miconia curta (Gleason) Wurdack, Mem. New York Bot. Gard. 10(5): 170. 1964. —Graffenrieda curta Gleason, Fieldiana, Bot. 28: 431. 1952. Shrub or tree 2–6 m tall, the branchlets, primary veins on the lower leaf surface, inflorescences, and hypanthia sparsely covered by a stellate, appressed pubescence; leaf blades oblong-elliptic to lance-elliptic, apex acute or shortly acuminate, base acute to rounded, subrigid and entire but soon deciduously appressed-ciliate, 10–18 × 4–7 cm, at first very sparsely stellate-puberulent on both surfaces but becoming glabrescent, shortly (to 1 cm) 5-pliveined or basally 5veined, with the veinlets barely elevated-reticulate on lower surface; petioles 2–4 cm long; panicles 8–12 cm long; flowers 4- or 5merous and mostly shortly (2–3 mm) pedicellate; petals white and glabrous, ca. 3 × 1.2– 1.5 mm, obovate-oblong; ovary 3- or 4locular, the apex sparsely stellate-scurfy. Endemic. Key to the Subspecies of M. curta 1. Apex of the ovary without setae ............... ........................................ subsp. curta 1. Apex of the ovary with 10 setulae 0.8–1.7 mm long .................. subsp. ptariensis

410

M ELASTOMATACEAE

M. curta subsp. curta Tepui-slope and streamside forests, 1100– 1800 m; Bolívar (Acopán-tepui, Auyán-tepui, Cerro Guaiquinima, Cerro Jaua), Amazonas (headwaters of Caño Iguapo, Cerro Aracamuni, slopes of Cerro Duida, Cerro Huachamacari, slopes of Cerro Marahuaka). Endemic. ◆Fig. 335. M. curta subsp. ptariensis Wurdack, Mem. New York Bot. Gard. 10(5): 170. 1964. Forested tepui slopes; Bolívar (slopes of Ptari-tepui). Endemic. Miconia decurrens Cogn., Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 253. 1909. Shrub or small tree 3–6 m tall, the branchlets, lower leaf surfaces, inflorescences, and hypanthia covered by an amorphous appressed indument (or delicately irregularly stellate); leaf blades ovate-elliptic, apex caudate-acuminate (1.5–2 cm), base acute, 11–19 × 4.5–8.5 cm, membranous and entire to obscurely denticulate, upper surface glabrous, strongly 5(7)-pliveined; petioles ca. 1 cm long, winged to the base due to the decurrent leaf base; panicle 6–18 cm long, many-flowered; flowers 5-merous, sessile; petals white, ca. 4 × 1.6–1.8 mm, narrowly obovate-oblong with an obtuse apex; ovary 3-locular, moderately stellate-puberulous. Evergreen lowland to lower montane forests, 200–400 m; Bolívar (Río Nichare), Amazonas (upper Río Matacuni). Barinas; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 340. Miconia dioica Wurdack, Mem. New York Bot. Gard. 10(5): 173. 1964. Erect or scandent shrub 2.5–5 m tall; branchlets rounded-quadrangular, especially near the leaf nodes; leaf blades narrowly ovate to oblong-ovate, apex acuminate, base obtuse to rounded, 4–12 × 2–5 cm, subcoriaceous and entire, shortly 3-pliveined; petioles 1–2.5 cm long, magenta; panicle 4– 10 cm long, pyramidal, the branches reddish; flowers 5-merous, unisexual, pedicels ca. 1 mm long; petals white, 0.7–0.8 × 0.7–0.9 mm, rotund; staminate flowers with abortive ovary, pistillate flowers with 3-locular ovary and abortive stamens. Tepui-summit thickets, along streamsides, 1800–2200 m; Bolí-

var (Auyán-tepui, Macizo del Chimantá [Apacará-tepui, Toronó-tepui]). Endemic. ◆Fig. 345. Miconia dispar Benth., Hooker’s J. Bot. Kew Gard. Misc. 2: 241. 1850. —Boyuyo palo de viejo. Tree 4–13 m tall, the obtusely quadrangular branchlets, petioles, and inflorescences densely puberulous with stellate-dendritic hairs 0.2–0.5(–1) mm long; leaf blades elliptic-oblong, apex rather abruptly shortacuminate, base broadly acute to obtuse, 20– 35 × 8–18 cm, firmly chartaceous and obscurely undulate-serrulate, upper surface glabrous, lower surface (densely to) moderately puberulous with short-stalked stellulate-dendritic hairs 0.6–0.8 mm diameter, basally 5-veined with the veinlets on lower surface somewhat elevated-reticulate; petioles 2.5–5 cm long; panicle 20–30 cm long, narrowly oblong, many-flowered; flowers 5merous, sessile, secund on the short side branchlets; petals white, 1.5–2 mm long, obovate-oblong, obscurely granulose; ovary 3-locular, the apex sparsely stellulatepuberulous. Evergreen lowland to lower montane forests, 100–1000 m; Bolívar (base of Auyán-tepui, Río Erebato, Río Hacha, Río Suapure), Amazonas (Río Casiquiare, Río Cataniapo, Río Cuao, Río Puruname, San Carlos de Río Negro to Solano, San Fernando de Atabapo, Serranía de Los Pijiguaos). French Guiana, Amazonian Peru, Brazil, and Bolivia. ◆Fig. 347. Miconia dodecandra (Desr.) Cogn. in Mart., Fl. Bras. 14(4): 243. 1887. —Melastoma dodecandra Desr. in Lam., Encycl. 4: 46. 1797. Shrub or small tree (3–)5–15 m tall, the branchlets, lower leaf surfaces, bracts, and hypanthia densely stellulate-puberulous; leaf blades elliptic-ovate, apex gradually short-acuminate, base obtuse to truncate, mostly 10–17 × 5–8 cm, thin-coriaceous and entire, upper surface glabrous, lower surface discolorous, basally 5-veined; petioles mostly 2–3.5 cm long; panicle 10–15 cm long and submultiflorous, the primary branches mostly 4(–6) per node; flowers (5)6-merous, subtended by a pair of deciduous, broadly elliptic, pubescent bracts ca. 6 mm long, the pedicels 2–5 mm long; petals white to

Miconia 411

salmon-green, mostly 8–8.5 × 3–3.8 mm, narrowly obovate-oblong, pruinose-granulose, reflexed; ovary (3)4(5)-locular, moderately stellulate-puberulous apically. Lower to upper montane forests, 200–2100 m; Bolívar (Auyán-tepui, Betania on Santa Elena de Uairén to Icabarú, Gran Sabana, La Escalera, Peray-tepui west of Santa Elena, base of Ptari-tepui, upper Río Caroní, Río Tabaro in Río Nichare basin, Roraima-tepui, base of Uaipán-tepui), Amazonas (Cerro Aracamuni, Cerro Huachamacari, Sierra de la Neblina, Sierra Parima). Venezuelan Coastal Range and Andes; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 337.

deciduously asperous with hairs 0.15–0.3 mm long; leaf blades elliptic to oblong-elliptic, apex obtusely acute to obtuse, base acute, 3–4 cm × 1–1.7 cm, firmly membranous and sparsely serrulate, glabrous on both surfaces, inconspicuously (to 0.4 cm) 3-pliveined, with the veinlets on lower surface laxly reticulate; petioles 0.7–1 cm long; panicle 3–5 cm long, submultiflorous; flowers 5-merous, shortly (ca. 1 mm) pedicellate; petals white, glabrous, ca. 2.2 × 2.1 mm, suborbicular; ovary 3-locular, the conical apex glabrous. Tepui-summit thickets and rocky areas, along streams, 2500–2700 m; Amazonas (Cerro Marahuaka). Mérida, Táchira; Colombian Andes. ◆Fig. 348.

Miconia egensis Cogn. in Mart., Fl. Bras. 14(4): 374. 1887. Miconia panicularis Gleason, Bull. Torrey Bot. Club 52: 384. 1925. Shrub or small tree 3–8 m tall, the rounded-quadrate young branches, petioles, primary leaf veins on the lower surface, and inflorescence axis densely puberulous with dendritic-stellulate hairs 0.1–0.2 mm long and sparsely to moderately setulose with minutely roughened hairs to 0.5 mm long; leaf blades elliptic to obovate-elliptic, apex subabruptly to abruptly short-acuminate, base narrowly to broadly acute, mostly 16– 25 × 7–10 cm, fragile and entire to obscurely crenulate-serrulate, upper surface glabrous, lower surface sparsely and deciduously puberulous with minute dendritic-stellulate hairs, 5-veined (including the tenuous marginal veins); petioles 1–2 cm long; panicle 8– 15 cm long, many-flowered, the primary branches often 4 per node; flowers 5-merous and shortly (ca 0.5 mm) pedicellate; petals white, 1.8–2.5 × 1–1.3 mm, obovate-oblong, moderately granulose; ovary 3-locular, glabrous. Rocky montane streamsides, 300–500 m; Bolívar (Río Paragua), Amazonas (Río Coro Coro west of Cerro Yutajé). Guyana, Suriname, French Guiana, Brazil.

Miconia elata (Sw.) DC., Prodr. 3: 182. 1828. —Melastoma elata Sw., Prodr. 70. 1788. Tree 10–15 m tall, the young growth and lower surface of leaves completely covered by minute stellulate-lepidote, appressed hairs; branchlets 4-angled, becoming terete with age; leaf blades ovate-elliptic to elliptic, apex shortly acuminate, base rounded to obtuse or acute, 15–25 × 8–17 cm, thin-coriaceous, entire to obscurely undulate, basally 5-veined, glabrous on upper surface, lower surface whitish tan-pubescent; petioles 4–7 cm long; panicles 12–20 cm long and nearly as broad, many-flowered; flowers 5, 6(7)-merous, sessile, interrupted-glomerulate on the branches; petals white and obscurely granulose, 2.8–3 × 1.6–1.9 mm, oblong-obovate; ovary 3-locular, apex stellulate-puberulent. Riparian forests, lower montane forests, 100–800 m; Bolívar (Río Tabaro in Río Nichare basin), Amazonas (Sierra Parima). Barinas, Lara, Mérida, Portuguesa, Zulia; southern Mexico, Central America, West Indies, Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 349.

Miconia elaeoides Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 223. 1850. Miconia pallida Gleason, Bull. Torrey Bot. Club 57: 72. 1930. Shrub 0.5–2 m tall, the inconspicuously 4angled branchlets, primary veins on lower leaf surface and inflorescence sparsely and

Miconia eugenioides Triana, Trans. Linn. Soc. London 28: 109. 1871. Oxymeris cuspidata Triana, Trans. Linn. Soc. London 28: 93. 1871. —Leandra cuspidata (Triana) Cogn. in Mart., Fl. Bras 14(4): 175. 1886. Shrub or small tree 1–8 m tall, the branchlets, primary veins on lower surface of leaves, and inflorescences sparsely to moderately and deciduously stellate-puberulent;

412

M ELASTOMATACEAE

leaf blades lance-oblong, apex long-acuminate, base broadly acute to rounded, firmly membranous and entire, 6–13 × 2.5–5 cm, upper surface glabrous and nitidulous, lower surface sparsely punctate, basally 3-veined with the inconspicuous veinlets plane and laxly reticulate; petioles 0.8–1.8 cm long; panicle 4–12 cm long, submultiflorous, the primary branches usually 4 per node; flowers 5-merous, pedicels 1.5–2 mm long; petals white and minutely granulose, 2.8–3.2 × 1– 1.4 mm, lance-oblong and acute at the apex; ovary 3-locular, glabrous. Seasonally flooded riparian forests, disturbed areas, secondary forests, low forests over bauxite deposits, 100–800 m; Bolívar (headwaters of Río Túriba), Amazonas (Caño Cotua, Caño Cupaven, Caño San Miguel, Cerro Vinilla, Maroa, Río Baría, Río Casiquiare, Río Cuao, Río Mawarinuma, Río Pasimoni, San Carlos de Río Negro, San Fernando de Atabapo, Yavita). Adjacent Colombia and Brazil. ◆Fig. 339. Miconia fallax DC., Prodr. 3: 181. 1828. Shrub 0.5–3 m tall, similar to M. stenostachya, except for the terete to rounded-quadrangular branchlets, the leaves 7–15 × 4–8 cm, almost sessile (petioles to 0.5 cm long) and with a cordulate base; flowers 5-merous, sessile. Lowland to upland savannas, 100– 900 m; Delta Amacuro (10 km east-southeast of Los Castillos), Bolívar (Cerro Bolívar, Guri area, Río Carrao, Río Parguaza, San Félix, Santa Elena), Amazonas (Puerto Ayacucho). Sucre; Guyana, Suriname, Brazil, Paraguay. ◆Fig. 343. Miconia fallax is doubtfully distinct from M. stenostachya. Miconia fragilis Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 171. 1850. Tree 10–20 m tall, the quadrangular young branchlets, inflorescences, and hypanthia moderately but rather deciduously puberulous with minute stellulate hairs 0.05 (–0.1) mm diameter; branchlets subnodose, with distinct narrow interpetiolar ridges; leaf blades oblong-lanceolate, apex acuminate to caudate, base acute to obtuse, 6–10 × 2–3(–4) cm, thin and entire, upper surface glabrous, lower surface finely granuloseglaucous as well as sparsely and deciduously stellulate-puberulous, obscurely (1–3 mm) 3-

pliveined; petioles 0.8–1.5 cm long; panicle mostly 5–7 cm long and rather few-flowered; flowers 4-merous and shortly pedicellate; petals white, 1.9–2 × 1 mm, obovate-oblong, minutely granulose; ovary 3-locular, the apex glabrous and slightly 8-ridged. Evergreen lowland forests and lower montane forests over bauxitic soils, 100–600 m; Delta Amacuro (Río Toro), Bolívar (Serranía de Los Pijiguaos). Guyana, French Guiana. Miconia fragrans Cogn. in Urb., Symb. Antill. 7: 526. 1913. Shrub or small tree 2–7 m tall, essentially glabrous, the compressed young branchlets becoming terete with age; leaf blades elliptic to ovate-elliptic, apex acute to abruptly short-acuminate, base broadly acute to obtuse, 11–30 × 6–14 cm, chartaceous and entire, 5-pliveined with the inner pair of primary veins diverging subalternately 1–3.5 cm above the base; petioles 1–3 cm long; panicle 8–13 cm long, few-branched (usually with 2 long lateral branches ca. 1 cm above the base), submultiflorous; flowers 5-merous, sessile, interrupted-verticillate; petals white, ca. 2.5 × 1.3 mm, obovate-oblong, obscurely pruinose-granulose; ovary 3-locular, the apex sparsely granulose. Montane forests, 1000– 1300 m; Bolívar (Sierra de Lema), Amazonas (Cerro Coro Coro, Cerro Marahuaka, Sierra de la Neblina). Sucre; Trinidad, Tobago, Guyana (Kaieteur), Brazil (Serra da Neblina). ◆Fig. 350. Miconia gratissima Benth. ex Triana, Trans. Linn. Soc. London 28: 101. 1871. —Kuepiyek (Arekuna). Small tree (2–)4–10 m tall, the branchlets, inflorescences, and hypanthia moderately to densely covered with delicate dendritic or dendritic-stellulate hairs; leaf blades elliptic to lance-elliptic, apex rather shortly (to ca. 2 cm) acuminate or acute, base acute, 10–30 × 4–10 cm, upper surface glabrous, lower surface completely covered by appressed cobwebby pubescence, basally 5veined (including the tenuous marginal veins) or pseudopliveined; petioles 0.5–2(–3) cm long; panicle 8–10(–15) cm long, submultiflorous, with paired branchlets; flowers very fragrant, 5-merous, essentially sessile to shortly (to 3 mm) pedicellate (pedicels ar-

Miconia 413

ticulate at hypanthium base), umbellate on terminal branchlets; hypanthium 7–8 mm long; calyx lobes 3.5–4.5 mm long; petals white to pinkish white, 5–5.5 × 4 mm, obovate, glabrous; ovary (4)5-locular, apically moderately glandular-puberulous. Evergreen lowland to montane forests, 100–1800 m; Bolívar (Kilómetro 88, Mazivaca 17 km south of Los Castillos, Río Urayaparutá, north of El Paují), Amazonas (Caño Colorado in Cerro Sipapo massif, Gavilán, Río Sipapo, San Carlos de Río Negro to Solano). Táchira; Central America, Guyana, Suriname, French Guiana, Brazil, Bolivia. Miconia grossidentata Wurdack, Phytologia 50: 289. 1982. Shrub 1–5 m tall; branchlets, primary veins on lower leaf surface, inflorescences, and hypanthia with fairly dense stellate hairs 0.1–0.2 mm long and 0.3–0.4 mm diameter; leaf blades elliptic, apex gradually acuminate, base acute, 7.5–15 × 2.5–5.5 cm, chartaceous and conspicuously bluntly toothed (teeth ca. 5 mm apart and 1.5–2 mm deep in the sinuses), upper surface glabrous, lower surface with scattered stellate hairs, basally 3-veined; petioles 0.6–2 cm long; inflorescence 3–6 cm long, few-flowered, subspicate; flowers 5-merous, sessile; petals 3.1–3.3 × 1.5–1.6 mm, oblong-obovate; ovary 3-locular, apex sparsely glandulose-puberulent. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (45 km east of Los Pijiguaos in headwaters of Río Túriba), Amazonas (Puerto Ayacucho to Gavilán, Río Cataniapo, Río Sipapo). Endemic. ◆Fig. 351. Miconia guaiquinimae Wurdack, Mem. New York Bot. Gard. 10(4): 41. 1961. Miconia guaiquinimae subsp. angustifolia Wurdack, Mem. New York Bot. Gard. 10(4): 42. 1961. Shrub or tree 3–15 m tall, the young growth completely covered by appressed, stellate hairs; young stems obtusely quadrangular, becoming terete; leaf blades elliptic to oblong-elliptic, apex gradually or subabruptly short-acuminate, base broadly acute or narrowly rounded-obtuse, 6–15 × 2.5–5 cm, coriaceous and entire, becoming glabrous on upper surface, densely stellatepubescent on lower surface, basally 3-veined

(excluding the faint marginal veins); petioles 1–2 cm long; panicles 7–12 cm long, manyflowered, the branches 4-verticillate; flowers 5-merous, sessile; petals white and minutely granulose, ca. 2.2 × 1 mm, obovate-oblong; ovary 3-locular, the apex granulose. Tepui scrub, montane and streamside forests, 1000–1700 m; Bolívar (Cerro Guaiquinima, Ilú-tepui, La Escalera). Endemic. ◆Fig. 342. Miconia holosericea (L.) DC., Prodr. 3: 181. 1828. —Melastoma holosericea L., Sp. P1. 390. 1753. —Boyuyo palo de vieja. Small tree 2–10 m tall, the branchlets, veins on the lower leaf surface, and inflorescence densely covered by appressed stellate hairs; leaf blades ovate-elliptic to oblong-elliptic, apex abruptly short-acuminate, base broadly acute to rounded, 12–25 × 6–12 cm, subcoriaceous and entire to very obscurely undulate, basally 5-veined or shortly (to 1 cm) 5-pliveined; petioles 2–4 cm long; panicle 3–8 cm long, few-flowered, terminal or axillary by overtopping growth; flowers 6-merous, subsessile and glomerate at branchlet ends, subtended by caducous bracts 3–5 mm long; hypanthium 5–6 mm long; calyx lobes ca. 3 mm long and oblong, deciduous at anthesis; petals white to pale pink, 7–8 × 3–4 mm, obovate-oblong, externally very sparsely and deciduously stellulate-puberulous; ovary usually 4-locular, sparsely glandular-puberulous on the stylar collar; fruits large, to ca. 1 cm diameter. Gallery forests, evergreen lowland to lower montane forests, savannas, 100– 1200 m; Bolívar (base of Auyán-tepui, 65 km southeast of Los Pijiguaos), Amazonas (El Pozo southeast of San Fernando de Atabapo, Río Cuao, Río Pamoni, San Carlos de Río Negro to Solano, Yavita). Monagas, Táchira, Zulia; Mexico, Central America, Trinidad, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil, Bolivia. ◆Fig. 352. Miconia huberi Wurdack, Mem. New York Bot. Gard. 50: 103. 1989. Shrub ca. 2 m tall; branchlets, primary veins on lower leaf surface, and inflorescence moderately and deciduously puberulent with dendritic hairs 0.1–0.2 mm long with stipitate, dendritic hairs 0.3–0.4 mm long intermixed; young branches obtusely sulcate-qua-

414

M ELASTOMATACEAE

drangular, becoming terete with age; leaf blades oblong-elliptic or ovate-oblong, apex narrowly acute or subacuminate, base broadly acute, 9–11(–15) × 3–6 cm, subcoriaceous and entire, surfaces at first sparsely dendritic-puberulent but becoming glabrescent, primary veins on lower surface obscurely setulose (hairs ca. 0.8 mm long), basally 5-veined, lower-order veins 3–5 mm apart; petioles 1.3–2.7 cm long; panicle 8–10 cm long, with 2–4 primary branches in each node, sparsely glandular-puberulent; flowers 4-merous, pedicels 2–5 mm long; hypanthium ca. 3.5 mm long; petals 7–7.3 × 4.3–4.5 mm, oblong-obovate, slightly emarginate; ovary 2-locular, glabrous. In rock crevices on tepui summit, 2600–2700 m; Bolívar (Ilútepui). Endemic. Miconia hypoleuca (Benth.) Triana, Trans. Linn. Soc. London 28: 119. 1871. —Chaenopleura hypoleuca Benth., J. Bot. (Hooker) 2: 315. 1840. Chaenophora ferruginea Crueg., Linnaea 20: 112. 1847. —Miconia cruegeriana Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 245. 1850. —Chaenopleura ferruginea (Crueg.) Griseb., Fl. Brit. W. I. 1: 259. 1860. Tree mostly 5–12 m tall, the young branchlets, lower surface of leaf blades, inflorescence, and hypanthia completely covered with an appressed cobwebby indument; young branchlets compressed, later hexagonal; leaf blades elliptic to ovate-oblong, apex subabruptly short-acuminate, base broadly acute to obtuse, 15–30 × 5–12 cm, chartaceous and undulate-serrulate, upper surface glabrous, basally 5-veined; petioles 2–5 cm long; panicle 9–18 cm long, many-flowered, ± pyramidal; flowers 5-merous, sessile; petals white, 2–2.3 × 1.2 mm, oblong-obovate, minutely granulose; ovary 3-locular, glabrous. Evergreen lowland and lower montane forests, 100–800 m; Bolívar (Bochinche, Río Cuyuní, Sierra de Maigualida, Sierra Imataca). Sucre; Trinidad, Guyana, Suriname, French Guiana, Brazil. Miconia ibaguensis (Bonpl.) Triana, Trans. Linn. Soc. London 28: 110. 1871. —Melastoma ibaguense Bonpl. in Humb. & Bonpl., Monogr. Melast. 1: 105, t. 45. 1815. —Tschudya ibaguensis (Bonpl.) Griseb., Fl. Brit. W. I. 250. 1860.

Clidemia maculata Benth., J. Bot. (Hooker) 2: 310. 1840. Clidemia miconioides Benth., J. Bot. (Hooker) 2: 310. 1840. Clidemia virgata Pittier, Bol. Soc. Venez. Ci. Nat. 11: 24. 1947. Shrub 1–2.5 m tall, the young growth moderately to densely setulose with smooth brownish hairs ca. 1(–3) mm long underlain by stellulate pubescence; leaf blades lanceolate to oblong-lanceolate, apex narrowly acute to short-acuminate, base acute to rounded, 7–13 × 2.5–5 cm, chartaceous and ciliolate-serrulate, upper surface sparsely fine-setulose to glabrous, lower surface sparsely to moderately fine-setulose, shortly (0.2–1 cm) 5-pliveined with lax veinlet areoles; petioles mostly 0.5–1 cm long; panicle 4–10 cm long, many-flowered; flowers 5merous, sessile, crowded-secund on the ultimate short branchlets; petals white, 2.5–3 × 1.8–2.3 mm, obovate-oblong, glabrous, apically obscurely crenulate; ovary 3-locular, sparsely setulose apically. Savannas, edges of Mauritia palm swamps and lowland to lower montane forests, 100–1300 m; Bolívar (Altiplanicie de Nuria, northwest of El Manteco, Hato La Vergareña, Río Kanarakuni, San Ignacio de Yuruaní, Santa Elena de Uairén), Amazonas (Ocamo). Anzoátegui, Aragua, Carabobo, Distrito Federal, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia; southern Mexico through Central America, West Indies, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. Miconia iluensis Wurdack, Mem. New York Bot. Gard. 10(5): 165. 1964. Shrub or small tree 3–15 m tall, the young growth sparsely and deciduously stellatescurfy; branchlets angled; leaf blades elliptic, apex gradually narrow-acuminate, base broadly acute to obtuse, 10–14 × 3.5–6 cm, entire and rigid-membranous, sparsely and persistently stellate-scurfy on the primary veins on lower surface, basally 5-veined; petioles 2–3.5 cm long; panicles 10–12 cm long, paniculate-umbellate; flowers 6merous, pedicels 6–12 mm long; petals white-pink, reflexed, glabrous except for the marginal glands, 8.5–9 × 4.5–5 mm, obovateoblong; ovary 4-locular, glabrous. Tepui-slope and summit forests, 1800–2300 m; Bolívar (Cerro Jaua, Ilú-tepui). Endemic. ◆Fig. 346.

Miconia 415

Miconia impetiolaris (Sw.) D. Don ex DC., Prodr. 3: 183. 1828. —Melastoma impetiolaris Sw., Prodr. 70. 1788. Southern Mexico, Central America, West Indies, Colombia, Venezuela; 3 varieties, 1 in Venezuela. M. impetiolaris var. spruceana Cogn. in Mart., Fl. Bras. 14(4): 272. 1877. Small tree 3–8 m tall, the young growth moderately to densely covered by short claviform hairs 0.1–0.3 mm long, these apically asperous; leaves sessile, blades elliptic-oblong to obovate-oblong, apex shortly acuminate, base cordate-amplexicaul (ca. 1 cm long), 20– 35 × 7–12 cm, subcoriaceous and apically undulate-denticulate, upper surface glabrous, lower surface becoming glabrescent with age, basally 3- or 5-veined; panicle 20–30 cm long, many-flowered; flowers 5-merous, sessile, interrupted-glomerulate along the rachis. Riparian forests, 100–300 m; Bolívar (Río Apacará, Río Caura, Río Hacha), Amazonas (El Pozo southeast of San Fernando de Atabapo, Río Casiquiare, Río Cataniapo, Río Cunucucuma, Río Guainía, Río Mawarinuma, Río Puruname, San Carlos de Río Negro, San Fernando de Atabapo, Tobogán de la Selva south of Coromoto). Colombia (Vaupés), Brazil (Amazonas). ◆Fig. 353. Miconia inaequalifolia Triana, Trans. Linn. Soc. London 28: 109. 1871. Shrub, the young growth moderately scurfy with very short hairs; leaves in each pair differently sized (to 1:2.5); blades lanceelliptic, apex acuminate, base auriculate, 5– 14 × 1.8–3.5 cm, membranous and crenulatedenticulate, upper side glabrous, lower surface sparsely scurfy but becoming glabrescent, basally 3- or 5-veined; petioles 2–5 mm long; panicle 7–8 cm long; flowers 5merous; pedicels ca. 1 mm long; petals white, minutely granulose, 2.1–2.5 × 1–1.2 mm; ovary 3-locular. Riparian forests, 100–200 m; Amazonas (Caño Catirico below mouth of Río Paciba, Río Casiquiare, Río Orinoco 5 km east of San Fernando de Atabapo). Amazonian Colombia and Brazil. Miconia kavanayensis Wurdack, Phytologia 18: 152. 1969. Tree to 12 m tall, the tetragonal-winged branchlets, primary veins on the lower sur-

face of leaves, and inflorescence completely covered by stellate-dendritic hairs; leaf blades oblong-elliptic, apex shortly acuminate, base obtuse, 25–42 × 10–15 cm, subcoriaceous and undulate-denticulate, upper surface glabrous, lower surface completely covered by stellate hairs ca. 0.4 mm diameter, basally 5-veined, the veinlets on the lower side elevated-reticulate; petioles 4–7 cm long; panicle 20–32 cm long, many-flowered, with branches 4–8-verticillate; flowers 5-merous, sessile, interrupted-glomerate on the branches; petals white and inconspicuously granulose, 1.6–1.7 × 1.1–1.3 mm, obovate-oblong and rounded to emarginate at the apex; ovary 3-locular, the conical apex densely stellate-puberulent. Montane forests, 1100–1400 m; Bolívar (Oparuma near Kavanayén). Endemic. Miconia laevigata (L.) DC., Prodr. 3: 188. 1828. —Melastoma laevigata L., Sp. P1. ed. 2, 1: 559. 1762. Shrub 0.5–3 m tall, the branchlets, veins on the lower leaf surface, inflorescence, and hypanthia sparsely to moderately and deciduously stellate-scurfy; leaf blades broadly lanceolate to oblong-ovate, apex gradually acuminate, base obtuse to rounded, 9–18 × 3–8 cm, firmly membranous and entire or inconspicuously serrulate, deciduously ciliolate, upper surface glabrous, lower surface glabrous on the surface, basally 5-veined with the veinlets plane and reticulate on lower surface; petioles 1–3 cm long; panicle 3–10 cm long, many-flowered; flowers 5merous, mostly sessile, and secund on the terminal branches; petals white and minutely granulose, ca. 3 × 1.5–2 mm, oblongobovate; ovary 3-locular, apex minutely granulose. Semideciduous lower montane forests, 300–500 m; Bolívar (Altiplanicie de Nuria). Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Nueva Esparta, Sucre, Táchira; Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil. Miconia lasseri Gleason, Bol. Soc. Venez. Ci. Nat. 15: 103. 1954. Shrub 2–4 m tall, essentially glabrous; leaf blades elliptic to oblong-elliptic, apex short-acuminate, base acute, 10–25 × 6–11 cm, fragile and entire, 3-pliveined 1–2 cm above the base; petioles 1–3 cm long; panicle

416

M ELASTOMATACEAE

(6–)10–14 cm long, narrowly oblong and fewflowered; flowers 6-merous, sessile; petals white, 3–5 × 2–5 mm, spathulate, glabrous; ovary 3-locular, glabrous. Lower montane forests, along streams, 400–1000 m; Bolívar (Río Caura, Río Kukenán, Río Uairén in upper Río Caura basin), Amazonas (Río Ventuari). Guyana, Suriname, Brazil. ◆Fig. 355.

Evergreen lowland and lower montane forests, riparian forests, near sea level to 500 m; Delta Amacuro (Río Amacuro, Río Toro), Bolívar (Altiplanicie de Nuria, upper Río Caura, Río Nichare, Río Oris, Río Paramichi, Río Toro, Salto Pará, Santa Maria de Erebato). Zulia; other distribution as in species. ◆Fig. 354.

Miconia lateriflora Cogn., Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 255. 1909. Small shrub 1–3 m tall, the young growth very sparsely and deciduously stellulatescurfy; branchlets terete; leaf blades elliptic, apex subabruptly short-acuminate, base acute to cordulate, 8–16(–25) × 3.5–10 cm, fragile and entire or inconspicuously undulate-serrulate, deciduously ciliolate, glabrous on both surfaces (rarely very sparsely strigulose above), basally 3(5)-veined; petioles 0.8–2(–3.5) cm; panicle small and fewflowered, rachis red; flowers 4-merous and mostly in 3-flowered dichasia; calyx lobes 0.2 mm long and oblate, the acute external teeth projecting 0.3–1.2 mm; petals white, 1.6–1.8 × 1.4–1.5 mm, oblong-obovate, minutely granulose; ovary 3- or 4-locular, glabrous; fruiting hypanthium sometimes sparsely glandular-setulose. Mexico, Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil; 2 subspecies, both in the flora area.

M. lateriflora subsp. monticellensis Wurdack, Phytologia 16: 171. 1968. Montane forests, 800–900 m; Bolívar (Santa Elena de Uairén). Guyana, Suriname.

1. Leaf blades acute to narrowly obtuse at base; lateral flowers of dichasia sessile ................................ subsp. lateriflora 1. Leaf blades slightly cordulate at base; lateral flowers of dichasia on pedicels 2–3 mm long ........... subsp. monticellensis

Miconia lepidota DC., Prodr. 3: 180. 1828. Miconia schomburgkii Benth., J. Bot. (Hooker) 2: 312. 1840. Miconia bifrons Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 178. 1850. Tree mostly 6–12 m tall, the alternately blunt-ancipital young branchlets, lower leaf surface, inflorescences, and hypanthia completely covered with lepidote hairs; leaf blades ovate-elliptic to oblong-elliptic, apex shortly and subabruptly acuminate, base obtuse to broadly acute, mostly 10–20 × 5–10 cm, firmly chartaceous and entire, upper surface glabrous, basally 3-veined; petioles mostly 2–3 cm long; panicle 10–25 cm long, many-flowered; flowers 5-merous, sessile, secund on the ultimate branchlets; petals white, 2–2.2 × 1–1.1 mm, obovate-oblong, minutely granulose; ovary 3-locular, glabrous. Evergreen lowland to montane forests, 50–1200 m; Bolívar (Gran Sabana, near Kavanayén, Río Parguaza, Río Suapure, Sierra de los Pijiguaos), Amazonas (near Culebra, Río Cataniapo, Río Cuao, Río Mawarinuma, Río Pasimoni, Río Sipapo, San Carlos de Río Negro, Yavita). Barinas, Lara; Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 344.

M. lateriflora subsp. lateriflora Miconia tessmannii Markg., Notizbl. Bot. Gart. Berlin-Dahlem 9: 266. 1925. Octopleura ciliata Triana, Trans. Linn. Soc. London 28: 146. 1871. —Ossaea ciliata (Triana) Cogn. in A. DC. & C. DC., Monogr. Phan. 7: 1067. 1891. Ossaea disparilis Standl., Contr. Arnold Arbor. 5: 120. 1933. —Miconia disparilis (Standl.) R.O. Williams, Fl. Trinidad 1: 388. 1934.

Miconia longifolia (Aubl.) DC., Prodr. 3: 184. 1828. —Melastoma longifolia Aubl., Hist. Pl. Guiane 432, pl. 170. 1775. —Chaenopleura longifolia (Aubl.) Griseb., Fl. Brit. W. I. 260. 1860. Miconia lambertiana DC., Prodr. 3: 185. 1828. Shrub (or small tree) 1–3(–10) m tall, the 4–6-angled young branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia sparsely and deciduously stellu-

Key to the Subspecies of M. lateriflora

Miconia 417

late-scurfy; leaves mostly 3- or 4-whorled, blades oblong-elliptic, apex acuminate, base acute, 8–20 × 2.5–7 cm, chartaceous and entire or obscurely undulate, glabrous on both surfaces, shortly 3-pliveined with lax venule reticulation; petioles 0.2–0.7(–1) cm long; panicle 7–15 cm long, multi-flowered, ± pyramidal, the inflorescence branches mostly 4 per node, reddish; flowers 5-merous on pedicels 0.5–2 mm long; petals white, 1.5 × 0.7–0.9 mm, obovate-oblong, densely granulose; ovary 3- or 4-locular, the apex minutely granulose. Riparian forests, lower montane forests, near sea level to 1000 m; Delta Amacuro (Ibaruma, Río Acure, Río Amacuro, Sacupana), Bolívar (Río Kukenán, Río Toro, Sierra Imataca), Amazonas (Ocamo, Río Siapa, Simarawochi). Táchira, Zulia; southern Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Miconia longispicata Triana, Trans. Linn. Soc. London 28: 117. 1871. Tree 4–9 m tall, the branchlets, lower surface of leaves, inflorescences, and hypanthia completely covered by minute dendriticstellulate hairs; leaf blades ovate-oblong to elliptic-oblong, apex narrowly and gradually acuminate, base subcordate, mostly 15–30 × 7–18 cm, firmly chartaceous and entire or obscurely undulate-subdenticulate, upper surface glabrous, basally 5-veined; petioles mostly 2–4 cm long; inflorescence 15–40 cm long and spicate, usually erect at anthesis; flowers 5-merous, sessile; petals white, 2–2.5 × 1.3 mm, glabrous; ovary 3-locular, the apex very sparsely stellulate-scurfy. Evergreen lowland to lower montane forests, 100–600 m; Amazonas (Río Asisa, Río Mawarinuma, Río Siapa, 25 km southwest of San Antonio del Orinoco, San Carlos de Río Negro to Solano). Colombia (Guainía), French Guiana, Brazil, Bolivia. ◆Fig. 357. Miconia lourteigiana Wurdack, Phytologia 20: 380. 1970. Shrub 3–8 m tall, the branchlets, lower surface of leaves, inflorescences, and hypanthia completely and persistently covered by an appressed, arachnoid tomentum; leaf blades oblong-elliptic, apex gradually acuminate, base acute, 10–20 × 5–8 cm, subcoriaceous and entire, upper surface glabrous and

nitid, basally 3-veined (excluding the faint marginal veins), with the flat veinlets densely reticulate on the lower surfacec but completely covered by the pubescence; petioles 1–2 cm long; panicle 6–12 cm long, many-flowered, the branches mostly long and bifurcate; flowers 5-merous, sessile, secund; petals white, minutely granulose, ca. 2 × 1.4–1.7 mm, obovate-oblong; ovary 3-locular, the apical collar glabrous. Lower montane and riparian forests, 200–500 m; Bolívar (Río Paramichi). Apure; Amazonian Colombia, Ecuador, Peru, Brazil, Bolivia. Miconia macrothyrsa Benth., J. Bot. (Hooker) 2: 312. 1840. Miconia pseudorubiginosa Cogn. in Urb., Symb. Antill. 7: 311. 1912. Shrub or small tree 0.5–2(–4) m tall, the branchlets, lower surface of leaves, inflorescences, and hypanthia completely covered with minute dendritic hairs underlain by a granulose-cobwebby indument; leaf blades broadly ovate-elliptic to elliptic, apex acute to rounded, base cordulate, 6–14 × 3.5–10 cm, rigid and densely but minutely ciliolateserrulate, upper surface centrally glabrous and marginally sparsely fine-setulose, basally 5- or 7-veined; petioles mostly 1–1.5 cm long; panicle mostly 10–20 cm long and narrowly oblong, many-flowered; flowers 5merous, sessile, crowded-secund on the short lateral branches; petals white, 3 × 1.7–2 mm, obovate-oblong, glabrous; ovary usually 5locular, the apex inconspicuously granulose. Forest edges, savannas, edge of Mauritia palm swamps, 100–1100 m; Bolívar (Altiplanicie de Nuria, El Palmar, Guayapo on Río Caura, Hato La Vergareña, Icabarú, San Ignacio de Yuruani, Santa Elena de Uairén, Sierra de Imataca, Tumeremo), Amazonas (Ocamo, Puerto Ayacucho to Samariapo, Sierra Parima). Anzoátegui, Miranda, Sucre, Trujillo; Colombia, Trinidad, Guyana, Suriname, French Guiana, eastern Peru, Brazil. ◆Fig. 359. Miconia macrotis Cogn. in DC., Monogr. Phan. 7: 749. 1891. Small tree ca. 5 m tall, the branchlets, primary veins on lower surface of leaves, and inflorescences densely tomentose with fine stipitate-stellate hairs (stipes 1–1.5 mm long); leaf blades oblanceolate-oblong, apex

418

M ELASTOMATACEAE

gradually long-acuminate, cordate-auriculate for 3–5 cm at the base, 35–75 × 12–25 cm, subrigid and prominently irregularly denticulate, upper surface glabrous and plane with discolored margins when dry, lower surface moderately stellate-puberulent, 7, 9(11)-veined with the veinlets on the lower surface densely elevated–reticulate; petioles 1–3 cm long, robust; panicle 10–25 cm long, oblong and compact, many-flowered; flowers mainly 6-merous, essentially sessile (the thick pedicels ca. 0.5 mm long); petals white, densely stellate-puberulent externally, ca. 7 × 3.5 mm, obovate-oblong; ovary 4-locular, densely stellate-puberulent apically. Montane forests, 900–1000 m; Bolívar (Cerro Ceitá in Hato Divina Pastora). Trinidad, Ecuador. ◆Fig. 360. Miconia marginata Triana, Trans. Linn. Soc. London 28: 110. 1871. Shrub 1–4 m tall, glabrous except for the petals and ovary; leaf blades elliptic, apex rather abruptly short-acuminate, base attenuate, 9–30 × 3–9 cm, thinly coriaceous and entire, dark-margined when dry, shortly 3-pliveined with lower order venation essentially not visible; petioles 0.5–1 cm long; inflorescence 6–10 cm long, racemiform or bifurcate, rather few-flowered; flowers 5merous, sessile, interrupted-verticillate; petals white, 0.8–1 × 0.5 mm, oblong-obovate, densely granulose; ovary 3-locular, the apex minutely glandular-puberulous. Gallery forests, 100–600 m; Bolívar (La Escalera, Río Apacará, Río Aparamán, Río Cuyuní, Río Nichare, Salto Pará), Amazonas (Río Autana, Río Mawarinuma, Río Yatua). Amazonian Colombia, Guyana, northern Brazil. ◆Fig. 358. Miconia mariae Wurdack, Mem. New York Bot. Gard. 10(5): 168. 1964. Shrub ca. 3 m tall, the branchlets at first compressed but soon terete, the petioles, primary leaf veins on lower surface, and inflorescences sparsely and deciduously stellulate-puberulous; leaf blades elliptic, apex shortly blunt-acuminate, base obtuse to rounded, 12–22 × 5–13 cm, firmly chartaceous and entire to inconspicuously crenulate, distantly appressed-ciliolate, glabrous on both surfaces, basally 5-veined with laxly reticulate veinlets; petioles 1–4 cm long; panicle 6–12 cm long, rather few-flowered;

flowers 5-merous and sessile or obscurely (to 0.5 mm) pedicellate; calyx apiculate-conical in bud, opening irregularly at anthesis; petals white, ca. 3 × 1.2–1.5 mm, oblong-obovate, glabrous; ovary 3-locular, inconspicuously glandular-setulose on the collar apex. Montane and lower montane forests, 500– 1300 m; Bolívar (Amurí-tepui, El Paují, headwaters of Río Caroní). Guyana. Miconia maroana Wurdack, Phytologia 16: 176. 1968. Shrub 2–4 m tall, the slightly compressed branchlets, lower surface of leaf blades, inflorescence, and hypanthia completely covered by dendritic, orangish hairs 0.2–0.5 mm long; leaf blades oblong-elliptic to elliptic, apex obtusely acute to shortly acuminate, base broadly acute, subcoriaceous and entire, 14–20 × 5–9 cm, upper surface glabrous, basally 3(5)-veined with the veinlets densely reticulate on lower surface but hidden by the pubescence; petioles 2–3 cm long; panicle 8– 15 cm long, subspicate or flowers glomerulate on short basal side branches 1–5 cm long; flowers 5-merous, sessile; petals white and minutely granulose, 2.4–2.5 × ca. 1.5 mm, obovate and apex obtuse; ovary 3-locular, the apex densely strigulose. Riparian forests, Río Negro caatinga, 100–200 m; Amazonas (Colón, Maroa to Yavita, Río Guianía, Río Negro, San Carlos to Solano). Brazil (Amazonas). ◆Fig. 356. Miconia matthaei Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 176. 1850. Small tree 5–8 m tall, the young growth densely incurved-setose with essentially smooth reddish hairs 1–2(–3) mm long underlain by a sparse stellulate indument; leaf blades elliptic, apex acuminate, base acute to obtuse, 10–20 × 4–7 cm, thinly coriaceous and entire to obscurely undulate-serrulate, upper surface sparsely appressed-setose at the bases of the primary veins but otherwise glabrous, lower surface sparsely to moderately setulose with fine hairs ca. 1 mm long, basally 3-veined or shortly 3-pliveined with fine venule areolation below; petioles 1–2.5 cm long; panicle 7–11 cm long, many-flowered; flowers 5-merous, sessile, multiglomerulate at short branchlet ends; petals white, 2.7–3 × 1.4–1.5 mm, obovate-oblong, granulose; ovary 3-locular. Understory of evergreen lowland and lower montane forests,

Miconia 419

100–900 m; Bolívar (Asukupa, Río Asa, Río Mawela in Río Erebato basin, Santa Elena de Uairén, near Urimán), Amazonas (Río Casiquiare, near Simarawochi, 10 km east of Tamatama). Apure, Táchira, Zulia; southern Mexico, Central America to Costa Rica, Colombia, Trinidad, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Miconia minutiflora (Bonpl.) DC., Prodr. 3: 189. 1828. —Melastoma minutiflora Bonpl. in Humb. & Bonpl., Monogr. Melast. 1: 50. 1809. Glossocentrum collinum Crueg., Linnaea 20: 111. 1847. —Miconia glossocentra Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 243. 1850. Cremanium trinitatis Crueg., Linnaea 20: 111. 1847. —Miconia trinitatis (Crueg.) Naudin, Ann. Sci. Nat. Bot. sér. 3.16: 246. 1850. Shrub or tree 2–8(–15) m tall, the branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia (basally) very sparsely dendritic-puberulous but soon glabrescent, otherwise glabrous; leaf blades lance-oblong, apex long-acuminate, base obtuse, 5–12 × 1.5–4 cm, rigid-chartaceous and entire, basally 3- or 5-veined with lax venule reticulation on lower surface; petioles 0.5–1 cm long; panicle 5–15 cm long, many-flowered; flowers 5-merous on pedicels 1-1.5 mm long; petals white, ca. 2 × 1 mm, ovate-elliptic, glabrous; ovary 3-locular, glabrous. Lower montane forests, 400–900 m; Bolívar (Cerro Bolívar, Cerro Marutaní, Río Suapure, 7 km east of Santa Elena de Uairén), Amazonas (Sierra Parima). Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Central America, Colombia, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 361. Miconia mirabilis (Aubl.) L.O. Williams, Fieldiana, Bot. 29: 574. 1963. —Fothergilla mirabilis Aubl., Hist. Pl. Guiane 441, t. 175. 1775. —Pomarrosa. Tree mostly 5–20 m tall, the branchlets, lower surface of leaves, inflorescences, and bracts moderately to completely covered with minute stellulate or lepidote-stellulate hairs; leaf blades elliptic to ovate-elliptic, apex gradually acuminate, base acute to rounded, mostly 10–15 × 4–7 cm, thin-rigid

and entire, upper surface glabrous, lower surface in the primary vein axils usually sparsely barbellate with hairs to 1 mm long, basally 5(7)-veined (rarely obscurely pliveined); petioles 2.5–5 cm long; panicle mostly 9–12 cm long, submultiflorous; flowers 5(6)-merous on pedicels 2–5 mm long, subtended by a pair of deciduous broadly elliptic to obovate-elliptic bracts 5–8(–15) mm long, umbellately arranged on the terminal branchlets; hypanthium 3–4.2 mm long, glabrous; petals white, 7–9 × 2.5–3.5 mm, narrowly obovate-oblong, pruinose-granulose; ovary 3(4, 5)-locular, apically moderately stellulate-puberulous. Evergreen lowland to montane forests, 50–1300 m; Delta Amacuro (Caño Jota-Sabuca near Caño Mariusa, Río Toro), Bolívar (La Escalera, Luepa, Río Chicanán northeast of Upata), Amazonas (Río Mawarinuma at base of Sierra de la Neblina). Anzoátegui, Carabobo, Distrito Federal, Falcón, Mérida, Miranda, Sucre, Táchira, Yaracuy; Mexico to Costa Rica, West Indies, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 363. Miconia mituana Wurdack, Phytologia 13: 75. 1966. Small tree 5–10 m tall, the young growth densely stellulate-scurfy; young branches quadrangular but becoming terete with age; leaf blades ovate-elliptic to suborbicular, apex subabruptly acuminate, base rounded or inconspicuously cordulate, 10–16 × 6.5–11 cm, subcoriaceous and indistinctly sinuatedenticulate, basally 7-veined, upper surface at first sparsely and shortly setose then becoming glabrous, lower surface densely stellulate; petioles 3–6.5 cm long; panicle 9– 13 cm long, main branches 4–6-verticillate; flowers 6-merous, pedicels 8–10 mm long at anthesis; petals pale pink, ca. 8.5 × 4 mm, obovate-oblong, ± retuse at apex; ovary 4locular, apex silvery-strigulose. Seasonally flooded riparian forests, 100–200 m; Amazonas (Río Marawinuma). Colombia (Amazonas, Vaupés). Miconia myriantha Benth., J. Bot. (Hooker) 2: 314. 1840. Shrub or small tree 2.5–12 m tall, vegetatively like M. minutiflora but the leaf blades on the lower surface usually noticeably glaucous; inflorescences and hypanthia as in M. minutiflora; calyx lobes 0.4–0.5 mm long,

420

M ELASTOMATACEAE

deltoid; petals white, 2–2.3 × 1–1.2 mm, oblong-triangular, granulose. Secondary forests, forest edges, riversides, 100–1000 m; Delta Amacuro (Koboime, Río Grande), Bolívar (Cerro Pitón, El Paují, Río Apacará, Río Ikabarú, Río Suapure, Río Tírica, Urimán), Amazonas (base of Cerro Sipapo, Isla Ratón, Río Cataniapo, Río Cuao, Río Mavaca, Río Mawarinuma, Río Negro, Río Padamo, San Carlos de Río Negro to Solano, Yavita). Amazonian Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 364. Miconia neblinensis Wurdack, Mem. New York Bot. Gard. 18(2): 288. 1969. Compact shrubs ca. 0.4–1 m tall; branchlets quadrangular, becoming terete with age; leaf blades elliptic to ovate-elliptic, apex acute or narrowly obtuse, base obtuse or rounded, 2–3 × 1.3–2 cm, rigid and obscurely ciliolate-serrulate, upper surface glabrous, lower surface with scattered minute glands, basally 3-veined, the second-order veins ca. 1.5 mm apart and impressed on upper surface; petioles 0.4–0.7 cm long; panicle 2–3 cm long, pyramidal; flowers 5-merous, pedicels ca. 1 mm long; petals white, 1.7–1.9 × 1.7–1.9 mm, suborbicular and emarginate; ovary 3(4)-locular, the conical apex glabrous. Forming low masses on open slopes up to cliff bases at tepui summit, 2600–2700 m; Amazonas (expected). Brazil (Serra da Neblina). Miconia nervosa (Sm.) Triana, Trans. Linn. Soc. London 28: 111. 1871. —Melastoma nervosa Sm. in Rees, Cycl. 23: Melastoma no. 31. 1819 [1812]. Miconia septuplinervis Pittier, Bol. Soc. Venez. Ci. Nat. 11: 25. 1947. Shrub or small tree 1–5 m tall, the young growth densely strigulose to appressed-setose with fine smooth hairs; leaf blades elliptic, apex gradually acuminate, base narrowly acute, 15–40 × 5–14 cm, thin and obscurely undulate-serrulate, upper surface sparsely fine-strigulose, lower surface sparsely to moderately appressed-setulose with fine hairs (and sometimes purple-flushed), notably 5- or 7-pliveined (the innermost primary veins diverging 3–10 cm and the second pair 1–4 cm above the blade base) with the veinlets on lower surface rather densely reticu-

late; petioles 0.5–3(–4) cm long; inflorescence 4–12(–17) cm long, narrowly oblong and rather few-flowered; flowers 5-merous, sessile, bracteate-glomerulate at ends of very short lateral branches; petals white, 3.5–4.5 × 1.5–1.8 mm, oblong, glabrous; ovary 3-locular, the apex sparsely glandular-strigulose. Evergreen lowland to lower montane forest understories, 50–900 m; Delta Amacuro (Río Amacuro), Bolívar (22 km south of El Dorado, El Palmar, El Paují, Maripa, Río Caura, Río Nichare, Río Paragua, Río Paramichí, Santa María de Erebato, base of Sierra de Maigualida), Amazonas (Río Cunucunuma, base of Sierra de la Neblina, Sierra Parima). Mérida, Monagas, Táchira, Zulia; southern Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 367. Miconia perobscura Wurdack, Mem. New York Bot. Gard. 10(4): 42, fig. 32F–M. 1961. Shrub 2–3 m tall, the young growth densely stellate-puberulent; branchlets grooved-quadrangular; leaf blades broadly elliptic, 15–23 × 10–17 cm, apex abruptly short-acuminate, base obtuse to truncate, membranous and inconspicuously undulateserrulate, both surfaces sparsely and somewhat deciduously stellate-puberulent, 7pliveined; petioles 4–13 cm long; panicle to 20 cm long and wide; flowers 5-merous, shortly (1–3 mm) pedicellate; petals white to pale pink, glabrous, 5–5.5 × 2.3–2.6 mm, oblong-obovate; ovary 3-locular, apex sparsely stellate-puberulent. Tepui-slope forests, 1700–1900 m; Amazonas (slopes of Sierra de la Neblina). Endemic, but likely in adjacent Brazil. ◆Fig. 390. Miconia perturbata Wurdack, Mem. New York Bot. Gard. 10(5): 172. 1964. Tree 4–18 m tall, the branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia sparsely stellate-scurfy but becoming glabrescent; young stems obtusely quadrangular, becoming terete with age; leaf blades elliptic, apex shortly obtuse-acuminate, base broadly acute to obtuse, 8–14 × 5– 7 cm, thinly coriaceous and entire, glabrous on both surfaces, basally 5-veined (including the faint marginal veins), with the plane

Miconia 421

veinlets laxly reticulate on the lower side; petioles 1–2.5 cm long; panicle 8–12 cm long, many-flowered, the branches usually 4(–6)verticillate; flowers 5-merous, pedicels 1–2 mm long; petals white, densely granulose, 1.7–1.8 × 0.9–1 mm, oblong; ovary 3-locular, the apex minutely papillose. Montane forests, 1100–1800 m; Bolívar (Cerro Sarisariñama), Amazonas (Cañon Grande of Sierra de la Neblina, Cerro Sipapo). Endemic. Miconia phaeophylla Triana, Trans. Linn. Soc. London 28: 113. 1871. Miconia fendleriana Cogn. in A. DC. & C. DC., Monogr. Phan. 7: 822. 1891. Shrub or tree 3–10(–15) m tall, the young growth moderately to densely covered with minute sessile-stellulate hairs, glabrescent with age; young branchlets compressed, without interpetiolar ridges; leaf blades oblong-elliptic to ovate-elliptic, apex shortly and rather abruptly blunt-acuminate, base broadly acute to rounded, 9–16 × 3–7 cm, subcoriaceous and entire, upper surface glabrous, lower surface usually somewhat glaucescent, basally 5-veined or shortly pseudopliveined with lax venule reticulation; petioles 1–2.5 cm long; panicle 6–16 cm long, many-flowered, often with 4 branches per node; flowers 5-merous, pedicels 0.7–1.3 mm long; petals white, 3–3.3 × 1.7–1.9 mm, obovate, granulose; ovary 3-locular, glabrous. Savannas, shrublands, forest edges, 100–1700 m; Bolívar (Altiplanicie de Nuria, Cerro Pitón, El Paují, Parupa, Río Asa, middle Río Caroní at sabana Arekuna, Río Caruay, Río Karaurín, Río Paragua, Santa Elena de Uairén), Amazonas (Cerro Duida, Cerro Parú, La Esmeralda, Sierra Maigualida, Sierra Parima, Tobogán de la Selva near Coromoto). Aragua, Carabobo, Falcón, Lara, Miranda, Táchira, Trujillo; Panama, Amazonian Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 362. Miconia pileata DC., Prodr. 3: 180. 1828. Shrub 1–3 m tall, the young branchlets, leaf nodes, and petioles with long hispid hairs 2–6 mm long; leaf blades elliptic-oblong, apex acute or subacuminate, base subrounded, 6–10 × 2.5–4 cm, margins subdenticulate and long-ciliate, basally 5veined, primary veins impressed on upper surface, upper surface very sparsely fine-

setulose, lower surface fine-setose on primary veins but glabrous on surface and lower-order veins; petioles 0.5–1.5 cm long; panicle erect, pubescent, 5–10 cm long, narrowly paniculate with short branches bearing secund flowers, peduncle and inflorescence branches reddish purple; flowers 5merous, sessile; petals pink, obovate, ca. 2 × 1.5 mm, apex subtruncate, glabrous; ovary 3locular, glabrous. Savannas with Mauritia palm swamps, forest edges, 100–300 m; Bolívar (30–40 km east of Ciudad Piar, Guri dam area, Sierra Imataca), Amazonas (base of Cerro Duida, Maroa, Santa Rosa de Amanadona). Amazonian Ecuador, Peru, and Brazil. Miconia pilgeriana Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 363. 1915. —Sacauyek (Arekuna). Shrub or tree 6–15 m tall, the terete branchlets, primary leaf veins on lower surface, and inflorescences densely covered by tan stellate hairs; leaf blades oblong-elliptic, apex caudate-acuminate (1–2.5 cm), base broadly acute, 12–20 × 3–5(–8) cm, rigidmembranous and entire, upper surface initially sparsely stellate-puberulent but soon becoming glabrescent, lower surface sparsely stellate-puberulent (the hairs sessile or very shortly (to 0.1 mm) stipitate and 0.2–0.3 mm diameter) but subglabrous with age, basally 5-veined with the inconspicuous veinlets plane and reticulate on the lower surface; petioles 1–2 cm long; panicle 7–10 cm long, many-flowered, the thin primary branches usually 4-verticillate; flowers 5-merous, shortly (0.5–1 mm) pedicellate; petals white and minutely granulose, 1.1–1.2 × 0.9–1 mm, subrounded; ovary 3-locular, glabrous. Riparian and montane forests, 400–1100 m; Bolívar (Cerro Guaiquinima, Kavanayén, Río Caura, Río Ikabarú, Río Uaiparú, Sierra de los Pijiguaos), Amazonas (Caño Yureba in lower Río Ventuari basin, Río Cuao). Anzoátegui; Amazonian Colombia, Peru, and Brazil. Miconia plukenetii Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 116. 1850. Tree mostly 3–10 m tall, the branchlets, lower surface of leaves, and inflorescences completely covered by tawny sessile stellatelepidote hairs; leaves sessile, the blades obovate-spatulate, apex rather abruptly short-

422

M ELASTOMATACEAE

acuminate, base cordulate-clasping, mostly 30–60 × 10–25 cm, chartaceous and entire, upper surface glabrous, 5-pliveined with the inner pair of primary veins diverging subalternately 1/4–1/3 the blade length above the base; panicle 15–35 cm long, many-flowered; flowers 5-merous, nearly sessile; petals white or pinkish, 5.5–7.9 × 3–4 mm, obovateoblong, deciduously granulose; ovary 3-locular, apex glandular-puberulous. Evergreen lowland to montane forests, 100–1100 m; Bolívar (base of Aprada-tepui, south of El Dorado, La Escalera, Urimán), Delta Amacuro (Río Toro). Trinidad, Guyana, French Guiana, Suriname. Miconia poeppigii Triana, Trans. Linn. Soc. London 28: 107. 1871. Miconia congesta Cogn., Repert. Spec. Nov. Regni Veg. 8: 2. 1910. Miconia surinamensis Gleason, Recueil Trav. Bot. Néerl. 32: 212. 1935. Tree mostly 15–33 m tall, the ± quadrate young branchlets, primary leaf veins on lower surface, inflorescences, and hypanthia sparsely to moderately stellulate-puberulous; leaf blades mostly elliptic, apex shortly blunt-acuminate, base acute, 10–15 × 3–5 cm, firmly chartaceous and entire, glabrous on both surfaces, shortly 3(5)-pliveined with veinlets laxly reticulate on lower surface; petioles 0.7–1.5 cm long; panicle 7–12 cm long, many-flowered, the branches, pedicels, and hypanthia glandular-setulose with slender hairs; flowers 5-merous on pedicels 0.5–2 mm long; petals white, 2–3 × 1–1.6 mm, obovate-oblong, glabrous; ovary 3-locular, glabrous. Riparian forests, 100–300 m; Bolívar (Río Chiguao, Río Parguaza), Amazonas (Río Coro Coro west of Cerro Yutajé, Río Majagua, Río Manapiare, Río Puruname). Apure, Barinas, Guárico, Mérida, Táchira, Zulia; Central America, Colombia, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. Miconia polita Gleason, Bull. Torrey Bot. Club 52: 379. 1925. Shrub 1–4 m tall, with terete branchlets, glabrous except for the leaf margins, inflorescence branches, and hypanthia; leaf blades elliptic to oblong-elliptic, apex shortacuminate, base broadly acute to obtuse, 10– 24 × 4–9 cm, subrigid and entire but remotely ciliolate, upper surface nitid and with

margins discolored when dry, basally 3- or 5veined with laxly reticulate veinlets; petioles 1–3.5 cm long; panicle 5–10 cm long and submultiflorous; flowers 5-merous on pedicels 1.5–3 mm long; petals white, 3.2–3.4 × 1.5–2 mm, obovate-oblong, glabrous; ovary 3-locular, glabrous. Montane forests and streamsides, 400–1100 m; Bolívar (Río Acanán, Río Apacará, Río Caruay). Guyana. ◆Fig. 368. Miconia prasina (Sw.) DC., Prodr. 3: 188. 1828. —Melastoma prasina Sw., Prodr. 69. 1788. Miconia pteropoda Benth., J. Bot. (Hooker) 2: 314. 1840. Miconia revoluta Benth., J. Bot. (Hooker) 2: 313. 1840. Miconia cristulata Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 177. 1850. Miconia mucronulata Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 358. 1915. Shrub or small tree 1.5–12 m tall, the young branchlets, primary veins on lower leaf surface, inflorescences, and hypanthia very sparsely to moderately but deciduously stellulate-puberulous; leaf blades elliptic to oblong or obovate-elliptic, apex acute to short-acuminate, base acute, 6–25 × 2.5–10 cm, chartaceous to subcoriaceous and entire to repand-denticulate, glabrous on both surfaces, 3- or 5-pliveined with very lax venule reticulation (1–3 mm); petioles 0.3–2 cm long; panicle 5–15 cm long, many-flowered; flowers 5-merous, sessile, secund on terminal branches; petals white, 2–3 × 1–1.5 mm, obovate-oblong, densely granulose; ovary 3locular, the apex sparsely stellulate-puberulous. Riparian forests, savannas, edges of Mauritia palm swamps, 100–1400 m; Delta Amacuro (Río Toro), Bolívar (Altiplanicie de Nuria, Corozal, El Tigre, La Prisión, Río Ariza, Río Bonita, Río Paragua, Río Parguaza, Río Tírica, Río Venamo, Salta Pará, Sierra Imataca, Tumeremo, Uaiparú), Amazonas (La Esmeralda, Río Coro Coro west of Cerro Yutajé, Río Mawarinuma, San Carlos de Río Negro, Sierra Parima). Aragua, Barinas, Cojedes, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Zulia, Yaracuy; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 372.

Miconia 423

Miconia pseudoaplostachya Cogn. in Mart., Fl. Bras. 14(4): 263, pl. 53, fig. 1. 1887. Shrub 2.5–4 m tall, the branchlets, lower surface of leaves, and inflorescences densely stellate-scurfy; leaf blades oblong-lanceolate, apex gradually long-acuminate, base obtuse to rounded, mostly 12–18 × 3–5 cm, subrigid and entire, upper surface glabrous, basally 3-veined; petioles 1–2(–4) cm long; inflorescence and flowers as in Miconia aplostachya. Riparian forests, 100–300 m; Bolívar (Río Caura, Río Paragua, Río Toronó, Salto Pará), Amazonas (base of Cerro Duida, Río Casiquiare, Río Cunucunuma, Río Padamo, Río Pasimoni). French Guiana. ◆Fig. 371. Miconia pseudocapsularis Wurdack, Mem. New York Bot. Gard. 10(4): 32. 1961. Shrub or tree 2–10 m tall, glabrous except for the inflorescence; leaf blades oblong-elliptic to ovate-elliptic, apex subabruptly obtuseacuminate, base cuneate, 12–23 × 5.5–12 cm, thin and entire or somewhat crenulate, shortly 5-pliveined with the veinlets very laxly reticulate; petioles 1–7 cm long; panicle sparsely and inconspicuously scurfy, submultiflowered; flowers 5-merous and strongly scorpioid-secund on the branchlets, the pedicels ca. 0.5 mm long; petals white, glabrous, 1.8–2 × 1.1–1.2 mm, obovate; ovary 3locular, glabrous; fruit 10-ribbed when dry. Riparian forests, tepui-slope forests, 100– 1700 m; Bolívar (Amaruay-tepui, Cerro Jaua, Sierra de Maigualida, Toronó-tepui), Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Huachamacari, Sierra de la Neblina). Brazil (Amazonas, Roraima). ◆Fig. 373. Miconia pubipetala Miq., Stirp. Surinam. Select. 50. 1850 [1851]. Diplochita parviflora Benth., J. Bot. (Hooker) 2: 302. 1840. —Miconia parviflora (Benth.) Cogn. in Mart., Fl. Bras. 14(4): 219. 1887. Small tree (2–)4–10 m tall, the young branchlets, lower surface of leaves, inflorescences, and hypanthia densely covered with appressed stellulate or lepidote-stellulate hairs; leaf blades oblong-elliptic to ovate-elliptic, apex subabruptly acuminate 1–3 cm, base broadly acute to rounded, (9–)15–25 × (5–)8–13 cm, rather firm and entire to ob-

scurely sinuate-denticulate, upper surface glabrous, basally 3(5)-veined; petioles 2–4 cm long; panicle 9–12 cm long, many-flowered; flowers 5-merous and in umbels, the pedicels 2–3 mm long; petals white or pinkflushed, 5–7.5 × 2–2.5 mm, obovate-oblong, externally densely scurfy; ovary 3-locular, the apical collar stellulate-strigulose. Seasonally flooded and riparian forests, near sea level to 400 m; Delta Amacuro (Koboina, Sacupana, San Victor), Bolívar (Río Ichún, Río Paragua, Río Venamo, Urimán), Amazonas (Río Cuao, Río Siapa). Trinidad, Guyana, Suriname, French Guiana, Brazil. Miconia punctata (Desr.) D. Don, ex DC., Prodr. 3: 184. 1828. —Melastoma punctata Desr. in Lam., Encycl. 4: 50. 1797. —Eurychaenia punctata (Desr.) Griseb., Fl. Brit. W. I. 259. 1860. Tree 5–18 m tall, the sharply compressedquadrangular branchlets, lower surface of leaves, inflorescences, and hypanthia completely covered with appressed and marginally fringed red-brown scales; leaf blades oblong-elliptic, apex very shortly acuminate, base broadly acute, subrigid and entire, 10– 30 × 5–13 cm, upper surface glabrous, basally 3-veined (excluding the tenuous marginal veins); petioles 1–3 cm long; panicle mostly 12–20 cm long, many-flowered; flowers 5-merous, sessile, secund on the ultimate branchlets; petals white, 2.3–2.8 × 1.1–1.5 mm, oblong-obovate, glabrous; ovary 3-locular, stellulate-puberulous apically. Evergreen lowland to montane forests, riparian forests, 100–1300 m; Bolívar (Acanán-tepui, Apacará-tepui, Río Yuruani), Amazonas (Cerro Yapacana, Río Casiquiare, Río Cuao, Río Cunucunuma, Río Pasimoni, San Carlos de Río Negro, Yavita). Apure, Aragua, Barinas, Carabobo, Distrito Federal, Mérida, Monagas, Sucre, Táchira, Yaracuy, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 369. Miconia pyrifolia Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 164. 1850. —Palo de agua. Tree (6–)10–25 m tall, the young branchlets, primary leaf veins on lower surface, and inflorescences sparsely and deciduously stellulate-scurfy; young branchlets obtusely

424

M ELASTOMATACEAE

quadrangular but terete with age, with distinct interpetiolar ridges; leaf blades oblongovate, apex shortly blunt-acuminate, base broadly acute to obtuse, 10–20 × 4–8 cm, thin-chartaceous and entire, glabrous on both surfaces, shortly (3)5-pliveined with lax venule areoles on lower surface; petioles 1–2 cm long; panicle mostly 8–10 cm long, manyflowered, with primary branches mostly 4 per node; flowers 5-merous and pedicellate 0.6–1.2 mm; petals white, 2.4–2.6 × 1.1–1.4 mm, obovate-oblong, glabrous; ovary 3-locular, truncate and glabrous. Evergreen lowland to lower montane forests, 100–1000 m; Delta Amacuro (Río Toro, San Victor), Bolívar (slopes of Auyán-tepui and Aparamán-tepui). Hispaniola, Guyana, Suriname, Peru, Brazil. Miconia racemosa (Aubl.) DC., Prodr. 3: 869. 1828. —Melastoma racemosa Aubl., Hist. Pl. Guiane 406, pl. 156. 1775. Shrub or small tree 2–5 m tall, with the general floral features of Miconia ciliata; branchlets glabrous in internodes, densely fine-setose at nodes; leaf blades obovate-elliptic to elliptic-oblong, apex acute to shortacuminate, base broadly acute to obtuse, 12– 23 × 6–11 cm, firmly chartaceous and densely ciliolate-serrulate, upper surface deciduously fine-setulose on the primary veins and glabrous or sparsely barbellate-strigulose and stellate-puberulous at the extreme margins, lower surface deciduously finesetulose on the primary veins and glabrous on the surface, basally 5-veined; petioles (1–) 2–6 cm long; inflorescence broadly oblong, with branches to 5 cm long. Evergreen lowland and secondary forests, near sea level to 100 m; Delta Amacuro (Caño Güiniquina, Caño Joba-Suburu east of Caño Sacupana, Curiapo, Jotajana, Manoa, Río Amacuro), Bolívar (Río Cuyuní). Anzoátegui, Aragua, Falcón, Lara, Miranda, Monagas, Sucre, Táchira, Zulia; West Indies, Guyana, Suriname, French Guiana, Brazil. Miconia radulaefolia (Benth.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 243. 1850. —Clidemia radulaefolia Benth., J. Bot. (Hooker) 2: 309. 1840. —Boyuyo morado. Miconia scrobiculata Cogn. in Mart., Fl. Bras. 14(4): 334, t. 66. 1887. Shrub 1–4 m tall, the young growth branchlets densely incurved-setose to stri-

gose with slender smooth hairs ca. 2 mm long; leaf blade elliptic to oblong-elliptic, apex gradually acuminate, base narrowly to broadly acute, 10–25 × 5–10 cm, firmchartaceous and minutely undulate-denticulate, upper surface moderately appressedsetulose with swollen-based hairs, lower surface moderately to densely fine-setulose, shortly 5-pseudopliveined; petioles 1–2 cm long; inflorescence spiciform or few-branched and mostly 8–18 cm long, heavily pubescent and reddish, with bracteate few-flowered glomerules; flowers 5-merous, sessile; petals white, 3–4 × 1.3–1.8 mm, obovate-oblong, glabrous; ovary 3-locular, apically strigulose. Disturbed areas of evergreen lowland and lower montane forests, forests along granitic outcrops, edges of rivers, 100–200(–700) m; Amazonas (Capibara, Cerro Yapacana, Río Mawarinuma, Río Pasimoni, Río Yatua, San Carlos de Río Negro, slopes of Sierra de la Neblina, Yavita). Amazonian Colombia, Guyana, Ecuador (Napo), Peru (Loreto), Brazil (Amazonas, Pará). ◆Fig. 370. Miconia rhytidophylla Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 139. 1850. —Clidemia desmantha Benth., J. Bot. (Hooker) 2: 309. 1840. Shrub 1–3 m tall, the terete branchlets, primary veins on lower leaves, and inflorescences densely puberulous with dendritic hairs 0.1–0.2 mm long and sparsely to moderately glandular-setulose with smooth hairs 1–1.5 mm long; leaf blades lance-elliptic, apex gradually acuminate, base broadly acute to obtuse, mostly 8–12 × 3–5 cm, subrigid and entire or very obscurely crenulate and ciliate on the margin, upper surface rugulose and deciduously stellulate-puberulous, lower surface densely puberulous with dendritic-stellulate hairs, basally 5-veined (including the tenuous marginal veins) with the veinlets elevated-reticulate on lower surface; petioles mostly ca. 1 cm long; panicle 10–15 cm long and few-branched from near the base (type II on Fig. 314); flowers 5merous, sessile, interrupted-glomerulate; petals white, 2.4–2.5 × 1–1.2 mm, obovate-oblong, glabrous; ovary 3-locular, the stylar collar stellulate-puberulous; fruit when dry strongly 10-ribbed, the ribs lighter colored than the rest of the fruit. Riparian forests, 100–200 m; Amazonas (San Carlos de Río Negro). Guyana(?), Brazil (Amazonas). ◆Fig. 376.

Miconia 425

Miconia roraimensis Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 362. 1915. Glabrous shrub or small tree 3–10 m, with tetragonal young branchlets; leaf blades ovate-elliptic to elliptic, apex subabruptly acuminate, base obtuse to rounded, 6–10 × 2.5–5 cm, subcoriaceous and entire, shortly 3-pliveined with dense venule reticulation; petioles 1–2.5 cm long; panicle 4–9 cm long and submultiflorous; flowers 5-merous on pedicels 2–3 mm long; petals white, 1.7–2 × 1.7–2 mm, subrotund, minutely granulose; ovary 4-locular, gland-edged on the apical collar. Upper tepui-slope forests, 1000–2400 m; Bolívar (Roraima-tepui), Amazonas (Cerro Aracamuni, Cerro Autana, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Guyana (Mount Roraima), Brazil (Amazonas: Serra da Neblina). ◆Fig. 375. Miconia rubiginosa (Bonpl.) DC., Prodr. 3: 183. 1828. —Melastoma rubiginosa Bonpl., Monogr. Melast. 1: 109, t. 47. 1816. Miconia rubiginosa var. platyura Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 166. 1850. Shrub or small tree 1–4(–10) m tall, the branchlets, lower leaf surfaces, and inflorescences densely puberulous with reddish short-stalked stellate hairs; leaf blades oblong-elliptic to ovate-oblong, apex acute to short-acuminate, base rounded to cordulate, 6–14 × 2.5–6 cm, rigid and entire, above deciduously stellulate-puberulous, basally 3(5) -veined; petioles 0.3–1 cm long; panicle (5–) 10–15(–25) cm long and submultiflorous; flowers 5-merous, sessile, fragrant; petals white, 2.3–3 × 1–1.7 mm, obovate-elliptic, densely granulose; ovary 3-locular, glabrous. Shrublands, savannas, 100–1200 m; Bolívar (Cerro Bolívar, Cerro Perro in upper Río Paragua basin, Chivatón, Gran Sabana, Guayaraca, Guri area, 12 km south of Kavanayén, Río Asa, base of Sororopán-tepui, Túriba, Uarama, Urimán, Uonquén), Amazonas (Galipero, La Esmeralda, Río Cunucunuma, upper Rio Ventuari, San Carlos de Río Negro, Sierra Parima, above Yutajé). Barinas, Distrito Federal, Falcón, Lara, Miranda, Monagas, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Costa Rica, Panama, Puerto Rico, Hispaniola, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia. ◆Fig. 374.

Miconia rufescens (Aubl.) DC., Prodr. 3: 180. 1828. —Melastoma rufescens Aubl., Hist. P1. Guiane 408, pl. 157. 1775. Miconia impetiginosa DC., Prodr. 3: 183. 1828. Miconia rufescens var. grandifolia Cogn. in Mart., Fl. Bras 14(4): 274. 1887. Shrub 0.5–1.5(–3) m tall, the young growth densely covered with smooth or sparsely barbellate hairs 2–5 mm long underlain by sessile or short-stipitate stellate hairs; leaves sessile or subsessile (petioles to 0.5 cm long), blades ovate, apex broadly acute to obtuse, base subcordate, 6–13(–19) × 4–8(–13) cm, rigid or subrigid and irregularly dentate, upper surface rugose and deciduously puberulous with short-stipitate stellate hairs, lower surface moderately puberulous with sessile and short-stipitate stellate hairs, 5- or 7-veined with the veinlets prominently elevated-reticulate on lower surface; panicle 8–15 cm long, subspicate, the short lateral branches with many-flowered glomerules; flowers 5-merous, sessile; petals white, 4–5 × 2.5 mm, oblong-obovate, glabrous; ovary 3-locular, inconspicuously granulose apically. Open areas, mostly on sparse or poor soils, 100–1200 m; widespread in Bolívar and Amazonas. Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Monagas, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 378. Miconia ruficalyx Gleason, Brittonia 1: 181. 1932. —Copedesma nitens Gleason, Bull. Torrey Bot. Club 52: 331. 1925. Tree 8–15(–20) m tall, the young growth densely stellulate-scurfy with appressed reddish hairs; leaf blades oblong-elliptic to lance-oblong, apex acute to short-acuminate, base attenuate-rounded, 7–18 × 2.5–5 cm, firm-chartaceous and entire, upper surface glabrous and nitidulous, lower surface sparsely to very sparsely and deciduously stellulate-scurfy (hairs ca. 0.1 mm diameter), basally 3-veined (excluding the tenuous marginal veins) with lax venulation; petioles 1–3 cm long; panicle 10–20 cm long, narrowly oblong; flowers 5-merous, sessile, crowded-secund on the short branchlets; petals white, 3.5–4.5 × 2–2.5 mm, obovate-oblong, glabrous; ovary 3(4)-locular, glabrous. Seasonally flooded riparian forests, near sea level to

426

M ELASTOMATACEAE

600 m; Delta Amacuro (Río Amacuro), Bolívar (El Abismo, El Samay, Los Pijiguaos, upper Río Paragua), Amazonas (Mahorima, Río Coro Coro west of Cerro Yutajé, Río Yudi). Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil (Amapá, Pará), Bolivia. Miconia rugosa Triana, Trans. Linn. Soc. London 28: 106. 1871. Miconia plumosa Gleason, Bull. Torrey Bot. Club 52: 381. 1925. Small tree 2–8 m tall, the young growth densely to moderately covered with flexuouserect basally roughened hairs 3–8 mm long underlain by stellulate pubescence; leaves sessile or short-petiolate (to 1 cm), blades elliptic to obovate-elliptic, apex acuminate, base attenuate and cordulate, 15–35 × 6–18 cm, subrigid and entire or obscurely undulate-denticulate, upper surface smooth to bullate-rugose and glabrescent, lower surface moderately stellate- or dendriticpuberulous, 2–6 cm subalternately 3- or faintly 5-pliveined; panicle 10–15 cm long, subracemiform; flowers 5-merous, sessile; petals usually white, ca. 5 × 3 mm, obovateoblong, minutely granulose; ovary 3-locular, apex sparsely glandular-puberulous. Understory of evergreen lowland to lower montane forests and disturbed areas, 100–700 m; Bolívar (El Paují, Río Abacapá, Río Acanán, Río Tírica), Amazonas (slopes of Cerro Aracamuni, slopes of Cerro Marahuaka, La Esmeralda, Río Casiquiare, Río Mawarinuma, base of Sierra de la Neblina, San Carlos de Río Negro to Solano). Amazonian Colombia, Guyana, Suriname, Amazonian Brazil. ◆Fig. 379. The specimens grouped under this species vary considerably in the degree to which the upper surface of leaves is strongly bullate or not. Those with strongly bullate surfaces tend to have more paniculate inflorescences and dendritic hairs on the veins on the lower surface of leaves. The smoother-leaved specimens have scurfy, long-reddish hairs on the stems. Both kinds occur in the area of San Carlos de Río Negro. Further study may show that two different species are involved. Smaller-leaved and less pubescent specimens from lowland Amazonas that were previously identified under this species more likely belong under Miconia matthaei. Finally, some specimens similar to Miconia rugosa in southern Amazonas state

have been tentatively assigned to Miconia cinchonaefolia DC., a species originally described from Brazil. These specimens differ in the denticulate leaf margins, rather spicate inflorescences, and sessile, ribbed fruits. They include Liesner 6552 (MO), along the road from San Carlos de Río Negro to Solano, and Liesner 16598 (MO), from the Río Mawarinuma at the base of Sierra de la Neblina. Miconia rupestris Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 363. 1915. Shrub 1–5 m tall, the sulcate-quadrate young branchlets, foliage, inflorescences, and hypanthia sparsely to moderately but deciduously scaly-scurfy; branchlet internodes sparsely to very sparsely setulose with rather stout smooth hairs to 1 mm long; leaf blades elliptic to oblong-elliptic, apex acuminate, base obtuse, 5–10 × 1.5–4 cm, subcoriaceous and distantly callose-serrulate, glabrous at maturity, shortly (3–5 mm) 3pliveined; petioles 1–2.5 cm long; panicle 3–8 cm long; flowers 5-merous, pedicels ca. 2 mm long; petals white, glabrous, 2 × 1–1.6 mm, broadly obovate; ovary 3-locular, glabrous. Dwarf forests on tepui summits and slopes, 2000–2700 m; Bolívar (Aparamán-tepui, Aprada-tepui, Auyán-tepui, Ilú-tepui, Kukenán-tepui, Macizo del Chimantá [Apacarátepui], Ptari-tepui, Roraima-tepui), Amazonas (Cerro Marahuaka). Guyana (Mount Roraima). Miconia rupticalyx Wurdack, Mem. New York Bot. Gard. 10(4): 35. 1961. Tree 5–8 m tall, the branchlets and inflorescences moderately stellate-puberulent with sessile hairs; leaf blades ovate, apex subabruptly short-acuminate, base rounded, 12–28 × 6–19 cm, membranous and undulate-serrulate (each tooth with a deciduous seta), upper surface glabrous, lower surface sparsely to moderately stellate-puberulent on the primary veins but the rest of the surface glabrous, basally (3)5- or 7-veined, the somewhat inconspicuous veinlets densely reticulate on lower surface; petioles (2–)4–11 cm long; panicle 10–12 cm long and wide, many-flowered; flowers 4-merous, sessile; petals white, glabrous, 2.2–2.3 × 1.3–1.4 mm, oblong-ovate; ovary (2)3-locular, inconspicuously glandular-setulose on the apical collar. Montane forests, 1000–1200 m; Ama-

Miconia 427

zonas (slopes of Sierra de la Neblina). Guyana, Ecuador, Bolivia. Miconia serialis DC., Prodr. 3: 182. 1828. Miconia tomentella Cogn. in Mart., Fl. Bras. 14(4): 284. 1887. Shrub or small tree 2–8 m tall, the branchlets, lower surface of leaves, inflorescences, and hypanthia completely covered with an appressed cobwebby indument; branchlet nodes with distinct raised interpetiolar lines; leaves in each pair somewhat different in size; leaf blades oblong-lanceolate to oblong-elliptic, apex acute to shortacuminate, base acute and decurrent, 6–20 × 2–7 cm, subrigid and entire or obscurely crenulate, upper surface glabrous, shortly 3pliveined (excluding the tenuous marginal veins); petioles 0.2–0.5 cm long; panicle 6–12 cm long and multiflorous; flowers 5-merous, sessile, secund on the branchlets; petals white, 3.2–3.3 × 2–2.2 mm, obovate, glabrous; ovary 3-locular, the apical cone very sparsely glandular. Evergreen lowland forests, 50–200 m; Delta Amacuro (Los Castillos), Bolívar (Las Trincheras, Río Paragua, upper Rio Paramichí, Sierra Imataca), Amazonas (Piedra Parhueña). Miranda; Suriname, French Guiana, Brazil. Miconia serrulata (DC.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 118. 1850. —Diplochita serrulata DC., Prodr. 3: 177. 1828. —Miconia macrophylla var. serrulata (DC.) Cogn. in Mart., Fl. Bras. 14(4): 241, t. 49. 1887. Shrub or small tree 1.5–6(–15) m tall, the branchlets, primary leaf veins on lower surface, and inflorescences densely scurfy with short dendritic or stellate-dendritic hairs; leaf blades ovate-oblong to ovate, apex acute to shortly acuminate, base rounded to cordulate, (15–)20–35 × (5–)10–20 cm, thin-rigid and regularly serrulate with blunt teeth, upper surface glabrous, lower surface (as the bracts and hypanthia) completely covered with stellulate hairs, basally (5)7(9)-veined; petioles 3–9 cm long; panicle 20–40 cm long and multiflorous, flowers predominately 6merous and sessile, subtended by deciduous bracts 6–11 × 4–7 mm; hypanthium 5–7 mm long, 12-ridged; calyx 2.5–3.5 mm long; petals white, 6–9 × 3.5–6 mm, obovate-oblong, externally densely stellulate-lepidote; ovary mostly 4- or 5-locular, apically sericeous-

strigulose. Evergreen lowland to lower montane forests, riparian forests, 100–900 m; widely scattered in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Lara, Mérida, Miranda, Táchira, Trujillo, Yaracuy, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 381. Miconia silicicola Gleason, Bull. Torrey Bot. Club 58: 428. 1931. Tree (or shrub) mostly 5–15 m tall, the young growth completely covered with subdendritic or very short-stalked stellate hairs; leaves sessile, the blades oblong-elliptic to elliptic-obovate, apex obtuse to short-acuminate, base cordulate-clasping, 10–30 × 5–10 cm, rigid and entire, upper surface glabrate, 5-pliveined with the inner pair of primary veins diverging 1/3–1/2 the blade length above the base; inflorescence paniculate, 8–12 cm long, narrowly oblong and rather few-flowered; flowers 5-merous, sessile; ovary 3-locular, apex sparsely glandular-setulose. Montane forests, 1200–2000 m; Bolívar (Arabopó, Auyán-tepui, Cerro Jaua, Gran Sabana, Ilútepui, Macizo del Chimantá [Chimantátepui], Uarama-tepui), Amazonas (Cerro Aratitiyope, Cerro Coro Coro, Cerro Duida, Cerro El Sol, Cerro Guanay, Cerro Yutajé, Sierra de Maigualida, Sierra Uasadi). Adjacent Guyana. ◆Fig. 383. Miconia solmsii Cogn. in Mart., Fl. Bras. 14(4): 393. 1887. Shrub or small tree 4–14 m tall; branchlets compressed, becoming terete with age; leaf blades elliptic-ovate to ovate-oblong, apex long-acuminate, base obtuse, 15–35 × 5–12 cm, subcoriaceous and entire to inconspicuously undulate, upper side glabrous (stellate-puberulous basally on the primary veins), lower surface sparsely puberulous with shortly stipitate stellate hairs (stipe 0.1–0.2 mm long, the apex 0.4–0.5 mm diameter), basally 5-veined; petioles 1.5–2.5 cm long; panicle 10–20 cm long, broad, manyflowered; flowers 5-merous, sessile or very shortly (to 0.3 mm) pedicellate; petals white and minutely granulose, ca. 1.5 × 0.8 mm, oblong-obovate; ovary 3-locular, glabrous. Evergreen lowland and riparian forests, 100–600 m; Bolívar (El Paují, Río Nichare), Amazonas (La Esmeralda, Río Mawarinuma,

428

M ELASTOMATACEAE

Río Orinoco between Río Ocamo and Piedra Mapoya). Lara, Mérida, Táchira, Trujillo; Trinidad. ◆Fig. 384. Miconia splendens (Sw.) Griseb., Fl. Brit. W. I. 256. 1860. —Melastoma splendens Sw., Prodr. 70. 1788. Miconia obovalis Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 183. 1850. Shrub or small tree 3–8 m tall, the branchlets, inflorescences, and hypanthia sparsely to moderately stellulate-scurfy; leaf blades obovate-oblong to elliptic-oblong, apex subabruptly acuminate, base acute, mostly 12–30 × 5–12 cm, firmly chartaceous and sinuate-denticulate to entire, upper surface glabrous, lower surface sparsely stellulatepuberulous on the main veins and glabrous on the surface, shortly (0.5–2 cm) 3-pliveined (excluding the tenuous marginal veins) with veinlets on lower surface densely plane-reticulate (areoles 0.2–0.3 mm wide); petioles 0.5–1.5 cm long; panicle 12–20 cm long and multiflorous; flowers 5-merous, subsessile; petals white, 3.5–3.7 × 1.8–2 mm, obovate, externally densely granulose; ovary usually 3-locular, apically moderately stellulategranulose. Evergreen lowland to lower montane and riparian forests, 100–1000 m; Delta Amacuro (Río Toro), Bolívar (Gran Sabana, Icabarú, Río Abacapá, Río Caura, Río Nichare, Río Paragua, Río Parguaza, Río Toronó), Amazonas (Cerro Sipapo, La Esmeralda, Mavaca, Río Cataniapo, Río Coro Coro west of Cerro Yutajé, Río Cuao, Río Manapiare, Río Padamo, Río Pasimoni, Río Siapa, Río Yatua, San Carlos de Río Negro). Apure, Barinas, Mérida, Miranda, Táchira, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 386. Miconia sprucei Triana, Trans. Linn. Soc. London 28: 108. 1871. Small tree 3–6 m tall; young branchlets and petioles densely scurfy, glabrescent with age; leaf blades obovate-oblong or ellipticlanceolate, apex abruptly long-acuminate, base long-attenuate and decurrent on the petiole, 10–15 × 4–6 cm, entire to more often undulate-margined, upper surface glabrous, lower surface glabrous except for the scurfypuberulent primary veins, 3-pliveined; petioles 0.5–1 cm long; panicle suberect, 2–3 cm long, the short branches divaricate, subfili-

form, 1–3-flowered; flowers 5-merous, pedicels 1–2 mm long; petals white, obovatespatulate, subreflexed, 2.5–3 × 1.5–2 mm, externally densely scurfy-puberulent; ovary 3-locular, glabrous or finely velutinous under magnification. Evergreen lowland forests, secondary forests, open areas, seasonally flooded riparian areas, 100–300 m, Amazonas (Río Emoni tributary of Río Casiquiare, Río Yatua, San Carlos de Río Negro). Brazil (Amazonas: Rio Negro). ◆Fig. 385. Miconia stenostachya DC., Prodr. 3: 181. 1828. Miconia hypargyrea Miq., Linnaea 18: 622. 1845. Shrub 0.5–2 m tall, the branchlets, lower leaf surfaces, inflorescences, and hypanthia completely covered with a white arachnoid indument; young branchlets sharply quadrangular; leaf blades elliptic-oblong, apex acute, base rounded, 8–16 × 4–9 cm, firmly chartaceous and entire or obscurely crenulate, upper surface glabrous and nitidulous, basally 5-veined (including the attenuated marginal veins); petioles 1–3 cm long; panicle 5–15 cm long, many-flowered; flowers 5-merous, sessile, secund on the branchlets, with persistent bracteoles; hypanthium 2.5– 3 mm long; petals white, 3.5–4 × 2 mm, oblong to obovate-oblong, densely granulose and glandular-ciliolate; ovary 3-locular, apically minutely granulose. Borders of lowland and upland savannas, 100–1300 m; Bolívar (Arabopó, Maripa, Río Asa, Río Karaurín, Río Paragua, Río Parguaza, Santa Elena de Uairén), Amazonas (La Esmeralda, Puerto Ayacucho to Samariapo, Santa Barbara, Sierra Parima, Tobogán de la Selva south of Coromoto). Anzoátegui, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Zulia, Táchira; southern Mexico, Central America, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia. ◆Fig. 382. Miconia stephananthera Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 359. 1915. Miconia palmetorum Pittier, Bol. Soc. Venez. Ci. Nat. 8: 314. 1942. Shrub 1–3 m tall, with the general aspect of M. alternans, the branchlets and inflorescences essentially glabrous, the leaf blades generally larger (12–29 × 6–14 cm) and firm-

Miconia 429

chartaceous with the base decurrent on the petiole almost to the base and the venule areoles somewhat finer (ca 0.3 mm, rather than 0.5–0.7 mm), the calyx limb only undulately lobed, the anther thecae 3–3.5 mm long. Savanna-forest ecotones, gallery forests, edges of Mauritia palm swamps, 100–1500 m; Bolívar (Altiplanicie de Nuria, base of Auyán-tepui, 14 km south of Caicara, Corozal 6 km from Maniapure, Hato La Vergareña, Kavanayén, Piedra Marimare in Río Orinoco, slopes of Ptari-tepui, Río Cuyuní, Santa Elena de Uairén, Yuruaní), Amazonas (Cerro Mahedi in middle Río Ocamo basin). Anzoátegui, Carabobo, Guárico, Portuguesa; eastern Colombia, Guyana, Suriname, Amazonian Brazil. Miconia steyermarkii Gleason, Fieldiana, Bot. 28: 435. 1952. Shrub or small tree 1.5–8 m tall, the rounded-quadrate branchlets, inflorescence, and hypanthia sparsely stellate-puberulent; leaves subsessile (petioles thick and 2–5 mm long), the leaf blades ovate-oblong, apex obtusely acute or inconspicuously obtuseacuminate, base cordulate to auriculate, 5– 7.5 × 2.5–3.5 cm, subcoriaceous and entire, very sparsely and deciduously stellate-puberulent on both sides, nitidulous on upper surface, basally 3(5)-veined; panicle 5–7 cm long, submultiflowered; flowers 5-merous, pedicels 1.5–2 mm long; petals white and minutely granulose, 3.2–3.5 × 1.5–1.6 mm, oblong-obovate; stigma truncate; ovary 3-locular, glabrous. Tepui-summit and -slope thickets, 800–1700 m; Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina). Endemic. ◆Fig. 377. Miconia strigosa (Triana) Wurdack, Phytologia 22: 404. 1972. —Heterotrichum strigosum Triana, Trans. Linn. Soc. London 28: 134. 1871. Shrub 1–2 m tall; young growth densely curved-bristly with fine, soft hairs 2–3 mm long; leaf blades elliptic-lanceolate, apex gradually long-acuminate, base broadly acute, (6–)9–17 × (3–)4–7 cm, firmly membranous and densely ciliate-serrulate on upper surface and moderately short-bristly, moderately fine-bristly on the lower surface, shortly (to 1.5 cm) 5(7)-pliveined with the veinlets on the lower side densely reticulate; petioles 1.5–5 cm long; panicle sub-multi-

flowered; flowers 6-merous, sessile; calyx 2– 3 mm long, the outer teeth linear-bristly and extending 6–8 mm; petals white, glabrous, 6–10 mm × 4–7 mm, oblong-obovate; ovary 4locular, quite nearly superior and moderately glandular-setulose. Seasonally flooded riparian forests, 100–200 m; Amazonas (Río Baría, Río Yatua). Brazil (Amazonas: Rio Negro basin). Miconia superba Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 356. 1915. Miconia pachypoda Gleason, Fieldiana, Bot. 28: 435. 1952. Tree 5–15 m tall, the branchlets, primary leaf veins on lower surface, inflorescences, and bracts covered with short dendritic hairs; branchlets, petioles, and inflorescences moderately to densely setose with robust smooth hairs 5–12 mm long; leaf blades ovate-elliptic to oblong-elliptic, apex abruptly shortacuminate, base broadly acute to obtuse, 20–35 × 10–22 cm (much larger in sterile shoots), coriaceous and entire, upper surface glabrous, lower surface completely covered with dendritic-stellate hairs, basally 5veined; petioles 3–9 cm long, robust; panicle 20–30 cm long, many-flowered, narrowly pyramidal; flowers 5-merous, sessile, subtended by deciduous bracts 7–11 × 6 mm; hypanthium ca. 3 mm long, glabrous; calyx 3.5 mm long and 1–1.5 mm lobed; petals pink, 10–11 × 5.5–6 mm, glabrous; ovary 3-locular, glabrous. Montane forests and tepui slopes, (200–)900–1400(–1900) m; Bolívar (slopes of Aprada-tepui near Urimán, Cerro Irama 30 km northeast of Uonquén, Kilómetro 88, La Escalera, Macizo del Chimantá [Apacarátepui, Churi-tepui], Río Aponguao, Roraimatepui and other tepuis). Adjacent Guyana. ◆Fig. 387. Miconia tetraspermoides Wurdack, Phytologia 18: 155. 1969. Tree (or shrub) mostly 2–8 m tall, the terete branchlets, primary leaf veins on the lower surface, inflorescences, and hypanthia sparsely to densely stellulate-puberulous; leaf blades oblong-lanceolate, apex shortly blunt-acuminate, base broadly acute, 8–14 × 3.5–5 cm, firmly chartaceous and distantly undulate-serrulate, glabrous on both surfaces, basally 3-veined with lax (1–2 mm) leaf venule areoles; petioles 1–2 cm long; panicle 5–9 cm long and submultiflorous, the

430

M ELASTOMATACEAE

branches 4–6 per node; flowers 5-merous on pedicels 1.5–3.5 mm long; petals white, 1.9– 2.2 × 1–1.1 mm, ovate-oblong with broadly acute apex, granulose; ovary usually 3-locular, granulose at the apex. Lower montane forests, forest edges, savannas, 400–1100 m; Bolívar (Gran Sabana, Oparuma near Kavanayén, Río Curutú in upper Río Paragua basin, Sierra de los Pijiguaos, near Urimán). Guyana, Suriname, French Guiana, Amazonian Brazil. Miconia theaezans (Bonpl.) Cogn. in Mart., Fl. Bras. 14(4): 419. 1888. —Melastoma theaezans Bonpl. in Humb. & Bonpl., Monogr. Melast. 1: 17, t. 9. 1807. Miconia multinervulosa Cogn., Monogr. Phan. 7: 926. 1891. Small tree (or shrub) 3–10 m tall, the quadrate young branchlets, leaves, and inflorescences glabrous or deciduously scaly; leaf blades oblong-elliptic to ovate-elliptic, mostly 5–10 × 2–4 cm, chartaceous to subcoriaceous and distantly callose-serrulate, basally 3veined or shortly pliveined with rather dense venule reticulation; petioles mostly 0.8–2 cm long; panicle mostly 4–8 cm long, many-flowered; flowers 5-merous on pedicels 0.8–1.2 mm long; petals white, 0.8–1.3 × 0.7–1.3 mm, suborbicular, glabrous; ovary 2- or 3locular, glabrous. Tepui-summit thickets, 1800–2600 m; Bolívar (Auyán-tepui, Ilútepui, Macizo del Chimanta, Roraima-tepui, Sierra de Maigualida), Amazonas (Cerro Yutajé, Sierra de la Neblina). Aragua, Barinas, Distrito Federal, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Central America, Jamaica, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 388. Miconia tillettii Wurdack, Mem. New York Bot. Gard. 10(5): 167. 1964. Shrub or small tree 1–3(–8) m tall, glabrous except for the inflorescence; leaf blades elliptic to obovate-elliptic, apex shortly and subabruptly blunt-acuminate, base cuneate, 10–30 × 4–10 cm, thin and entire, 5pliveined with very laxly reticulate veinlets; petioles 1.5–5 cm long; panicle 6–13 cm long and submultiflorous, sparsely and deciduously scurfy; flowers 5-merous on pedicels 1– 1.2 mm long; petals white, ca. 2 × 0.8 mm, obovate-oblong, glabrous; ovary 3-locular,

sparsely puberulous apically. Montane forests and tepui-slope forests, 800–1800 m; Bolívar (Carrao-tepui, Santa Elena de Uairén, Sierra de Lema, Sororopán-tepui), Amazonas (Cerro Cuao, Cerro Duida, Cerro Huachamacari, Sierra de la Neblina, Sierra Parima). Guyana, Suriname, French Guiana, Brazil (Roraima). ◆Fig. 380. Miconia tinifolia Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 225. 1850. Miconia tinifolia var. roraimensis Wurdack, Phytologia 22: 408. 1972. Shrub 2–10(–15) m tall, the young growth glabrous or moderately puberulous; young branchlets quadrangular; leaf blades elliptic to oblong-elliptic, apex shortly blunt-acuminate, base acute to obtuse, 3–8 × 1.5–3 cm, subrigid and obscurely ciliolate-serrulate, basally 3-veined or shortly 3-pliveined; petioles 0.5–1.5 cm long; panicle mostly 4–6 cm long; flowers 5-merous on pedicels 1–2 mm long; petals white, 1.5–2.3 × 1.5–2.2 mm, obovate-suborbicular, minutely granulose; ovary 3-locular, glabrous. Tepui-summit scrub, 2300–2800 m; Bolívar (Auyán-tepui, Los Testigos, Ptari-tepui, Roraima-tepui, Tramen-tepui), Amazonas (Cerro Marahuaka). Widespread in the Venezuelan Andes and Coastal Cordillera, Anzoátegui, Zulia; Colombia, Guyana, Ecuador. ◆Fig. 365. Miconia tomentosa (Rich.) D. Don ex DC., Prodr. 3: 183. 1828. —Melastoma tomentosa Rich., Actes Soc. Hist. Nat. Paris 1: 109. 1792. —Jucunda tomentosa (Rich.) Benth., J. Bot. (Hooker) 2: 302. 1840. —Diplochita tomentosa (Rich.) Griseb., Fl. Brit. W. I. 252. 1860. —Sakau (Arekuna). Pogonorhynchus amplexans Crueg., Linnaea 20: 107. 1847. —Miconia amplexans (Crueg.) Cogn. in Mart., Fl. Bras. 14(4): 256. 1887. Small to fairly large tree (2–)4–10(–20) m tall, the young growth densely covered with dendritic-stellulate hairs; leaves (sub)sessile, the blades elliptic to pandurate, apex acuminate, base attenuate and ± cordateclasping, 20–40 × 10–25 cm, chartaceous to subcoriaceous and entire to obscurely undulate-serrulate, upper surface glabrous, lower surface sparsely to moderately puberulous with dendritic-stellate hairs, prominently (to 15 cm) 3-pliveined; panicle 12–30 cm long,

Miconia 431

oblong, many-flowered; flowers 5-merous, sessile; petals white to pink-purple, 5–8 × 2– 4 mm, obovate-oblong, minutely pruinosegranulose; ovary 3(4)-locular, apex glandular-puberulous. Evergreen lowland and lower montane forest understories and forest edges, 50–900 m; widespread in Bolívar and Amazonas. Apure, Barinas, Mérida, Monagas, Táchira, Zulia; Central America, Cuba, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Miconia traillii Cogn. in Mart., Fl. Bras. 14(4): 242. 1887. Small tree 3–8 m tall, vegetatively pubescent as in M. serrulata; leaf blades oblongelliptic, apex short-acuminate, base subrounded-obtuse, 15–25 × 6–12 cm, rather firm and obscurely undulate-serrulate, basally 3-veined (excluding the tenuous marginal veins), upper surface glabrous, lower surface light tan-brown with distinct stellate, lepidote, or dendritic hairs; petioles 3–6 cm long; panicle 10–20 cm long, oblong, submultiflorous, with dichasially branched side branches; flowers predominantly 6merous and subsessile, subtended by deciduous elliptic bracts 2–3 mm long; petals white, 6–9 × 3–6 mm, obovate-oblong; ovary 4- or 5locular, very shortly sericeous. Lower montane forests, 300–700 m; Amazonas (slopes of Cerro Huachamacari, upper Río Siapa). Guyana, Suriname, French Guiana, Peru (Loreto), Brazil (Amapá, Amazonas, Roraima). Miconia trinervia (Sw.) D. Don ex Loudon, Hort. Brit. 174. 1830. —Melastoma trinervia Sw., Prodr. 69. 1788. Melastoma scorpioides Schltdl. & Cham., Linnaea 5: 564. 1830. —Miconia scorpioides (Schltdl. & Cham.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 243. 1850. Miconia anceps Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 150. 1850. Miconia caudiculata Pittier, J. Wash. Acad. Sci. 14: 448. 1924. Shrub or tree 2–15 m tall, the young growth moderately puberulent with minute lepidote-stellulate hairs; brachlets ancipital (flattened), the nodes with a well-developed interpetiolar ridge; leaf blade elliptic-oblong, apex acute to short-acuminate, base acute and decurrent, 13–30 × 5–10 cm, firmchartaceous and entire, upper surface glabrous, lower surface ± glabrescent, 3-pli-

veined with lax venule areoles; petioles 0.5– 1.5 cm long; panicle 15–25 cm long, manyflowered; flowers 5-merous, sessile, secund on the curved branchlets; petals white, 2.7– 3.1 × 1.6–1.8 mm, oblong-obovate, minutely granulose; ovary 3-locular, apex sparsely stellulate-scurfy. Evergreen lowland forests, riparian and seasonally flooded forests, 100– 500 m; Bolívar (Caño Colorado, Río Maniapure, lower slopes of Sierra de Maigualida), Amazonas (Caño Yureba in lower Río Ventuari basin, Río Casiquiare, Río Mawarinuma, upper Río Orinoco, Río Siapa, Río Yatua, Santa Bárbara, Yutajé). Apure, Barinas, Táchira, Zulia; southern Mexico through Central America, Jamaica, Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 389. Miconia truncata Triana, Trans. Linn. Soc. London 28: 104. 1871. Shrub to small tree 2.5–6 m tall, the branchlets and lower leaf surfaces covered by a pale, narrowly appressed, amorphous and lepidote indument; leaf blades elliptic-ovate, apex slightly acuminate, base obtuse, 8–18 × 3.5–7 cm, entire and firm-membranous, upper surface glabrous, very slightly (to 0.7 cm) 5-pliveined; petioles 1–2(–3) cm long; panicles 3–10 cm long, few-flowered; flowers 6-merous, pedicels 2–5 mm long; petals white or pink, glabrous, 7.5–8 × 5.5–6 mm, oblong-obovate and rounded at the apex; ovary 5(?)-locular, apex conical and inconspicuously glandular-puberulent. River banks of black-water rivers, 100–200 m; Amazonas (Caño San Miguel, Río Negro near San Carlos, Río Sipapo, Río Yatua). Colombia (Vaupés). ◆Fig. 366. Miconia undata Triana, Trans. Linn. Soc. London 28: 107. 1871. —Boyuyo palo de vieja. Shrub 1–2 m tall; stems and leaves completely glabrous, leaf blades narrowly elliptic to narrowly obovate, apex caudate (3–4 cm), base attenuate, 15–22 × 4–6 cm, 3-pliveined, nitid on upper surface with sulcate secondary veins, the same veins prominent and raised on lower surface; petioles 1–2 cm long; inflorescence 5–8 cm long, few-flowered, with up to twice-branched side branches. Evergreen lowland forest understory near streams, 100–200 m; Bolívar (Río Caura), Amazonas (Río Mawarinuma, San Carlos de

432

M ELASTOMATACEAE

Río Negro to Solano). Brazil (Amazonas, Rondônia). Miconia wittii Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 367. 1915. Miconia micrantha Pilger, Verh. Bot. Vereins Prov. Brandenburg 47: 173. 1905, non Cogn. 1896. Shrub 1.5–6 m tall, the young growth completely covered by sessile, lepidote hairs, the pubescence initially rufous then becoming rufo-cinereous; young branches strongly compressed but soon becoming terete; leaf blades lanceolate to elliptic, apex narrowly acute or shortly acuminate, base broadly acute to rounded, 8–20 × 3–8 cm, subcoriaceous and entire, glabrous on the upper surface, 3- or 5-veined with the veinlets on the lower surface completely covered by the pubescence; petioles 0.7–3 cm long; panicle 8– 15 × 5–14 cm, many-flowered, the flowers 5merous, sessile, with few glomerules at the tips of the short side shoots; petals white, glabrous, 2.7–3.3 × 1.5–1.7 mm, obovate and emarginate; ovary (2)3-locular, apex moderately lepidote. White-sand savannas, edge of forests and granitic outcrops, seasonally flooded riparian areas of black-water rivers, 100–200 m; Amazonas (Caño Cupaven oppo-

site San Fernando de Atabapo, Caño San Miguel, base of Cerro Yapacana, Río Baria, Río Casiquiare, Río Pasimoni, Río Sipapo, Río Yatua, near San Carlos de Río Negro to Solano). Colombia (Vaupés), Brazil (Amazonas). Miconia yatuensis Wurdack, Mem. New York Bot. Gard. 10(4): 33, fig. 32A–E. 1961. Shrub 1–2 m tall, the branchlets, petioles, and inflorescences densely scurfy with dendritic hairs; leaves sometimes very unequal in size in pairs or even pseudoalternate, leaf blades elliptic, apex subabruptly short-acuminate to caudate, base cuneate, chartaceous, clearly undulate-crenate, 15–23 × 6–9 cm, moderately scurfy along the central vein on the lower surface, the rest of the surface glabrous, 3-pliveined with the veinlets laxly reticulate on the lower surface; petioles 4–11 cm long; inflorescence ca. 7 cm long, the branches densely flowered; flowers 4merous, sessile; petals white and slightly granulose externally, ca. 1.6 × 1–1.1 mm, obovate; ovary 3-locular, apex glandularciliolate. Seasonally flooded forests along black-water rivers, 100–200 m; Amazonas (Río Mawarinuma, Río Yatua). Endemic.

Fig. 315. Miconia aguitensis

Miconia 433

Fig. 316. Miconia affinis

Fig. 317. Miconia acinodendron

434

M ELASTOMATACEAE

Fig. 318. Miconia biglomerata

Fig. 319. Miconia aplostachya

Fig. 320. Miconia acutifolia

Miconia 435

Fig. 321. Miconia alata

Fig. 322. Miconia ceramicarpa

436

M ELASTOMATACEAE

Fig. 323. Miconia albicans

Fig. 324. Miconia alternans

Miconia 437

Fig. 325. Miconia alborufescens

Fig. 326. Miconia argyrophylla

438

M ELASTOMATACEAE

Fig. 327. Miconia ampla

Miconia 439

Fig. 328. Miconia borjensis

Fig. 329. Miconia centrodesma

440

M ELASTOMATACEAE

Fig. 330. Miconia bracteata

Fig. 331. Miconia carassana

Miconia 441

Fig. 332. Miconia campestris

Fig. 334. Miconia bubalina

Fig. 333. Miconia brevipes

442

M ELASTOMATACEAE

Fig. 335. Miconia curta subsp. curta

Fig. 336. Miconia cautis

Fig. 337. Miconia dodecandra

Miconia 443

Fig. 338. Miconia chrysophylla

Fig. 340. Miconia decurrens

Fig. 339. Miconia eugenioides

444

M ELASTOMATACEAE

Fig. 341. Miconia ciliata

Fig. 342. Miconia guaiquinimae

Fig. 343. Miconia fallax

Miconia

Fig. 344. Miconia lepidota

Fig. 345. Miconia dioica

Fig. 346. Miconia iluensis

445

446

M ELASTOMATACEAE

Fig. 347. Miconia dispar

Miconia 447

Fig. 348. Miconia elaeoides

Fig. 349. Miconia elata

448

M ELASTOMATACEAE

Fig. 350. Miconia fragrans

Fig. 351. Miconia grossidentata

Miconia 449

Fig. 352. Miconia holosericea

Fig. 353. Miconia impetiolaris var. spruceana

450

M ELASTOMATACEAE

Fig. 354. Miconia lateriflora subsp. lateriflora

Fig. 355. Miconia lasseri

Miconia 451

Fig. 356. Miconia maroana

Fig. 357. Miconia longispicata

452

M ELASTOMATACEAE

Fig. 358. Miconia marginata

Fig. 359. Miconia macrothyrsa

Miconia 453

Fig. 360. Miconia macrotis

454

M ELASTOMATACEAE

Fig. 361. Miconia minutiflora Fig. 362. Miconia phaeophylla

Fig. 363. Miconia mirabilis

Miconia 455

Fig. 364. Miconia myriantha

Fig. 366. Miconia truncata

Fig. 365. Miconia tinifolia

456

M ELASTOMATACEAE

Fig. 367. Miconia nervosa

Fig. 368. Miconia polita

Miconia 457

Fig. 369. Miconia punctata

Fig. 370. Miconia radulaefolia

458

M ELASTOMATACEAE

Fig. 371. Miconia pseudoaplostachya

Fig. 372. Miconia prasina

Fig. 373. Miconia pseudocapsularis

Miconia

Fig. 374. Miconia rubiginosa

Fig. 375. Miconia roraimensis

Fig. 376. Miconia rhytidophylla

459

460

M ELASTOMATACEAE

Fig. 377. Miconia steyermarkii

Fig. 378. Miconia rufescens

Miconia 461

Fig. 379. Miconia rugosa

462

M ELASTOMATACEAE

Fig. 380. Miconia tillettii

Fig. 381. Miconia serrulata

Miconia 463

Fig. 382. Miconia stenostachya

Fig. 383. Miconia silicicola

464

M ELASTOMATACEAE

Fig. 384. Miconia solmsii

Miconia 465

Fig. 385. Miconia sprucei

Fig. 386. Miconia splendens

466

M ELASTOMATACEAE

Fig. 387. Miconia superba

Miconia 467

Fig. 388. Miconia theaezans

Fig. 389. Miconia trinervia

468

M ELASTOMATACEAE

Fig. 390. Miconia perobscura

28. MICROLICIA D. Don, Mem. Wern. Nat. Hist. Soc. 4: 301. 1823. by Paul E. Berry Usually ericoid shrubs. Leaves isomorphic, tightly decussate. Flowers 5merous and terminal on leafy branchlets. Hypanthium terete; calyx lobes persistent; petals usually pink, obovate, often cuspidate-tipped, usually eciliate. Stamens 10, dimorphic, glabrous; anthers oblong and rostrate, 1-pored; connective well prolonged below the thecae and with a ventral, often expanded appendage. Ovary 3locular, superior, glabrous; style glabrous; stigma punctiform. Capsule 3-valved. Seeds numerous, oblong to ovoid, slightly curved, foveolate. Colombia, Venezuela, Guyana, Peru, Brazil (most species on the Brazilian Shield), Bolivia; ca. 130 species, 2 in Venezuela, both in the flora area. Key to the Species of Microlicia

1. 1.

Stems, leaves, and hypanthia glabrous ................................ M. benthamiana Stems, leaves, and hypanthia strigulose or puberulent .......... M. guanayana

Microlicia benthamiana Triana ex Cogn. in Mart., Fl. Bras. 14(3): 77. 1883. Microlicia bryanthoides Oliver, Timehri 5: 192. 1886. Microlicia steyermarkii Gleason, Fieldiana, Bot. 28: 425. 1952. Densely branched, glabrous shrub 0.3–1 m tall; leaf blades 4–9 × 0.7–3 mm, elliptic to oblanceolate, apex rounded to acute, base acute, subrigid, ± punctate on both sides, slightly 3-veined, petiole to 0.8 mm long; petals 6.8–20 × 3.5–7 mm, not setulose. Open tepui scrub, 1000–2700 m; Bolívar (most tepui slopes and summits, Gran Sabana), Amazo-

nas (Cerro Parú, Cerro Yutajé, Sierra Parima). Guyana, Brazil (Roraima). ◆Fig. 391. Microlicia guanayana Wurdack, Mem. New York Bot. Gard. 10(1): 95. 1958. Densely branched shrub, the stems and hypanthia puberulent; leaf blades oblanceolate, apex abruptly acute, base attenuate, 6–9 × 1.2–1.5 mm, subrigid and setuloseciliolate, 1-veined, subsessile; pedicels 0.5– 1.5 mm long; petals 10–11 × 4–5 mm, with a terminal seta 1–1.5 mm long. Tepui scrub, 1700–1800 m; Amazonas (Cerro Guanay). Endemic.

Monochaetum 469

Fig. 391. Microlicia benthamiana

29. MONOCHAETUM (DC.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 4: 48. 1845, nom. cons. —Arthrostemma sect. Monochaetum DC., Prodr. 3: 138. 1828. Arthrostemma sect. Monocentrum DC., Coll. Mem. I. Mélast. 51. 1829. Grischowia H. Karst., Auswahl Gew. Venez. 15. 1848. Loevigia H. Karst. & Triana, Linnaea 28: 434. 1856. Roezlia Regel, Trudy Imp. S.-Petersburgsk. Bot. Sada 1: 98. 1871. by Frank Almeda Shrubs or subshrubs. Leaves petiolate, entire, strigose to hirtellous. Inflorescences terminal, simple or compound cymes. Flowers 4-merous. Hypanthium terete, 8-veined; calyx lobes persistent or deciduous after anthesis; petals obovate, usually ciliate. Stamens 8 (in flora area) or 4, dimorphic; thecae subulate with a solitary, dorsally inclined apical pore; connective prolonged dorso-basally into upturned appendages. Ovary superior, 4-locular, free, setose apically. Fruits 4-locular capsules. Seeds cochleate, smooth or with a barely evident surface pattern. Mexico, Central America, Colombia, Venezuela, Guyana, Ecuador, Peru; 45 species; 15 in Venezuela, 1 of these in the flora area.

470

M ELASTOMATACEAE

Monochaetum bonplandii (Kunth) Naudin, Ann. Sci. Nat. Bot. sér. 3, 4: 51. 1845. —Rhexia bonplandii Kunth in Humb. & Bonpl., Monogr. Melast. 2: 159. 1823 [1808]. Rhexia canescens Bonpl. in Humb. & Bonpl., Monogr. Melast. 2: 47. 1823 [1808]. Rhexia incana Spreng., Syst. Veg. 2: 309. 1825. Chaetogastra rhynchanthera Benth., Pl. Hartw. 180. 1839 [1845]. Loevigia sericea H. Karst. & Triana, Linnaea 28: 434. 1856. Monochaetum sericeum Naudin ex Linden, Belgique Hort. 8: 169. 1858. Shrub to 2 m tall, the branches, lower surface of leaves, and hypanthia moderately to densely hirtellous with smooth, spreading hairs 1–2 mm long; leaves 1.5–4 × 0.7–3.2 cm, elliptic to ovate-elliptic, 5(–7)-veined, the upper surface moderately to densely strigose; petals 9–13 × 5–8 mm, pink; stamens 8, the larger anthers fertile, gently arcuate, 8– 11 mm long with appendages 2–3 mm long, the small anthers sterile and filiform, 3–5 mm long with appendages 0.5–1.7 mm long. Open tepui summit areas, forest margins, 1800–2800 m; Bolívar (most tepuis). Anzo-

Fig. 392. Monochaetum bonplandii

átegui, Lara, Mérida, Portuguesa, Táchira, Trujillo; Colombia, Guyana (Mount Roraima). ◆Fig. 392.

30. MOURIRI Aubl., Hist. Pl. Guiane 452. 1775. Petaloma Sw., Prodr. 4. 1788. Olisbea DC., Prodr. 3: 31. 1828. Guildingia Hook., Bot. Misc. 1: 122. 1830. by Thomas Morley Trees or shrubs, evergreen or sometimes deciduous. Blades entire, pinnateveined, mostly glabrous; stipules represented by an interpetiolar line or a series of small scales or awns; petioles short or none. Inflorescence a small thyrse or dichasium, sometimes a solitary flower, sometimes umbelloid or fasciculate, axillary, ramiflorous, or cauliflorous; bracts small, often early deciduous. Flowers bisexual, 5merous or seldom 4-merous. Hypanthium usually well developed, sometimes none; calyx lobed or truncate, the lobes often separating further at anthesis, sometimes the lobes fused and partly or fully enclosing the petals before anthesis, then splitting regularly or irregularly at anthesis; petals purple, pink, white, or yellow, rarely red, lanceolate to ovate, triangular, trullate, elliptic, or obovate. Filaments usually folded inward in bud and straight at anthesis, rarely short in bud and not unfolding, the antesepalous ones usually shorter than the antepetalous ones; anthers usually isomorphic, the thecae straight or curved, adaxial at anthesis, dehiscent by two lengthwise slits or apical slits or pores or rarely a single pore, connective usually ± caudate below on the abaxial side, an elliptic, depressed gland usually present on the abaxial

Mouriri 471

side of the connective. Ovary 1–5-locular, the placentation free-central, axile, axilebasal, basal, or parietal, the locules from close together to widely separated. Fruit a berry with 1–12 seeds, often crowned by the hypanthium and calyx, or sometimes the fruit consisting of 2–5 widely separated roundish 1-seeded lobes. Seeds polished at least on the enlarged outer face of the ovule, often all over except for the hilum; testa often adhering to the locule wall; radicle short and straight, the cotyledons thick, fleshy, and planoconvex. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 85 species, 26 in Venezuela, 24 of these in the flora area. Key to the Species of Mouriri 1.

1.

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

4. 5(4).

5. 6(5).

6. 7(6).

7. 8(7).

Calyx fully closed until anthesis, then splitting regularly or irregularly, if calyx lobes regular then either the flowers 4-merous or the calyx circumscissile ............................................................................................. 2 Calyx not closed as above, the petals protruding from the calyx before anthesis, the calyx lobes usually recognizable before anthesis, regular, the flowers 5-merous and the calyx persistent ..................................... 4 Calyx splitting into a few irregular pieces at anthesis; calyx including inferior ovary 10–15 mm long ....................................... M. rhizophorifolia Calyx splitting into 4 or 5 regular ovate-triangular lobes; calyx including ovary ca. 4–6 mm long ........................................................................... 3 Flowers 4-merous, 1 or 2 in the leaf axils; peduncles 0.2–1 mm long (excluding pedicel); calyx persistent .......................................... M. latihila Flowers 5-merous, 1–25 in clusters, axillary or not, 4–20 mm long to base of farthest pedicel; calyx circumscissile ...................................... M. huberi Flowers sessile or nearly so on the peduncles, the true pedicels < 1 mm long (the true pedicel is that part of the flower stalk between the ovary base and the first bract pair or joint below the flower) ........................ 5 Flowers pedicellate, some or all pedicels 1 mm or more long .................. 9 Flowers 1–13 per inflorescence, some or all peduncles or inflorescences 4–15 mm long when measured to base of farthest pedicel or flower; gland on anther 1.4–2 mm long ............................................... M. vernicosa Flowers consistently single on short-bracted peduncles 1.3–4 mm long; gland on anther 0.2–0.7(–1.1) mm long ................................................ 6 Leaves with the primary, secondary, and marginal veins grooved on upper surface and prominent on lower surface; fruit with 7–10 grooves extending 1/3 to all the way down the sides from the apex ........ M. sagotiana Leaves with only the primary vein grooved on upper surface and prominent on lower surface; fruits not grooved ............................................. 7 Upper bracts triangular to broadly ovate-triangular, acute or abruptly so; ovary plus calyx 3.9–5 mm long; thecae 1.6–2.7 mm long; ovary locules 2 ...................................................................................... M. brevipes Upper bracts broadly rounded and often apiculate; ovary plus calyx 1.7–3 mm long; thecae 0.6–1.2 mm long; ovary locule 1 ...................... 8 Petals white to greenish white; primary vein strongly 2-angled below .............................................................................................. M. angulicosta

472

M ELASTOMATACEAE

8. 9(4).

9.

10(9).

10.

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

13.

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

15. 16(15).

16. 17(9).

Petals yellow; primary vein rounded below ............................... M. myrtifolia Midvein on upper surface of leaf flat or rounded, sometimes grooved along each side so that the vein is prominent (xylem of midvein tubular, visible with a hand lens when the leaf is broken 1/4–1/2 of the way above the base); flowers never consistently 1 per peduncle ............................... 10 Midvein on upper surface of leaf distinctly grooved along the center (xylem usually open, forming a U or semicircle with the edges turned inward as seen in section, if tubular then the flowers consistently axillary and single on the peduncles) ............................................................... 17 Locules of the ovary 5, widely separated, appearing as if in the base of the hypanthium, rising at least 1/3 of their height above the style base, set at least 1/3 of their diameter out from the side of the style, often causing bulges on the outside of the apparent hypanthium; ovules parietalbasal ..................................................................................................... 11 Locules of the ovary 1–5, close together, their tops sometimes about level with the style base, their inside edges about even with the side of the style or nearer the center, bulging outwardly or not; ovules axile to basal ..................................................................................................... 13 Calyx including ovary 5–8 mm long; lateral veins in the dried leaf inconspicuous or invisible ................................................................ M. cauliflora Calyx including ovary 3–5 mm long; lateral veins in dried leaf usually fully visible above or below or on both sides ....................................... 12 Leaf blades 9–18 × 2.1–7 cm; calyx lobes 0.4–0.6 mm long, 0.9–1.4 mm long when measured from the stamen attachment .................. M. nervosa Leaf blades 5–12 × 1.5–3.1 cm; calyx lobes 0.8–1.3 cm long, 1.2–1.9 mm long when measured from the stamen attachment ........... M. angustifolia Thecae of anther 4.2–7 mm long, the anthers 4.2–8 mm long, the calyx including inferior ovary 8–17 mm long; calyx splitting between the lobes at anthesis for 1.3–3 mm ............................................ M. grandiflora Thecae 1.7–3.7 mm long, the anthers 2.1–4.1 mm, calyx plus interior ovary 3.6–7(–8.5) mm, if > 7 mm then the splitting distance between the lobes at anthesis 0–0.5 mm ........................................................... 14 Thecae 1.7–2.3 mm long, 0.7–0.8 times the length of the anther; calyx plus inferior ovary 3.6–5.2 mm long; leaf blades 3–13 cm long ..... M. guianensis Thecae 2.5–3.7 mm, 0.8–1 times the length of the anther; calyx plus inferior ovary 4.2–8.5 mm long; leaf blades 4–29 cm long ....................... 15 Calyx lobes 3.8–4.1 mm long when measured from the stamen attachment; pedicels 7–15 mm long; petioles 4–6 mm long; blades 11–29 cm long .......................................................................................... M. longifolia Calyx lobes 0.9–2.6 mm long when measured from the stamen attachment; pedicels 1–10 mm long; petioles 1–5 mm; blades 4–18 cm ...... 16 Leaf blades usually broadly acute to rounded at base, very rarely cordate; calyx lobes 0.9–1.3 mm long when measured from the stamen attachment; ovary locules 2 or 3 .............................................................. M. nigra Leaf blades consistently cordate at base; calyx lobes 1.3–2.6 mm long from the stamen attachment; ovary locules 4 or 5 ...................... M. sideroxylon Anther thecae curved over the apex of the anther and extending at least 0.7–0.8 mm down the other side, dehiscing by lengthwise slits ........ 18

Mouriri 473

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

21(17). 21. 22(21).

22.

23(21). 23.

24(23).

24.

25(24).

25.

Anther thecae not curved over the apex and down, dehiscent by lengthwise slits or apical pores ...................................................................... 21 Calyx including inferior ovary 8–10 mm long ................................ M. ficoides Calyx including inferior ovary 2.7–4.1 mm long ..................................... 19 Petioles 2.5–6 mm long; leaf blades acute and attenuate at base ........................................................................................... M. subumbellata Petioles 0–1.8 mm long; leaf blades broadly acute to rounded or cordate at base ....................................................................................................... 20 Leaves sessile and cordate, (3.5–)6–10 cm long, ovate to oblong-ovate ................................................................................................. M. uncitheca Leaves with petioles 0.8–1.8 mm long, the blades 2.2–6 cm long, elliptic, elliptic-oblong, or slightly ovate-elliptic, cordate at base or rarely truncate or broadly acute ............................................................ M. densifoliata Flowers consistently 1 per peduncle ....................................................... 22 Flowers usually several to many per peduncle, seldom 1 and never consistently so ................................................................................................ 23 Leaf blades 2.7–6.1 cm long, cordate at base with a small notch; calyx lobes triangular-acuminate, 1.4–2.4 mm long before anthesis; petals 5–7 mm long ........................................................................ M. myrtilloides Leaf blades 4–12 cm long, rounded to acute at base; calyx lobes truncate or nearly so, 0–0.4 mm long (during anthesis the lobes split apart an additional 1.2–3 mm); petals 7–12 mm long ......................... M. pauciflora Petals white, pink, purple, or yellow; anther thecae 2.5–3.7 mm long; midrib xylem tubular (see first lead of couplet 9) ............................... M. nigra Petals usually yellow, rarely white (in M. vernicosa); anther thecae 1.5– 2.4 mm long; midrib open above as seen in section (see second lead of couplet 9) .............................................................................................. 24 Calyx lobes at anthesis not splitting apart or doing so only to 0.2 mm; anther gland 1.4–2 mm long, narrow, appearing as a slit in the edge of the anther; true pedicels 0.6–2.5 mm long; bracts of the inflorescence triangular, acute, persistent ............................................................ M. vernicosa Calyx lobes at anthesis splitting apart for 0.9–1.6 mm; anther gland 0.7– 1.4 mm long, not narrow and slit-like; pedicels 2–7.5 mm long; bracts usually deciduous by anthesis, if persistent then the upper ones rounded to broadly acute ..................................................................... 25 Anthers dehiscing by slits widest at the apex but extending nearly to the base of the theca; internodes of inflorescence just below pedicels (0.5–) 0.7–4 mm long; petals mostly drying yellow to yellow-brown; ovary 1or 2-locular .............................................................................. M. acutiflora Anthers dehiscing by apical slits; internodes just below pedicels 0.1–0.7 (–1.2) mm long; petals mostly drying brownish to black; ovary 3–5locular ..................................................................................... M. collocarpa

Mouriri acutiflora Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 284. 1852. —Flor de mosquito amarillo. Shrub or tree 3–30 m tall; petals yellow to orange. Along streams and in moist primary and secondary forests above or below flood

level, ca. 50–300 m; Bolívar (Río Parguaza), Amazonas (Río Casiquiare, Río Guaviarito on Cerro Guanay, Río Guayapo, Río Pasiba). Southeastern Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 394.

474

M ELASTOMATACEAE

Mouriri angulicosta Morley, Phytologia 22: 424. 1972. Tree to 30 m tall; petals white to greenish white. Moist forests above flood level, 500– 700 m; Bolívar (Cerro Ichún). Suriname, French Guiana, Brazil (Amazonas, Pará). The Venezuelan specimen differs from other members of the species in having a hypodermis in the leaf. Mouriri angustifolia Spruce ex Triana, Trans. Linn. Soc. London 28: 155. 1871. Small tree to 6 m tall; petals white or pinkish. Moist forests, caatingas, white-sand campina forests, 100–200 m; Amazonas (San Carlos de Río Negro). Colombia (Guainía), Brazil (northern Amazonas). Mouriri angustifolia resembles M. nervosa but is smaller in stature. Mouriri brevipes Hook., J. Bot. (Hooker) 2: 24. 1840. Shrub or small tree 1.5–15 m tall; flowers very fragrant; petals yellow. Moist, often secondary forests usually near water, seasonally flooded riparian forests of black-water rivers, 100–300 m; Bolívar (Río Oris in Río Paragua Basin), Amazonas (Caño San Miguel, Río Atacavi, Río Baría, Río Pasimoni). Guyana, Suriname, northern Brazil. ◆Fig. 393. Mouriri cauliflora Mart. ex DC., Prodr. 3: 7. 1828. Shrub or tree to 15 m tall; petals white to pink or rose-violet. Moist, well-drained, usually primary forests, sometimes caatingas, often on white-sand soil; 100–200 m; Amazonas (Yavita). Southeastern Colombia, Brazil (Amazonas), Peru. Mouriri collocarpa Ducke, Anais Reunião Sul-Amer. Bot. 3: 69. 1940. —Dereyo. Mouriri acutiflora var. oligantha Gleason, Recueil Trav. Bot. Néerl. 32: 212. 1935. Tree to 30 m tall; petals yellow. Moist primary or secondary forests, on clay or sand soils; 50–200 m; Amazonas (middle Río Orinoco). Suriname, French Guiana, Brazil (Amazonas, Pará, Rondônia). Mouriri densifoliata Ducke, Trop. Woods 76: 25. 1943. Small tree to 5 m tall; petals usually white, varying to yellow or light yellowish

tinged with rose. Savannas, dominant in thickets, 100–200 m; Amazonas (81 km above the mouth of Río Guayapo). Brazil (Amazonas, Pará, Roraima). Mouriri ficoides Morley, Phytologia 22: 425. 1972. —Derello hoja grande. Tree to 35 m tall; petals white to yellow or partly light orange to rose; fruits edible. Moist primary forests, mostly above flood level, reportedly in Leopoldinia piassaba palm stands, 100–200 m; southwestern Amazonas. Brazil (Amazonas). ◆Fig. 395. Mouriri grandiflora DC., Prodr. 3: 8. 1828. —Derello, Môlôkoto kahi. Mouriri princess Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 283. 1852. Usually a small tree, sometimes to ca. 30 m tall, often shrubby, often weak and somewhat vine-like; petals white to rose or yellow or marked with red or orange; fruit edible. Moist primary or secondary forests, above or below flood level, commonly near streams, sometimes at edges of savannas, 100–400 m; Amazonas (scattered). Eastern and southern Colombia, Guyana, Suriname, French Guiana, Brazil, Amazonian Ecuador, Peru, and Bolivia. ◆Fig. 397. Mouriri guianensis Aubl., Hist. Pl. Guiane 452. 1775. —Cascarito, Cometure. Petaloma mouriri Sw., Prodr. 73. 1788. Mouriri polyantha Miq., Linnaea 18: 290. 1844. Mouriri weddellii Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 286. 1852. Mouriri ulei Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 184. 1905. Typically a small, often bushy tree to 10 m tall, less often to 30 m tall; petals white to pink or sometimes yellow; fruit edible. Moist secondary and apparently primary forests, savannas, open places, and sea shores, usually near water, often in low places subject to flooding, reported on sand and clay soils, near sea level to 200 m; Delta Amacuro (widespread), Bolívar (Río Caura), Amazonas (Río Orinoco above Río Atabapo). Apure, Monagas; Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil (widespread). Mouriri huberi Cogn., Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 255. 1909. —Guarataro, Guayaba paujicera.

Mouriri 475

Tree to 30 m tall; petals white to rose or yellow; fruits edible. Moist primary or secondary forests above flood level on clay or sand soil, 100–500 m; Delta Amacuro (Río Toro, Serranía de Imataca), northeastern Bolívar. Guyana, French Guiana, Peru, Brazil (Amapá, Amazonas, Pará). Mouriri latihila Morley, Ann. Missouri Bot. Gard 76: 434. 1989. Tree to 3 m tall. Evergreen lowland forests, 100–200 m; Amazonas (between Yavita and Pimichín). Endemic. Mouriri longifolia H.B.K. (Morley), Phytologia 22: 427. 1972. —Myrtus longifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 258. 1825. —Derello, Hoja grande. Tree to 10 m tall; petal color unknown. Nonflooded lowland forests, 100–200 m; Amazonas (middle and upper Río Orinoco). Endemic. Mouriri myrtifolia Spruce ex Triana, Trans, Linn. Soc. London 28: 154. 1871. Shrub or usually a tree to 11 m tall; petals yellow; fruits edible. Moist forests, often near streams, above or below flood level, 100–1300 m; central Bolívar, Amazonas (Boca Mavaca). Apure, Mérida; Ecuador, Peru, Brazil (Amazonas, Mato Grosso, Rondônia, Roraima). Mouriri myrtilloides (Sw.) Poir. in F. Cuvier, Dict. Sci. Nat. ed. 2, 33: 163. 1824. —Petaloma myrtilloides Sw., Prodr. 833. 1788. Shrub or tree to 6 m tall; petals white to yellow. Neotropics. 4 subspecies, 2 in Venezuela, 1 of these in the flora area. M. myrtilloides subsp. orinocensis Morley, Brittonia 23: 423. 1971. Thickets or low woods near streams or gallery forests, 50–200 m; Bolívar (Río Suapure), Amazonas (Isla Ratón, Raudale de Atures). Apure; adjacent Colombia. Mouriri nervosa Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 183. 1905. —Guaratarillo. Shrub or tree to 12 m tall; petals white to rose. Understories of moist primary or secondary forests, open woods or savannas, mostly on sandy soil, above flood level, 200–

700 m; Bolívar (lower Río Suapure, vicinity of Upata). The identity of the two collections from the flora area is uncertain. Both are sterile; leaf shapes of Mouriri nervosa and M. sideroxylon overlap, and although the leaves of these collections best match those of M. nervosa, the plants are out of the usual range of that species and are within the range of M. sideroxylon, and so may belong to that species. Mouriri nigra (DC.) Morley, Phytologia 22: 428. 1972. —Eugenia nigra DC., Prodr. 3: 268. 1828. —Derello, Dereye, Macho, Tereyo. Mouriri plasschaerti Pulle, Recueil Trav. Bot, Néerl. 6: 283. 1909. Small tree to 15 m tall; petals white, pink, purple, or yellow. Moist primary forests, often near water, above or below flood level, sometimes at the savanna edges, on sand or clay soil, 50–700 m; western Bolívar (Río Parguaza, lower Río Suapure), western Amazonas (Culebra, Río Casiquiare, Río Cataniapo, lower Río Ventuari, road between San Carlos de Río Negro and Solano). Colombia (Vaupés), Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil. Mouriri pauciflora Spruce ex Cogn. in Mart., Fl. Bras. 14(4): 581. 1888. Tree to 10 m tall; petals usually yellow, sometimes rose-violet. Moist forests usually near water, above or below flood level, 100– 200 m; southern Amazonas (La Esmeralda, Río Casiquiare). French Guiana, Brazil (Amapá, Pará, Maranhão). Mouriri rhizophorifolia (DC.) Triana, Trans. Linn. Soc. London 28: 153. 1871, “rhizophoraefolia.” —Olisbea rhizophorifolia DC., Prodr. 3: 31. 1828, “rhizophoraefolia.” —Guarataro, Guavillo, Guayaba Pesjua. Guildingia psidioides Hook., Bot. Misc. 1: 122. 1830. Tree to 30 m tall; petals white; fruits edible. Tall to low and shrubby forests, moderately moist to dry, the ground level to sloping and rocky, 100–300 m; Bolívar (40 km south of Tumeremo). Anzoátegui, Aragua, Miranda, Monagas, Sucre, Yaracuy; Trinidad, Tobago. ◆Fig. 398.

476

M ELASTOMATACEAE

Mouriri sagotiana Triana, Trans. Linn. Soc. London 28: 155. 1871. Shrub or tree 2–5(–15) m tall; petals yellow to yellow-orange. Understories of moist or sometimes mesophytic primary or sometimes secondary forests, sometimes savannas, always above flood level, 300–600 m; Delta Amacuro (Río Toro), Bolívar (near Río Asa, Río Icabarú), Amazonas (Río Manaviche east of Boca Mavaca). Guyana, Suriname, French Guiana, Peru, Brazil. Mouriri sideroxylon Sagot ex Triana, Trans. Linn. Soc. London 28: 155. 1871. —Guaratarillo. Shrub or tree 3–28 m tall; petals pink, sometimes white, purple, light blue, or light yellow. Moist forests above flood level, often gallery forests along rivers and sometimes subject to flooding, sometimes dry forests, scrub, or caatinga or at the edges of savannas, 100–1100 m; Delta Amacuro (Río Toro), Bolívar (scattered), Amazonas (scattered). Colombia (Vaupés), Guyana, Suriname, French Guiana, Brazil. Mouriri subumbellata Triana, Trans. Linn. Soc. London 28: 154. 1871.

Shrub or tree 2–5(–25) m tall; petals white or yellowish white. Moist low to high primary forests, usually above flood level, sometimes bordering savannas or on sandy river banks, 100–200 m; Amazonas (Caño Cotúa on Río Temi, Río Guainía, Río Ventuari). Colombia (Guainía), French Guiana, Peru, Brazil. Mouriri uncitheca Morley & Wurdack, Mem. New York Bot. Gard. 10(4): 46. 1961. —Derello. Shrub or tree 1–4 m tall; petals white, cream-colored, or rose. Savannas, whitesand areas with shrubs and small trees, sometimes subject to flooding, 100–200 m; southwestern Amazonas. Endemic. ◆Fig. 396. Mouriri vernicosa Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 285. 1852. Tree to 25 m tall; petals yellow or orange, rarely white. Moist forests above or below flood level, commonly on stream banks, ca. 100–200 m; Amazonas (base of Cerro Sipapo, Río Cuao). Colombia (Vaupés), Suriname, French Guiana, Peru, Brazil.

Fig. 393. Mouriri brevipes

Mouriri 477

Fig. 394. Mouriri acutiflora

Fig. 395. Mouriri ficoides

478

M ELASTOMATACEAE

Fig. 396. Mouriri uncitheca

Fig. 397. Mouriri grandiflora

Myriaspora 479

Fig. 398. Mouriri rhizophorifolia

31. MYRIASPORA DC., Prodr. 3: 165. 1828. by Paul E. Berry Shrubs or small trees. Leaves isomorphic, opposite. Flowers 5-merous, 1–4-axillary in upper leaf nodes. Hypanthium terete; calyx calyptriform and rather irregularly dehiscent above the torus, the persistent base splitting into several (usually 5) lobes; petals white, obovate-lanceolate and acute, minutely granulose, outside cen-

480

M ELASTOMATACEAE

Fig. 399. Myriaspora egensis

trally strigose. Stamens 10, isomorphic, glabrous, thickly lance-oblong; anthers minutely l-pored and not appendaged. Ovary 10-locular, completely inferior, minutely puberulous at the apex; style glabrous; stigma capitate and 5-lobuled. Fruit a berry. Seeds numerous, oblong-elliptic, with lateral raphe and verrucose surface. Amazonian Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 1 species. Myriaspora egensis DC., Prodr. 3: 165. 1828. Myriaspora surinamensis Steud., Flora 27: 722. 1844. Myriaspora decipiens Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 101. 1851. Shrub or small tree 2–8 m tall; young growth and hypanthia densely fine-setose with hairs 2–5 mm long; leaf blades ovate-elliptic, mostly 9–20 × 3.5–9.5 cm, ovate-elliptic, apex gradually acuminate, base acute to rounded, chartaceous and ciliate-denticulate, on both surfaces sparsely to moderately

appressed-setose with fine hairs 1–2 mm long, shortly (1–2 cm) 5-pliveined with laxly reticulate veinlet; petioles 0.5–3 cm long; pedicels 3–7 mm long, with a pair of subpersistent bracteoles 1–2.5 mm long; petals 8–12 × 5–6 mm; stigma 2.5–3 mm diameter. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (Río Canaracuni, Río Caura, Río Paramichi, Serranía de Imataca, Urimán), Amazonas (Tamatama). Monagas, Zulia; Amazonian Colombia, Guyana, French Guiana, Suriname, Peru, Brazil. ◆Fig. 399.

Neblinanthera 481

32. NEBLINANTHERA Wurdack, Mem. New York Bot. Gard. 10(5): 153. 1964. by Paul E. Berry Small shrubs. Leaves opposite, isomorphic. Flowers (5)6(7)-merous, in terminal, multiflowered panicles. Hypanthium terete; calyx lobes barely developed, the external teeth projecting upwards; petals obovate-oblong, glabrous, nonciliate. Stamens similar in form but different in size, glabrous; anthers oblong, minutely 1porate; connective not prolonged, adnate ventrally to the thecae, dorsally with a minute, truncate basal tooth. Ovary superior, 3-locular, moderately glandularsetulose; style glabrous; stigma punctiform. Capsule 3-valved, each valve prominently bifid at the apex. Seeds not known. Endemic to Sierra de la Neblina, Cerro Aracamuni, and Serra Pirapucú in southern Venezuela and adjacent Brazil; 1 species.

Fig. 400. Neblinanthera cumbrensis

482

M ELASTOMATACEAE

Neblinanthera cumbrensis Wurdack, Mem. New York Bot. Gard. 10(5): 153. 1964. Shrub 1–2 m tall; lower surface of leaves, inflorescence, and hypanthia moderately finesetulose with hairs 0.1–0.7 mm long ending in a terminal gland; leaf blades ovate, apex gradually acuminate, base cordate, firmly membranous and ciliolate, 6–14 × 2.5–6.5

cm, 9(11)-veined, with lower order venation densely reticulate on lower surface; petioles 2–5 cm long; panicle 2–7 cm long, trifurcate from the base; pedicels 2–5 mm long; petals white, 3 × 1.5 mm; anthers orange; capsule 5–6 mm long. Montane elfin forests on tepui slopes, 1300–1800 m; Amazonas (Cerro Aracumuni, Sierra de la Neblina). Brazil (Amazonas: Serra Pirapucú). ◆Fig. 400.

33. NEPSERA Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 28. 1850. by Paul E. Berry Herb or subshrub. Stems 4-angled. Flowers 4-merous, in lax, terminal, multiflowered panicles. Hypanthium terete; calyx lobes ovate-subulate, persistent; petals white, oblong-lanceolate, acute, each tipped with a single glandular setula. Stamens 8, slightly dimorphic in size, glabrous; thecae subulate, minutely l-pored ventroapically; connective slightly prolonged, ventro-basally with a bilobed appendage

Fig. 401. Nepsera aquatica

Opisthocentra 483

(lobes bluntly acute), dorsally not appendaged. Ovary 3-locular, superior, glabrous. Capsule 3-valved. Seeds numerous, cochleate, foveolate. Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Brazil; 1 species. Nepsera aquatica (Aubl.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 28. 1850. —Melastoma aquatica Aubl., Hist. Pl. Guiane 430, t. 169. 1775. —Rhexia aquatica (Aubl.) Sw., Fl. Ind. Occid. 2: 650. 1798. —Spennera aquatica (Aubl.) Mart. ex DC., Prodr. 3: 116. 1828. Herb or subshrub 0.5–2 m tall, the branchlets, veins on the lower surface of leaves, inflorescences, and hypanthia sparsely and inconspicuously puberulous with partly gland-tipped hairs; leaf blades 2–5 × 1.5–3 cm, ovate to elliptic-ovate, apex acute, base

rounded to cordate, chartaceous and serrulate, upper surface glabrous or sparsely and deciduously strigulose, lower surface glabrous except for the veins, 5- or 7-veined; petiole 0.5–1 cm long; panicle to 30 cm long; pedicels mostly 3–5 mm long; petals 4.5–7 × 2.5–8 mm. Open, wet lowland areas, Mauritia palm swamps, near sea level to 200 m; Delta Amacuro (Caño Simoina west of Isla Cocuina, Jotajana, Río Cuyubini), Bolívar (Ciudad Bolívar, Río Venamo southwest of Anacoco). Anzoátegui, Monagas, Sucre; other distribution as in genus. ◆Fig. 401.

34. OPISTHOCENTRA Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 749. 1867. by Paul E. Berry Small shrubs. Leaves opposite, isomorphic. Flowers 4-merous, in short, axillary, scorpioid cymes. Hypanthium terete, funnel-shaped to campanulate; calyx lobes broadly oblate and persistent, with minute external teeth; petals elliptic-ovate, apex acute and mucronulate, minutely granulose, nonciliate. Stamens 8, subiso-

Fig. 402. Opisthocentra clidemioides

484

M ELASTOMATACEAE

morphic, glabrous; thecae subulate and strongly curved, with a ventrally inclined pore; connective not prolonged, ventrally with 2 abruptly acute lobes at the base, dorsally with a flagella parallel to the connective. Ovary superior, 3-locular, mostly glabrous; style glabrous; stigma punctiform. Fruit a 3-valved capsule. Seeds minute, pyramidal, minutely muricate, with a large raphe. Río Negro basin in Amazonian Colombia, Venezuela, and Brazil; 1 species. Opisthocentra clidemioides Hook. f. in Benth. & Hook. f., Gen. Pl. 1: 749. 1867. Small shrub 0.5–3 m tall, minutely granulose in the stems, leaves, and young hypanthia, but soon glabrescent; leaf blades elliptic or ovate-elliptic, apex gradually acuminate, base broadly acute to obtuse, entire, membranous, 10–21 × 5–12 cm, 3- or very weakly 5-veined; petioles 2–7 cm long; inflo-

rescence axis 5–10 mm long, ca. 6–8-flowered; pedicels 3–4 mm long; petals white, 7–9 × 5.5–7 mm. Evergreen lowland forests, mostly in black-water river basins, 100– 200 m; Amazonas (Capibara, Maroa, San Carlos de Rio Negro, San Fernando de Atabapo, San Simón de Cocuy, Yavita to Pimichín). Colombia, Brazil (Amazonas). ◆ Fig. 402.

35. OSSAEA DC., Prodr. 3: 168. 1828. by Paul E. Berry Shrubs or small trees with smooth hairs. Flowers 4–6-merous, in short lateral cymes or pseudolateral or in racemes in the upper leaf axils. Hypanthium terete; calyx with the external teeth well developed or inconspicuously lobed; torus glabrous within; petals usually white and lanceolate, acute at the apex, glabrous. Stamens essentially isomorphic (slightly dimorphic in size), glabrous; thecae oblong and with a ventro-terminal pore, the connective neither prolonged nor appendaged. Ovary 3-locular and partly to entirely inferior; style glabrous; stigma not expanded. Fruit a several-seeded berry, costate or terete. Seeds pyramidal to obovoid, smooth. Southern Mexico, Central America, West Indies (where most diverse), Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil; ca. 80 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Ossaea 1. 1.

Flowers 6-merous, sessile; fruits terete when dry; young growth finely bristly ...................................................................................... O. mavacana Flowers 4-merous, in racemose inflorescences; fruits 8-ribbed when dry; young growth granulose-puberulent ..................................... O. micrantha

Ossaea mavacana Wurdack, Phytologia 21: 127. 1971. Shrub with terete stems; leaves and inflorescences moderately to densely short-bristly with fine, smooth hairs; leaf blades 6– 10(–16) × 3–5(–7.5) cm, elliptical, apex shortly acuminate, base acute, shortly (0.5– 1.2 cm) 5-pliveined; petioles 1–1.6 cm long; flowers sessile and 6-merous, usually in pairs in the upper leaf axils, covered by 2 pairs of persistent, narrowly ovate bracts 5– 6 × 2.5–3 mm; calyx limbs ± entire and

ciliolate, the external teeth subulate and setulose, 2.7–3 mm long; petals ca. 3 × 1.2 mm, oblong-lanceolate; ovary 6-locular. Evergreen lowland forests, 100–200 m; Amazonas (Boca Mavaca). Brazil (Roraima). ◆Fig. 403. Ossaea micrantha (Sw.) Macfad., Fl. Jamaica 2: 49. 1850. —Melastoma micrantha Sw., Fl. Ind. Occid. 71. 1788. —Octopleura micrantha (Sw.) Griseb., Fl. Brit. W. I. 260. 1860.

Ossaea 485

Fig. 403. Ossaea mavacana

Fig. 404. Ossaea micrantha

486

M ELASTOMATACEAE

Sagraea neurocarpa Naudin, Ann. Sci. Nat. Bot. sér. 3, 18: 94. 1852. Shrub 2–3 m tall; stems ± quadrangular; young growth moderately covered by a scalystellate, soon dehiscent pubescence; leaf blades elliptic to ovate-elliptic, apex gradually acuminate, base acute to narrowly obtuse, 8–20 × 3.5–7(–10) cm, shortly (0.5–1.5 cm) and subalternately 5-pliveined; inflorescence 3–6 cm long, pseudolateral, the branches divergent; flowers 4-merous, pe-

dicels slender, 2–3 mm long (to 5 mm in fruit); calyx with minute external teeth; petals 3–3.5 × 1–1.2 mm, lanceolate, obtusely acute; ovary 4-locular, 2/3-inferior; fruit notably 8-costate when dry. Evergreen lowland forests, forest understories, Amazonas (Capibara, Tamatama, Río Atabapo, San Carlos de Río Negro). Aragua, Carabobo, Mérida, Portuguesa, Trujillo, Zulia; southern Mexico, Greater Antilles, Colombia, Ecuador, Peru. ◆Fig. 404.

36. PACHYLOMA DC., Prodr. 3: 122. 1828. Urodesmium Naudin, Ann. Sci. Nat. Bot. sér. 3, 15: 338. 1851. by Paul E. Berry and Navina Luckana Subshrubs or shrubs 0.2–3 m tall. Leaves opposite, shortly petiolate. Flowers terminal, solitary or in panicles, 4-merous. Hypanthium campanulate to urceolate; calyx lobes fleshy, inconspicuous; petals pink to violet, ovate or obovate, acute to obtuse at the apex, ciliate. Stamens 8, isomorphic or anisomorphic, glabrous or the filaments inconspicuously glandular-puberulent apically; anthers linear-subulate, arcuate, 1-pored; connective prolonged below the thecae into a bilobed ventral appendage, with a dorsal caudal appendage on the connective between the thecae and the filament insertion. Ovary free, glabrous or apically puberulent or setose, 4-locular; style thin, exserted; stigma punctiform. Fruit a 4-valved capsule. Seeds numerous, cochleate, strongly muriculate. Endemic to the Guayana Shield in Colombia (Vaupés), Venezuela, Guyana, Brazil (Amazonas); 4 species, 3 in Venezuela, all in the flora area. The fourth species (Pachyloma coriaceum DC.) occurs in Colombia, Brazil, and Guyana. A number of intermediate forms between the species covered below have been collected and may represent hybrids or distinct new taxa. Key to the Species of Pachyloma 1. 1. 2(1). 2.

Leaf surface conspicuously glandular-setose; apex of ovary densely setose ..................................................................................................... P. setosum Leaf surface glabrous; apex of ovary glabrous .......................................... 2 Leaves 3(5)-veined, basally rounded or subcordate, generally > 15 mm wide ....................................................................................... P. huberioides Leaves 1-veined or weakly 3-veined, basally acute or attenuate, < 12 mm wide ............................................................................................ P. pusillum

Pachyloma huberioides (Naudin) Triana, Trans. Linn. Soc. London 28: 64. 1871. —Urodesmium huberioides Naudin, Ann. Sci. Nat. Bot. sér. 3, 15: 338. 1851. Pachyloma scandens Ducke, Arq. Inst. Biol. Veg. 2: 65. 1935. Low shrub 0.2–2.5 m tall or occasionally scandent-lianoid; leaves lanceolate to ovate-

elliptic, apex narrowly acute to acuminate, base rounded to subcordate, 4–10 × 1.5–5 cm; petioles 2–8 mm long; petals pink to purple. White-sand savannas, Mauritia palm swamps, open areas in cracks of granitic outcrops, 50–800 m; Amazonas (Caño Chimoni in Río Casiquiare basin, Caño Cupaven opposite San Fernando de Atabapo, Caño

Pachyloma 487

Fig. 405. Pachyloma huberioides

Fig. 406. Pachyloma setosum

Fig. 407. Pachyloma pusillum

488

M ELASTOMATACEAE

Pimichín, Cerro Aracamuni, base of Cerro Duida, Cerro Vinilla, savannas at base of Cerro Yapacana, savannas of La Esmeralda, Río Atabapo basin, Río Baría, Río Pasimoni 50 km above mouth, Río Siapa, Río Sipapo). Colombia (Guainía), Brazil (Amazonas, Roraima). ◆Fig. 405. Pachyloma pusillum Wurdack, Mem. New York Bot. Gard. 10(5): 145, fig. 66a–c. 1964. Pachyloma nanum Wurdack, Phytologia 48: 238. 1981. Subshrub 0.2–0.8 m tall, generally xylopodial (with thick woody underground stem); leaves lance-elliptic, acute at both ends, 2.5– 5.5 × 0.3–1.1 cm, conduplicate when young; petioles 0–3 mm long; petals reddish purple.

White-sand savannas, 100–200 m; Amazonas (Canaripó, Caño Yagua, base of Cerro Moriche, savanna at base of Cerro Yapacana, Cucurital near middle Caño Yagua, Río Puruname, 25 km southeast of Santa Bárbara). Endemic. ◆Fig. 407. Pachyloma setosum Wurdack, Mem. New York Bot. Gard. 10(1): 107. 1958. Shrub 0.2–2 m tall; leaves oval-shaped, rounded at both ends, 3.0–8.5 × 1.5–3.5 cm; petiole 2–4 mm long; petals lavender. Savanna-shrubland on white sand (sabanetas), 100–200 m; Amazonas (western base of Cerro Yapacana, outskirts of Maroa, near San Antonio on Río Sipapo, environs of San Carlos de Río Negro). Endemic (probably in adjacent Guainía, Colombia). ◆Fig. 406.

37. PHAINANTHA Gleason, Bull. Torrey Bot. Club 75: 539. 1948. by Navina G. Luckana and Paul E. Berry Terrestrial herbs, scandent shrubs, lianas, or epiphytes, usually rooting at the nodes, sometimes appressed to tree trunks. Leaves opposite or in whorls of 3, but more often alternate by abortion of leaves on one side of the stem. Inflorescences axillary or terminal, cymose-paniculate or umbelliform. Flowers 4-merous. Hypanthium terete, conical; calyx hyaline and calyptriform, dehiscing at anthesis above the torus; petals white or pink, obovate. Stamens 8, isomorphic or slightly anisomorphic, glabrous; thecae linear-subulate and curved, dehiscent by a small pore; connective not prolonged or only shortly so, with a cordiform or panduriform basal appendage. Ovary free, 4-locular, glabrous or minutely glandular; style glabrous; stigma punctiform. Capsule 4-locular; seeds numerous, linear. Southeastern Venezuela, western Guyana, Ecuador; 5 species, 4 in Venezuela, all in the flora area. Recently a fifth, undescribed species of Phainantha was discovered in southernmost Ecuador, in a montane area on sandstone (Palacios et al. 8565, MO, US). Previously, this genus was considered a Guayana Shield endemic. Key to the Species of Phainantha 1. 1. 2(1). 2. 3(2). 3.

Terrestrial or epiphytic herbs with opposite and ternate leaves; petioles 5–11 cm long ......................................................................... P. steyermarkii Lianas or epiphytic, woody-stemmed plants with opposite or alternate leaves and petioles < 5 cm long ............................................................. 2 Leaves opposite, sessile or subsessile, the blades < 1.5 cm long, apically rounded or emarginate ........................................................ P. myrteoloides Leaves alternate, petiolate, the blades 6–20 cm long, apically acuminate ................................................................................................................ 3 Lower surface of leaves setulose at anthesis; anther connectives not prolonged .................................................................................... P. laxiflora Lower surface of leaves glabrous at anthesis; anther connectives prolonged 0.3–1 mm ........................................................................ P. maguirei

Phainantha 489

Fig. 408. Phainantha maguirei

Fig. 409. Phainantha myrteoloides

Fig. 410. Phainantha steyermarkii

490

M ELASTOMATACEAE

Phainantha laxiflora (Triana) Gleason, Bull. Torrey Bot. Club 75: 539. 1948. —Adelobotrys laxiflora Triana, Trans. Linn. Soc. London 28: 68. 1871. Liana or woody epiphyte. Moist montane forests, 400–1200 m; Bolívar (La Escalera, Río Apacará, Río Tírica, Sierra de Lema, 53 km southwest of Urimán). Guyana. Phainantha maguirei Wurdack, Mem. New York Bot. Gard. 10(5): 147, fig. 65a– h. 1964. Liana or woody-stemmed epiphyte; calyx red, corolla white. Moist montane forests, 1100–1700 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Venamo, Kamarkawarai-tepui, La Escalera, Macizo del Chimantá [Abacapá-tepui, Sarvén-tepui]). Guyana. ◆Fig. 408.

Phainantha myrteoloides Wurdack, Bol. Soc. Venez. Ci. Nat. 23: 74, fig. 6. 1962. Wiry- or slightly woody-stemmed epiphyte appressed to tree trunks, or lithophyte; petals white. Shady streamsides in moist montane forests, 800–1600 m; Bolívar (Cerro Venamo, La Escalera). Guyana. ◆Fig. 409. Phainantha steyermarkii Wurdack, Bol. Soc. Venez. Ci. Nat. 32: 367, fig. 18. 1976. Terrestrial to epiphytic herb; stems thick, to 22 mm diameter; leaves 5-veined, long-petiolate, glabrous to shaggy-pubescent; inflorescence scapose, peduncle 12–16 cm long, flowers 4—10 in an umbellate cluster; petals pink. Shady understory of forest inside giant sinkholes, streambanks on tepui summits, 1100– 1400 m; Bolívar (Cerro Sarisariñama in and around the Sima Mayor). Endemic. ◆Fig. 410.

38. POTERANTHERA Bong., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci, Math., Seconde Pt. Sci. Nat. 6: 137. 1838. by Paul E. Berry Tiny herbs 5–25 cm tall. Stems quadrangular. Leaves opposite, linear, sessile, enervate, glandular-ciliate. Flowers solitary in upper leaf axils, shortly pedunculate, 5-merous. Calyx persistent, with lanceolate-acuminate lobes; petals orbicular-obovate or oblong, the apex rounded or rarely acute, with a terminal glandular seta. Stamens 10, the larger 5 fertile, the smaller abortive; fertile anthers short, obovoid or subglobose, dehiscent by a large pore; connective below the thecae shortly prolonged and bilobed, bicalcarate, or bituberculate at the articulation with the filament. Style elongate, terete, and straight, slightly thickened toward the apex; stigma punctiform or capitellate. Ovary and capsule 3-locular. Seeds numerous, ovoid or subreniform, areolate or foveolate. Venezuela, Brazil; 2 species, 1 in Venezuela. Poteranthera pusilla Bong., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6: 138, t. 8, fig. 1. 1838. Miniature herb 5–10 cm tall; leaves linear, 5–9 mm long, ca. 1 mm wide; flowers subsessile, white to pink, anthers yellow, with glandular trichomes. Open sandy areas and seepages on granitic outcrops, moist savannas, 50–200 m; Bolívar (Cerro El Médano 23 km southwest of Caicara, lower Río Suapure), Amazonas (Puerto Ayacucho). Guárico; Brazil (Goiás, Mato Grosso). ◆ Fig. 411.

Fig. 411. Poteranthera pusilla

Pterogastra 491

39. PTEROGASTRA Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: pl. 15, fig. 8. 1849, and 3, 13: 97. 1850 [text]. by Paul E. Berry and Susanne S. Renner Erect herbs, subshrubs, or shrubs 0.2–1.5 m tall. Branching dichotomous, the branchlets quadrangular or 4-winged. Leaves decussate and isomorphic, elliptic to lanceolate, subsessile or petiolate, with 1 or 2 pairs of longitudinal primary veins arising at the base of the blade. Inflorescences terminal dichotomous cymes. Flowers 4- or 5-merous, pedicellate. Hypanthium narrowly campanulate, 4- or 5-winged, the wings dorsally spinose-ciliate; calyx lobes lanceolate, persistent; petals obovate with

Fig. 412. Pterogastra divaricata

Fig. 413. Pterogastra minor

492

M ELASTOMATACEAE

a short claw, the margins finely ciliolate. Stamens 8 or 10, dimorphic; filaments thin, glabrous; anthers subulate and opening by a single pore, the connective prolonged below the thecae and arcuate, ventrally bilobed, dorsally exappendiculate. Ovary 4or 5-locular, apically setose with glandular trichomes surrounding the style base. Fruit a loculicidal capsule. Seeds numerous, cocheate. Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil?; 2 species, both in Venezuela. This treatment is extracted in large part from S. S. Renner, Nordic J. Bot. 14: 65–71. 1994. Key to the Species of Pterogastra 1. 1.

Flowers 5-merous (visible in both flower and fruit) ................... P. divaricata Flowers 4-merous ................................................................................ P. minor

Pterogastra divaricata (Bonpl.) Naudin, Ann. Sci. Nat. Bot. sér. 3, 13: 98. 1850. —Rhexia divaricata Bonpl. in Humb. & Bonpl., Monogr. Melast. 2: 59, t. 22. 1812 [1813]. Pterogastra major Triana, Trans. Linn. Soc. London 28: 40, t. 2, fig. 30b. 1871. Pterogastra glabra Gleason, Bull. Torrey Bot. Club 52: 326, fig. 1. 1925. Erect, sparingly branched herb to shrub 0.3–1.5 m tall. Savannas, Mauritia palm swamps, weedy places, 50–800 m; Bolívar (ca. 20 km northeast of Cerro Guaiquinima, Upuigma-tepui), Amazonas (La Esmeralda, Puerto Ayacucho to Samariapo, Río Casi-

quiare basin, San Carlos de Río Negro). Apure, Táchira, Zulia; Colombia, Guyana, Ecuador, Peru, Brazil(?). ◆Fig. 412. Pterogastra minor Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: pl. 15, fig. 8. 1849, and 3, 13: 98. 1850 [text]. Annual erect herb 0.2–0.6 m tall. Curatella savannas, depressions in granitic outcrops, 50–200 m; Bolívar (Río Parhueña), Amazonas (5 km northeast of Galipero, Isla Ratón, Puerto Ayacucho, Raudale de Atures, Río Autana, Río Orinoco between Santa Rosa and mouth of Río Vichada, Samariapo). Adjacent Colombia (Meta). ◆Fig. 413.

40. PTEROLEPIS (DC.) Miq., Comm. Phytogr. 72. 1840, nom. cons. —Osbeckia sect. Pterolepis DC., Prodr. 3: 140. 1828. Brachyandra Naudin, Ann. Sci. Nat. Bot. sér. 3, 2: 143. 1844. by Paul E. Berry and Susanne S. Renner Herbs or shrubs, annual or perennial. Leaves decussate, isomorphic, 3- or 5veined. Flowers (3)4- or 5-merous, in terminal cymes or glomerules or solitary in the upper leaf axils. Hypanthium free from the ovary, campanulate, slightly ribbed, with at least some stellate-penicillate trichomes in the intercalycine sinuses; calyx lobes triangular, erect, persistent, ciliate along the margin; petals obovate with a minute claw, pink, reddish purple, or white. Stamens 6, 8, or 10, subisomorphic or dimorphic; anthers yellow or basally pink, subulate or truncate, the pore apical or ventrally inclined, the connective barely or distinctly prolonged below the anthers to the filament insertion, ventrally with a yellow, lobed appendage. Ovary free, (3)4- or 5locular, apically setose; style terete; stigma truncate or punctiform. Fruit a loculicidal capsule. Seeds numerous, small, cochleate with a regularly tuberculate seed coat. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, naturalized on Hawaii; 14 species, 2 in Venezuela, both in the flora area. This treatment is extracted in large part from S. S. Renner, Nordic J. Bot. 14: 73–104. 1994

Pterolepis 493

Key to the Species of Pterolepis 1. 1.

Flowers clustered at the branch tips and upper leaf axils; calyx lobes at anthesis 3–5 mm long .............................................................. P. glomerata Flowers solitary in the upper leaf axils or terminal; calyx lobes at anthesis 1–3 mm long ............................................................................ P. trichotoma

Pterolepis glomerata (Rottb.) Miq., Comm. Phytogr. 2: 78. 1840. —Rhexia glomerata Rottb., Descr. Rar. Pl. Surin. 8. 1776. —Osbeckia glomerata (Rottb.) DC., Prodr. 3: 141. 1828. —Arthrostemma glomerata (Rottb.) Cham., Linnaea 9: 454. 1835. —Chaetogastra glomerata (Rottb.) Benth., J. Bot. (Hooker) 2: 290. 1840. Rhexia angusturensis Rich. in Humb. & Bonpl., Monogr. Melast. 2: 77, pl. 29. 1823 [1813]. —Arthrostemma angusturenese (Rich.) Cogn. in Mart., Fl. Bras. 14(3): 276. 1885. Chaetogastra callichaeta Benth., J. Bot. (Hooker) 2: 291. 1840. Heavily branched annual herb 20–60 cm tall; flowers 4-merous, sessile; petals pink to purple or white. Savannas, near sea level to 200 m; Bolívar (Ciudad Bolívar, Tumeremo to Anacoco road). Anzoátegui, Guárico, Miranda, Monagas, Sucre; West Indies, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Pterolepis trichotoma (Rottb.) Cogn. in Mart., Fl. Bras. 14(3): 261. 1885. —Rhexia trichotoma Rottb., Descr. Rar. Pl. Surin. 9. 1776. Rhexia pumila Bonpl. in Humb. & Bonpl., Monogr. Melast. 2, pl. 35. 1823 [1813]. —Pterolepis pumila (Bonpl.) Cogn. in Mart., Fl. Bras. 14(3): 263. 1885. Pterolepis pumila var. procera Cogn. in Mart., Fl. Bras. 14(3): 264. 1885. Regularly branched annual herb 10–50 cm tall; flowers 4-merous, subsessile; petals pink or white. Disturbed areas, savannas, 50–200 m; Bolívar (6 km from Maniapure to Caicara), Amazonas (Raudale de Atures). Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Guárico, Lara, Mérida, Miranda, Monagas, Portuguea, Táchira, Yaracuy; Mexico, Central America, West Indies, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 414.

Fig. 414. Pterolepis trichotoma

494

M ELASTOMATACEAE

41. RHYNCHANTHERA DC., Prodr. 3: 106. 1828, nom. cons. by Paul E. Berry and Susanne S. Renner Shrubs or subshrubs 0.3–3 m tall; branchlets subterete, angular, or 4–6-angled, hirsute with glandular hairs. Leaves decussate, broadly ovate to linear, petiolate, subsessile or sessile, thin, margins serrulate, serrate, or rarely appearing entire, with 1–4 pairs of longitudinal lateral veins arising at the base of the blade. Inflorescence terminal, thyrsoid, with few- to many-flowered biparous or uniparous cymes; bracts leaf-like and gradually reduced in size distally. Flowers bisexual, 5-merous. Hypanthium campanulate; calyx lobes linear-subulate to triangular, spreading, persistent; petals obovate, thin, reddish pink, purple, magenta, or white. Antesepalous stamens 5, fertile, alternating with 5 abortive antepetalous staminodia, the fertile stamens isomorphic or more often dimorphic with one stamen longer than the others; filaments thin, terete; anthers subulate with a pronounced beak (rostrum), single-pored, the connective greatly prolonged below the thecae to the filament insertion and with a small anterior simple or bilobed appendage. Ovary subglobose, free, perigynous, 3–5-locular, apically setose or glabrous; style thin, terete; stigma punctiform. Fruit a 3–5-loculicidal capsule. Seeds numerous, 0.75–1 mm long, nearly straight, hilum basal. Mexico, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay; 15 species, 4 in Venezuela, 3 of these in the flora area. This treatment was largely extracted from S. S. Renner, Nordic J. Bot. 9: 601– 630. 1990. Key to the Species of Rhynchanthera 1. 1. 2(1). 2.

The 5 fertile stamens dimorphic, one much longer than the others ............................................................................................... R. grandiflora The 5 fertile stamens isomorphic .............................................................. 2 Leaf blades ovate to narrowly ovate, base cordate, rounded or subacute, petiolate, with 2, 3, or 4 pairs of lateral primary veins ........ R. dichotoma Leaf blades lanceolate to linear, base acute, subessile or sessile, with 1 or 2 pairs of lateral primary veins ................................................ R. serrulata

Rhynchanthera dichotoma (Desr.) DC., Prodr. 3: 107. 1828. —Melastoma dichotoma Desr. in Lam., Encycl. 4: 41. 1797. Shrub or subshrub 0.3–1.5 m tall. Open wet places, near sea level to 100 m; Delta Amacuro (Sacupana). Trinidad, Guyana, French Guiana, Peru, Brazil. Rhynchanthera grandiflora (Aubl.) DC., Prodr. 3: 107. 1828. —Melastoma grandiflora Aubl., Hist. Pl. Guiane 414, t. 160. 1775. —Osbeckia aubletiana Spreng., Syst. Veg. 2: 311. 1825. Rhynchanthera ambigua Naudin, Ann.

Sci. Nat. Bot. sér. 3, 12: 210. 1849. Rhynchanthera orinocensis Sprague, Trans. & Proc. Bot. Soc. Edinburgh 22: 431. 1905. Rhynchanthera cardonae Wurdack, Phytologia 14: 263. 1967. Shrub or subshrub 0.5–2 m tall. Open places, 50–400 m; Bolívar (Caicara, Canaima, between Ciudad Bolivar and Río Pao, lower Río Caroní, upper Río Paragua), Amazonas (base of Cerro Yapacana, La Esmeralda, Puerto Ayacucho, Río Asisa). Apure, Guárico, Monagas; Mexico, Panama, Colombia, Guyana, Suriname, French Guiana, Peru, Bolivia, Brazil. ◆Fig. 415.

Salpinga 495

Rhynchanthera serrulata (Rich.) DC., Prodr. 3: 108. 1828. —Rhexia serrulata Rich. in Humb. & Bonpl., Monogr. Melast. 2: 74, t. 28. [1813] 1823. Rhynchanthera modesta Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: 213. 1849. Subshrub 0.3–0.8 m tall. Savannas, 50– 200 m; Bolivar (Caicara to Puerto Ayacucho road, Serranía Carichana near Río Orinoco). Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Brazil.

Fig. 415. Rhynchanthera grandiflora

42. SALPINGA DC., Prodr. 3: 112. 1828. Saccolena Gleason, Bull. Torrey Bot. Club 52: 373. 1925. by Paul E. Berry Herbs or subshrubs. Leaves isomorphic. Flowers 5-merous, arranged in pedunculate, unilateral cymes. Hypanthium ± 10-ridged, narrowly campanulate; calyx lobes triangular to lanceolate, persistent; petals pink or white, obovate to oblongobovate and rounded at the apex, glabrous or sparsely glandular-ciliolate. Stamens 10, ± dimorphic, glabrous; thecae subulate, minutely 1-pored; connective not or slightly prolonged, dorsally with a blunt ascending appendage and sometimes with a short, blunt, basal spur. Ovary 3-locular, glabrous, with an umbilicate apex; style glabrous; stigma punctiform. Fruit a 10-ridged capsule. Seeds numerous, clavate, tuberculate. Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil; 10 species, 4 in Venezuela, all restricted to the flora area. Species of Salpinga usually grow in shady, moist habitats such as streamside boulders, cliff seeps, or mossy tree trunks. They generally have tough root systems that can withstand temporary flooding.

496

M ELASTOMATACEAE

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

3.

Large herbs 50–150(–200) cm tall; petioles 1–5 cm long, leaf blades 5–15 × 3–7 cm ......................................................................................... S. secunda Small herbs 5–30 cm tall; petioles 0–2 cm long, leaf blades 1–7.5 × 0.8–5 cm ................................................................................................. 2 Flowers axillary in upper leaf axils or clustered in pseudo-umbels, not scapose ........................................................................................... S. pusilla Flowers several to numerous on a scape ................................................... 3 Leaves clustered at branch tips; stems tumid-succulent, constricted at the leaf nodes, 5–8 mm diameter, with brownish-lepidote scales ................................................................................................... S. maguirei Leaves basal or intermittent along the stem, not grouped at the tips; stems thinner and less succulent than above, not constricted at the nodes, without brownish scales ............................................. S. glandulosa

Salpinga glandulosa (Gleason) Wurdack, Phytologia 14: 267. 1967. —Macrocentrum glandulosum Gleason, Bull. Torrey Bot. Club 58: 422. 1931. Terrestrial or lithophytic herb 5–25 cm tall, with numerous sessile glands; leaf blade orbicular to broadly elliptic, 1–3 × 0.8–2 cm, petiole 5–17 mm long; petals white or pink. Rocks and caves near waterfalls, escarpments and rock ledges, mossy montane forest undergrowth, (500–)700–1700 m; Bolívar (Cerro Guaiquinima, Cerro Guanacoco, Cerro Ichún, Cerro Pauo in upper Río Caura basin, Cerro Sarisariñama, La Escalera, Macizo del Chimantá, Sierra de Lema), Amazonas (Cerro Autana, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Brazil (Amazonas: Serra Pirapucú). ◆Fig. 416.

Salpinga secunda Schrank & Mart. ex DC., Prodr. 3: 113. 1828. —Shú-she (Yekwana). Large terrestrial to epiphytic herb or small shrub 0.5–1.5(–2) m tall; stems ± succulent, quadrangular; petals white to pink. Shady streamside boulders or on moist tree trunks or boulders in evergreen lowland and lower montane forests, 50–300 m; Amazonas (Piedra Arauicaua, Piedra Cucuy, Pimichín, Río Baría, Río Casiquiare, Río Cunucunuma basin, Río Negro, Yavita). Amazonian Colombia, Ecuador, Peru, Brazil. ◆Fig. 418.

Salpinga maguirei Gleason, Mem. New York Bot. Gard. 8: 137. 1953. Terrestrial herb 10–30 cm tall, stem thick but constricted at the nodes. Montane forest understory, moist rocks, 900–1400 m; Amazonas (Cerro Parú, Cerro Sipapo, Cerro Yapacana). Endemic. ◆Fig. 417. Salpinga pusilla (Gleason) Wurdack, Phytologia 14: 267. 1967. —Macrocentrum pusillum Gleason, Bull. Torrey Bot. Club 58: 422. 1931. Terrestrial or epiphytic herb 5–20 cm tall. Moist mossy areas on rocks or on moist tree trunks in montane forests, 1100–2800 m; Amazonas (Cerro Aracamuni, Cerro Duida, Sierra de la Neblina). Brazil (Amazonas). ◆Fig. 419.

Fig. 416. Salpinga glandulosa

Salpinga 497

Fig. 417. Salpinga maguirei

Fig. 418. Salpinga secunda

Fig. 419. Salpinga pusilla

498

M ELASTOMATACEAE

43. SANDEMANIA Gleason, Bull. Misc. Inform. Kew 1939: 480. 1939. by Paul E. Berry and Susanne S. Renner Shrub 1–2 m tall. Branchlets angled. Leaves opposite, petiolate, exstipulate; blades 4.5–10 × 1.5–4 cm, ovate, 5(–9)-veined. Inflorescence a many-flowered, terminal panicle to 18 cm long. Flowers 4-merous. Hypanthium narrowly campanulate; pedicels ca. 1 mm long; calyx lobes triangular, 1–1.2 mm long; petals narrowly elliptic, apically acute, pale purple with darker blotches at the base, 3–4.5 × 1.2–2 mm. Stamens slightly dimorphic; filaments 3.4–3.8 or 3–3.6 mm long, glabrous; connective prolonged below the pollen sacs and modified at the base into 2 blunt, ventral lobes. Ovary free from the hypanthium wall, globose, 2-locular; style filiform, sigmoid; stigma punctiform. Fruit a loculicidal capsule. Seeds numerous, cochleate. Venezuela, Peru, Brazil; 1 species. This treatment is extracted in large part from S. S. Renner, Brittonia 39: 441– 446. 1987. Sandemania hoehnei (Cogn.) Wurdack, Phytologia 20: 370. 1970. —Comolia hoehnei Cogn., Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 5, Bot. 3: 9. 1912. Sandemania glandulosa Wurdack, Phytologia 5: 53. 1954. Shrub 1–2 m tall; leaf blades stiff, brittle. On rocky slopes above river in steep canyon, open vegetation on tepui slopes, 300–800 m; Amazonas (Sierra de la Neblina). Peru, Brazil. ◆Fig. 420.

Fig. 420. Sandemania hoehnei

Siphanthera 499

44. SIPHANTHERA Pohl ex DC., Prodr. 3: 121. 1828. by Frank Almeda and Orbelia R. Robinson Annual or suffrutescent perennial herbs, 2–60(–100) cm tall. Leaves petiolate to sessile, entire to serrate, occasionally revolute, glabrous to copiously pubescent. Inflorescences terminal, modified simple or compound dichasia, lax to congested, occasionally umbelliform. Petals 4, frequently clawed. Stamens 4 (or 8 and dimorphic in S. cowanii), antesepalous, sometimes with 1–4 antepetalous staminodia; anthers linear-subulate to suborbicular, connective thickened dorsally, and variously modified ventrally into sessile or prolonged, poorly or well-developed, bilobed appendages. Ovary superior, 2-locular, glabrous, ellipsoid to globose, occasionally compressed laterally. Capsule globose, rounded, emarginate or notched at summit. Seeds tear-shaped to reniform, areolate. Colombia, Guyana, Suriname, Peru, Brazil, Bolivia; 15 species, 7 in Venezuela, all in the flora area. Key to the Species of Siphanthera 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

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

Branchlet internodes completely glabrous ..................................... S. cowanii Branchlet internodes variously pubescent ................................................ 2 Cauline pubescence consisting (at least in part) of appressed malpighiaceous hairs ............................................................................ S. fasciculata Cauline pubescence consisting of simple, appressed or spreading hairs ................................................................................................................ 3 Cauline leaves superficially appearing quaternate-verticillate due to the proximity of the internodes (the 4 leaves of comparable size) .... S. duidae Cauline leaves paired at each node (the 2 leaves of comparable size) ..... 4 Cauline leaves linear-oblong; thecae of staminodia hatchet-shaped in profile view, with a well-defined terminal pore ................................. S. foliosa Cauline leaves elliptic, ovate, cordate, suborbicular, or rhombic; staminodia (when present) linear-oblong, filamentous and lacking a well-defined pore, or with a rudimentary elliptic anther sac that is distinctly rostrate with a minute terminal pore ................................................... 5 Cauline leaves sessile or subsessile, subcordate at the base ...... S. cordifolia Cauline leaves petiolate, if subsessile then not subcordate at the base .. 6 Inflorescence a simple or umbelliform dichasium; flowers pedicellate .................................................................................................. S. cordifolia Inflorescence a bracteate, glomerulate head; flowers sessile or subsessile ................................................................................................................ 7 Hypanthium at least twice the length of the enclosed capsule .. S. hostmannii Hypanthium and enclosed capsule ± equal in length, or the hypanthium rarely exceeding the enclosed capsule by 1/3 of its length ....... S. dawsonii

Siphanthera cordifolia (Benth.) Gleason, Bull. Torrey Bot. Club 56: 400. 1929. —Meisneria cordifolia Benth., J. Bot. (Hooker) 2: 299. 1840. Meisneria microlicioides Naudin, Ann. Sci. Nat. Bot. sér. 3, 12: 204. 1849. —Siphanthera microlicioides (Naudin)

Cogn. in Mart., Fl. Bras. 14(3): 194. 1883. Siphanthera jenmanii Gleason, Brittonia 1: 137. 1932. Siphanthera cordifolia var. glomerata Gleason, Fieldiana, Bot. 28: 426. 1952. Strict, suffrutescent perennial herb 10–43

500

M ELASTOMATACEAE

cm tall; leaves 1–3-veined or obscurely 3veined, ovate to rhombic-ovate, 4–7(–9) × 2– 4(–7) mm; densely to moderately glandularsetose intermixed with a shorter denser underlayer of setulose hairs, the upper surface sparsely to moderately setulose, the lower surface glandular-punctate, sparsely glandular-setose, setulose, or glabrate; stamens 4, with 4 antepetalous staminodes present per flower; appendages not developed; mature capsules with a rounded summit. Soil pockets in open rocky ground, savannas, and wet meadows, 700–2200 m; Bolívar (Gran Sabana), Amazonas (Cerro Parú). Colombia (Vaupés), Guyana. ◆Fig. 423. Siphanthera cowanii Wurdack, Mem. New York Bot. Gard. 10(1): 97. 1958. Strict annual herb 7.5–61 cm tall, unbranched or loosely branched; leaves sessile, 1-veined, linear, 5–40 × 0.3–2 mm, glabrous, but obscurely punctate on lower surface; stamens 8, weakly dimorphic, alternately differing in size; antesepalous anthers tapered distally into a rostrum ± 0.75 mm long with a truncate pore; capsule apex emarginate. White-sand savannas, 100–200 m; Amazonas (base of Cerro Moriche, Guarinuma, 5 km southeast of Santa Bárbara del Orinoco, Yapacana savannas). Endemic. Siphanthera dawsonii Wurdack, Los Angeles County Mus. Contr. Sci. 28: 8. 1959. Unbranched or loosely branched, annual herb 2.5–24 cm tall; basal leaves 1–3-veined, broadly elliptic to ovate, 2–6 × 2–4 mm; cauline leaves elliptic to ovate, occasionally suborbicular, 1.25–8 × 0.5–6 mm; stamens 4, occasionally with 1–4 antepetalous, linear staminodes ca. 2 mm long; antesepalous anthers ovoid, the pore typically inclined dorsally, but sometimes emarginate to truncate apically, the connective thickened dorsally and dilated basally into a yellow, ventrally bilobed appendage that superficially appears to encircle the filament; mature capsules emarginate at the summit. Sandy, rocky places near rivers, 500–900 m; Bolívar (Ciudad Piar). Brazil (Goiás, Mato Grosso, Minas Gerais). Siphanthera duidae (Gleason) Wurdack, Fieldiana, Bot. 29: 541. 1963. —Po-

teranthera duidae Gleason, Bull. Torrey Bot. Club 58: 415. 1931. Sparingly branched or fastigiate, occasionally suffrutescent perennial herb 2.5–11 cm tall; basal leaves (when present) obscurely 1–veined, elliptic to spatulate, sometimes ovate, ca. 2 × 1 mm; stamens 4; anthers broadly ellipsoid, constricted distally into a rostrum ca. 0.1 mm long; connective prolonged 0.1–0.2 mm below the thecae, and modified into a ventrally bilobed appendage, ca. 0.75 mm long, 0.75–1 mm wide, each lobe bluntly hook-like and markedly upturned; mature capsules emarginate at the summit. Rock crevices and sandy soils, 1000–1600 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 421. Siphanthera fasciculata (Gleason) Almeda & O.R. Rob., Novon 9: 131. 1999. —Farringtonia fasciculata Gleason, Fieldiana, Bot. 28: 426. 1952. Fastigiate, sparingly branched, erect, suffrutescent, perennial herb 5–21 cm tall; basal leaves rosulate, sparsely covered with malpighiaceous trichomes to glabrate, the blades ovate-elliptic, 3–7 × 1.5–2.5 mm, the upper surface glabrous, the lower surface punctate; cauline leaves semisucculent, 1veined, ovate to ovate-lanceolate, 1.5–6 × 0.5–1 mm, with a spine-like hair apically; stamens 4; anthers tapering to a rostrum 0.75–1 mm long; connective thickened dorsally, prolonged 0.1–0.2 mm below the thecae, and modified into a ventro-basally bilobed appendage, 0.5–1 × ± 0.75 mm, each lobe bluntly hook-like and markedly upturned; mature capsules rounded at the summit. Savannas, periodically flooded areas, by rocks, sandy areas, 100–200 m; Amazonas (Caño Pimichín, base of Cerro Yapacana, Río Atabapo). Colombia (Vaupés), Brazil (Amazonas, Pará). ◆Fig. 424. Siphanthera foliosa (Naudin) Wurdack, Mem. New York Bot. Gard. 10(1): 97. 1958. —Tulasnea foliosa Naudin, Ann. Sci. Nat. Bot. sér. 3, 2: 143. 1844. —Poteranthera foliosa (Naudin) Cogn. in A. DC. & C. DC., Monogr. Phan. 7: 121. 1891. Poteranthera minor Gleason, Bull. Torrey Bot. Club 52: 339. 1925. Branched or unbranched, erect, occasionally suffrutescent herb 6–57 cm tall; leaves

Siphanthera 501

Fig. 421. Siphanthera duidae

Fig. 423. Siphanthera cordifolia

Fig. 422. Siphanthera hostmannii

Fig. 424. Siphanthera fasciculata

Fig. 425. Siphanthera foliosa

502

M ELASTOMATACEAE

mostly 1-veined, linear, 4–20 × 0.3–3.5 mm, glabrous to sparsely hirtellous; stamens 4, with 4 antepetalous staminodes present per flower; antesepalous anthers 0.75–1 × 0.5– 0.75 mm; connective thickened dorsally, prolonged ventrally at the base into a yellow, linear to globose, bilobed appendage, ± 0.25 mm long; antepetalous filament 0.75–1 mm long; staminodia 0.5–1 mm long, hatchetshaped in profile view, the reduced thecae terminated by a truncate or laterally inclined pore; mature capsules rounded at the summit. Palm savannas, ca. 100 m; Bolívar (Serranía Carichana). Colombia (Meta), Brazil, Bolivia. ◆Fig. 425. Siphanthera hostmannii Cogn. in Mart., Fl. Bras. 14(3): 199, pl. 48, fig. 2. 1883. Siphanthera paraensis Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr. 2: 510. 1898. Siphanthera capitata Gleason, Brittonia 1: 137. 1932. Siphanthera tatei Gleason, Brittonia 3:

177. 1939. Strict annual herb 2–52 cm tall, laxly branched or fastigiate; basal leaves early deciduous, glabrate to sparsely hirtellous, 1veined, broadly elliptic to elliptic-oblong, occasionally rhombic, 1–3 × 1–2 mm; cauline leaves glabrate to hirtellous, 1–3-veined, suborbicular to elliptic, occasionally rhombic, 1.5–9 × 1–6 mm; stamens 4, with 4 antepetalous linear staminodes, 1–1.75 mm long per flower; antesepalous anthers 0.5– 0.75 × ± 0.5 mm; connective prolonged basally 0.1–0.2 mm, and modified ventrally below the thecae into a bilobed appendage; mature capsules emarginate at the summit. River banks, shallow bogs, moist sandy soil, dry open sandstone, 400–1700 m; Bolívar (Arekuna, Auyán-tepui, Cerro Guaiquinima, Cerro Ichún, Gran Sabana, Macizo del Chimantá, Salto Uraima, Serranía Marutaní), Amazonas (Cerro Duida, Sierra de la Neblina). Colombia (Vaupés), Guyana, Suriname, Peru, Brazil (Amapa, Amazonas, Pará, Roraima). ◆Fig. 422.

Fig. 426. Tateanthus duidae

Tibouchina 503

45. TATEANTHUS Gleason, Bull. Torrey Bot. Club 58: 424. 1931. by Paul E. Berry Shrubs or small trees. Leaves opposite, 5- or 7-pliveined, resinous-glandular. Inflorescences terminal cymose panicles. Flowers 5-merous. Hypanthium 5-winged; calyx lobes broadly triangular, persistent, carinate; petals yellow, obovate, rounded and retuse at the apex, minutely glandular-ciliate. Stamens 10, isomorphic; filaments moderately glandular-puberulous; thecae oblong, the apex rostrate and a small ventrally inclined pore; connective not prolonged, exappendiculate. Ovary 3/4inferior, 5-locular, the apex densely granulose-glandular; style densely glandularpuberulous; stigma punctiform. Fruit an apically dehiscent capsule. Seeds numerous, narrowly fusiform, densely and finely granulate. Southern Venezuela, northern Brazil; 1 species. Tateanthus duidae Gleason, Bull. Torrey Bot. Club 58: 424, pl. 34. 1931. Shrub 0.8–5 m tall. Rocky tepui-summit ridges, edges of tepui forests, streamsides, 800–2000 m; Bolívar (Cerro Marutaní, Cerro

Sarisariñama), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina). Brazil (Amazonas: Serra Pirapucú). ◆Fig. 426.

46. TIBOUCHINA Aubl., Hist. Pl. Guiane 445. 1775. by Paul E. Berry Herbs, shrubs, or small trees; stems and foliage covered by scales or hairs. Leaves opposite or whorled, isomorphic. Flowers 5-merous (in Venezuelan species), often showy, sometimes involucrate-bracteate, usually in terminal panicles. Hypanthium terete; sepals usually persistent; petals obovate to elliptic, ciliolate, usually magenta or pink. Stamens twice as many as the petals, often dimorphic, filaments glabrous to puberulent; anthers linear-subulate to narrowly oblong, with a single pore; connective usually prolonged below the thecae, ventrally bilobed beyond the insertion of the filament and dorsally not usually appendaged. Ovary 5-locular, apex pubescent; stigma usually punctiform. Fruit a capsule, 5-locular. Seeds numerous, cochleate, minutely tuberculate. Mexico, Central America, and Antilles south to northern Argentina; ca. 350 species, 23 in Venezuela, 17 of these in the flora area. Some species of Tibouchina, such as T. catharinae and T. striphnocalyx, have branched trichomes between the calyx lobes that are similar to those that characterize the genus Pterolepis. Key to the Species of Tibouchina 1. 1.

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

Leaves small, 0.5–3(–5) cm long; upper surface of leaves apparently 1veined; restricted to tepui summits ...................................................... 2 Leaves variable in size, from smaller to much larger than above; upper surface of leaves 3- or 5-veined; from tepui summits to lowland habitats ................................................................................................................ 4 Upper surface of leaf covered with scales ........................................ T. duidae Upper surface of leaf without obvious scales ............................................ 3 Scales on lower surface of leaf with rounded apex, imbricate, and com-

504

M ELASTOMATACEAE

3.

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

9.

10(9). 10. 11(4). 11. 12(11).

12.

13(11). 13. 14(13). 14. 15(14). 15.

pletely covering the surface; only known from summit of Cerro Sipapo ................................................................................................. T. sipapoana Scales on lower surface of leaf with acute apex, not imbricate, and incompletely covering the surface; only known from summit of Cerro Jaua .............................................................................................. T. steyermarkii Hypanthium and stem internodes covered by scales ............................... 5 Hypanthium and stem internodes covered by hairs ............................... 11 Leaves mostly 7–10 × 2.5–4 cm; ventral lobes of the staminal connectives densely covered by fine hairs ................................................................ 6 Leaves smaller than above, mostly 1.5–4(–8) × 1–2(–3.5) cm; ventral lobes of the staminal connectives glabrous .................................................... 7 Scales of the hypanthium 1.5–2 mm long, dense, not appressed ... T. llanorum Scales of the hypanthium ca. 1 mm long, loose, appressed .... T. bipenicillata Leaves 5-veined; floral bracts united 1/3–2/3 their length ................. T. aspera Leaves 3-veined; floral bracts free or united as above .............................. 8 Inner floral bracts almost completely fused; lowlands of Amazonas state ................................................................................................. T. spruceana Floral bracts all free to the base; uplands and highlands of Amazonas and Bolívar states ......................................................................................... 9 Leaf blades thick-coriaceous, elliptic to suborbicular, 1–2.5 × 0.8–1.5 cm, the upper surface with a conspicuously raised pseudovein between the midvein and the lateral veins; flowers solitary .................... T. dissitiflora Leaf blades coriaceous, ovate-elliptic to lance-elliptic, (1.5–)2–4 × 1–2 cm, the upper surface without a raised pseudovein between the midvein and lateral veins; flowers paniculate or solitary ................................ 10 Flowers paniculate; calyx lobes 4–5 mm long; petals 11–14 × 6.5-7.5 mm ..................................................................................................... T. fraterna Flowers solitary; calyx lobes 5.5–6.5 mm long; petals 16 × 10-11 mm ........................................................................................................ T. huberi Intersepalar sinuses with branched-pediculate, setose appendages ..... 12 Intersepalar sinuses without pediculate appendages ............................ 13 Trichomes on hypanthium usually with a pedicel 1–1.3 mm long and a terminal tuft of many setae; calyx lobes narrowly triangular, 2–2.5 mm wide at the base; in upland to highland habitats, 600–1800 m elevation ................................................................................................. T. catharinae Trichomes on hypanthium usually lacking a pedicel and with fewer setae; calyx lobes narrowly linear-subulate, 0.6–1 mm wide at the base; in mostly lowland habitats, 50–300(–800) m elevation ........ T. striphnocalyx Calyx lobes caducous after anthesis .......................................... T. cryptadena Calyx lobes persistent after anthesis ...................................................... 14 Trichomes on hypanthium glandular-tipped; restricted to the summit of Cerro Sipapo .......................................................................... T. kunhardtii Trichomes on hypanthium not glandular-tipped; species more widely distributed ................................................................................................. 15 Inflorescence a narrow panicle of bracteate glomerules; calyx lobes 6– 8 mm long; petioles 1–5 mm long ................................................ T. gracilis Inflorescence laxly paniculate and few-flowered; calyx lobes 2.5–5 mm long; petioles 3–20 mm long ................................................................ 16

Tibouchina 505

16(15). Petals pink to purple; staminal connective prolonged 1.5–3.5 mm, the ventral lobes swollen; usually above 800 m elevation .......... T. geitneriana 16. Petals white; staminal connective prolonged 0.5 mm or less, the ventral lobes not swollen; usually below 800 m elevation ................... T. longifolia Tibouchina aspera Aubl., Hist. Pl. Guiane 446, pl. 177. 1775. Shrub 0.3–3 m; leaves 2–8 × 1–3.5 cm, 5veined, rough (asperous); panicles few-flowered; petals purple, 9–15 × 7–9 mm. Honduras, Nicaragua, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 2 varieties, both in the flora area. Key to the Varieties of T. aspera 1. Stem pubescence tightly appressed .......... ........................................... var. aspera 1. Stem pubescence lax ..... var. asperrima T. aspera var. aspera Savannas, Mauritia palm swamps, 100– 1000 m; Bolívar (Arabopó, El Paují, near Icabarú, Represa Guri area), Amazonas (Canaripó, Cerro Asisa, Cerro Yutajé, Coromoto, La Esmeralda, Puerto Ayacucho, Río Autana, Río Manapiare basin, near San Carlos de Río Negro, Santa Bárbara). Barinas, Monagas, Sucre, Trujillo, Zulia; Central America to Amazonian Bolivia and Brazil. ◆Fig. 429. T. aspera var. asperrima Cogn. in Mart., Fl. Bras. 14(3): 374. 1885. Tibouchina brachyanthera Pittier, J. Wash. Acad. Sci. 13: 386. 1923. Upland savannas, ca. 1000 m; Bolívar (Río Kukenán). Trujillo; Guyana, Suriname, Brazil. Wurdack (Phytologia 5: 131. 1955) states that there are intermediate forms between these two varieties. Tibouchina bipenicillata (Naudin) Cogn. in Mart., Fl. Bras. 14(3): 385. 1885. —Lasiandra bipenicillata Naudin, Ann. Sci. Nat. Bot. sér 3, 13: 156. 1850. —Pleroma bipenicillata (Naudin) Triana, Trans. Linn. Soc. London 28: 45. 1871. Shrub 1.5–6 m tall; leaves 6–11 × 2.5–5 cm; panicle few- to many-flowered, petals purple, 13–20 × 8–11 mm. Granitic outcrops, 100–400 m; Bolívar (Cerro San Borja along lower Río Suapure), Amazonas (areas both

north and south of Puerto Ayacucho). Barinas, Lara, Portuguesa, Táchira; Colombia. ◆Fig. 431. Tibouchina catharinae Pittier, Bol. Soc. Venez. Ci. Nat. 11: 18. 1947. Shrub 1–3 m; leaves 7–11 × 1.5–5 cm, 3veined; panicle densely flowered; hypanthium trichomes branched, petals purple, 4– 7 × 3–4 mm. Montane shrublands on both granite and sandstone, 600–1800 m; Amazonas (Cerro Autana, Cerro Cuao, Cerro Sipapo). Endemic. ◆Fig. 434. This species is very close to T. striphnocalyx and differs mainly in its larger floral bracts and flowers, with denser hypanthial pubescence. The leaves tend to be relatively larger and broader. Tibouchina cryptadena Gleason, Fieldiana, Bot. 28: 426. 1952. Shrub 1.5–4 m tall; stems quadrangular; leaves 6–15 × 1.5–5 cm, 5-veined; panicles lax and few-flowered, petals purple, 15–25 × 11–16 mm. Shrubby upper montane slopes, 1200–1900 m; Bolívar (Auyán-tepui, Kamarata, Uaipán-tepui), Amazonas (Sierra Parima). Barinas, Mérida. Táchira. ◆Fig. 430. Tibouchina dissitiflora Wurdack, Mem. New York Bot. Gard. 10(1): 101. 1958. Shrub 0.5–1.5 m tall; branches covered by flattened, inconspicuously ciliate trichomes; leaves 1–2.5 × 0.8–1.5 cm; flowers solitary, petals purple, 12–19 × 8–17 mm. Tepui shrublands, 1500–2200 m; Amazonas (Cerro Avispa, Cerro Coro Coro, Cerro Guanay, Cerro Yutajé, Sierra de la Neblina). Endemic. Tibouchina duidae Gleason, Fieldiana, Bot. 28: 428. 1952. Subshrub 0.1–1 m tall; leaves 1–2 × 0.8–1 cm, 3-veined; flowers solitary, petals purple, ca. 12 × 8 mm. Tepui shrublands, sandy streambanks, 1200–2100 m; Bolívar (Cerro Jaua), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Parú). Endemic. ◆Fig. 432.

506

M ELASTOMATACEAE

Tibouchina fraterna N.E. Br., Trans. Linn. Soc. Bot. 6: 28. 1901. Shrub 0.5–4 m tall; leaves 2–3 × 1–2 cm, 3-veined, with 4–8 irregular long rows of fine scale tips between veins; panicle few-flowered, petals purple, 11–14 × 6.5–7.5 mm. Venezuela, Guyana; 2 subspecies, both in the flora area. Key to the Subspecies of T. fraterna 1. Hypanthium scales ca. 1 mm long; calyx lobes 1.5–2 mm wide, shortly ciliate and with short scales ........ subsp. fraterna 1. Hypanthium scales 2–5 mm long; calyx lobes 1 mm wide, long-ciliate and with long scales ................. subsp. paruana T. fraterna subsp. fraterna Slopes by waterfalls, shrub islands, swampy areas, 600–2000 m; Bolívar (Auyántepui, Cerro Venamo, Murisipán-tepui, Ptaritepui, Roraima-tepui, Uei-tepui, Uaramatepui in northeastern Gran Sabana), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Guyana. ◆Fig. 435. T. fraterna subsp. paruana Wurdack, Mem. New York Bot. Gard. 10(1): 101. 1958. 1100–2000 m; Amazonas (Cerro Parú). Endemic. Tibouchina geitneriana (Schltdl.) Cogn. in DC., Monogr. Phan. 7: 255. 1891. —Chaetogastra geitneriana Schltdl., Linnaea 27: 523. 1856. Tibouchina pseudomollis Gleason, Bull. Torrey Bot. Club 56: 400. 1929. Tibouchina impressa Gleason, Fieldiana, Bot. 28: 428. 1952. Shrub 0.3–1.5 m tall; leaves 2.5–5.5 × 0.6–1.8 cm; inflorescence lax and few-flowered; petals pink to purple, 9–12 × 4–5 mm. Cliff bases, rocky shrub islands on tepuis, 800–2300 m; Bolívar (Auyán-tepui, Cerro Venamo, Macizo del Chimantá [Toronótepui], Perai-tepui west of Santa Elena de Uairén, Roraima-tepui, Sororopán-tepui), Amazonas (Cerro Guanay, Cerro Huachamacari, Cerro Yaví). Widespread in northern Venezuela; Guyana. Tibouchina gracilis (Bonpl.) Cogn. in Mart., Fl. Bras. 14(3): 386. 1885.

—Rhexia gracilis Bonpl., Rhex. 138, pl. 52. 1823. Subshrub 0.3–1.5 m tall, usually singlestemmed, with long internodes and shaggy, rufous trichomes on leaves and stems; leaves 3–12 × 0.7–4 cm, 5-veined; inflorescence a narrow panicle of bracteate glomerules, petals purple, 15–25 × 10–15 mm. Mauritia palm swamps, grassy savannas, 300–1300 m; Bolívar (Ciudad Piar, near Parupa, San Ignacio de Yuruaní). Anzoátegui, Mérida, Monagas, Zulia; Colombia to northern Argentina. ◆Fig. 437. Tibouchina huberi Wurdack, Phytologia 69: 318. 1990. Small shrubs 0.3–1 m tall; leaves 1.5– 2.5(–4.5) × 1–2 cm; flowers usually solitary; petals purple, 15–16 × 10–11 mm long. Tepui shrublands, on rocks, 1400–2200 m; Bolivar (Sierra de la Maigualida), Amazonas (Sierra de Maigualida, Cerro Ualipano). Endemic. Tibouchina kunhardtii Gleason, Phytologia 3: 242. 1950. Shrub 1–2 m tall; leaves coriaceous, 4.5– 10 × 1.5–3 cm, subsessile; panicles broad and loosely flowered, flowers with loose recurved, gland-tipped hypanthial scales; petals purple. Bogs, streambanks, and scrub on tepui slopes and summit, 1000–1800 m; Amazonas (Cerro Cuao, Cerro Sipapo). Endemic. ◆Fig. 436. Tibouchina llanorum Wurdack, Mem. New York Bot. Gard. 10(5): 142. 1964. Shrub 1–1.5 m tall; stems 4–6-angled; leaves 7–12 × 4–6 cm, 5-veined; narrow panicle of compact glomerules, 4–20 cm long; hypanthium scales dense-scurfy, 1.5–2 mm long, not appressed; petals purple, 13–15 × 11–13 mm. Savannas, Mauritia palm swamps, 50—200 m; Bolívar (Río Cuchivero, Río Parguaza). Carabobo, Cojedes, Lara, Portuguesa; Colombia (Meta), Brazil (Amazonas, Pará), Bolivia (Beni, La Paz, Santa Cruz). ◆Fig. 438. Tibouchina longifolia (Vahl) Baill., Adansonia 12: 74. 1877. —Rhexia longifolia Vahl, Eclog. Amer. 1: 39. 1796 [1797]. Subshrub 0.3–1 m tall; leaves 3.5–10 × 1– 3.5 cm; inflorescence lax and few-flowered, petals white, 5–7 × 3–4 mm. Lower montane

Tibouchina 507

Fig. 427. Tibouchina sipapoana

Fig. 429. Tibouchina aspera var. aspera

Fig. 428. Tibouchina spruceana

508

M ELASTOMATACEAE

Fig. 430. Tibouchina cryptadena

Fig. 431. Tibouchina bipenicillata

Tibouchina 509

Fig. 432. Tibouchina duidae

Fig. 433. Tibouchina steyermarkii

Fig. 434. Tibouchina catharinae

510

M ELASTOMATACEAE

Fig. 435. Tibouchina fraterna subsp. fraterna

Fig. 436. Tibouchina kunhardtii

Tibouchina 511

Fig. 437. Tibouchina gracilis

Fig. 438. Tibouchina llanorum

512

M ELASTOMATACEAE

forests and openings, 300–900 m; Bolívar (Perai-tepui west of Santa Elena de Uairén, Santa Maria de Erebato). Widespread throughout northern Venezuela; Mexico, West Indies, Colombia to Bolivia. Tibouchina sipapoana Gleason, Phytologia 3: 240. 1950. Weak shrub to 50 cm tall; leaves 1–2(–5) × 0.5–0.8(–1.2) cm, 1-veined; flowers solitary, petals purple, ca. 17 × 13 mm. Open tepui summit, ca. 1900 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 427. Tibouchina spruceana Cogn. in Mart., Fl. Bras. 14(3): 376, pl. 86, fig. 2. 1886. Shrub 0.5–2.5 m tall; leaves 2–5 × 0.7–2 cm, 3-veined; flowers solitary or ternate at branch tips; petals purple, 15–20 × 8–15 mm. Savannas, sandy streambanks, 50—200 m; Bolívar (expected), Amazonas (base of Cerro Yapacana, La Esmeralda, along Río Orinoco, San Juan de Ucata). Apure; Colombia, Brazil, Bolivia. ◆Fig. 428. Tibouchina steyermarkii Wurdack, Mem. New York Bot. Gard. 23: 873. 1972.

Shrub 0.5–1 m tall; leaves 0.7–1.5 × 0.3– 0.7 cm, 1-veined; flowers terminal on short branchlets, petals purple, ca. 12 mm long. Tepui shrublands, streamsides, 1700–2100 m; Bolívar (Cerro Jaua). Endemic. ◆Fig. 433. Tibouchina striphnocalyx (DC.) Gleason, Phytologia 3: 241. 1950. ––Osbeckia striphnocalyx DC., Prodr. 3: 140. 1828. —Pterolepis striphnocalyx (DC.) Cogn. in Mart., Fl. Bras. 14(3): 286. 1885. Tibouchina yavitensis Pittier, Bol. Soc. Venez. Ci. Nat. 11: 19. 1947. Shrub 1–3 m tall; leaves 4–12 × 1–5 cm; panicle many-flowered, compact, calyx and hypanthium with prominent long-branched trichomes; petals purple, 4–6 × 2.5–3 mm. Shrubby white-sand savannas, moist depressions, 50–300(–800) m; Amazonas (Caño Cotúa near base of Cerro Yapacana, Maroa, Río Atabapo basin, Salto Huá along Brazilian border, San Carlos de Rio Negro, near San Juan de Ucata, Yavita). Southeastern Colombia, Brazil (Amazonas).

47. TOCOCA Aubl., Hist. Pl. Guiane 437. 1775. Truncaria DC., Prodr. 3: 106. 1828. Myrmidone Mart., Nov. Gen. Sp. Pl. 3: 149. 1842. Microphysa Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 99. 1851, non Schrenk 1844. Sphaerogyne Naudin, Ann. Sci. Nat. Bot. sér. 3, 15: 331. 1851. Hormocalyx Gleason, Phytologia 1: 174. 1935. by Fabián A. Michelangeli Shrubs or small trees, rarely vines. Leaves opposite, often with a swollen ant domatium at the base of the blade or the apex of the petiole, blades isomorphic or differently sized, sometimes a domatium on only one of each leaf pair. Young stems and petioles glabrous or with long glandular trichomes. Inflorescence a terminal panicle or raceme, or a pseudoterminal cyme or panicle. Flowers 4(5)- or 6-merous. Hypanthium terete, occasionally winged; calyx lobed, the outer teeth truncate or developed, the inner teeth truncate, rarely developed; petals usually white or pink, occasionally red, yellow, or brown, emarginate at the apex, glabrous or pruinose. Stamens isomorphic (in the flora area), glabrous; anthers yellow-white or blue, stout, with a single apical pore oriented ventrally, upright, or dorsally, connective not prolonged except for a dorso-basal minute tooth (rarely absent). Ovary (3–)5-locular, 1/3to completely inferior, apex conical to rounded, with or without a corona of glandular setae; stigma usually expanded (funnelform or capitate), rarely truncate, glabrous, rarely glandular-puberulous. Fruit baccate, blue or black at maturity, usually with many small pyramidal or oblong seeds. Southern Mexico, Central America, Colombia, Venezuela, Trinidad, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 45 species, 26 in Venezuela, 23 of these in the flora area.

Tococa 513

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

4. 5(4). 5. 6(3). 6. 7(6). 7. 8(7). 8. 9(6). 9. 10(9). 10. 11(9). 11. 12(1). 12.

13(12).

13.

14(13). 14.

Leaves without ant domatia ...................................................................... 2 Leaves with ant domatia ......................................................................... 12 Petals 15–21 mm long ...................................................................... T. obovata Petals 4–12 mm long .................................................................................. 3 Calyx outer teeth conspicuous and projecting vertically above the inner teeth ....................................................................................................... 4 Calyx outer teeth inconspicuous or barely projecting horizontally ......... 6 Leaves 10–25 cm long; petioles 10–55 mm long; calyx outer teeth stout and projecting vertically; anthers ca. 7 mm long; stigma extended or slightly capitate ....................................................................... T. subciliata Leaves 1.5–6 cm long; petioles 0.2–0.3 mm long; calyx outer teeth flat; anthers 3.5–5.5 mm long; stigma truncate ........................................... 5 Leaves 1.5–3 cm long; petals pink; apex of the ovary rounded to conical and with a corona of setae ............................................................. T. ciliata Leaves 3–9 cm long; petals white; apex of the ovary truncate and glabrous .......................................................................................................... T. liesneri Stigma truncate ......................................................................................... 7 Stigma expanded or capitate ..................................................................... 9 Leaves 6–15 cm long, lanceolate to elliptic, apex acute; anther pore dorsally inclined .................................................................................. T. nitens Leaves 2–6 cm long, broadly elliptic to obovate, apex obtuse to round; anther pore ventrally inclined ................................................................... 8 Leaves 3-veined; flowers on thin pedicels 1–3 mm long .......... T. bolivarensis Leaves 5-veined; flowers sessile or on short (to 0.5 mm long) and thick pedicels ...................................................................................... T. oligantha Vine or small shrub; anther pore dorsally inclined ................................ 10 Tree or tall shrub; anther pore ventrally inclined .................................. 11 Vine .......................................................................................... T. caryophyllea Shrub ....................................................................................... T. cinnamomea Leaves 5- or 7-pliveined, lower surface green, 9–22 cm wide; petioles 6–17 cm long; hypanthium 6.5–8 mm long ............................. T. tepuiensis Leaves basely veined, lower surface crimson to red, 2–8 cm wide; petioles 1–4 cm long; hypanthium 4–5 mm long ............................ T. erythrophylla Ant domatia free from the leaf blade and at least twice as wide as the constriction between the ant domatia and the leaf blade ........................ 13 Ant domatia totally or partially impressed in the base of the leaf blade, if free from the leaf, the constriction between the ant domatia and the blade at least 2/3 of the width of the ant domatia ................................ 17 Ant domatia with an apical ascending lobe on each side; inflorescence terminal, but quickly becoming lateral as the lateral branch outgrows it; outer teeth of the calyx projecting 3–5 mm above the torus; petals white .................................................................................... T. macrophysca Ant domatia round or obovate, without ascending lobes; inflorescence always terminal; outer teeth of the calyx not or barely projecting beyond the torus (to 1.5 mm); petals white, pink, or yellow ........................... 14 Petals yellow; apex of ovary without a corona around the union with the style, glabrous ............................................................................. T. aristata Petals pink to white; apex of the ovary with a corona around the union

514

M ELASTOMATACEAE

with the style, usually with short setae .............................................. 15 15(14). Leaf trichomes 0.3–0.6 mm long; hypanthium with 5 small puberulous ridges ....................................................................................... T. pauciflora 15. Leaf trichomes 2–20 mm long; hypanthium terete ................................. 16 16(15). Inflorescence a single-branched panicle ........................................ T. bullifera 16. Inflorescence a multiple-branched panicle ................................ T. guianensis 17(12). Flowers 6-merous; the outer sepals fused into a skirt that surrounds the hypanthium ........................................................................ T. macrosperma 17. Flowers 5-merous; the outer sepals not fused into a skirt that surrounds the hypanthium ................................................................................... 18 18(17). Ant domatia present in some of the leaves, usually only one leaf of each pair, and when present > 1/3 of the total length of the leaf ................. 19 18. Ant domatia present in most of the leaves, usually in both leaves of each pair and never more than 1/4 of the total length of the leaf ................ 20 19(18). Leaves ovate, apex acute to obtuse, with short stellate trichomes on the veins of both surfaces; inflorescence a terminal panicle with 5 or more flowers; hypanthium glabrous or with a few stellate hairs but without clavate glands; calyx with the outer teeth projecting upward 0.5–1 mm above the inner teeth; petals white with a subapical glandular setae; anther pore ventrally inclined; ovary apex conical with a corona of glandular trichomes, style puberulous at the base ...................... T. ciliata 19. Leaves oblong to rounded, apex obtuse to round, with long glandular trichomes on the upper surface; inflorescence a terminal or pseudoterminal cyme with 3–5 flowers; hypanthium with stellate hairs and many minute clavate glands; calyx with the outer teeth laterally compressed and projecting downward along the hypanthium, giving the appearance of a winged hypanthium, with many long glandular trichomes; petals red to pink, glabrous; anther pores dorsally inclined; ovary apex truncate and with a few clavate glands, style glabrous ............................................................................................... T. rotundifolia 20(18). Anthers 2.5–4 mm long ............................................................................ 21 20. Anthers 5.5–8 mm long ............................................................................ 22 21(20). Inflorescence a cyme, flowers on pedicels 6–11 mm long; calyx outer teeth free, projecting vertically, but not beyond the broad and flat inner teeth, 4–5 mm long; petals white; anther connective with a small dorsobasal tooth; stigma truncate, ovary 3-locular, the apex truncate and glabrous ............................................................................................ T. hirta 21. Inflorescence a raceme, with the central pedicel thickened, flowers sessile; calyx outer teeth connate and with two apical setae projecting vertically, inner teeth 1–1.5 mm long; petals orange to brownish; anther connective without a small dorso-basal tooth; stigma expanded, ovary 4-locular, the apex with a ring around the base of the style ............................................................................................ T. pachystachya 22(20). Leaves lanceolate, the ant domatia ≥ 1/2 the total width of the leaf ................................................................................................... T. lancifolia 22. Leaves elliptic to ovate, the ant domatia < 1/4 the width of the leaf ....... 23 23(22). Young inflorescences with bracts to 3 cm long; calyx outer teeth flat, not or barely projecting vertically; style glabrous ................................. T. cordata 23. Young inflorescences without bracts; calyx outer teeth projecting vertically above the inner teeth; style pubescent ............................. T. coronata

Tococa 515

Tococa aristata Benth., J. Bot. (Hooker) 2: 305. 1840. Shrub 1–3 m tall; leaves elliptic to broadly elliptic, 10–25 × 6–15 cm; ant domatia 1/3–1/5 impressed in the leaf; flowers orange to brownish yellow. Evergreen lowland to montane forests, 400–1600 m; Bolívar (south of Caicara, El Paují, Icabarú, Kavanayén, Macizo del Chimantá, Ptari-tepui, Sierra de Lema, Urimán). Guyana, Brazil. ◆Fig. 440. Tococa bolivarensis Gleason, Fieldiana, Bot. 28: 438. 1952. Shrub 0.7–2 m tall; flowers pale red to pink. Shrublands on tepui summits, 1600– 2400 m. Venezuela; 2 subspecies, both endemic. Key to the Subspecies of T. bolivarensis 1. Branches, both surfaces of leaves, and hypanthium with short glandular trichomes ................. subsp. bolivarensis 1. Branches, leaves, and hypanthium glabrous ..................... subsp. occidentalis T. bolivarensis subsp. bolivarensis Bolívar (Auyán-tepui, Carrao-tepui, Karaurín-tepui, Sierra de Lema, Sierra de Maigualida, Uei-tepui). Endemic. ◆Fig. 441.

caryophyllea DC., Prodr. 3: 106. 1828. —Miconia caryophyllea (DC.) Triana, Trans. Linn. Soc. London 28: 106. 1871. Vine 6–15 m long, with a few adventitious roots originating in the internodes; mature inflorescences pendent. Evergreen forests, usually near small creeks, 100–300 m; Amazonas (Río Negro). Peru, Brazil. Tococa caryophyllea is the only vine in the genus. Many of the specimens of T. caryophyllea have been incorrectly assigned to T. cinnamomea, a shrubby species that has similar flowers. Tococa ciliata Triana, Trans. Linn. Soc. London 28: 133. 1871. Shrub 0.5–3.5 m tall; leaves 1.5–6 cm, pale green, border ciliolate, occasionally with ant domatia. White-sand savannas, Mauritia palm swamps, 100–300 m; Amazonas (Caño Caname, Caño Cotúa near base of Cerro Yapacana, Río Pasimoni). Brazil (upper Río Negro). ◆Fig. 443. Although Tococa ciliata has not yet been collected in adjacent Colombia, it surely occurs there.

T. bolivarensis subsp. occidentalis Wurdack, Mem. New York Bot. Gard. 10(4): 45. 1961. Bolívar (southwest of Entrerios), Amazonas (Cerro Guanay, Cerro Yaví, Cerro Yutajé, Río Asita). Endemic. The differences in the two subspecies, along with their distribution, may warrant the recognition of these two taxa as separate species.

Tococa cinnamomea Triana, Trans. Linn. Soc. London 28: 133. 1871. Shrub 0.7–2.5 m tall; petals pink. Evergreen lowland forests, usually near small creeks, 100–300 m; Amazonas (Río Negro). Colombia, Brazil. Tococa cinnamomea is known in Venezuela only from the type collection. Specimens of T. caryophyllea are usually incorrectly determined as this species because the flowers and leaves are very similar. However, they differ in habit, T. caryophyllea growing as a vine, and T. cinnamomea as a shrub.

Tococa bullifera Mart. & Schrank ex DC., Prodr. 3: 165. 1828. Tococa spruceana Cogn. in Mart., Fl. Bras. 14(4): 443. 1888. Tococa ulei Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 177. 1905. Shrub 0.5–2 m tall; inflorescence a terminal single-branched panicle; ovary apex with a corona of setae. Evergreen lowland forests, 50–200 m; Amazonas (mouth of Río Matacuni, Río Negro). Peru, Brazil. See note under Tococa guianensis.

Tococa cordata O. Berg ex Triana, Trans. Linn. Soc. London 28: 133. 1871. Shrub 0.7–3 m tall; ant domatia completely impressed in the blade. Seasonally flooded and disturbed forests, 50–200 m; Amazonas (Piedra Cocuy, Río Casiquiare, San Carlos de Río Negro). Colombia?, Brazil. Vegetatively, Tococa cordata is similar to T. coronata, but with thinner and deeply cordate leaves. The floral characters are sufficient to distinguish them as two different species.

Tococa caryophyllea (DC.) S.S. Renner, Biollania 6: 497. 1997. —Truncaria

Tococa coronata Benth., J. Bot. (Hooker) 2: 303. 1840.

516

M ELASTOMATACEAE

Tococa subnuda Benth., J. Bot. (Hooker) 2: 303. 1840. Tococa castrata Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 93. 1851. Tococa egensis Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 92. 1851. Tococa longisepala Cogn. in Mart., Fl. Bras. 14(4): 449. 1888. Tococa traillii Cogn. in Mart., Fl. Bras. 14(4): 626. 1888. Tococa glandulosa Gleason, Bull. Torrey. Bot. Club. 53: 252. 1931. Shrub 2–6 m tall; leaves lanceolate to elliptic, 10–30 × 6–10 cm; petals pink; fruits blue. Riparian and seasonally flooded forests, 50–200 m; Bolívar (Río Horeda near Ciudad Bolívar, Río Parguaza), Amazonas (Río Baría, Río Casiquiare, Río Cataniapo, Río Cunucunuma, Río Manapiare, Río Negro, Río Orinoco, Río Temi, Río Ventuari). Anzoátegui, Apure, Guárico; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 445. Due to the large amount of variability among different specimens, it seems more appropriate to group Tococa egensis, T. subnuda, and T. longisepala under one species as suggested by Wurdack [Melastomataceae, in Lasser, Flora de Venezuela, 8(1): 1–737. 1973]. Tococa coronata is the oldest name in the complex, and therefore is the one here applied, even though in some herbaria T. egensis has been more widely used. Tococa erythrophylla (Ule) Wurdack, Mem. New York Bot. Gard. 10(5): 176. 1964. —Miconia erythrophylla Ule, Notizbl. Königl. Bot. Gart. Berlin 6: 357. 1915. Small to medium-sized tree 1–7 m tall; lower surface of leaves red to dark purple, glabrous; petals white. Tepui slope and summit forests, 1100–2300 m; Bolívar (Auyántepui, Cerro Jaua, Macizo del Chimantá [Acopán-tepui, Churí-tepui], Roraima-tepui), Amazonas (Cerro Marahuaka). Guyana. ◆Fig. 444. Tococa guianensis Aubl., Hist. Pl. Guiane 437, pl 174. 1771. Melastoma physophora Vahl, Eclog. Amer. 1: 45. 1796 [1797]. Melastoma tococo Desr. in Lam., Encycl. 4: 39. 1797. Tococa aubletii D. Don, Mem. Wern. Nat. Hist. Soc. 4: 304. 1823.

Tococa roreimi Benth., J. Bot. (Hooker) 2: 305. 1840. Tococa acuminata Benth., Bot. Voy. Sulphur 94. 1844. Tococa cardiophylla Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 93. 1851. Tococa didymophysca Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 90. 1851. Tococa occidentalis Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 92. 1851. Tococa erythrostachya Spruce ex Triana, Trans. Linn. Soc. London 28: 132. 1871. Tococa coriaceae S. Moore, J. Bot. 18: 3. 1880. Tococa ferruginea G. Nicholson, Dict. Gard. 4: 49. 1887. Miconia erioneura Cogn., Mem. Torrey Bot. Club 6: 38. 1896. —Tococa erioneura (Cogn.) Wurdack, Mem. New York Bot. Gard. 10(5): 176. 1964. Tococa discolor Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 176. 1905. Tococa juruensis Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 176. 1905. Tococa loretensis Ule, Notizbl. Königl. Bot. Gart. Berlin. 6: 365. 1915. Maieta glandulifera Standl., Proc. Biol. Soc. Wash. 37: 52. 1924. Shrub 1–5 m tall; petals pink; ovary with an apical corona of short setae; mature fruits black. Common in gaps and borders of lowland forests, 50–1600 m; widespread in southern Delta Amacuro and throughout Bolívar and Amazonas. Anzoátegui, Apure, Barinas, Guárico, Mérida, Táchira, Trujillo, Zulia; widespread from Southern Mexico to the Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 447. Species considered here as synonyms have been traditionally considered by other authors as a complex of closely related species. However, the large amount of continuous variation, both within and across populations, makes it impossible to give species status to most of the names recognized in this complex. Variation in the pubescence of the leaves and hypanthium, setae in the calyx teeth, and the size and shape of the ant domatia can give very different aspect to specimens at the extreme of the variation. Perhaps the only species clearly distinguishable from Tococa guianensis is T. bullifera, due to its unbranched inflorescences. This complex is in need of careful revision. The

Tococa 517

list of synonyms presented here only contains names applied to specimens from the Venezuelan Guayana or neighboring regions and does not pretend to be complete. Tococa hirta O. Berg ex Triana, Trans. Linn. Soc. London 28: 132. 1871. Shrub 0.5–2 m tall; ant domatia round and partially impressed in the leaf blade; leaves elliptic with an acute apex, 5–13 × 3–8 cm, with abundant nonglandular trichomes to 3 mm long. White-sand savannas, Río Negro caatinga, 50–200 m; Amazonas (Maroa, Pimichín, San Carlos de Río Negro, Yavita). Colombia, Brazil. Tococa lancifolia Spruce ex Triana, Trans. Linn. Soc. London 28: 133. 1871. Shrub 0.7–2 m tall; leaves lanceolate, 12– 25 × 3–5 cm. Seasonally flooded forests, especially along black-water streams or rivers, 50–200 m; Amazonas (base of Cerro Yapacana, south of Puerto Ayacucho, Río Atabapo, Río Baría, Río Casiquiare, Río Orinoco, Río Pasimoni, San Carlos de Río Negro, Tamatama). Endemic (but most likely in adjacent Colombia). ◆Fig. 448. Tococa liesneri Wurdack, Ann. Missouri Bot. Gard. 76: 961. 1989. Shrub 0.4–0.7 m tall; leaves ovate-oblong, 3-veined, 1.5–2.5 × 0.8–1.5 cm; petals pink. Tepui meadows, dwarf forests on tepui summits, 1400–1700 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Tococa macrophysca Spruce ex Triana, Trans. Linn. Soc. London 28: 132. 1871. Shrub 0.5–4 m tall; ant domatia shiny, with apical, ascending lobes; petals white. Evergreen lowland forests, sometimes seasonally flooded forests, Río Negro caatinga, 50–300 m; Amazonas (Capibara, base of Cerro Sipapo, base of Cerro Yapacana, Maroa, San Carlos de Río Negro, San Juan de Puruname, base of Sierra de la Neblina, Yavita). Colombia, Brazil. ◆Fig. 449. Tococa macrosperma Mart., Nov. Gen. Sp. Pl. 3: 148. 1832. —Myrmidone macrosperma (Mart.) Mart., Nov. Gen. Sp. Pl. 3: 149. 1832. Myrmidone lanceolata Cogn. in Mart., Fl. Bras. 14(4): 468, pl. 98, fig. 2. 1888.

Hormocalyx hirsutus Gleason, Phytologia 1: 174. 1935. Shrub 0.3–3 m tall; leaves anisophyllous, the larger leaves elliptic-oblong, 8–22 × 4–10 cm, apex acuminate, base cordate, the smaller leaves ovate to oblong, 2–6 × 1–3.5 cm; petals red, turning white after anthesis. Evergreen lowland to montane forests, 100– 1300 m; Bolívar (Canaima, Cerro Guaiquinima, Cerro Ichún, Cerro Venado, Icabarú, Río Aonda on Auyán-tepui, Río Caroní, Río Carrao, Río Churún near Guarimba, Río Las Ahallas near El Paují, Santa Maria de Erebato), Amazonas (Caño Caname, Cerro Aracamuni, Cerro Autana, Cerro Huachamacari, Piedra Arauicaua, Piedra Cocuy, Río Cunucunuma, Río Mawarinuma, Río Negro, Río Puruname, San Fernando de Atabapo, Sierra de la Neblina). Colombia, Guyana, Peru, Brazil, Bolivia. ◆Fig. 439. Tococa macrosperma was originally described by Martius in the genus Tococa, and on the following page segregated into Myrmidone on the basis of its unique calyx and large seeds. Several researchers have agreed that Myrmidone is closely related to Tococa and even that it is solely a segregate. Wurdack (personal communication) kept Myrmidone as a separate genus not only on the basis of the calyx and seeds, but also due to the lack of the dorsal tooth in the anther connective. However, T. macrosperma is polymorphic for this character, and therefore should not be used as a diagnostic character, or to keep the two genera separate. The differences between Myrmidone lanceolata and T. macrosperma involve characters that are highly variable across populations, and do not warrant the recognition of the former taxon as a separate species or even at the infraspecific level. Tococa macrosperma is often confused with species of Maieta. Even though these two taxa are similar, they can be easily distinguished based on the floral characters given above. Additionally, the anthers of Maieta are incurved, with the thecae extending basally beyond the point of attachment with the connective. Vegetatively, they can be distinguished by the base of leaves and the shape of the ant domatia. In T. macrosperma, the leaves are cordate and the ant domatia are scrotiform, while in Maieta the leaves are attenuate at the base with the ant domatia

518

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narrowly elliptic to fusiform and completely impressed in the leaf blade. Tococa nitens (Benth.) Triana, Trans. Linn. Soc. London 28: 133. 1871. —Miconia nitens Benth., J. Bot. (Hooker) 2: 313. 1840. Tococa wedellii Naudin, Ann. Sci. Nat. Bot. sér. 3, 16: 94. 1851. Shrub 0.5–2 m tall; leaves broadly elliptic, 5–14 × 3–6 cm, the upper surface shiny, the margin ciliolate to entire; petals pink. Whitesand savannas, savanna-forest ecotones, 100–1600 m; Bolívar (Canaima, Guayaraca, Icabarú, base of Ilú-tepui, Kamarata, Kavanayén, Luepa, Río Aponguao, Río Carrao, Río Parú), Amazonas (Guarinuma, La Esmeralda). Guyana, Brazil. ◆Fig. 442. Tococa obovata Gleason, Bull. Torrey Bot. Club 58: 429. 1931. Shrub 0.3–2 m tall; petals pinkish to red, 15–21 mm long. Tepui summit scrub, 1400– 2200 m. Venezuela; 2 subspecies, both endemic. Key to the Subspecies of T. obovata 1. Upper surface of leaves blistered, lower surface foveolate ... subsp. neblinensis 1. Upper and lower surface of leaves smooth .................................... subsp. obovata T. obovata subsp. neblinensis Wurdack, Mem. New York Bot. Gard. 10(5): 175. 1964. Ant domatia occasionally developed; if so, impressed in the leaf blade, to 0.5–1.2 cm long. Amazonas (Sierra de la Neblina). Endemic. T. obovata subsp. obovata Tococa montana Gleason, Bull. Torrey Bot. Club 58: 430. 1931. Ant domatia always absent. Bolívar (Cerro Jaua), Amazonas (Cerro Duida). Endemic. ◆Fig. 453. Tococa oligantha Gleason, Bull. Torrey Bot. Club 58: 431. 1931. Miconia setimarginata Pittier, Bol. Soc. Venez. Ci. Nat. 11: 27. 1947. Shrub 0.5–1.5 m tall; leaves oblong to ovate, 3–8 × 2.5–6 cm; petals pink. Forests

and open areas on the summits and slopes of tepuis, 1300–2200 m; Bolívar (Cerro Jaua), Amazonas (Cerro Coro Coro, Cerro Duida, Cerro Guanacoco, Cerro Guanay, Cerro Parú, Cerro Sipapo, Cerro Yutajé, Sierra de Parima). Brazil (Amazonas: Serra la Neblina). ◆Fig. 451. Tococa pachystachya Wurdack, Mem. New York Bot. Gard. 10(4): 43, fig. 33C– G. 1961. Shrub 0.3–3 m tall; ant domatia 2/3 to totally impressed in the leaf blade, with short and thick trichomes. Tepui summit and slope forests, 1400–2100 m; Amazonas (Cerro Aracamuni, Sierra de la Neblina). Brazil (Amazonas: Serra la Neblina). ◆Fig. 450. The status of Tococa pachystachya in the genus Tococa is questionable, since some characters in the calyx and the ant domatia suggest that this species might be more closely related to Miconia than to some species of Tococa. Tococa pauciflora Spruce ex Triana, Trans. Linn. Soc. London 28: 131. 1871. Shrub 2.5–3 m tall; leaves lanceolate to oblong, apex acute, 12–25 × 5–12 cm; ant domatia free from the leaf, to 4 cm long; calyx outer teeth projecting vertically, with an apical glandular trichome. Evergreen lowland forests, 50–200 m; Amazonas (Río Guainía). Colombia. Tococa pauciflora is known only from two collections. Its affinities are unclear, since it shares characters with both the T. coronata group and some Peruvian species with winged calyces. Tococa rotundifolia (Triana) Wurdack, Phytologia 19: 193. 1969. —Microphysa rotundifolia Triana, Trans. Linn. Soc. London 28: 141. 1871, “Microphysca.” Small shrub 0.2–1.2 m tall; leaf and stem trichomes terminating in a gland 4–18 mm long, red or fuchsia; hypanthium red, with the calyx outer teeth projecting downward, forming 5 wings on the surface of the hypanthium; petals red. Río Negro caatinga, white-sand scrub (bana), 50–200 m; Amazonas (Caño Cotúa near base of Cerro Yapacana, Maroa, Pimichín, San Carlos de Río Negro to Solano). Colombia, Brazil. ◆Fig. 446.

Tococa 519

Tococa subciliata (DC.) Triana, Trans. Linn. Soc. London 28: 133. 1871. —Miconia subciliata DC., Prodr. 3: 187. 1828. Tococa planifolia Benth., J. Bot. (Hooker) 2: 305. 1840. Tococa lasiostyla Cogn. in Mart., Fl. Bras. 14(4): 455, pl. 97, fig. 1. 1888. Shrub 0.5–2.5 m tall; leaves lanceolate to oblong-lanceolate, 12–27 × 3–8 cm, apex acute, base round to cordate; calyx outer teeth stout, projecting upward beyond the inner teeth; petals pink to white. Seasonally flooded riparian forests, usually in the lower section of the river bank, 50–200 m; Amazonas (Brazo Casiquiare, Caño Pimichín, Río Guainía, Río Negro, Río Pasiba, Río Pasimoni). Colombia, Peru, Brazil. ◆Fig. 452. Tococa subciliata is very closely related to T. coronata and T. lancifolia (as well as to other species outside the flora area), but it lacks the ant domatia present in T. lancifolia, possibly due to the seasonal flooding it must endure.

Tococa tepuiensis Wurdack, Mem. New York Bot. Gard. 10(5): 176. 1964. Tree 4–14 m tall; hypanthium glabrous, ca. 7 mm long; petals white to pink. Tepuislope forests, 1100–2100 m. Venezuela, Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of T. tepuiensis 1. Leaves glabrous, apex obtuse ................... ..................................... subsp. glabrata 1. Leaves pubescent, with long glandular trichomes, apex acute .............................. ................................. subsp. tepuiensis T. tepuiensis subsp. glabrata Wurdack, Mem. New York Bot. Gard. 10(5): 177. 1964. Amazonas (Sierra de la Neblina, Sierra Parima). Brazil. T. tepuiensis subsp. tepuiensis Bolívar (Aparamán-tepui, Macizo del Chimantá [Toronó-tepui], Río Asisa). Endemic. ◆Fig. 454.

Fig. 439. Tococa macrosperma

520

M ELASTOMATACEAE

Fig. 440. Tococa aristata

Fig. 441. Tococa bolivarensis subsp. bolivarensis

Fig. 442. Tococa nitens

Tococa 521

Fig. 443. Tococa ciliata

Fig. 444. Tococa erythrophylla

Fig. 445. Tococa coronata

522

M ELASTOMATACEAE

Fig. 446. Tococa rotundifolia

Fig. 447. Tococa guianensis

Tococa 523

Fig. 448. Tococa lancifolia

Fig. 449. Tococa macrophysca

524

M ELASTOMATACEAE

Fig. 450. Tococa pachystachya

Fig. 452. Tococa subciliata

Fig. 451. Tococa oligantha

Tococa 525

Fig. 453. Tococa obovata subsp. obovata

Fig. 454. Tococa tepuiensis subsp. tepuiensis

526

M ELASTOMATACEAE

48. TOPOBEA Aubl., Hist. Pl. Guiane 476. 1775. by Paul E. Berry Shrubs or trees, sometimes vining, often epiphytic. Flowers 6-merous, axillary and pedicellate from upper leafy branchlet nodes, 4(6?)-bracteate. Hypanthium terete; calyx ± 6-lobed, usually persistent; petals pink (in Venezuela), minutely retrorse-ciliolate but otherwise glabrous. Stamens 12, isomorphic, glabrous; anthers usually laterally coherent, narrowly oblong, with a dorso-apical pore; connective usually with a dorso-basal spur. Ovary 4–6-locular and at least partly inferior; style glabrous. Fruit a berry. Seeds numerous. Southern Mexico, Central America, Colombia, Guyana?, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia; ca. 50 species, 3 in Venezuela, 1 of these in the flora area. Topobea ferruginea Gleason, Bull. Torrey Bot. Club 58: 434. 1931. Well-branched tree 5–15(–30) m tall; stems 4-angled; leaf rigid, 3-veined, blades elliptic-oblong, apex obtuse to rounded, base broadly acute to obtuse, 5–11(–14) × 3–6(–9) cm, the lower surface ferruginous-lepidote; petioles 1.5–4.5 cm long; petals pink, 12–13 × 7–7.5 mm. Tepui-slope forests, shrub islands in upland savannas, 900–2000 m; Bolívar (Cerro Guaiquinima, Ilú-tepui, Macizo del Chimantá, Ptari-tepui, Uei-tepui), Amazonas (Cerro Duida, Cerro Guanay, Cerro Marahuaka, Cerro Parú, Cerro Yutajé). Brazil (Amazonas). ◆Fig. 455.

Fig. 455. Topobea ferruginea

Votomita 527

49. VOTOMITA Aubl., Hist. Pl. Guiane 90. 1775. Meliandra Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 156. 1825. Coryphadenia Morley, Amer. J. Bot. 40: 248. 1953. by Thomas Morley Evergreen shrubs or trees. Petioles short; stipules none or represented by a minute interpetiolar fringe; blades entire, pinnate-veined, glabrous or pubescent. Inflorescence a 1–3(4)-flowered cyme; bracts small and usually early deciduous. Flowers pedicellate, bisexual, 4-merous. Free hypanthium none; calyx limb bowlshaped, usually with 4 small teeth that are free early in the bud; petals 4, white, whitish, pink, or violet. Stamens 8, equal or nearly so, sometimes laterally fused; filaments short, not inflexed in bud; anthers straight, erect, the connective produced above the thecae, usually bearing an apical, elliptic, introrse gland, the introrse thecae dehiscing by lengthwise slits. Ovary 1–4-locular, the placentation axile or free-central or the ovules in a whorl around an axile or basal placenta in each locule; ovules 5–48 per ovary; style 1, persistent or early deciduous. Fruit a 1-seeded berry

Fig. 456. Votomita orinocensis

528

M ELASTOMATACEAE

in the five examples known, crowned by the persistent calyx. Seeds smooth but unpolished on the enlarged outer face of the ovule; radicle short and straight, the cotyledons thick, fleshy, and planoconvex. Panama, Cuba, Colombia (likely), Guyana, Suriname, French Guiana, Peru, Brazil; 10 species, 2 in Venezuela, both in the flora area. Key to the Species of Votomita 1. 1.

Filaments separate; ovules 10 or fewer per ovary .................... V. orinocensis Filaments monadelphous; ovules 22–25 per ovary ................. V. ventuarensis

Votomita orinocensis Morley, Bull. Torrey Bot. Club 90: 14. 1963. Shrub ca. 2.5 m tall; petals white. Edges of granitic dome, 100–200 m; Amazonas (along Caño Cupaven, opposite mouth of Rio Atabapo). Endemic. ◆Fig. 456.

Votomita ventuarensis Morley, Novon 9: 241. 1999. —Cometure tierra firmero. Tree to 15 m tall; known only from one fruiting specimen. Seasonally flooded riparian forests, 100–200 m; Amazonas (Río Macubana between Río Yureba and Río Marueta on lower Río Ventuari). Endemic.

MELIACEAE by Terence D. Pennington and K. S. Edwards Trees, treelets, or shrubs, rarely with sympodial branching, rarely herbs with woody rootstocks; indument usually simple, less frequently of bifid or stellate hairs or of peltate scales. Bark sometimes with a milky exudate. Leaves spirally arranged, very rarely decussate, usually pinnate, with or without a terminal leaflet, sometimes with a dormant terminal bud, less frequently simple, rarely 1- or 3-foliolate, very rarely bipinnate; leaflets usually entire, rarely lobed, dentate, serrate, or crenate; buds protected by a luster of scale leaves or naked. Inflorescence usually axillary, less frequently terminal, rarely ramiflorous, cauliflorous, or epiphyllous, usually paniculate with cymose branchlets (thyrsoid), less frequently racemose, fasciculate, or spicate, or flowers in pairs or solitary. Flowers bisexual or unisexual (plants monoecious, dioecious, or polygamous); rudiments of opposite sex usually well developed in unisexual flowers. Calyx usually shallowly or deeply (2)3–5(–7)-lobed, less frequently truncate or with free sepals, rarely closed in bud and circumscissile at the base; aestivation open or imbricate; petals 3–6(–14), free or less frequently united below, often partially fused to the staminal tube; aestivation imbricate or contorted, less frequently valvate. Filaments rarely completely free, usually partly or completely united to form a staminal tube, with or without appendages; staminal tube 5–14 cm long, globose, urceolate, campanulate, cyathiform, or cylindrical and then sometimes curved or inflated distally, rarely ribbed; throat open or constricted; margin entire, crenate, lobed, or toothed, or bearing appendages; appendages very variable in shape, free or partly or completely united, 2(3–10) times as many as the anthers; anthers (3–)5–10(–23), hairy or glabrous, sometimes locellate (locules divided by transverse membranes into a row of smaller chambers), inserted apically on the filaments or on the margin of the staminal tube, or within the throat of the staminal

Carapa

529

tube and partially or completely included, usually alternate with the appendages, rarely opposite them; usually in a single whorl, rarely in 2 alternating whorls; connective sometimes produced to form a short or long appendage; antherodes in pistillate flowers smaller, not dehiscing or producing pollen; disk intrastaminal, stipitate, patelliform, cyathiform, tubular, or absent, free from or partly or completely fused to the base of the staminal tube or ovary. Ovary (1)2–6(–20)-locular; locules 1–manyovulate; placentation axile or very rarely parietal; ovules collateral, superposed, or biseriate; style head capitate or discoid, less frequently conical, pileate, or deeply lobed, or forming an ovoid, globose, fusiform, cylindrical, conical, or obconical pollen receptacle; pistillode in staminate flowers narrower with a longer, more-slender style, less-glandular style head, and very small abortive ovules. Fruit a loculicidal or septifragal capsule, berry, drupe, or very rarely a nut. Seed unwinged or winged and usually attached to a large woody columnella; endosperm usually lacking. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; 51 genera and ca. 550 species; 5 genera and 30 species in the flora area. Key to the Genera of Meliaceae 1. 1. 2(1). 2.

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

Locules with 3–many ovules, these biseriate; fruit a septifragal capsule (subfamily Swietenioideae) ................................................................... 2 Locules with 1 or 2 ovules; fruit a loculicidal capsule or drupe (subfamily Melioideae) ............................................................................................. 3 Staminal tube of completely united filaments; anthers 8–10(–12), inserted within throat of staminal tube; seeds unwinged ............. 1. Carapa Stamens 5, filaments free but adnate to an androgynophore below; anthers inserted apically on filaments; seeds with a thin papery wing ...................................................................................................... 2. Cedrela Leaves 2- or 3-pinnate ........................................................................ 4. Melia Leaves 1-pinnate, rarely 1–3-foliolate ....................................................... 4 Anthers inserted within throat of staminal tube ............................ 3. Guarea Anthers inserted at apex of filaments or on margin of staminal tube .................................................................................................... 5. Trichilia

1. CARAPA Aubl., Hist. Pl. Guiane 2(suppl.): 32, t. 387. 1775. Deciduous or evergreen trees or treelets. Leaves even-pinnate with a dormant glandular leaflet at apex, or exceptionally odd-pinnate; leaflets 6–10 pairs, glabrous or with simple hairs. Flowers 4- or 5(6)-merous, unisexual, but with well-developed vestiges of opposite sex present, borne in large erect or spreading, much-branched, many-flowered, thyrses, either axillary or subterminal, subtended by sterile bracts. Calyx 4- or 5-lobed ± to base, the lobes rounded or broadly ovate, imbricate; petals free, 4 or 5(6), slightly contorted, imbricate, longer than calyx in bud, spreading in open flowers. Staminal tube cup-shaped, cylindrical, or urceolate, divided at apex into 8–10 entire, short-bifid, or crenulate-lobed appendages, and bearing inside alternately with appendages 8–10(–12) ± sessile, included anthers or antherodes. Nectary well developed, cushion-shaped, surrounding ovary and free from staminal tube. Ovary 4- or 5(6)-locular, partially sunken into nectary, with (2)3–8 ovules in each locule; style short; style head discoid, ± blocking entrance to staminal tube,

530

M ELIACEAE

Fig. 457. Carapa guianensis

Cedrela 531

receptive surface thickened and glandular; pistillode similar to pistil but with a longer, more slender style and thinner style head; locules well developed but ovules rudimentary. Fruit a large, pendulous, subwoody, subglobose, or oblong-cylindrical, septifragal capsule, opening by 4 or 5 valves from base and apex simultaneously, valves becoming leathery on drying; columella reduced, ultimately breaking down. Seeds large, 8–35, angular due to mutual compression, with outer surface opposite hilum rounded; sarcotesta woody, thick, smooth; endosperm absent; embryo with large fused cotyledons; radicle lying above hilum; germination cryptocotylar; cataphylls spirally arranged; eophylls simple, entire, the later ones trifoliate, bright red. Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, West Africa, Asia, Oceania; 2 variable species, 1 in Venezuela. Carapa guianensis Aubl., Hist. Pl. Guiane 2(suppl.): 32, t. 387. 1775. —Carapa. Deciduous or semi-evergreen tree to 55 m tall with a large, very dense crown or of several erect or arching branches; buttresses to 2 m tall; bark smooth, light gray to grayish brown, sometimes reddish, with shallow fissures, scaling in flat, nearly square plates or horizontal strips; inner bark pinkish brown or red, fibrous; wood pinkish, flowers white or creamy white, slightly musk-scented. Swampy ground along river banks, low-lying seasonally or

permanently flooded forests, near sea level to 400(–1400) m; Delta Amacuro (widespread), Bolívar (Cerro Venamo, El Dorado, El Palmar). Anzoátegui, Monagas, Sucre, Yaracuy; widely distributed along the Atlantic coast of Central America (also on the Pacific slope in Costa Rica), West Indies, Colombia, Trinidad-Tobago, French Guiana, Ecuador, Peru, Brazil (Amazon basin, Maranhão). ◆Fig. 457. Timber from Carapa guianensis is heavily exploited. It is used in building furniture, houses, and boats.

2. CEDRELA P. Browne, Civ. Nat. Hist. Jamaica 158, t. 10, fig. 1. 1756. Deciduous trees. Leaves even-pinnate, rarely odd-pinnate; leaflets entire, glabrous or with simple hairs. Flowers 5-merous, unisexual, but with well-developed vestiges of opposite sex present, borne in much-branched thyrses. Calyx lobed ± to base, shallowly dentate, or cup-shaped and split down 1 or 2 sides; petals 5, free, longer than calyx in bud, imbricate and adnate 1/3–1/2 their length to a long columnar androgynopore (nectary) by a median carina thereby preventing them from spreading in open flowers. Stamens 5, free, adnate to androgynophore below; filaments expanded at base, flattened; anthers in staminate flowers large, yellow, and dehiscing; antherodes in pistillate flowers small, shrivelled, brownish, and not producing pollen; staminodes absent. Ovary 5-locular, borne at apex of gynophore, each locule with 8–14 ovules; style short; style head discoid with glandular stigmatic papillae; pistillode in staminate flowers more slender, with well-developed locules and a longer style, vestigial ovules very small. Fruit a pendulous or erect, woody, obovoid, cylindroid, or claviform, septifragal capsule, opening from apex by 5 valves; columella woody, broadly winged, extending to apex of capsule, columnar apex with a sterile area; seed scars conspicuous. Seeds with a terminal wing attached by seed end to distal part (apex) of columella and winged toward base of capsule, with residual endosperm; cotyledons collateral, flattened and leaf-like; radicle laterally exserted; germination phanerocotylar; eophylls opposite, trifoliate; leaflets sinuate, entire.

532

M ELIACEAE

Fig. 458. Cedrela odorata

Guarea 533

Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina; ca. 8 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Cedrela 1.

1.

Leaflet blade generally with varying amounts of indument, either hirsute, pilose, tomentose, or villous; terminal cymules of inflorescence crowded and congested; petals with a pinkish tinge outside; capsule 3.5–15 cm long .............................................................................. C. fissilis Leaflet blade generally glabrous on the lower surface, sometimes with scattered hairs along midrib and secondary veins; terminal cymules of inflorescence open, lax; petals greenish white; capsule 2–4 cm long ..................................................................................................... C. odorata

Cedrela fissilis Vell., Fl. Flumin. 72. 1825 [1829]; Fl. Flumin. Icon. 2: pl. 68. 1827 [1831]. —Cedro. Tree to 30 m tall, with straight cylindrical bole; bark grayish brown to gray-black, regularly and evenly furrowed or fissured longitudinally; flower with agreeable scent; petals greenish white, yellowing with age. Lowland to lower montane forests, mostly on nonflooded ground, near sea level to 800 m; Delta Amacuro (near Altiplanicie de Nuria, Caño Güiniquina, Río Acure), Bolívar (Guayapo on lower Río Caura, Sierra de Maigualida). Barinas; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina. Timber from Cedrela fissilis has limited commercial use. Cedrela odorata L., Syst. Nat. ed. 10, 940. 1759. —Cedro, Cedro colorado, Cedro rojo. Cedrela brownii Loefl. ex Kuntze, Gen. Pl.

1: 111. 1891. —Surenus brownii (Loefl. ex Kuntze) Kuntze, Revis. Gen. Pl. 1: 111. 1891. Deciduous tree 30–60 m tall, to 1.5 m diameter, with dense, rounded or ± flat crown, buttresses to ca. 1 m tall; bark grayish brown to gray-black, regularly and evenly furrowed or fissured longitudinally; flowers greenish white or off-white, with unpleasant odor. Semideciduous to evergreen lowland and montane forests, 50–1200 m; Delta Amacuro (east-northeast of El Palmar, near Río Aguirre, near Río Cuyubini, Río Guanamo east of Campamento Río Grande, upper Río Toro), Bolívar (near Cerro Venamo, Chiguao, 55 km northeast of Ciudad Piar, Gran Sabana, Represa Guri). Widespread elsewhere in Venezuela; throughout lowland Central America, West Indies, and South America to northern Argentina. ◆Fig. 458. Cedrela odorata, Spanish cedar, is used commercially and is highly regarded for woodworking.

3. GUAREA F. Allam. ex L., Mant. Pl. 150, 228. 1771, nom. cons. Trees or treelets; indument of simple hairs. Leaves nearly always pinnate with a terminal bud usually showing intermittent growth, rarely without terminal bud, very rarely unifoliate; leaflets sometimes glandular-punctate and -striate; bud scales absent. Inflorescence axillary, ramiflorous, or cauliflorous, a panicle, raceme, or spike. Flowers (where known) unisexual (plants dioecious). Calyx with an almost entire margin or shallowly to deeply 3–7-lobed, aestivation open; petals (3)4–6(7), free, nearly always valvate, rarely slightly imbricate. Staminal tube cylindrical, sometimes contracted at throat, margin entire, crenate, or with short truncate or emarginate lobes; anthers (7)8–12(–14), glabrous, inserted within throat of staminal tube, completely included or partly exserted, alternate with lobes; antherodes simi-

534

M ELIACEAE

lar, smaller, without pollen. Nectary short- to long-stipitate, nearly always expanded to form a collar at base of ovary. Ovary 2–10(–14)-locular, locules with 1 or 2 superposed ovules; style head discoid; pistillode smaller, less swollen, with well-developed nonfunctional ovules. Fruit a 2–10(–14)-valved loculicidal capsule, locules 1- or 2seeded; pericarp leathery or woody; endocarp thin, cartilaginous. Seed often shaped like segment of an orange, with a thin fleshy sometimes vascularized sarcotesta; embryo with thick plano-convex, superposed or rarely collateral cotyledons; radicle abaxial, extending to surface or when cotyledons collateral then radicle apical; endosperm absent. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Africa; ca. 35 species, 10 in Venezuela, all of these in the flora area. Key to the Species of Guarea 1. 1. 2(1). 2. 3(2).

3.

4(2).

4. 5(4).

5.

6(5). 6. 7(6). 7. 8(6). 8. 9(8).

Ovary 2–13-locular, locules with 2 superposed ovules ............................. 2 Ovary 4(5)-locular, locules with 1 ovule .................................................. 10 Ovary glabrous ........................................................................................... 3 Ovary pubescent ......................................................................................... 4 Leaf without a terminal bud; ovary 2 or 3(4)-locular; fruit testiculate, constricted between the seeds, smooth, without lenticels; cotyledons collateral ..................................................................................... G. silvatica Leaf with a terminal bud; ovary (3)4-locular; fruit ellipsoid to globose, or irregularly shaped, usually with large pale lenticels; cotyledons superposed ....................................................................................... G. kunthiana Leaflets strongly bullate, tertiary venation very prominent on the lower surface; lower blade densely golden pubescent; petals 14–15 mm long ...................................................................................................... G. sprucei Leaflets not bullate; lower blade usually without golden pubescence; petals 11–14 mm long ................................................................................. 5 Flowers subtended by 1–3 prominent lanceolate bracteoles 4–6 mm long; young parts densely golden tomentose; lower blade uniformly puberulous with crisped hairs ............................................... G. trunciflora Flowers not subtended by prominent bracteoles (bracteoles if present then inconspicuous and much smaller); young parts densely golden tomentose or not; lower blade not uniformly puberulous with crisped hairs ........................................................................................................ 6 Indument of young parts and lower surface of blade prominent, villose, tomentose (often golden), or coarsely pubescent .................................. 7 Indument of young parts fine, minute; lower surface of blade minutely puberulous or glabrous .......................................................................... 8 Fruit < 3.5 cm long, winged, ribbed, or ± smooth .................. G. macrophylla Fruit > 3.5 cm long, ± smooth ................................................... G. grandifolia Fruit > 3.5 cm long; lower surface of leaf blade moderately to densely puberulous or ± glabrous ...................................................... G. grandifolia Fruit ≤ 3.5 cm long; lower surface of leaf blade ± glabrous ...................... 9 Leaflets ca. 4 times as long as wide; inflorescence to 40 cm long; petals 8– 12 mm long; anthers 1–1.5 mm long; ovary 4–6(7)-locular; fruit 2.5–

Guarea 535

9.

10(1). 10. 11(10). 11. 12(11). 12. 13(11). 13. 14(13).

14.

15(13). 15. 16(15). 16. 17(16).

17.

3.2 cm long .................................................................................. G. gomma Leaflets ca. 3 times as long as wide; inflorescence < 25 cm long; petals 6– 9 mm long; anthers usually < 1 mm long; ovary nearly always 4-locular; fruit usually < 2.5 cm long .................................................. G. macrophylla Fruit ca. 3.5 cm diameter ............................................................... G. silvatica Fruit usually 1.5–2.5 cm diameter, never > 3.5 cm diameter ................. 11 Fruit ribbed, tuberculate, or winged ....................................................... 12 Fruit smooth ............................................................................................. 13 Fruit obscurely or prominently torulose or verrucose, if ribbed then ribs torulose or verrucose .......................................................... G. macrophylla Fruit narrowly ribbed, ribs not torulose or verrucose ............... G. pubescens Fruit glabrous ........................................................................................... 14 Fruit tomentose, puberulous, or papillose .............................................. 15 Lower surface of blade with tuft of hairs in vein axils, otherwise glabrous; secondary veins often convergent; leaflets usually elliptic or oblanceolate; fruit usually rough-surfaced, pale brown, if dark then without prominent lenticels ....................................................................... G. glabra Lower surface of blade glabrous; secondary veins usually parallel; leaflets often oblong; fruit nearly always shining brown, smooth, with prominent pale lenticels ..................................................................... G. guidonia Lower surface of blade puberulous to tomentose ....................... G. pubescens Lower surface of blade ± glabrous ........................................................... 16 Fruit coarsely pubescent, depressed-globose or globose ............ G. pubescens Fruit finely puberulous or papillose, usually pear-shaped ..................... 17 Leaves without intermittent apical growth; inflorescence to 40 cm long; petals 8–12 mm long; anthers 1–1.5 mm long; ovary 4–6(7)-locular; fruit 2.5–3.2 cm long ................................................................... G. gomma Leaves with intermittent apical growth; inflorescence < 25 cm long; petals 6–9 mm long; anthers usually < 1 mm long; ovary nearly always 4-locular; fruit usually < 2.5 cm long ........................................... G. macrophylla

Guarea glabra Vahl, Eclog. Amer. 3: 8. 1807. —Carapillo. Evergreen tree to 25 m tall, larger specimens with small buttresses; bark shallowly fissured or scaling, pale gray or grayish brown; flowers white or whitish green, fragrant; fruit pale brown to dark red-purple. Evergreen lowland forests, near sea level to 700 m; Delta Amacuro (near Río Aguirre, east to Río Grande), Bolívar (Altiplanicie de Nuria, Río Arisa, near Río Cuyuní, 12 km from Upata). Aragua, Distrito Federal, Falcón, Sucre, Táchira, Yaracuy; Mexico, Central America, Puerto Rico, Jamaica, Lesser Antilles Colombia, Ecuador, Peru, Brazil. ◆Fig. 459. Guarea gomma Pulle, Recueil Trav. Bot. Neérl. 6: 271. 1909.

Tree to 40(–55) m tall, sometimes buttressed with bole fluted to 3 m; bark middle brown, soft, slightly corky and scaling in irregular patches; corolla cream-colored; fruit red. Lowland unflooded forests, 100–400 m; Bolívar (Río Paramichi near Serranía Marutaní). Colombia, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. Guarea grandifolia DC., Prodr. 1: 624. 1824. Tree to 50 m tall, larger specimens with buttresses to 3–4 m tall; bark pale brown, scaling in irregular plates; calyx reddish; corolla cream-colored; mature fruit reddish brown. Evergreen lowland and lower montane forests, 50–800 m; Amazonas (near Ocamo, near Sierra de la Neblina). Miranda;

536

M ELIACEAE

Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil (Amapá, Amazonas, Pará, Rondônia). ◆Fig. 461. Guarea guidonia (L.) Sleumer, Taxon 5: 194. 1956. —Samyda guidonia L., Sp. Pl. 443. 1753. —Acamosebi (Warao), Cabimbo, Mayjama (Yekwana), Peonía, Trompillo. Guarea puberula Pittier, Bol. Soc. Venez. Ci. Nat. 4: 357. 1938. Tree to 25 m tall, often branched from near the base; young tree bark smooth, brown, becoming fissured with age; flowers white or cream-colored; fruit smooth and shining brown or reddish brown. Riparian forests, lowland forests, near sea level to 500 m; Delta Amacuro (Caño Araguao, Caño Araguabisi, near Guarucara between Caño Araguao and Caño Güiniquina, Tucupita to Los Güires), Bolívar (near Canaima, near La Unión, Río Caura, Río Cuchivero, Río Nichare), Amazonas (near Ocamo, near Río Yureba, Sierra Parima). Widespread elsewhere in Venezuela; Costa Rica, Panama, Greater Antilles, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina. ◆Fig. 460. Guarea kunthiana A. Juss., Mém. Mus. Hist. Nat. 19: 241, 290. 1830. Tree to 14 m tall; bark grayish brown; flowers sometimes fragant; corolla creamcolored or pinkish red ; mature capsule from brown to reddish brown or rarely maroon, nearly always with conspicuous white lenticels. Evergreen nonflooded forests, ca. 200 m; Amazonas (lower Río Marutaní). Apure, Distrito Federal, Mérida, Táchira; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. Guarea macrophylla Vahl, Eclog. Amer. 3: 8. 1807. Lesser Antilles, Amazonian Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina, Uruguay; 5 subspecies, 2 in Venezuela, both in the flora area. Key to the Subspecies of G. macrophylla 1. Inflorescence a short (to 30 cm), densely flowered to long, slender, laxly flowered thyrse; fruit usually fig-shaped and tapering to base ...... subsp. pachycarpa

1. Inflorescence a long (to 50 cm), slender, densely flowered thyrse; fruit usually globose and abruptly contracted into a short stipe .......... subsp. pendulispica G. macrophylla subsp. pachycarpa (C. DC.) T.D. Penn., Fl. Neotrop. 28: 298. 1981. —Guarea trichilioides var. pachycarpa C. DC. in Mart., Fl. Bras. 11(1): 184. 1878. Tree to 25 m tall; calyx red; corolla white to coral pink; mature capsule deep redpurple. Unflooded and seasonally flooded forests, 100–1600 m; Bolívar (near Macizo del Chimantá) Amazonas (Caño Iguapo southeast of La Esmeralda, Cerro Sipapo, Río Casiquiare, Río Mavaca, Río Puruname, Río Siapa). Zulia; French Guiana, Ecuador, Peru, Brazil (Amapá, Bahia, Espírito Santo, Pará), Bolivia. G macrophylla subsp. pendulispica (C. DC.) T.D. Penn., Fl. Neotrop. 28: 290. 1981. —Guarea pendulispica C. DC., Repert. Spec. Nov. Regni Veg. 7: 59. 1909. Tree to 10 m tall; flowers white; fruits red. Riparian or flooded forests, 100–200 m; Bolívar (Caño Fatua, upper Río Caura), Amazonas (Caño Iguapo). Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia. Guarea pubescens (Rich.) A. Juss., Mém. Mus. Hist. Nat. 19: 241, 286. 1830. —Trichilia pubescens Rich., Actes Soc. Hist. Nat. Paris 1: 108. 1792. Colombia, Guyana, Suriname, French Guiana, Brazil, (Amapá, western and central Amazon basin); 2 subspecies, both in the flora area. Key to the Subspecies of G. pubescens 1. Leaflets usually broadly elliptic or oblanceolate, often < 3 times as long as wide; midrib usually sunken and slightly puberulous; upper blade with minute raised dots; petiole and rachis unwinged; capsule often globose and minutely tuberculate, ribbed or not ... subsp. pubescens 1. Leaflets narrowly elliptic or lanceolate, usually > 3 times as long as wide; midrib prominent on upper surface, glabrous; upper blade without raised dots; petiole and rachis narrowly winged; capsule depressed-globose, irregularly ribbed ........ .................................. subsp. pubiflora

Guarea 537

Fig. 459. Guarea glabra

Fig. 460. Guarea guidonia

538

M ELIACEAE

Fig. 461. Guarea grandifolia

Guarea 539

Fig. 462. Guarea silvatica

Fig. 463. Guarea pubescens subsp. pubiflora

540

M ELIACEAE

G. pubescens subsp. pubescens. —Catarajaca, Tortolito blanco. Treelet to 6 m tall; bark pale gray or grayish white, fissured; calyx red or pink; petals cream-colored or white; fruit dull dark red or purple. Nonflooded lowland forests, open savannas, 50–300 m; Delta Amacuro (near Río Acure), Bolívar (50 km west of Cerro Guaiquinima, upper Río Caura), Amazonas (Culebra, Ocamo, Río Casiquiare, Río Padamo). Apure; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amapá, Amazonas, Pará, Roraima). G. pubescens subsp. pubiflora (A. Juss.) T.D. Penn., Fl. Neotrop. 28: 298. 1981. —Guarea pubiflora A. Juss., Mem. Mus. Hist. Nat. 19: 241, 287. 1830. Treelet to 10 m tall, often producing aerial adventitious roots when subject to flooding; calyx deep red-purple; corolla pink-creamcolored; fruit with fleshy reddish purple pericarp. Riverbanks in periodically and permanently flooded forests, 50–200 m; Bolívar (near Río Parguaza), Amazonas (Caño Yagua, Caño Yapacana, Isla Ratón, near Quiratare, Pamoni, lower Río Casiquiare). Colombia, Brazil (Amazonas, Pará), Bolivia. ◆Fig. 463.

Guarea silvatica C. DC. in Mart., Fl. Bras. 11(1): 195, t. 57. 1878. Tree to 20 m tall; bark smooth, grayish green, becoming dark gray and scaling in thin, narrow, longitudinal pieces; flowers cream-colored, scentless; fruit with thin, leathery, reddish brown pericarp. Evergreen lowland forests, common in secondary forests, 100–200 m; Amazonas (Río Yatúa). Guyana, French Guiana, Peru, Brazil (Acre, Amazonas, Espírito Santo, Maranhão, Pará). ◆Fig. 462. Guarea sprucei C. DC. in Mart., Fl. Bras. 11(1): 196, t. 58. 1878. Tree to 33 m tall, with golden pubescence on most parts; leaves to 40 cm long, prominently bullate, the lower surface of leaflets with numerous minute red papillae. Evergreen lowland forests, 100–200 m; Amazonas (Río Mawarinuma). Brazil. Guarea trunciflora C. DC., Monogr. Phan. 1: 571. 1878. —Juasudu (Yekwana). Tree to 18 m tall; flowers scentless, yellowish. Lowland nonflooded forests, 50–1000 m; Bolívar (upper Río Nichare, Río Parguaza, Río Tabaro near Río Nichare). Colombia, Peru.

4. MELIA L., Sp. Pl. 384. 1753. Monopodial trees or treelets; indument a mixture of simple and tufted-stellate hairs. Leaves 2- or 3-pinnate. Inflorescence a many-flowered axillary panicle, ultimate branchlets often cymose. Flowers bisexual and staminate on same individual (plants polygamous). Calyx 5(6)-lobed to near base, the sepals sometimes imbricate at base; petals 5(6), free, imbricate. Staminal tube 0.4–1 cm long, narrowly cylindrical, slightly expanded at mouth, 10–12-ribbed, terminated by slender, filiform, truncate or 2–4-lobed appendages, as many as or twice as many as the anthers; anthers 10(–12), hairy or glabrous, inserted on margin of staminal tube or just inside, surrounding base of ovary. Ovary 4–8-locular; locules with 2 superposed ovules; style head capitate to coroniform, with 4–8 short, erect or incurved stigmatic lobes. Fruit a 3–8-locular drupe; endocarp thick, bony, hollowed at base and apex; locules 1(2)seeded. Seed oblong, laterally compressed; aril lacking; testa leathery, sometimes slightly swollen and fleshy around hilum; embryo embedded in thin endosperm; cotyledons flat, collateral, radicle superior, short, projecting from cotyledons. Old World tropics, 1 species introduced and naturalized in the New World; ca. 5 poorly defined species, 1 in Venezuela. Melia azedarach L., Sp. Pl. 384. 1753. Tree or shrub to 10 m tall; flowers pink, lilac, pale blue, or dark violet; fruit a berry. Cultivated and naturalized in Delta Amacuro, Bolívar, and Amazonas. Mérida; Old

World tropics, cultivated and naturalized in Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 464.

Melia 541

Fig. 464. Melia azedarach

542

M ELIACEAE

5. TRICHILIA P. Browne, Civ. Nat. Hist. Jamaica 278. 1756. Trees or treelets; indument usually of simple hairs, less frequently of malpighiaceous, dibrachiate, or stellate hairs, or of peltate scales. Leaves usually pinnate, less frequently 3- or 1-foliolate, rarely digitate; leaflets sometimes glandular-punctate and -striate; buds usually naked, rarely subtended by a cluster of small scale leaves. Flowers usually unisexual (plants dioecious), less frequently bisexual (plants polygamous), in axillary thyrsoid panicles, rarely fasciculate or corymbose, or reduced to a few-flowered raceme. Calyx usually shallowly to deeply (3)4–6-lobed, less frequently sepals free, aestivation usually open, less frequently imbricate, rarely quincuncial; petals (3)4 or 5(6), free or partially united, imbricate, valvate, or rarely quincuncial. Filaments completely united to form a cyathiform, urceolate, or short-cylindrical staminal tube, margin usually toothed or lobed, or rarely entire, or filaments partly (rarely completely) free and then with or without 2 terminal lobes or appendages; anthers (4)5–10(11), hairy or glabrous, inserted between teeth or lobes on margin of staminal tube, or apically on filaments; antherodes narrower than anthers, not dehiscing, without pollen. Nectary usually a fleshy annulus surrounding base of ovary, less frequently broadly stipitate, patelliform, or cyathiform, occasionally partly fused to base of staminal tube, or absent. Ovary 2 or 3(4)-locular, locules with 1 or 2 collateral or less frequently superposed ovules; pistillode smaller but often with well-developed abortive ovules; style head usually capitate, less frequently conical or columnar, with or without lobes. Fruit a 2- or 3(4)-valved loculicidal capsule; valves leathery or woody, locules 1- or 2-seeded. Seed ± plano-convex, partly or completely surrounded by a thin or fleshy arillode, rarely with a complete sarcotesta; usually without endosperm, rarely with copious endosperm; embryo with plano-convex, or less frequently thin and flat, collateral or rarely superposed cotyledons; radicle superior, included or occasionally exserted; rarely with superposed cotyledons and central included radicle. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa, Indo-Malayan area; ca. 100 species, 24 in Venezuela, 16 of these in the flora area. Key to the Species of Trichilia 1.

1.

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

3.

Petals imbricate or quincuncial, nearly always free, if subvalvate then plant with indument of stellate hairs; filaments partially or completely fused, rarely free; nectary present or absent (section Trichilia) .......... 2 Petals valvate, free or united; indument not stellate or lepidote; staminal tube of completely united filaments; nectary absent or rarely represented by an obscure swelling around base of ovary (section Moschoxylum) ........................................................................................ 8 Indument of stellate hairs or peltate scales .............................................. 3 Indument not of stellate hairs or peltate scales ....................................... 4 Leaflets opposite; stellate or peltate indument conspicuous; sepals united, with open aestivation; fruit globose to broadly ovoid, 1–2.2 cm long ..................................................................................................... T. lepidota Leaflets alternate or only subopposite; indument minute; sepals ± free, imbricate; fruit narrow, usually ellipsoid or obovoid, 2.5–3.8 cm long ............................................................................................... T. mazanensis

Trichilia 543

4(2). 4. 5(4). 5. 6(5). 6. 7(6). 7. 8(1). 8. 9(8). 9. 10(9). 10. 11(10). 11. 12(11). 12. 13(12). 13. 14(9). 14. 15(14). 15. 16(15). 16. 17(8). 17. 18(17). 18. 19(18). 19. 20(19). 20. 21(20).

Locules with 2 superposed ovules ................................................... T. pallida Locules with 1 or 2 collateral ovules ......................................................... 5 At least some locules with 2 collateral ovules ................................. T. elegans Locules always uniovulate ......................................................................... 6 Sepals free, strongly imbricate ............................................ T. septentrionalis Sepals free or united, not or only slightly imbricate at base .................... 7 Ovary glabrous; outer surface of filaments pubescent; fruit 0.7–1.3 cm long .......................................................................................... T. micrantha Ovary pubescent or strigose; outer surface of filaments glabrous; fruit (2–) 2.5–3.5 cm long ............................................................................... T. rubra Leaflets dimorphic or heteromorphic, lowest pairs much reduced and often a different shape, sometimes reduced to vestigial scales .............. 9 Leaflets not dimorphic or heteromorphic ................................................ 17 Petals free or united only at base ............................................................ 10 Petals united 1/3–2/3 their length .............................................................. 14 Reduced basal leaflets represented by a pair of minute caducous scales 1–2 mm long ........................................................................... T. quadrijuga Reduced basal leaflets with at least some expanded blades .................. 11 Fruit nearly always verrucose ......................................................... T. pleeana Fruit smooth, sericeous, or tomentose .................................................... 12 Young branches, leaf rachis, and inflorescence coarsely pubescent; fruit tomentose ........................................................................... T. inaequilatera Indument of young branches, leaf rachis, and inflorescence puberulous or shortly pubescent; fruit puberulous or finely sericeous ..................... 13 Largest leaflets 4.7–6.8 cm long; petals 2–2.5 mm long ........... T. tetrapetala Largest leaflets 8.5–22 cm long; petals 2.5–5 mm long ........ T. schomburgkii Indument of young parts and leaf rachis prominent, densely pubescent, tomentose, or villose ............................................................. T. gamopetala Indument of young parts and leaf rachis minute, usually appressedpuberulous or strigose ......................................................................... 15 Reduced basal leaflets represented by 1 pair of subulate scales ................................................................................................ T. quadrijuga Reduced basal leaflets usually > 1 pair, always with an expanded blade .............................................................................................................. 16 Petals 1.5–2.5 mm long .............................................................. T. tetrapetala Petals ≥ 2.5 mm long .............................................................. T. schomburgkii Leaves 1–3(4)-foliolate or -digitate ............................................. T. acuminata Leaves pinnate, the majority with at least 5 leaflets, rarely with a few 3or 4-foliolate, never digitate ................................................................ 18 Ovary 2-locular ....................................................................................... T. cipo Ovary 4-locular ......................................................................................... 19 Lower surface of leaf with scattered to moderate, appressed, medifixed, or dibrachiate hairs .................................................................... T. quadrijuga Lower surface of leaf glabrous, or if pubescent, then hairs not appressed, medifixed, or dibrachiate ..................................................................... 20 Arillode strongly developed at apex and along adaxial surface ............. 21 Arillode completely enveloping seed ......................................... T. quadrijuga Secondary veins (10)11–13(–16), strongly arcuate, ascending and conver-

544

21.

M ELIACEAE

gent; leaflets usually oblanceolate; whole plant subglabrous, only young growth with minute, appressed, dibrachiate hairs .......... T. surinamensis Secondary veins (13–)16–18(–21), shallowly ascending, slightly convergent or parallel; leaflets oblong or elliptic; indument denser and more persistent, most hairs basifixed ........................................................ T. cipo

Trichilia acuminata (Humb. & Bonpl. ex Roem. & Schult.) A. DC. & C. DC., Monogr. Phan. 1: 704. 1878. —Odontandra acuminata Humb. & Bonpl. ex Roem. & Schult., Syst. Veg. 5: 511. 1819. —Mezclo. Odontocandra karstenii Triana & Planch., Ann. Sci. Nat. Bot. 15: 373. 1872. —Trichilia karstenii (Triana & Planch.) A. DC. & C. DC., Monogr. Phan. 1: 704. 1878. Tree to 20 m tall; flowers white; mature fruits reddish. Ca. 600 m; Bolívar (lower Río Suapure). Panama, Colombia. Trichilia cipo (A. Juss.) C. DC. in Mart., Fl. Bras. 11(1): 214. 1878. —Moschoxylum cipo A. Juss., Mém. Nus. Hist. Nat. 19: 239. 1830. Tree to 15 m tall; flowers cream-colored, unscented; fruit grayish. Riparian forests, 200–300 m; Bolívar (El Cácaro between Río Caura and Río Paragua, Salto Pará). Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil. Trichilia elegans A. Juss. in A. St.-Hil. et al., Fl. Bras. Merid. 2: 79, t. 98. 1829. Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil; 2 subspecies, 1 in Venezuela. T. elegans subsp. elegans Tree to 20 m tall; flowers white or greenish white; fruit maroon or wine red. Riparian flooded forests, 200–300 m; Bolívar (Represa Guri). Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, northern Argentina. Trichilia gamopetala T.D. Penn., Fl. Neotrop. 28: 177. 1981. Tree to 20 m tall; flowers creamy white to grayish cream-colored. Amazonas (Cerro Huachamacari, near Ocamo, Río Cunucunuma). Endemic. Trichilia inaequilatera T.D. Penn., Fl. Neotrop. 28: 209. 1981. —Cedril. Tree to 25 m tall, with smooth gray bark;

flowers whitish yellow; young fruit puberulent, tan-orange-colored. Periodically flooded lowland forests, river banks, 100–300 m Bolívar (La Ceiba, Río Oris in Río Paragua basin, near San Salvador de Paúl), Amazonas (near Río Atacavi, Santa Bárbara). Colombia, Peru, Brazil (Amazonas). ◆Fig. 466. Trichilia lepidota Mart., Flora 22(1) Beibl.: 54. 1839. Tree 15–25 m tall; trunk slightly fluted at base; flowers white, sweet-scented; fruit maroon. Eastern Venezuela, Guyana, Suriname, Brazil (Bahía and along the eastern coast from Río Grande do Sul to Minas Gerais); 3 subspecies, 1 in Venezuela. T. lepidota subsp. leucastera (Sandwith) T.D. Penn., Fl. Neotrop. 28: 40. 1981. —Trichilia leucastera Sandwith., Kew Bull. 1933: 328. 1933. —Biscochuelo, Kerosene. Deciduous to evergreen lowland and lower montane forests, 100–600 m; Bolívar (near El Descanso, near Río Botanamo, near San Salvador de Paúl). Guyana, Suriname. ◆Fig. 467. Trichilia mazanensis J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3): 742. 1949. —Cedrito. Tree to 20 m tall; flowers whitish or greenish yellow; fruit greenish yellow. Periodically flooded forests, river margins, 50–200 m; Bolívar (near El Palmar, La Paragua, Río Caroní, Río Caura near mouth of Río Nichare, near San Salvador de Paúl), Amazonas (upper Caño Parucito, Casiquiare, La Esmeralda, near Ocamo, Río Emoni in lower Río Siapa basin, Río Guayapo, near Sierra de la Neblina). Colombia, Peru, Brazil (Amapá, Amazonas). Trichilia micrantha Benth., Hooker’s J. Bot. Kew Gard. Misc. 3: 369. 1851. —Caoba. Tree to 20 m tall, with rounded but-

Trichilia 545

tresses; outer bark light brown, almost white, soft, smooth; inner bark cream-colored, soft; latex honey-colored; flowers yellow-green or whitish; fruit yellowish. Evergreen lowland to montane forests, seasonally flooded riparian forests, 50–1100 m; Amazonas (Caño Mesaque in Río Atacavi basin, near Cerro Sipapo, 25 km southeast of La Esmeralda, near Piedra Cocuy, Río Cunucunuma basin, 12 km southeast of San Fernando de Atabapo, near Sierra de la Neblina). Monagas, Sucre; Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil (Amazonas, Mato Grosso Pará, Rondônia), Peru, Bolivia. Trichilia pallida Sw., Prodr. 67. 1788. Trichilia montana var. fendleriana C. DC. in A. DC. & C. DC., Monogr. Phan. 1: 654. 1878. Trichilia subsimplex Steyerm., Fieldiana, Bot. 28(2): 279. 1952. Tree to 25 m tall; bark of smaller specimens smooth or dimpled, pale gray; flowers cream-colored or white; fruit with greenish yellow pericarp. Lowland and montane rainforests, to 2000 m; Bolívar (near Cerro Venamo, near Río Botanamo, upper Río Caura), Amazonas (Cerro Duida, Río Cataniapo). Anzoátegui, Barinas, Carabobo, Distrito Federal, Lara, Sucre, Zulia; widespread in Mexico, Central America, Antilles, and South America. ◆Fig. 468. Trichilia pleeana (A. Juss.) C. DC. in Mart., Fl. Bras. 11(1): 215. 1878. —Moschoxylum pleeanum A. Juss., Mém. Mus. Hist. Nat. 19: 239, 281. 1830. —Colorado, Mijarro. Guarea adenocarpa Pittier, Bol. Soc. Venez. Ci. Nat. 4: 359. 1938. Guarea delgadoi Pittier, Bol. Soc. Venez. Ci. Nat. 4: 356. 1938. Tree to 30 m tall; bole slightly fluted with small buttresses at the base of larger specimens; bark gray, scaling off in long, thin, irregular sheets; flowers greenish yellow; fruit dark glossy green at maturity. Unflooded and seasonally flooded evergreen forests, 50–300 m; Delta Amacuro (Río Toro), Bolívar (near Cerro Yutajé, near Chiguao, near Río Botanamo). Widespread elsewhere in Venezuela; Costa Rica, Panama, Colombia, Trinidad-Tobago, Guyana, Ecuador, Peru, Amazonian and southern Brazil, Bolivia. ◆Fig. 465.

Trichilia quadrijuga H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 215. 1821 [1822]. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, 1 in Venezuela. T. quadrijuga subsp. quadrijuga. —Biscochuelo amarillo. Tree to 30 m tall; bole slightly fluted at the base in older specimens; bark graybrown, scaling in long irregular pieces; flowers yellowish green to white. Usually unflooded lowland forests, frequently riverbanks, rarely on periodically flooded sites, 50–400 m; Delta Amacuro (near Río Acure, Río Toro), Bolívar (near Chiguao, La Paragua, mouth of Río Caura to Salto Pará, Tumeremo to Bochinche), Amazonas (Río Ocamo basin). Panama, Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Amazonian and southern Brazil, Bolivia. ◆Fig. 470. Trichilia rubra C. DC. in Mart., Fl. Bras. 11(1): 203. 1878. —Biscochuelo negro, Dejaka (Yekwana). Tree to 25 m tall; bark light brown to pale gray, smooth; flowers cream-colored or yellow-green, vanilla-scented; fruit brown. Seasonally flooded riparian forests, evergreen lowland forests, near sea level to 200 m; Delta Amacuro (near Río Acure, Río Toro, near San José de Amacuro), Amazonas (upper Río Cunucunuma, near Río Machicare). Colombia, Guyana, Peru, Amazonian Brazil, Bolivia. ◆Fig. 471. Trichilia schomburgkii C. DC. in A. DC. & C. DC., Monogr. Phan. 1: 695. 1878. Eastern Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Amapá, Pará, western Amazon basin); 2 subspecies, both in the flora area. Key to the Subspecies of T. schomburgkii 1. Leaflets (8)9–12; reduced basal leaflets (3)4 pairs; secondary veins 15–18 on either side of midrib; petals fused 1/2–3/4 their length ........... subsp. javariensis 1. Leaflets 5–7(8); reduced basal leaflets 2 or 3 pairs; secondary veins 9–14 on either side of midrib; petals free or rarely fused near base, strongly reflexed .................... ........................... subsp. schomburgkii

546

M ELIACEAE

T. schomburgkii subsp. javariensis T.D. Penn., Fl. Neotrop. Monogr. 28: 169, fig. 30B, 31C-D. —Dajaka (Yekwana), Iwakrete-piji (Yanomami). Tree to 20 m tall; flowers sweet-scented, greenish cream-colored; fruit grayish green. 200–600 m; Bolívar (El Cácaro between Río Caura and Río Paragua, 58 km southeast of Serranía de los Pijiguaos), Amazonas (Río Putaco 19 km north of the mouth of Río Ocamo). Colombia, Peru, Brazil. T. schomburgkii subsp. schomburgkii. —Biscochuelo negro, Sijanama (Yekwana), Suipo, Vomitivo. Tree to 20 m tall, with smooth, grayish brown to grayish green bark; flowers sweetscented; calyx and corolla green or greenish cream-colored; fruit grayish green. Unflooded lowland forests, periodically flooded forests, 100–600 m; Delta Amacuro (eastnortheast of El Palmar, Río Toro), Bolívar (near El Palmar, near Río Acure, Río Arisa, near Río Botanamo, near Río Yuruaní, 55 km south of Jabillal on Río Caura), Amazonas (San Carlos de Río Negro, Serranía Batata on Cuao-Sipapo massif). Falcón; Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará). ◆Fig. 469. Trichilia septentrionalis C. DC. in Mart., Fl. Bras. 11(1): 220. 1878. —Caramacate rebalsero. Trichilia moritzii C. DC. in A. DC. & C. DC., Monogr. Phan. 1: 707. 1878. Trichilia lanceolata Pittier, Arb. Arbust.

Venez. 1: 8. 1921, reprinted from Bol. Comerc. Industr. Venez. no. 13: 1921, nom. illeg., non C. DC. 1878. Trichilia grandis Lasser & Maguire, Bol. Soc. Venez. Ci. Nat. 15: 101, pl. s.n. 1954. Tree to 20 m tall, with smooth gray or gray-brown bark, finely longitudinally cracked or lenticellate; lower surface of leaflets pale green; flowers greenish cream-colored, sweet-scented; mature fruit red. Evergreen lowland to montane forests, 100-1200 m; Bolívar (near Cerro Venamo, near Los Testigos, lower Río Caura, near Santa Elena de Uairén), Amazonas (Río Matacuni, Río Putaco). Carabobo, Miranda; Costa Rica, Panama, Colombia, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. Trichilia surinamensis (Miq.) C. DC. in A. DC. & C. DC., Monogr. Phan. 1: 679. 1878. —Moschoxylum surinamense Miq., Stirp. Surinam. Select. 73. 1850 [1851]. Treelet or tree to 15(–35) m tall, with smooth bark; flowers white to greenish yellow; fruit gray-green. Evergreen lowland forests, riparian habitats, usually but not exclusively on unflooded land, 50–200 m; Amazonas (near Ocamo). Suriname, French Guiana, Brazil (Amapá, Pará, Roraima). Trichilia tetrapetala C. DC. in Mart. Fl. Bras. 11(1): 211. 1878. Tree ca. 16 m tall. Lower montane forests, 300–600 m; Bolívar (Cerro Ichún). Southern Brazil.

Fig. 465. Trichilia pleeana

Trichilia 547

Fig. 466. Trichilia inaequilatera

Fig. 467. Trichilia lepidota subsp. leucastera

548

M ELIACEAE

Fig. 468. Trichilia pallida

Fig. 469. Trichilia schomburgkii subsp. schomburgkii

Trichilia 549

Fig. 470. Trichilia quadrijuga subsp. quadrijuga

Fig. 471. Trichilia rubra

550

M ENDONCIACEAE

MENDONCIACEAE by Dieter C. Wasshausen Herbaceous or suffrutescent, counterclockwise-twining shrubs or vines. Stems articulated when young, glabrous or pubescent with simple or stellate, glandular and eglandular trichomes, cystoliths lacking. Leaves opposite, simple, entire, exstipulate, petiolate. Flowers solitary or clustered in the leaf axils, each pedunculate and subtended by 2 large, flat or keeled, spathe-like bracteoles (previously called bracts by this and other authors) green, variously shaped and vestured, often equaling the corolla tube, valvate, often partially connivent or connate, remaining closed around the flower, often widely spreading at the fruiting stage. Calyx inconspicuous, annular or cupular or truncate to irregularly dentate or lobed; corolla sympetalous, ± salverform, not inflated above, contorted, whitish, greenish, or reddish, often with purplish markings within, the tube cylindric to funnelform, the limb subequally 5-lobed or bilabiate, the upper lip 2-lobed, the lower lip 3-lobed, the lobes spreading or reflexed. Stamens 4, didynamous, included, attached to the corolla tube and alternate with the lobes; anthers bilocular, with parallel, subequal to unequal locules that are ± pubescent at the base, dehiscing longitudinally or by subapical pores or slits; pollen spherical, 5- or 6-colporate; staminode, if present, 1, inconspicuous. Gynoecium of 2 carpels united to form a compound, superior, bilocular ovary, or one locule ± reduced or even suppressed, a cupular nectar disk present around the base of the ovary; ovules 2 per locule, collateral, presumably unitegmic and tenuinucellar, the modified funiculi of the ovules lacking; style filiform; stigma shallowly and unequally bilobed. Fruit drupaceous, ovoid to ellipsoid; mesocarp fleshy; endocarp bony. Seeds 1 or 2, the cotyledons twice folded; endosperm absent. Pantropics; 2 genera and ca. 60 species, 1 genus and 10 species in the flora area. Traditionally the Mendonciaceae have been included in the Acanthaceae, but they lack both the cystoliths and the specialized mechanism of seed dispersal that characterize that family. Contrary to the Cronquist system followed here, most current specialists in the Acanthaceae, including myself, tend to include the family in the Acanthaceae. For a somewhat detailed examination of this relationship see R. K. Brummitt (Acanthus 5: 1–3. 1989). The other genus of Mendonciaceae is Anomacanthus R. Good (= Gilletiella De Wild. & T. Durand), a monotypic genus in westcentral tropical Africa. 1. MENDONCIA Vell. ex. Vand., Fl. Lusit. Bras. Spec. 43. 1788. Characters the same as for the family, except that in Anomacanthus the mature ovules are equally developed in two locules whereas in Mendoncia the mature ovules in one locule are retarded in their development. Southern Mexico, Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, tropical western Africa, Madagascar; ca. 60 species, 14 in Venezuela, 10 of these in the flora area. Lianas and vines are generally poorly known and collected in South America and then they often lack mature flowers or the corollas have dropped off, hence according to Daniel (Acta Bot. Mex. 17: 53–60. 1992), “most workers have distinguished species primarily on the basis of vegetative and bracteolar characters.”

Mendoncia 551

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

3. 4(3).

4.

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

6.

7(5). 7. 8(7). 8. 9(7).

9.

Bracteoles ovate to oblong-ovate ............................................................... 2 Bracteoles oblong, oblong-elliptic, oblong-lanceolate or oblong-obovate ................................................................................................................ 5 Flowers 4 or 5 in each axil; corollas very narrow, the upper half of the tube subcylindric, apically 4.5 mm diameter .................................. M. cardonae Flowers solitary or in pairs in each axil; corollas funnelform, apically ca. 8 mm diameter .......................................................................................... 3 Bracteoles to 20 × 10 mm, sparingly appressed-hirtellous without; leaf blades short-acuminate, the apex bearing a slender mucro to 2 mm long .............................................................................................. M. sprucei Bracteoles 23–30 × 13–17 mm, hirsute without; leaf blades aristate at apex, the awn 4–6 mm long ................................................................... 4 Bracteoles yellowish green to brownish green, 25–30 mm long, hirsute; leaf blades membranous, dark green and lustrous on the upper surface, glabrous; corollas 35 mm long .............................................. M. neblinensis Bracteoles dull green, 23–25 mm long, densely hirsute; leaf blades firmly membranous, dull dark green on the upper surface, sparingly hirsute; corollas 30 mm long ........................................................... M. steyermarkii Bracteoles narrowly oblong or oblong-lanceolate ..................................... 6 Bracteoles oblong, oblong-elliptic, or oblong-obovate ............................... 7 Leaf blades narrowly to broadly elliptic, rather thin, densely pubescent; bracteoles narrowly oblong, 35 × 8 mm, densely velvety-pubescent ................................................................................... M. hoffmannseggiana Leaf blades elliptic, chartaceous, minutely and sparingly hirtellous; bracteoles oblong-lanceolate, 20–25 × 10 mm, densely brown-strigose ................................................................................................ M. williamsii Bracteoles oblong or oblong-elliptic, coriaceous; leaf blades subcoriaceous to thick-coriaceous ................................................................................. 8 Bracteoles oblong-obovate to oblong-elliptic, not coriaceous; leaf blades membranous, not coriaceous ................................................................. 9 Leaf blades elliptic to oblong, 3–6 cm wide; bracteoles oblong, vinaceouslavender with green ................................................................... M. coriacea Leaf blades oblong-elliptic, 7.5–9 cm wide; bracteoles oblong-elliptic, green ......................................................................................... M. phalacra Bracteoles oblong-obovate, 10–13 mm wide, essentially glabrous; stems glabrous; leaf blades glabrous, chartaceous, the veins prominent on the lower surface .............................................................................. M. obovata Bracteoles oblong-elliptic, 15–18 mm wide, long-hirsute; stems hirsute with yellowish trichomes; leaf blades hirsute, membranous, the veins inconspicuous on the lower surface ........................................... M. bivalvis

Mendoncia bivalvis (L. f.) Merr., J. Arnold Arb. 29: 213. 1948. —Besleria bivalvis L. f., Suppl. Pl. 280. 1781 [1782]. Mendozia hirsuta Poepp. & Endl. in Poepp., Nov. Gen. Sp. Pl. 3: 10. 1845.

—Mendoncia hirsuta (Poepp. & Endl.) Nees in DC., Prodr. 11: 52. 1847. Vine; leaf blades 6–15 × 5.5–8 cm, ovate to oblong-ovate or elliptic, narrowed to an acuminate, mucronulate tip at the apex;

552

M ENDONCIACEAE

pedicels densely pilose; bracteoles 20–30 mm long, obtuse or rounded and mucronulate at the apex, thin, veiny; corollas cream white, tinged with yellow distally, ca. 25 mm long; drupe 15–18 × 9–12 mm, oblong to ovoid, compressed, oblique, deep purple when ripe. Lowland forests, 100–300 m; Bolívar (between San Félix and Upata, Tumeremo). Barinas; Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil, Bolivia. ◆Fig. 474. Mendoncia cardonae Leonard, Proc. Biol. Soc. Wash. 52: 18. 1939. Vine; leaf blades 5–14 × 2.5–6.5 cm, ovate to elliptic (sometimes asymmetrical), shortacuminate at apex; bracteoles 20–23 × 10–14 mm, acute to obtuse, mucronate at the apex, chartaceous, minutely strigose without; corolla red, 30 mm long; drupe 15 × 10 mm, minutely strigose. Lowland to montane forests, 100–1300 m; Bolívar (scattered), Amazonas (San Carlos de Río Negro, base of Sierra de la Neblina, Sierra Parima). Aragua, Portuguesa, Táchira. ◆Fig. 473. Mendoncia coriacea Wassh., Ann. Missouri Bot. Gard. 76: 1123. 1989. Vine; leaf blades 8–14 × 3.5–6 cm, abruptly acuminate (the tip blunt and terminating in a mucro 1 mm long), glabrous, dark green and minutely punctate on upper surface, dull green on lower surface; bracteoles coriaceous, 25–35 × 10–14 mm, rounded or obtuse at the apex, tipped by a mucro 1 mm long, glabrous and rugose without; corolla white with lavender stripes, glabrous, 45 mm long; drupe subglobose, dark winepurple to black, 18 × 15 mm, glabrous. Montane forests, 1200–1400 m; Bolívar (La Escalera-Cerro Venamo region). Endemic. Mendoncia hoffmannseggiana Nees in DC., Prodr. 11: 50. 1847. —Bejuco rojo. Mendoncia pubescens Hoffsgg. ex Nees in Mart., Fl. Bras. 9: 9. 1847, “Mendozia pubescens.” Mendoncia schomburgkiana Nees in DC., Prodr. 11: 50. 1847. Vine; leaf blades 5–12 × 3.7–8 cm, abruptly acuminate at the apex and terminating in an awn 2–4 mm long; pedicels densely yellowish brown-pilose; bracteoles subfalcate, abruptly acute or acuminate at the apex; corolla red or purplish, 35–55 mm

long, tubiform; drupe obliquely obovate, dark purple to blackish, 15 mm diameter, sparingly puberulous, succulent when ripe. Primary lowland forests along streams to cloud forests on summits of escarpments, 50–700 m; Delta Amacuro (Serranía de Imataca), Bolívar (scattered). Trinidad, Guyana, Suriname, French Guiana, Ecuador, Amazonian Brazil. ◆ 472. Mendoncia neblinensis Wassh., Ann. Missouri Bot. Gard. 76: 1120. 1989. Vine; leaf blades 5–11 × 3–5.5 cm, elliptic to ovate, abruptly acuminate at the apex; bracteoles ovate, apiculate, muricate; corolla bright to dark red, the lobes pink within; drupe fleshy, black, 17 × 8 mm, ellipsoid, glabrous. Lowland evergreen forests on hummocks along gravelly black-water rivers, 100–200 m; Amazonas (base of Sierra de la Neblina). Endemic. Mendoncia obovata Lindau, Bull. Herb. Boissier 5: 646. 1897. Vine; leaf blades to 18 × 8 cm, elliptic, acuminate at the apex, acute at the base, lateral veins curved toward the margins; flowers red (according to Spruce); bracteoles green, rotund, and briefly mucronate at the apex, to 30 × 12 mm, the margins agglutinate. Lowland forests, 100–200 m; Amazonas (Río Casiquiare, base of Sierra de la Neblina). Colombia (Valle), Brazil (Amazonas). Mendoncia phalacra Leonard, Contr. U.S. Natl. Herb. 31: 676. 1958. Vine; leaf blades to 12.5 cm long, shortacuminate and bluntly apiculate at the apex, glabrous; bracteoles to 30 × 15 mm, rounded and mucronulate at the apex, green, glabrous; corolla white, inside of the lobes tinted with purple-brown, 40 mm long; drupe purplish black, 17 × 10 × 7 mm, obtuse, glabrous. Forests on the edges of river banks, 100–300 m; Bolívar (base of Cerro Guaiquinima, Río Yudi), Amazonas (base of Sierra de la Neblina). Colombia (Amazonas, Vaupés). Mendoncia sprucei Lindau, Bull. Herb. Boissier 5: 647. 1897. Vine; leaf blades ovate to oblong-ovate, to 9 × 6 cm, sparingly appressed-puberulous; bracteoles green, ovate, rounded to acute and apiculate at the apex; corolla red, 30 mm long; drupe purplish black, slightly flat-

Mendoncia 553

Fig. 472. Mendoncia hoffmannseggiana

Fig. 473. Mendoncia cardonae

Fig. 474. Mendoncia bivalvis

554

M ENDONCIACEAE

tened, ca. 20 × 10 mm, glabrous. Dense lowland and montane forests especially along rivers, 100–1000 m; Bolívar (Divina Pastora), Amazonas (scattered). Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia. Mendoncia steyermarkii Wassh., Ann. Missouri Bot. Gard. 76: 1120. 1989. Vine; leaf blades 9–11 × 4–6.5 cm, ovate, abruptly acuminate at the apex; bracteoles ovate, apiculate, minutely muricate; corolla deep red, glabrous, upper lip ca. 5 mm long, lobes 3.5 mm wide; drupes subglobose, black, shining, 15–18 mm diameter, glabrous. Mon-

tane forests, 1100–1400 m; Bolívar (La Escalera-Cerro Venamo region). Endemic. Mendoncia williamsii Wassh., Ann. Missouri Bot. Gard. 76: 1121. 1989. Vine; leaf blades to 15 × 9 cm, apically short-acuminate with a cusp to 5 mm long, the upper surface drying blackish, hirtellous; pedicels densely brown-strigose; bracteoles gradually narrowed to an acute tip; drupes 15 × 6 × 4–5 mm, oblique at the tip, pubescent. Nonflooded evergreen lowland forests, 100–200 m; Amazonas (Capibara, Río Casiquiare). Endemic.

MENISPERMACEAE by Rupert C. Barneby Herbaceous vines or woody lianas, rarely small trees or subshrubs, sometimes sarmentose. Stems cylindric or if woody sometimes anomalously thickened on 2 sides and becoming ribbon-like; indument of simple trichomes or lacking. Leaves alternate; exstipulate; petiole often swollen or pulvinulate at each end; blade unlobed or all or some palmately lobed or parted, entire, either palmately veined from base or pinnately veined. Inflorescences either serially supra-axillary or cauliflorous, spicate, racemose, or paniculate, with or without leaves. Flowers bracteate, mostly small, greenish, whitish, or dull reddish, unisexual (plants dioecious). Perianth commonly imbricate; sepals 6–many, free or partly united; petals 6 or lacking (in Cissampelos the pistillate perianth of 1 sepal and 1 petal and the staminate perianth 4-merous). Androecium of staminate flower usually 3- or 6-merous, less often 12–more- or 1-merous; filaments free or united into a column; anthers bilocular, dehiscent by slits, the slits essentially vertical but often tilted by dilation or flexion of either filament or connective and appearing obliquely extrorse, obliquely introrse, or horizontal. Androecium of pistillate flower represented by staminodes or absent. Carpels of pistillate flower commonly 3 (in Sciadotenia 6 or more, in Cissampelos 1), sessile on a dilated torus or rarely stipitate or elevated on a stipe-like gynophore, each biovulate but 1-seeded; stigma tongue-shaped; ovary after fertilization commonly asymmetrically dilated on abaxial side only and becoming horseshoe-shaped or annular, the style scar remaining sub-basal, but in some genera symmetrical and straight. Fruit a drupe with membranous (red, yellow) or coriaceous exocarp; mesocarp often mucilaginous; endocarp crustaceous, bony, or woody, either smooth externally or often engraved or ornamented with papillae or processes, the adaxial face often intruded from various directions into the seed cavity as a septum or as an externally concave projection (condylus). Endosperm either continuous, ruminate, or absent; embryo conforming to the curvature of seed cavity and to the condylus when present, the radicle very short, the cotyledons either vermiform, or thick and horny and pressed face to face, or foliaceous and divaricate in 1 plane.

M ENISPERMACEAE 555

Pantropics, very few temperate; ca. 70 genera and 400 species, 12 genera and 27 species in the flora area. There is evidence of an undescribed species or genus of the family in the form of incomplete pistillate material from the base of Cerro Duida (Liesner 17838) and perhaps of a barren shoot (Gentry 46854) from Sierra de la Neblina base camp. Characters include leaves oblong-elliptic, acuminate, pinnately veined; pistillate inflorescences axillary, short, simply racemose, the flowers solitary on a short obese pedicel; perianth of 3 small outer and 3 larger sepals; petals none; staminodes linear, compressed, truncate, bearing sterile locules across the top; carpels 3, densely pallidpubescent. The foliage suggests Telitoxicum, but the pedicels and the apetalous flowers are discordant; lack of petals suggests Abuta, but the leaf venation would be discordant there. Pistillate flowering Menispermaceae are best identified by comparison with a well-curated collection. Key to the Genera of Menispermaceae 1. 1. 2(1).

Plant staminate .......................................................................................... 2 Plant pistillate and bearing fruit ............................................................. 13 Perianth and androecium 4-merous, the 4 petals connate into a membranous cup around a 4-merous synandrium .......................... 5. Cissampelos 2. Perianth and androecium in cycles of 3, the petals mostly free or absent, but if united of fleshy texture ................................................................ 3 3(2). Sepals imbricated in 4 or more cycles, forming a cone .......... 11. Sciadotenia 3. Sepals in 2 or 3 cycles of 3 ......................................................................... 4 4(3). Petals absent .................................................................................. 1. Abuta 4. Petals present ............................................................................................. 5 5(4). Leaf blades whitened beneath with a dense coat of minute, matted trichomes, these not individually perceptible without magnification ..................................................................................................... 6. Curarea 5. Leaf blades usually glabrous, or thinly pubescent, or loosely tomentulose with clearly visible trichomes ................................................................ 6 6(5). Flowers sessile on a simple primary axis ................................ 7. Disciphania 6. Flowers pedicellate, variously racemose or paniculate ............................ 7 7(6). Innermost cycle of sepals valvate in bud; petals fleshy (unknown in Caryomene), forming a pseudodisk around the androecium (genera more easily distinguished by their fruits) ............................................. 8 7. Innermost cycle of sepals imbricate in bud; petals membranous, not forming a pseudodisk .................................................................................. 10 8(7). Leaf blades broadly rounded at apex ........................................ 4. Caryomene 8. Leaf blades at least shortly acuminate at apex ........................................ 9 9(8). Stronger secondary veins 1 or 2 pairs, arising from midrib at the very base of the leaf blade ..................................................... 2. Anomospermum 9. Stronger secondary veins several pairs, arising from midrib distal to base of leaf blade .......................................................................... 10. Orthomene 10(7). Leaf blades pinnately veined, the lateral veins arising at insertion of petiole similar to succeeding pairs; inflorescence a simple raceme, arising from wood at least 1 year old, below leaves of the current year ............................................................................................. 12. Telitoxicum

556

M ENISPERMACEAE

10.

11(10). 11. 12(11). 12. 13(1). 13. 14(13). 14. 15(14).

15.

16(13). 16. 17(16). 17. 18(17). 18. 19(18).

19.

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

21.

22(17). 22.

Leaf blades palmately 3–7-veined from insertion of the petiole, the first lateral pair clearly longer and stronger than succeeding pairs; inflorescence wholly or in part axillary to leaves on branchlets of the current year or, if cauliflorous, then paniculate .............................................. 11 Inner sepals < 1 mm long ......................................................... 8. Hyperbaena Inner sepals > 1.5 mm long ...................................................................... 12 Filaments 6, free from base ........................................................ 3. Borismene Filaments 3 or 6, united into a synandrium .......................... 9. Odontocarya Seeds without endosperm; cotyledons fleshy or horny; embryo horseshoeshaped or annular; style scar contiguous to the pedicel .................... 14 Seeds with endosperm, this enveloping the small embryo; style scar distant from pedicel (except in Cissampelos) .......................................... 16 Carpels 6; drupes elevated on stipe-like gynophores coalescent proximally into a column ....................................................................... 11. Sciadotenia Carpels 3; drupes sessile or elevated on a column .................................. 15 Drupes elevated on a 3-rayed column, oblong-obovoid, pubescent; leaf blades minutely densely white-tomentulose on lower surface ..................................................................................................... 6. Curarea Drupes sessile on the torus, narrowly obovoid, cuneately narrowed at base, or compressed orbicular, glabrous; leaf blades glabrous or thinly pilosulous on lower surface .................................................. 8. Hyperbaena Carpel 1 .................................................................................... 5. Cissampelos Carpels 3, if some abortive, an attachment scar visible on torus .......... 17 Endosperm ruminate, formed of separable lamellae .............................. 18 Endosperm continuous ............................................................................ 22 Seed inversely U-shaped, molded over a septum vertically intruded from base of drupe ........................................................................................ 19 Seed either straight or J-shaped, folded or molded over a septiform condylus intruded from the drupe’s ventral side ................................ 21 Testa of endocarp unarmed internally; lamellae of endosperm separated by membranous integuments; inflorescence racemosely several-flowered ....................................................................................................... 20 Testa of heavily ligneous endocarp armed around interior wall of the cavity with rows of compressed teeth interdigitating with lamellae of endosperm, these lacking integument; inflorescence of solitary axillary flowers .................................................................................... 4. Caryomene Leaf blades pinnately veined .................................................. 12. Telitoxicum Leaf blades 3–5-veined from insertion of petiole .............................. 1. Abuta Condylus horizontal to the seed’s long axis, arising midway between pedicel and style scar and descending obliquely into the seed cavity, the embryo conformably J-shaped in vertical section; style scar lateral ........................................................................................ 2. Anomospermum Condylus vertical to the seed’s long axis, arising from the whole length of the cavity’s adaxial angle and intruded halfway across the cavity to occupy a narrow groove in the ventral face of the straight endosperm; style scar terminal ............................................................... 10. Orthomene Drupes sessile on a simple primary axis; endocarp 8-winged lengthwise, dorsiventrally symmetrical like the embryo ....................... 7. Disciphania Drupes pedicellate, either racemose or paniculate; endocarp variously

Abuta 557

armed or smooth externally, dorsiventrally asymmetrical, the testa intruded into the cavity from the ventral face as a septum or condylus .............................................................................................................. 23 23(22). Seed molded over a hollow condylus that appears externally as a saucer-, cup-, or trough-shaped depression, sometimes covered over as a hollow chamber ............................................................................... 9. Odontocarya 23. Seed folded lengthwise around a septiform condylus that appears externally as a shallow groove ........................................................ 3. Borismene 1. ABUTA Aubl., Hist. Pl. Guiane 619, t. 250. 1775. Woody lianas, some of great size, flowering in the forest canopy or in festoons hanging from riparian woods, a few shrubby but potentially sarmentose, the young growth either glabrous, strigulose-pilosulous, or densely tomentose. Leaf blades ovate, elliptic, or obovate, often acuminate, entire, 3- or 5-veined from the petiole and with 1 or more major pairs of secondary veins distally. Inflorescences axillary or serially supra-axillary on branches of current year, the fruits sometimes persistent below foliage, the staminate inflorescence branched, the flowers shortly racemose or subcymose on branchlets of second order, the pistillate inflorescence simply racemose. Staminate flower: sepals (disregarding accessory bracteoles) 6, in 2 heteromorphic cycles, the inner cycle imbricate in bud; petals absent; stamens 6, the outer cycle sometimes obsolescent, the filaments either free or partly connate, straight or incurved, linear or dilated distally, the anthers collateral or separated by thickened connective, either lateral and then either introrse or extrorse, or terminal and either vertically or obliquely dehiscent. Pistillate flower: perianth as in the staminate flower; staminodes (3)6; carpels 3. Drupe sessile, oblong-obovoid, subcompressed, with almost basal style scar; exocarp mealy-coriaceous; mesocarp watery or scanty; endocarp horseshoe-shaped, the testa either crustaceous or ligneous, finely 3-sulcate its whole length and engraved-reticulate, 0.2–1.5 mm thick. Embryo vermiform, embedded in laminate endosperm, folded over a straight vertical septiform condylus entering from the base. Tropical forests from southern Mexico to Bolivia and southeastern subtropical Brazil (most diverse in Amazonia and Guyana, Suriname, and French Guiana); ca. 25 species, 9–10 in Venezuela, 7 of these in the flora area. The taxonomy of Abuta has suffered from spasmodic, rather than systematic revision, from innate difficulty in procuring securely matched staminate and fruiting material, and from reckless identification of sterile specimens. The fruits are too uniform to provide strong specific characters, and the leaves are more variable in shape and pubescence than has been admitted. The sometimes distinctive androecia are relatively seldom available. A record from Venezuelan Guayana of A. mycetandra Krukoff & Barneby was based on sterile stems plausibly referable to Abuta pahni. A record of A. candollei is mentioned below under A. grisebachii. Abuta rufescens, A. pahni, and probably others furnish ingredients for curare dart poisons. Key to the Species of Abuta 1. 1.

Leaf blades glabrous on both surfaces ...................................................... 2 Leaf blades either pilose or densely tomentose on lower surface (species much confused and difficult to separate without staminate flowers) ..... 5

558

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

M ENISPERMACEAE

Leaf blades small, < 10 cm long, obovate to oblanceolate, obtuse ..................................................................................................... A. obovata Leaf blades either larger, some > 10 cm long, or sharply acuminate, or both ......................................................................................................... 3 Inflorescence of both sexes and carpels glabrous (even when young), pedicels filiform-ascending ................................................... A. grandifolia Inflorescence of both sexes and young carpels puberulent; pedicels short and recurved .......................................................................................... 4 Pedicel of staminate flowers recurved; young carpels puberulent or velutinous, ripe drupe glabrous .................................................... A. imene Pedicel of staminate flowers erect; drupe gray-velvety ................ A. velutina Upper surface of leaves finely minutely reticulate (the areoles about 0.1– 0.2 mm diameter) ........................................................................... A. pahni Upper surface of leaves plane between primary and secondary veins .... 6 Inner sepals of staminate flowers 2–3.5 mm long ....................... A. rufescens Inner sepals of staminate flowers ca.1.5 mm long .................... A. grisebachii

Abuta grandifolia (Mart.) Sandw., Bull. Misc. Inform. Kew 1937: 397. —Cocculus grandifolius Mart. in Buchner, Repert. Pharm. 36: 345. 1830. Abuta guyanensis Eichler in Mart., Fl. Bras. 13(1): 181, pl. 42, fig. 2. 1864. Weak bushy shrub 1.5–20 m tall, sometimes sarmentose or scandent; drupe yellowish when ripe, the thin mesocarp edible. Rocky river banks, islands in river rapids, cliffs, forest margins and tree islands in savannas, 100–900 m; western and northern Bolívar, Amazonas (frequent on upper Río Orinoco and Río Negro). Common and widespread throughout Amazon Basin, from eastern Colombia to Guyana, Suriname, French Guiana, and south to Bolivia and the Brazilian Planalto. ◆Fig. 475. Abuta grisebachii Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 47. 1862. Liana to 30 m or more, flowering in forest canopy. Lower montane forests, ca. 400–900 m; southern Bolívar (Sierra Pakaraima), Amazonas (upper Orinoco and slopes of Sierra Parima). Scattered over northern half of the Amazon Basin in southeastern Colombia, northern Brazil, northeastern Peru. The Venezuelan specimens, all sterile and therefore only tentatively referable to Abuta grisebachii, have been referred to species not yet certainly recorded from the Guayana: A. mycetandra Krukoff & Barneby (Steyermark 89029), and A. candollei Triana & Planch. (Steyermark 117774), this very similar vegetatively, but sharply distinguished by the

sepals of the staminate flower being all of nearly equal length but the outer cycle much narrower than the inner one. In A. grisebachii the sepals are in 2 very unequal cycles, the inner much long than the outer. Abuta imene (Mart.) Eichler, Flora 47: 389. 1864. —Cocculus imene Mart. in Buchner, Repert. Pharm. 36: 341. 1830. —Hoja de mono. Liana to 20–35 m high, flowering in the canopy. Primary evergreen and gallery forests, 100–1100 m; Bolívar (Río Nichare, Gran Sabana), Amazonas (upper Río Negro). Widespread in Amazon Basin, north into French Guiana. ◆Fig. 476. Abuta obovata Diels, Notizbl. Bot. Gart. Berlin-Dahlem 13: 20. 1936. —Capuyyen-cu-me-peu, Huatacurán. Liana to 15–25 m long; fruit yellow with sweet translucent pulp. Nonflooded evergreen and gallery forests, 100–1300 m; Bolívar (Gran Sabana), Amazonas (upper Río Negro and Río Orinoco headwaters). Discontinuously scattered through Guyana, Suriname, French Guiana, northern and western Brazilian Amazonia from Amapá just into Peru and southwest just into Bolivia. ◆Fig. 478. Abuta pahnii (Mart.) Krukoff & Barneby, Mem. New York Bot. Gard. 22(2): 43. 1971. —Cocculus pahnii Mart., Flora 24(Beibl. 2): 45. 1841. —Chinak. Woody liana, commonly 10–30 m; Nonflooded forests of valley lowlands and foot-

Abuta 559

hills, 100–1200 m; Bolívar (Gran Sabana), Amazonas (upper Orinoco valley). eastern Venezuela to Zulia; widespread in western Amazonia, from Ecuador to northeastern Bolivia and western Brazil. ◆Fig. 479. This species is closely akin to A. rufescens but distinguished by the characters in the key and by the much smaller staminate flowers. Abuta rufescens Aubl., Hist. Pl. Guiane 618. 1775. Abuta splendida Krukoff & Moldenke, Bull. Torrey Bot. Club 68: 241. 1941. Abuta wilson-brownei R.S. Cowan, Brittonia 7: 394. 1952. Woody liana attaining the canopy. Primary forests, both in nonflooded lowlands and gallery forests of upland savanna country, 50–800 m; northern and southern Bolívar (Río Acanán, Río Caura), southern Amazonas (Río Cunucunuma). Apure; Guy-

Fig. 475. Abuta grandifolia

ana, Suriname, French Guiana, throughout Amazon Basin. ◆Fig. 477. Abuta velutina Gleason, Bull. Torrey Bot. Club 58: 361. 1931. Liana. Evergreen lowland forests, 100– 400 m; Bolívar (mouth of Río Caura), Amazonas (La Esmeralda, Río Cunucunuma, Río Parú). Widespread in central and western Amazon Basin in Ecuador, Peru, and Brazil.

560

M ENISPERMACEAE

Fig. 476. Abuta imene

Abuta 561

Fig. 477. Abuta rufescens

Fig. 478. Abuta obovata

562

M ENISPERMACEAE

Fig. 479. Abuta pahnii

2. ANOMOSPERMUM Miers, Ann. Mag. Nat. Hist. ser. 3, 14: 101. 1864. High-climbing woody lianas, glabrous or yellowish setose, flowering from young growth in the forest canopy. Leaves chartaceous to coriaceous, 3–5-veined, distally pinnately veined, often reticulate, entire. Staminate inflorescence either cymose-paniculate or consisting of a succession of pseudoracemose, few-flowered units borne serially along either leafy or leafless branchlets. Staminate flower: sepals 6, the 3 outer small, the inner much larger, imbricate in bud; petals 6, either submembranous, or fleshy and forming a pseudodisk, partially encasing the 6 free stamens; filaments either erect or incurved; anthers either extrorse or introrse. Pis-

Anomospermum 563

tillate flowers solitary or paired in leaf axils, perianth as in the staminate flower, but androecium staminodal; carpels 3. Drupes (often solitary by abortion) sessile, obliquely obovoid or subglobose, but incurved through about 3/4 circle, the adaxial side short, the abaxial distended in a broad convex arc; exocarp mealy-coriaceous, orange when ripe; mesocarp thin or obsolete; endocarp crustaceous or ligneous, externally incised-venulose and often pitted, the inner wall either smooth or armed with intruded woody teeth corresponding with the external pits. Endosperm copious, ruminate, enclosing the vermiform embryo, molded over a laminar condylus descending into the seed cavity in a plane horizontal to the fruit’s axis, J-shaped in vertical section. Neotropics; 6 species, 2 in Venezuela, 1 of these in the flora area. Specimens from Delta Amacuro (Steyermark 114674, staminate, with immature leaves, and 114777, sterile) that have been referred to Anomospermum reticulatum (Mart.) Eichler, a species very widely dispersed to the west and south of Venezuelan Guayana, probably are juvenile Orthomene schomburgkii, difficult to distinguish without fruit. Anomospermum steyermarkii Krukoff & Barneby, Mem. New York Bot. Gard. 20(2): 30. 1970. Large vine. Wet, nonflooded forests, 500– 1300 m; Bolívar (Cerro Ichún, vicinity of Karaurín-tepui), Amazonas (Sierra de la Neblina, Sierra Parima). Brazil (Amapá, Amazonas, Roraima). ◆Fig. 480.

Fig. 480. Anomospermum steyermarkii

564

M ENISPERMACEAE

Fig. 481. Borismene japurensis

3. BORISMENE Barneby, Mem. New York Bot. Gard. 22(4): 144. 1972. Lianas flowering in forest canopy, glabrous throughout. Leaf blades ovate, entire, 3–5-veined from base, then reticulate. Inflorescences serially fasciculate in leaf axils or distally paniculate, the pedicellate staminate flowers borne mostly on secondary branchlets, the pistillate ones simply racemose. Staminate flower: sepals 9, membranous, strongly graduated, the inner imbricate in bud; petals 6, partially encasing the 6 free stamens; anthers vertically dehiscent. Pistillate flowers: perianth as in staminate flowers; carpels 3. Drupe sessile, obovoid; exocarp thinly coriaceous, red turning black; mesocarp a watery pulp; endocarp straight, the testa papery. Endosperm continuous, bony, folded lengthwise around a septiform condylus intruded from the ventral side of endocarp; embryo straight, the cotyledons foliaceous. Upper Amazon Basin in Brazil, Peru, and Bolivia, disjunct in Pacific lowland Colombia, in eastern Venezuelan Guayana, and in coastal Bahia, Brazil; 1 species. Borismene japurensis (Mart.) Barneby, Mem. New York Bot. Gard. 22(4): 145, fig. 2. 1972. —Cocculus japurensis Mart., Flora 24(2) Beibl.: 44. 1841. —Anomospermum japurense (Mart.) Eichler, Flora 47: 388. 1864. —Traga venado, Tragwena.

Hyperbaena cuatrecasasii Moldenke, Phytologia 2: 134. 1946. High-climbing liana. Lowland, nonflooded forests, 100–300 m; Delta Amacuro-Bolívar border (Sierra Imataca). Distribution elsewhere as in genus. ◆Fig. 481.

4. CARYOMENE Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 52. 1971. High-climbing woody lianas flowering in the forest canopy, glabrous throughout. Leaf blades simple, 3–5-veined from the base, the secondary veins from midrib incurved-ascending. Staminate inflorescence unknown; pistillate flowers solitary or paired in the axil of leaves. Drupe massive, subsessile, obliquely obovoid; exocarp stiffly coriaceous; mesocarp mucilaginous; endocarp thickly woody, smooth or dimpled externally, produced internally in the form of teeth or plates interdigitating with endosperm, this lamellate, but the lamellae lacking individual integument.

Cissampelos 565

Seed horseshoe-shaped, folded over a septiform condylus intruded from base of drupe; embryo linear-vermiform, terete. Southeastern Venezuela, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil; 5 or 6 species, 1 in Venezuela. Caryomene olivascens Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 57. 1971. —Li-wadi-wain-mue. Liana to 30 m. Tall riparian forests, ca. 200 m; Amazonas (Río Cunucunuma). French Guiana, northeastern Brazil. ◆Fig. 482.

Fig. 482. Caryomene olivascens

5. CISSAMPELOS L., Sp. Pl. 1031. 1753. Weak subherbaceous or slender, woody, twining vines and one subshrub with stiffly erect stems, setose, softly puberulent or tomentellous, or glabrous. Leaf blades either ovate-cordate to ovate-suborbicular and palmately veined from base or peltate and radially veined from insertion of petiole. Inflorescence either axillary to leaves or cauliflorous, solitary or commonly fasciculate, the staminate composed of lax corymbs in a compound panicle, the pistillate of condensed corymbs in a pseudoraceme, often leafy-bracteate. Staminate flower: actinomorphic; sepals 4, distinct; petals membranous, united into a campanulate or shallow cup; stamens 4, the filaments united into a column, this sometimes very short, the anthers sessile, extrorse, horizontally dehiscent. Pistillate flower: zygomorphic; carpel solitary, subtended on abaxial side by 1 sepal and 1 petal, these early deciduous. Drupe berrylike; exocarp thinly coriaceous; mesocarp juicy; endocarp crustaceous, horseshoeshaped or annular, laterally compressed, vertically 6-keeled on periphery and sides, the keels coarsely undulate or verrucose, the style scar basal. Embryo enveloped in endosperm, linear. Pantropics; ca. 20 species, 6 in Venezuela, 4 of these in the flora area. Some of the species in this genus are not satisfactorily defined. The twining species vary much in pubescence and leaf outline, and specimens are often named only with difficulty.

566

M ENISPERMACEAE

Fig. 483. Cissampelos ovalifolia

Fig. 484. Cissampelos pareira

Fig. 485. Cissampelos andromorpha

Curarea 567

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

3.

Stems stiffly erect from woody root-stock, mostly < 1 m tall; petioles stout, < 1.5 cm long ............................................................................ C. ovalifolia Stems twining or scandent, > 1 m tall; petioles slender, most of them ≥ 2 cm ................................................................................................................ 2 Leaf blades strongly peltate, the petiole entering the blade 5–30 mm from its base ................................................................................ C. tropaeolifolia Leaf blade basifixed or only shallowly peltate, the petiole entering the blade < 5 mm from its base.................................................................... 3 Staminate inflorescence elongate, the primary axis giving rise to a succession of cymose branchlets; pistillate inflorescence usually lacking foliaceous bracts; inflorescences of both sexes often (but not always) cauliflorous, either immediately or far below the leaves ........ C. andromorpha Staminate inflorescence short, often no longer than wide, the primary axis giving rise to lateral branchlets at 1–3 nodes; pistillate inflorescence commonly bearing foliaceous bracts; inflorescences of both sexes normally axillary to leaves .......................................................... C. pareira

Cissampelos andromorpha DC., Syst. Nat. 1: 1: 539. 1818 [1817]. —Barbasco lagartilla, Baruta de lagartillo, Mucumi, Pica pica. Cissampelos ramiflora Miers, Contr. Bot. 3: 163. 1871. Vine with trunk rarely to 5 mm diameter, over trees and bushes. Primary and disturbed forests, 100–1300 m; Delta Amacuro (Río Amacuro), Bolívar (Gran Sabana, Río Cuyuní basin), Amazonas (upper basins of Río Negro, Río Orinoco, and Río Ventuari). Common and widely dispersed from Guatemala to Bolivia, Guyana, Suriname, French Guiana, and extratropical southeastern Brazil. ◆Fig. 485. Cissampelos ovalifolia DC., Syst. Nat. 1: 1: 537. 1818 [1817]. —Aspuela-guara, Oreja de tigre. Functionally herbaceous subshrub densely pilose-tomentose throughout, the erect annual stems rising from a woody base resistant to fire. Savannas and forest openings, 100–1300 m; scattered but locally abundant in southeastern Delta Amacuro, Bolívar, and

Amazonas. Scattered from Colombia to Guyana, south to Bolivia and southeastern Brazil. ◆Fig. 483. Cissampelos pareira L., Sp. Pl. 1031. 1753. —Pareira-brava. Climbing over shrubs and low trees. Moist or seasonally dry shrub-woodlands, becoming weedy in disturbed scrub vegetation, 50– 500 m; Bolívar (Gran Sabana, lower Río Orinoco valley), northern and central Venezuela; circumtropical. ◆Fig. 484. This is the Pareira-brava of the pharmacopeias. Cissampelos tropaeolifolia DC., Syst. Nat. 1: 1: 532. 1818 [1817]. Climbing, variable in pubescence. Moist woodlands, savanna thickets, 100–300 m; northern and southeastern Bolívar. western Coastal Range; widely dispersed from southern Mexico to Brazil and Bolivia. An almost glabrous form from Upata has been mistaken for Cissampelos glaberrima St. Hil.

6. CURAREA Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 7. 1971. Woody lianas flowering in or below the forest canopy. Leaf blades ovate-subcordate, densely minutely white- or sordid-tomentellous on lower surface, palmately 3– 9-veined from base, the midrib giving rise to 1 or 2 pairs of major secondary veins. Inflorescence of both sexes either cauliflorous or subtended by leaves, either racemose-paniculate or the pistillate simply racemose. Staminate flower: sepals 6, firm,

568

M ENISPERMACEAE

silky-pubescent dorsally, the outer cycle minute, the inner cycle valvate in bud; petals 6, membranous; androecium 6-merous, the usually clavate filaments free, the anthers subvertically dehiscent. Pistillate flower: sepals as in staminate flowers, but petals commonly 3; staminodes absent; carpels 3. Drupes elevated on and deciduous from radially ascending, stoutly linear or drum-shaped gynophores, oblong-ellipsoid, horseshoe-shaped, the style scar almost basal; exocarp pubescent; mesocarp mealy or filamentous; testa of endocarp chartaceous, almost smooth. Endosperm absent; cotyledons horny. Neotropics; 4 species, 2 in Venezuela, both in the flora area. Key to the Species of Curarea 1.

1.

Staminate flowers with inner sepals 1.2–1.5 mm long and petals glabrous; gynophore of mature drupes drum-shaped, ca. 1 mm long, wider than long .......................................................................................... C. candicans Staminate flowers with inner sepals 1.5–2.3 mm long and petals puberulent externally; gynophore of mature drupes linear-claviform, 3–6 mm long, much longer than wide ................................................... C. toxicofera

Curarea candicans (Rich. ex DC.) Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 12. 1971. —Abuta candicans Rich. ex DC., Syst. Nat. 1: 1: 543. 1818 [1817]. —Chondrodendron candicans (Rich. ex DC.) Sandwith, Bull. Misc. Inform. Kew 1930: 342. 1930. —Shina-teu. Abuta limaciifolia Diels in Engl., Pflanzenr. IV. 94(46): 194. 1910. —Chondro-

Fig. 486. Curarea candicans

dendron limaciifolium (Diels) Moldenke, Brittonia 3: 20. 1938. Liana to 30 m or more, but often smaller. Lowland nonflooded forests, elevation not indicated; rare in Bolívar (near El Dorado) and

Disciphania 569

Amazonas (Río Cunucunuma, between Cerro Duida and Cerro Huachamacari, Steyermark 125686). Guyana, Suriname, French Guiana, northeastern Brazil (Pará, Amapá). ◆Fig. 486. Records for this species in the flora area need confirmation from fertile material. Stems and foliage are used in preparation of dart poison. Curarea toxicofera (Wedd.) Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 9, fig. 1. 1971. —Cocculus toxico-

ferus Wedd. in Castelnau, Exped. Part. Cent. Amer. Sud. 22. 1851. —Chondrodendron toxicoferum (Wedd.) Krukoff & Moldenke, Brittonia 3: 21. 1938. Cauliflorous liana. Nonflooded primary forests, disturbed woodlands, sometimes riparian, 50–400 m; Amazonas (near Puerto Ayacucho). Widely dispersed from Panama to northeastern Bolivia, most common in middle and upper Amazonia. Curarea toxicofera is an important ingredient of some curares.

7. DISCIPHANIA Eichler in Mart., Fl. Bras. 13(1): 168. 1864. Slender vines, the old trunks often clothed in furrowed corky bark, the foliage glabrous, pilosulous, or rarely setose. Leaf blades either unlobed, palmately lobed or 3–5-foliolate, a few species peltate, the venation primarily palmate or radiate. Inflorescence supra-axillary on branches of current year and simply spicate, less often cauliflorous, or branched, or racemose, but the flowers sessile in all but few North American species. Staminate flower: perianth of 6 subequal sepals, these either free or connate to middle, either ascending or rotate; petals 6, rarely 3, these polymorphic in shape, texture and orientation, in the flora area obdeltate, fleshy and crowded into a pseudodisk; androecium 3-merous, the filaments free, often dilated, the anthers vertically or obliquely dehiscent. Pistillate flower: perianth as in the staminate flower, sometimes smaller; staminodes absent; carpels 3. Drupes oblongellipsoid; exocarp coriaceous, red; mesocarp mucilaginous; endocarp ovoid-ellipsoid, straight, dorsiventrally compressed, the coriaceous or crustaceous testa 8-ribbed lengthwise and the ribs commonly winged or fringed, the cavity dorsoventrally symmetrical, without condylus; cotyledons foliaceous, divaricate in 1 plane between plates of continuous endosperm. Southern Mexico and Antilles to warm-temperate southeastern Brazil and Paraguay; ca. 25 species, 1 in Venezuela. The genus is well characterized by the straight, winged or ribbed endocarp but is polymorphic in leaves and perianth. Disciphania ernstii Eichler, Jahrb. Königl. Bot. Gart. Berlin 2: 329, pl. 2. 1883. Widely dispersed in northern and western South America; several varieties, 1 in Venezuela. Disciphania ernstii is sometimes cultivated for its edible drupes. D. ernstii var. ernstii Vine to 12 m high; leaves sometimes dimorphic, some cordate, entire, and green, others pentagonal or deeply lobed and whitemottled. Over trees, below 1000 m; presumably in Amazonas (collected once without precise locality [Croizat 386 (NY)] on the upper Orinoco). Venezuelan Coastal Cordillera; widespread from eastern Panama to southern Brazil, Paraguay. ◆Fig. 487.

Fig. 487. Disciphania ernstii var. ernstii

570

M ENISPERMACEAE

Fig. 488. Hyperbaena domingensis

Odontocarya 571

8. HYPERBAENA Miers ex Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 47. 1861. Trees, shrubs (some xeromorphic), and woody vines with twining branchlets, flowering in the forest canopy, either glabrous or pilosulous. Leaf blades ovate, obovate, or elliptic, entire or dentate, the venation either pinnate or 3-veined and thereafter pinnate. Inflorescences either axillary, or terminally and laterally paniculate, or rarely cauliflorous, the staminate of various complexity, the pistillate either racemose or spicate. Staminate flower: sepals 6, the 3 inner larger, imbricate in bud; petals usually 6, rarely absent; stamens 6, the filaments free. Pistillate flower: perianth as in the staminate flower; carpels usually 3, glabrous. Drupe sessile, obovoid or compressed orbicular, the style scar almost basal; exocarp green, orange-yellow, purplish red, or finally black, coriaceous or mealy, glabrous; mesocarp usually thin; endocarp either horseshoe-shaped or annular, the bony embryo folded over a vertical, either septiform or clavate condylus. Endosperm absent. Neotropics; 19 species, 1 in Venezuela. Hyperbaena domingensis (DC.) Benth., J. Proc. Linn. Soc., Bot. 5(suppl. 2): 50. 1861. —Cocculus domingensis DC., Syst. Nat. 1: 1: 528. 1818 [1817]. —Anelasma domingense (DC.) Miers, Ann. Mag. Nat. Hist. ser. 2, 7: 43. 1851. —Pachygone domingensis (DC.) Eichler in Mart., Fl. Bras. 13(1): 197. 1864. Hyperbaena graciliflora Miers, Contr. Bot. 3: 295. 1871. —Pachygone graciliflora (Miers) Sagot, Ann. Sci. Nat. Bot. sér. 4,

11: 141. 1881. Forest vine; leaves ovate, reticulate; inflorescences effuse cymose-paniculate; flowers minute. Forest canopy or draped over riparian shrubs, tropical lowlands, cloud forests, elevation not indicated; Bolívar (Altiplanicie de Nuria), Amazonas (upper Río Negro). Coastal Range; Greater Antilles, Colombia south to Bolivia, Guyana, Suriname, French Guiana, eastern and southeastern Brazil. ◆Fig. 488.

9. ODONTOCARYA Miers, Ann. Mag. Nat. Hist. ser. 3, 14: 99. 1864. Softly woody vines with terete stems, the old wood smooth, the papery epidermis often exfoliating, the foliage glabrous or pilosulous. Leaf blades entire, ovate, cordate, or angulate, 3–5-veined from base and thereafter pinnately veined, the undersurface sometimes with pockets or discolored areoles in angles of primary veins. Inflorescence either pseudoracemose or variously branched, either axillary to leaves or cauliflorous and remote from them, the flowers of both sexes pedicellate, the staminate usually fasciculate, rarely solitary, the pistillate always solitary. Staminate flower: sepals 6, membranous, the outer cycle short, the inner at least twice as long, imbricate in bud; petals 6, rarely 3, involute or ventrally grooved; androecium (1)3- or 6-merous, the filaments rarely all free, the inner 3 or all 6 commonly united part-way or up to the anthers into a synandrium; anthers dehiscent by vertical or rarely transverse slits. Pistillate flower: perianth as in the staminate flower; staminodes 3–6; carpels 3, glabrous. Drupes often solitary by abortion of 2 carpels, sessile, straight, globose-ellipsoid subterete; exocarp coriaceous, green becoming white, yellow, or red; mesocarp either mucilaginous or fibrous, either deciduous from or persistent on the crustaceous or woody endocarp, this dorsoventrally asymmetrical, dorsally rounded and vertically keeled, smooth, papillate or cristate, ventrally depressed and excavated into a shallow or deeply cupulate condylus, at each end 3-toothed. Embryo straight; cotyledons foliaceous, divaricate in one plane between plates of continuous endosperm. Neotropics, extratropical southeastern Brazil and northern Argentina; ca. 35 species, 7 in Venezuela, 3 of these in the flora area.

572

M ENISPERMACEAE

As currently understood, Odontocarya is precariously distinct from the Paleotropic Tinospora Miers. The largely Amazonian O. floribunda Diels, resembling O. mallosperma in twice-racemose staminate inflorescence associated with leaves, but differing in leaves cordate at base and staminate flowers solitary at each node of the panicle, has been collected in the western Orinoco basin in Apure and is to be sought in northwestern Bolívar or adjoining Amazonas. Key to the Species of Odontocarya 1.

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

3.

4(2).

4.

Lower surface of leaf blades differentiated in axils of primary veins into a discolored patch; flowers of both sexes borne on primary inflorescence axis, the staminate solitary and fasciculate, the pistillate all solitary; leaf blades cordate or sagittate ................................................ O. tamoides Lower surface of leaf blades not differentiated; flowers or some of them borne on secondary axes of inflorescence; leaf blades various ............. 2 Specimen with staminate flowers ............................................................. 3 Specimen with pistillate flowers or fruits ................................................. 4 Inflorescence arising from branchlets of current year, accompanied by scattered leaves, and the flowers borne on branchlets of second order; synandrium 6-merous ........................................................ O. mallosperma Inflorescence arising from leafless trunk distant from leaves in the canopy, and the flowers borne on branchlets of the third order; synandrium 3-merous ..................................................................... O. sp. A Inflorescence arising from branchlets of current year, the drupes racemose along its primary axis; leaf blades 4.5–7 cm long, cuneate at base; endocarp clothed in persistent matted fibers ................... O. mallosperma Inflorescence from leafless rope-like stem distant from leaves in canopy, the drupes racemose on secondary axes of a panicle; leaf blades unknown (but very likely > 10 cm); endocarp unknown (but very likely smooth) ............................................................................................ O. sp. A

Odontocarya mallosperma Barneby, Mem. New York Bot. Gard. 20(2): 102. 1970. Liana to 20 m. Humid riverine forests, ca. 200–400 m; scattered in Bolívar. Apure; disjunct in northeastern Brazil. ◆Fig. 489. Odontocarya tamoides (DC.) Miers, Ann. Mag. Nat. Hist. ser. 3, 14: 100. 1864. —Cocculus tamoides DC., Syst. Nat. 1: 1: 521. 1818 [1817]. Weak vine. River banks, seashores, edges of moist lowland forests, sometimes weedy in disturbed brush-woodlands, near sea level to 200 m; scattered in Delta Amacuro, Bolívar, Amazonas. Widely and discontinuously dispersed in Neotropics; 2 varieties, both in the flora area.

Key to the Varieties of O. tamoides 1. Leaf blades pubescent throughout or along veins of undersurface .................... ...................................... var. canescens 1. Leaf blades glabrous on both surfaces ....................................... var. tamoides O. tamoides var. canescens (Miers) Barneby, Mem. New York Bot. Gard. 20: 91. 1970. —Odontocarya hederifolia var. canescens Miers, Contr. Bot. 3: 64. 1871. Cocculus pauper Griseb., Abh. Konigl. Ges. Wiss. Göttingen 7: 162. 1857. —Odontocarya paupera (Griseb.) Diels, Pflanzenr. IV. 94(46): 172. 1910. Delta Amacuro, northern Bolívar (lower Río Orinoco), Amazonas (upper Río Negro).

Odontocarya 573

Fig. 489. Odontocarya mallosperma

Fig. 490. Odontocarya tamoides var. canescens

574

M ENISPERMACEAE

Portuguesa, Zulia; Mexico, Lesser Antilles, Guyana, Suriname, French Guiana, Paraguay. ◆Fig. 490. This is the more common and widespread variety of the species. O. tamoides var. tamoides Delta Amacuro. Guyana, Suriname, French Guiana, Brazil (Pará). Odontocarya sp. A Amazonas. Four flowering collections,

staminate and pistillate, of cauliflorous Odontocarya are known from the headwaters of Río Orinoco and Río Negro. Plausibly but by no means certainly representing a single species, they remain unidentifiable, as all lack the characteristic leaves and fruits. A staminate inflorescence, likewise without leaves, collected at 60 m in rainforest along Río Botanamo in northeastern Bolívar, may represent O. wullschlaegelii (Eichler) Barneby, but cannot be identified positively without foliage.

10. ORTHOMENE Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 79. 1971. Woody vines of moderate dimensions, with twining terminal branchlets, either glabrous, puberulent or setose. Leaf blades firm, lustrous, 3–5-veined from petiole. Inflorescences borne on growth of current year or in 1 species cauliflorous, efoliate or randomly foliate, all simply pseudoracemose with serially supra-axillary pedicels, or the staminate twice-racemose, with flowers subcymosely racemose on secondary branchlets, in the cauliflorous species the whole inflorescence contracted into a glomerule. Staminate flower: sepals (not including subtending bracteoles) 6, the 3 outer very small, the inner much larger, firm or fleshy, imbricate in bud; petals 6, firm or fleshy, each involute around a filament, often forming a triangular pseudodisk; anthers tilted forward toward center of flower, dehiscent by extrorse slits. Pistillate flower: perianth as in staminate flower, but petals smaller and planer, and stamens sterile; carpels 3. Drupes sessile, ellipsoid or oblong-ellipsoid, straight or almost so, minutely apiculate by the style scar; exocarp fleshy becoming coriaceous, becoming black; mesocarp thin or almost absent; endocarp crustaceous,

Fig. 491. Orthomene schomburgkii

Sciadotenia 575

externally either smooth or pitted, inverted from the drupe’s adaxial side as a vertical laminar septum traversing about 1/3 the width of the seed cavity. Endosperm ruminate, folded around the septum; embryo vermiform, straight. Eastern Panama, tropical South America; 4 species, 3 in Venezuela, all in the flora area. Key to the Species of Orthomene 1.

1.

2(1).

2.

Branchlets of current year and axes of inflorescences yellow-setose; leaf blades bullate by impression of secondary veins; endocarp dimpled ......................................................................................................O. hirsuta Branchlets of current year and axes of inflorescence glabrous or almost so; leaf blades plane, smooth or finely reticulate; testa of endocarp smooth .................................................................................................... 2 Flowers scattered along new leafy branchlets or on efoliate branches of current year, both staminate and pistillate ones slenderly pedicellate; drupe smooth .................................................................... O. schomburgkii Flowers subsessile, clustered on knots along branches at least one year old; drupe verruculose ......................................................... O. verruculosa

Orthomene hirsuta (Krukoff & Moldenke) Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 81. 1971. —Anomospermum hirsutum Krukoff & Moldenke, Bull. Torrey Bot. Club 70: 401. 1943. Vine. Nonflooded lowland forests, ca. 100– 200 m; Amazonas (near northern base of Sierra de la Neblina). Scattered, middle to upper Amazonian Brazil and Peru. Orthomene schomburgkii (Miers) Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 80. 1971. —Anomospermum schomburgkii Miers, Contr. Bot. 3: 71. 1871. —Huevo de chiguire, Traga venado. Slender vine. Creek banks, riparian for-

ests, flooded or not, 50–1300 m; common in Delta Amacuro, Bolívar, and Amazonas. Eastern Venezuela; eastern Panama, Colombia, south to northeastern Bolivia and in Brazil to Rio de Janeiro. ◆Fig. 491. This is the most frequent Menispermaceae in the region, after Abuta grandifolia. Orthomene verruculosa (Krukoff & Barneby) Barneby & Krukoff, Mem. New York Bot. Gard. 22: 81. 1971. —Abuta verruculosa Krukoff & Barneby, Mem. New York Bot. Gard. 20: 24. 1970. Vine with orange-yellow drupes on old wood. Evergreen lowland rainforests, 100–200 m; Amazonas (Río Guayapo 149 km above mouth). Colombia, French Guiana, Brazil.

11. SCIADOTENIA Miers, Ann. Mag. Nat. Hist. ser. 2, 7: 43. 1851. Woody vines and sarmentose bushes, either glabrous, strigulose, or velvetytomentellous throughout. Leaf blades entire, ovate, lanceolate, or flabellate, 3–7veined from insertion to petiole, the secondary venation subhorizontally scalariform. Inflorescences borne on branchlets of current year, the staminate either pseudoracemose or racemose-paniculate, the pistillate ones either racemose or spicate, few- or 1-flowered. Staminate flower: sepals imbricate in (3)4–8 cycles of 3, externally puberulent or silky, the innermost cycle valvate in bud; petals 6, sometimes appendaged ventrally; stamens 6, the filaments all free or the inner ones connate into a synandrium, the connective dilated or not, the anthers dehiscent by oblique slits. Pistillate flower: perianth as in the staminate flower, but the androecium staminodal; carpels commonly 6, in one species 9–15. Drupes elevated from the torus

576

M ENISPERMACEAE

Fig. 492. Sciadotenia cayennensis

Fig. 493. Sciadotenia sprucei

Telitoxicum 577

on silky-puberulent gynophores connate proximally into a column, in outline obovoid with subbasal or exceptionally lateral style scar; exocarp subcoriaceous or membranous; mesocarp varying from thinly to copiously mucilaginous; endocarp coriaceous or crustaceous, commonly horseshoe-shaped, in 1 species incurved through 3/4 circle. Embryo of 2 plumply dilated cotyledons folded over a vertical septiform condylus, or in 1 species over an umbonate condylus; endosperm absent. Eastern Panama, Venezuela, Guyana, Suriname, French Guiana, Amazonia; ca. 18 species, 2 in Venezuela, both in the flora area. Sciadotenia is distinguished from other genera in the flora area by small conelike staminate flowers and by 6–15-carpellate pistillate flowers that give rise to drupes elevated on proximally confluent gynophores. Key to the Species of Sciadotenia 1.

1.

Innermost 3 sepals 1/3 longer than those next below; pistillate flower solitary on flexibly pendulous peduncle, 9–12(–15)-carpellate; drupe evenly incurved through ± 3/4 circle over dilated condylus ............... S. cayennensis Innermost 3 sepals not or scarcely longer than those next below; pistillate flowers shortly racemose, 6-carpellate; drupe horseshoe-shaped, folded over a septiform condylus ............................................................ S. sprucei

Sciadotenia cayennensis Benth., J. Proc. Linn. Soc., Bot. 5, suppl. 2: 51. 1861. Wiry liana, reaching forest canopy or smaller in riparian and disturbed brushwoodlands; drupes orange when ripe, darkening and wrinkled when dried. Nonflooded lowlands, ca. 100 m; Amazonas (Orinoco-Negro divide, sources of Río Atacavi). Guyana, Suriname, French Guiana, northeastern Brazil. ◆Fig. 492.

Sciadotenia sprucei Diels in Engl., Pflanzenr. IV. 94(46): 84. 1910. Slender liana; leaves lustrous, yellowtinged when dry. Margins of nonflooded forests, regenerating woodlands, 100–200 m; Amazonas (San Carlos de Río Negro). Scattered through the eastern and middle Amazon basin in Brazil, ascending the Río Negro into Colombia (Vaupés). ◆Fig. 493.

12. TELITOXICUM Moldenke, Brittonia 3: 42. 1938. High-climbing woody cauliflorous vines, glabrous except for often pubescent inflorescences, in most respects resembling Abuta, but the leaves pinnately veined and 6 petals present in flowers of both sexes. Inflorescences fasciculate on old wood below the current foliage, the slender, sometimes pendulous staminate ones narrowly paniculate, the flowers subcymosely racemose on short, filiform secondary branchlets, the pistillate ones simply racemose, with axes lignescent in fruit. Staminate flower: sepals 6, submembranous, glabrous, blackening when dry; petals 6, membranous, partially encasing the free stamens, imbricate in bud; anthers separated by dilated connective, dehiscent by vertical extrorse slits. Pistillate flower: perianth as in the staminate flower; staminodes 6; carpels 3. Drupes subsessile on a clavate torus, oblong-ellipsoid, variably oblique at base; exocarp thick, mealy-coriaceous; mesocarp almost absent; horseshoe-shaped endocarp, ruminate endosperm, and vermiform embryo as in Abuta. Neotropics; ca. 6 species, 1 in Venezuela. Some Amazonian species contribute to Indian dart poisons.

578

M ENISPERMACEAE

Telitoxicum inopinatum Moldenke, Brittonia 3: 46. 1938. —Awadu-di-venadu-ilu. (Yekwana). Vine; leaves stiff, lustrous, pinnately veined, incipiently bullate. Forests, savanna bush-islands, 50–300 m; Amazonas (upper Río Cunucunuma, Río Cataniapo). Guyana, Suriname. ◆Fig. 494.

Fig. 494. Telitoxicum inopinatum

MENYANTHACEAE by Kay Yatskievych Aquatic or semi-aquatic perennial or rarely annual herbs. Stems with a well-developed system of intercellular canals and spaces. Leaves simple (3-foliolate in Menyanthes), alternate; stipules absent or represented by expanded winged margins of the petiole; petiole sheathing at base; blades cordate, reniform, or linear, rarely peltate. Inflorescence of diverse types. Flowers bisexual and often distylous or unisexual (plants dioecious), actinomorphic, usually 5-merous. Sepals 5, connate at the base; corolla sympetalous, 5-lobed, with a tube and valvate to induplicate-valvate lobes; margins fringed or crested. Stamens 5; filaments attached to corolla tube alternate with the lobes, sometimes scales (staminodes?) alternate with the filaments; anthers mostly sagittate basally, opening by longitudinal slits. Ovary 2-carpellate, 1-locular, superior to half-inferior, usually surrounded at the base by a nectary disk; style 1; stigma 2-lobed; ovules unitegmic, numerous. Fruit a variously

Nymphoides 579

dehiscent capsule or a berry. Seeds numerous to few, usually smooth; endosperm copious, firm, oily; embryo linear, axile. Cosmopolitan; 5 genera and ca. 45 species, 1 species in the flora area. 1. NYMPHOIDES Ség., Pl. Veron. 3: 121. 1754. Perennial rhizomatous herbs. Leaves floating, rounded to cordate at base, rarely peltate. Inflorescences umbel-like cymose clusters. Flowers bisexual and distylous or unisexual (plants dioecious), emergent, yellow or white. Distylous flowers: either styles long with well-developed 2-lobed stigmas and anthers borne below the level of the stigma on short filaments, or styles short with smaller stigmas and anthers borne above the stigmatic level on long filaments. Unisexual flowers: staminate flowers without a style; stigmas undeveloped; ovaries as large as those of pistillate flowers but nonfunctional. Pistillate flowers with short style; stigmas well-developed and fleshy. Nectar glands 5, borne at base of ovary opposite corolla lobes. Fruits maturing under water, usually small. Seeds nearly smooth to papillate. Southeastern Canada, eastern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, northern Argentina, Old World tropics (most diverse); 20 species, 1 in Venezuela. Nymphoides indica (L.) Kuntze, Revis. Gen. Pl. 2: 429. 1891, “indicum.” —Menyanthes indica L., Sp. Pl. 145. 1753. —Bora. Villarsia humboldtiana H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 187. 1818 [1819]. —Nymphoides humboldtiana (H.B.K.) Kuntze, Revis. Gen. Pl. 2: 429. 1891, “humboldtianum.” Aquatic herb; leaf blades ± ovate to orbicular, cordate at the base, the lower surface reddish, the upper surface green, frequently

with purplish blotches; flowers white with a small yellow mark in the center or wholly yellow, the lobes of the corolla fimbriate on the margins. Shallow water or muddy margins of lakes, 50–300 m; Bolívar (Altiplanicie de Nuria, 29 km southeast of Caicara, 17 km south of Tumeremo). Apure, Guárico, Monagas; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil, northern Argentina, Old World tropics. ◆Fig. 495.

Fig. 495. Nymphoides indica

580

M IMOSACEAE

MIMOSACEAE by Rupert C. Barneby, James W. Grimes, Paul E. Berry, David Brunner, Enrique Forero, Lourdes Cárdenas, Giovanna De Martino, Helen C. F. Hopkins, and Elêna Maria de Lamare Occhioni Trees, shrubs, subshrubs, and herbs, few monocarpic, many with root nodules harboring nitrogen-fixing bacteria, either unarmed or armed with spinescent stipules, or with axillary or intrastipular thorns, or with epidermal prickles, the leaf axes often bearing pitted, plane or mounded nectaries; pubescence of simple, or glandtipped, or granular, sometimes branched or rarely scale-like trichomes. Leaves alternate (exceptionally opposite), either spirally or distichously inserted, mostly bipinnate but simply pinnate in Inga and few species of Calliandra, Cojoba, and Macrasamanea, or modified into a phyllode; stipules foliaceous, setiform, spinescent, or inflated, sometimes fugacious or obsolete; leaf stalks (petiole and rachis combined), pinnae, and leaflets usually all pulvinate, the pulvini controlling sleep movements. Inflorescence of axillary, terminally paniculate, or cauliflorous capitula, umbelliform capitula, spikes, or spiciform racemes, the perianth of individual flowers variable in size, but often small with conspicuously exserted white, pink, red, or yellow androecium, the flowers of each unit of inflorescence either homomorphic or 2(3)-morphic, either all bisexual, or some staminate and some neuter (or both), or 1–few terminal flowers unlike the peripheral ones in form or function. Perianth actinomorphic, 3–6-merous; calyx campanulate or cylindric, the teeth valvate (rarely imbricate) in bud; petals usually united into a tube, sometimes free, valvate. Androecium haplostemonous, diplostemonous, or indefinite; filaments either free or united into a tube, this often united at base with the corolla to form a stemonozone (zone of concrescence between corolla tube and androecium), the flower then perigynous; anthers dorsifixed, 2-thecate, extrorsely dehiscent by slits, the connective sometimes gland-tipped, the pollen grains shed singly or in polyads; an intrastaminal nectariferous disk sometimes present. Carpels in flora area genera 1, ripening into a fruit highly diverse in dimension, compression, curvature (straight, decurved, coiled), in texture and ornamentation of valves, and in mode of dehiscence, either 1-locular or divided by adventitious septa into several 1-seeded locules; dehiscence either inert, or elastic, through either one or both sutures, or none. Seeds dispersed either by lomentiform disintegration of pod valves or by predators; seed funicles either filiform or variously dilated, sometimes into a fleshy aril; seeds either hard and long-lived, the bony or crustaceous testa then areolate (with a pleurogram or engraved margin of the areole) or not, or in few genera obese, with papery testa or a fleshy sarcotestal layer, or planocompressed with thin-papery, exareolate, sometimes winged testa; embryo straight, with basally cordate cotyledons and short radicle; endosperm sometimes present but seldom copious. Tropical and warm-temperate areas circumglobally; ca. 50 genera and 2500 species, 26 genera and 164 species in the flora area. Key to the Genera of Mimosaceae by Rupert C. Barneby 1. 1.

Specimen in flower ..................................................................................... 2 Specimen in fruit ...................................................................................... 33

M IMOSACEAE 581

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

6(5). 6. 7(6). 7. 8(6). 8.

9(8).

9.

10(8). 10. 11(10). 11. 12(11).

12. 13(12).

13.

Stamens of indefinite number, more than twice as many as the divisions of the perianth ....................................................................................... 3 Stamens as many or twice as many as the divisions of perianth ........... 21 Filaments free to base or shortly united at base without forming a tube ....................................................................................................... 2. Acacia Filaments distinctly united into a tube ..................................................... 4 Leaves simply even-pinnate ............................................................... 12. Inga Leaves bipinnate (the primary axis in some species of Zygia very short and easily overlooked) ........................................................................... 5 Inflorescence cauliflorous, arising from trunk or from year-old branches below foliage of current year ........................................................ 26. Zygia Inflorescence of capitula, either solitary, or fasciculate, or pseudoracemose, arising from axils of leaves of current growth, or terminally paniculate ................................................................................................................ 6 Leaflets of each pinna exactly 1 pair and of ample size, 5–12 cm long .... 7 Leaflets of distal pinnae 2–many pairs, sometimes in Calliandra leaflets of each pinna 3 (i.e., 11/2 pairs), tiny to ample ....................................... 8 Stems armed with stipular spines (but some flowering branchlets unarmed); flowers of each capitulum isomorphic ............. 20. Pithecellobium Stems unarmed; flowers of each head heteromorphic, a single terminal one thicker and commonly longer than the peripheral ones ......... 1. Abarema Petiolar nectaries absent; stipules commonly striately veined ................ 9 Petiolar nectaries on the primary leaf stalk 1 or more, either at base of petiole, near mid-petiole, or between the first pair of pinnae; stipules ordinarily not > 1-veined, or veinless externally, or evanescent ........ 10 Leaflets chartaceous to coriaceous; corolla at least somewhat striate; stigma obconical; pollen in 8-grained polyads; legume coriaceous, ligneous (more rigid than below) ................................................... 6. Calliandra Leaflets thin, membranaceous; corolla not striate; stigma cup-shaped; pollen in 16-grained polyads; legume membranaceous to coriaceous .................................................................................................. 25. Zapoteca Leaflets large, to 6–11 × 3–5.5 cm; tall emergent trees ............ 7. Cedrelinga Leaflets smaller (if approaching 6 cm in length then small trees or shrubs) .............................................................................................................. 11 Bracts of lower flowers of each capitulum with a concave nectary on the inner face ....................................................................... 14. Macrosamanea Bracts without nectary ............................................................................ 12 Flowers of each capitulum dimorphic, 1 (rarely 2 or 3) terminal ones at once thicker and longer than the peripheral ones and their androecial tube also longer .................................................................................... 13 Flowers of each capitulum isomorphic or almost so ............................... 20 Venation of leaflets palmate, the primary vein posterior to midrib evidently longer and stronger than the first secondary vein arising from posterior side of midrib; inflorescence either a terminal efoliate panicle of capitula or composed of efoliate pseudoracemes of capitula arising from coeval leaves ......................................................................... 3. Albizia Venation of leaflets either pinnate, the primary vein posterior to midrib no longer than first secondary vein, or reduced to the simple midrib; inflorescences various, but not as above, the capitula almost always arising directly from axils of coeval leaves ......................................... 14

582

M IMOSACEAE

14(13). Pedicel of outermost peripheral flowers of the capitulum < 1.5 mm long, often obsolete ....................................................................................... 15 14. Pedicel of outermost peripheral flowers 2–22 mm long ......................... 16 15(14). Pinnae of longer leaves 11–20 pairs and leaflets of longer pinnae 40– 80 pairs; largest leaflets 2.5–6 mm, simply costate ................................ ............................................................ 10. Enterolobium (E. schomburgkii) 15. Pinnae of leaflets fewer than above; leaflets either larger or pinnately veined or both ........................................................................... 1. Abarema 16(14). Ovary truncate at apex ............................................................................ 17 16. Ovary tapering into the style .................................................................. 19 17(16). Leaflets of longer pinnae 2–12 pairs .................................... 11. Hydrochorea 17. Leaflets of longer pinnae 14–35 pairs ..................................................... 18 18(17). Pinnae of larger leaves 6–10 pairs .................................................. 5. Balizia 18. Pinnae of larger leaves 2–6 pairs ............... 11. Hydrochorea (H. gonggrijpii) 19(16). Pedicel of outermost peripheral flowers of the capitulum 2–9.5 mm long ................................................................................................. 23. Samanea 19. Pedicel of outermost peripheral flowers 11–22 mm long ... 22. Pseudosamanea 20(12). Stems armed at random nodes with supra-axillary thorns (modified barren branchlets); leafy branchlets terminated by a perule of striate scales ................................................................................... 8. Chloroleucon 20. Stems unarmed; perules absent .......................................... 10. Enterolobium 21(2). Calyx lobes imbricate in bud ................................................................... 22 21. Calyx lobes valvate in bud ....................................................................... 23 22(21). Flowers densely aggregated into globose or clavate capitula, dimorphic, the upper bisexual and 10-androus, the lower neuter, with staminodal androecium .................................................................................. 17. Parkia 22. Flowers borne in elongate cylindric spikes, homomorphic, all bisexual, but the stamens dimorphic, 5 antesepalous, short, fertile, 5 antepetalous, elongate, without anther ................................... 18. Pentaclethra 23(21). Inflorescence of globose or ovoid-ellipsoid capitula less than twice as long as wide .................................................................................................. 24 23. Inflorescence racemose or spicate, the axis many times longer than the diameter of the inflorescence .............................................................. 27 24(23). Androecium of lower flowers of each capitulum sterile or petaloid ................................................................................................. 16. Neptunia 24. Androecium of all flowers perfect ............................................................ 25 25(24). Peduncle without involucre; anthers ovate or round in dorsal view, the sacs wrapped around a dilated connective; petiolar nectary relatively rare, and present only on serially prickly bush-lianas; herbs, vines, and shrubs of variable aspect; fruit a craspedium .......................... 15. Mimosa 25. Peduncle encircled below the capitulum with a small lobed involucre (early detached and sliding down peduncle); anthers ovate to oblong in dorsal view, the sacs parallel, attached to a narrow connective; petiolar nectary present; plant an unarmed tree or shrub; fruit a continuous legume ..................................................................................................... 26 26(25). Anthers glabrous; trees 10–25 m tall, the bark rugulose-verrucose; pinnae of larger leaves 12–30 pairs, leaflets of longer pinnae 35–80 pairs; pods subsessile ......................................................................... 4. Anadenanthera

M IMOSACEAE 583

26.

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

28.

29(28). 29.

30(29). 30. 31(30). 31. 32(31).

32.

33(1).

33.

34(33).

34.

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

Anthers hairy; shrubs to small trees 3–15 m tall, the bark smooth; pinnae of larger leaves 4–8 pairs, leaflets of longer pinnae 13–21 pairs; pods with stipes 4–14 mm long ....................................................... 13. Leucaena Inflorescence an exserted efoliate panicle of spikes; leaf stalk without nectary; unarmed vines ............................................................... 9. Entada Inflorescence of axillary spikes; leaf stalk (and often some pinna axes) with cupular or shield-shaped nectary ............................................... 28 Flowers haplostemonous, 5-merous, 5-androus; lower face of leaflets with minute, usually reddish, peltate trichomes; prickly vine with fruit a craspedium ................................................................................ 15. Mimosa Flowers diplostemonous, 4- or 5-merous, 8–10-androus; leaflets without specialized trichomes; trees, shrubs, and vines but if the latter the fruit a conventional compressed legume ..................................................... 29 Anthers ovate or round in dorsal view, the sacs wrapped around a dilated connective, this not gland-tipped; fruit a craspedium ............. 15. Mimosa Anthers oblong in dorsal view, the sacs straight and parallel, attached to narrow connective tipped with a caducous gland; fruit a legume with continuous valves ................................................................................. 30 Plant armed .............................................................................. 19. Piptadenia Plant unarmed ......................................................................................... 31 Leaflets alternate along pinna rachis .......................... 24. Stryphnodendron Leaflets in opposite pairs along pinna rachis ......................................... 32 Leaflets of longer pinnae < 15 pairs and the larger ones nearly always over 1 cm long; pod much < 20 cm long and deeply constricted between seeds, these lentiform, not winged ...................................... 19. Piptadenia Leaflets of longer pinnae 25–40 pairs and < 1 cm long; pod mostly 20– 40 cm long and not constricted between seeds, these discoid and winged ....................................................................... 21. Pseudopiptadenia Fruit a craspedium, the valves when ripe separating from the continuous, persistent frame of the sutures (the replum) and breaking up by transverse fracture into 1-seeded, free-falling articles ............................... 34 Fruit a legume with continuous valves, or (seldom) lomentiform, then breaking into 1-seeded articles both across the valves and the adherent part of the sutures ............................................................................... 35 Craspedium very large, ca. 25–45 × 5–8.5 cm, the thin exocarp exfoliating at maturity; inflorescence an asymmetrical (horizontal or pendulous) efoliate panicle of spikes; seeds without either pleurogram (engraved margin of areole on seed face) or endosperm .............................. 9. Entada Craspedium smaller than above, and the exocarp not exfoliating; inflorescences various, but if paniculate and efoliate then symmetrically so; seeds with pleurogram and at least thin endosperm .............. 15. Mimosa Leaves simply pinnate ........................................................................ 12. Inga Leaves bipinnate or reduced to phyllodia ............................................... 36 Leaves reduced to simple phyllodia; exotic trees, planted or naturalized ....................................................................................................... 2. Acacia Leaves bipinnate (sometimes with 1 pair of pinnae or of leaflets) ......... 37 Herbs and small subshrubs ........................................................ 16. Neptunia Shrubs, trees, and lianas ......................................................................... 38

584

M IMOSACEAE

38(37). Stems armed either at or between nodes with either epidermal prickles, spinescent stipules, or axillary thorns ................................................ 39 38. Stems unarmed ........................................................................................ 43 39(38). Stipules spinescent (absent from some flowering branchlets) ............... 40 39. Stipules flaccid or deciduous, the armament of axillary thorns or epidermal prickles .......................................................................................... 41 40(39). Pinnae and leaflets several pairs; valves of ripe pod not coiling, not red inside; seeds without aril .............................................................. 2. Acacia 40. Pinnae and leaflets each 1 pair; valves of pod separating and coiling, red or orange internally; seeds half enveloped in a fleshing aril ........................................................................................ 20. Pithecellobium 41(39). Armament of straight thorns, consisting of modified axillary branchlets; stems perceptibly thickened immediately below each node ............................................................................................. 8. Chloroleucon 41. Armament of epidermal prickles, these either scattered along stems and leaf stalks or paired at stem nodes; stems not thickened below nodes .............................................................................................................. 42 42(41). Pods arising from a globose, claviform, or rarely linear receptacular axis, this when linear < 2 cm long ........................................................ 2. Acacia 42. Pods arising from elongate pitted axis > 2 cm long ................ 19. Piptadenia 43(38). Pods arising from elongate pitted axis .................................................... 44 43. Pods arising from a globose, clavate, or short-cylindric receptacle ........ 48 44(43). No leaf nectary on petiole ...................................................... 18. Pentaclethra 44. A nectary on leaf stalk either between or below the first pair of pinnae ... 45 45(44). Leaflets alternate along pinna rachis .......................... 24. Stryphnodendron 45. Leaflets in opposite pairs along pinna rachis ......................................... 46 46(45). Leaflets few and ample, to 5 pairs per pinna, ovate-acuminate, the larger ones 6–12 cm; leaf nectary warty; pods usually numerous and crowed into an involved cluster, recurved through half a circle or more, the valves convexly raised over each seed, very tardily dehiscent .................................................... 24. Stryphnodendron (S. polystachyum) 46. Leaflets > 5 pairs per pinna and smaller, none over 4.5 cm long, mostly much smaller; leaf nectary shallowly cupular or shield-shaped; pods less curved, nearly plane, dehiscent through the ventral suture ...... 47 47(46). Pod undulately linear (mostly < 20 cm long), the sutures deeply recessed between successive seeds, these separated by narrow isthmi; seeds lentiform, not winged, with pleurogram ............................. 19. Piptadenia 47. Pod elongately linear (mostly 20–40 cm long), the sutures not recessed between seeds (unless randomly where an ovule aborts); seeds discoid, the lustrous thin testa wing-marginate and lacking pleurogram ................................................................................... 21. Pseudopiptadenia 48(43). Ripe pod elastically dehiscent along both sutures from apex to base, the valves coiled backward but not twisted on their long axis ................. 49 48. Ripe pod either indehiscent, or inertly dehiscent, or dehiscent along ventral suture only (follicular), or if the valves elastically recurved then also randomly twisted .......................................................................... 50 49(48). Valves of pod membranous .......................................................... 25. Zapoteca 49. Valves of pod coriaceous or ligneous .......................................... 6. Calliandra 50(48). Fruit a papery samariform loment 20–70 × 3.5–6 cm, pinched between the

M IMOSACEAE 585

50. 51(50).

51. 52(51). 52. 53(52). 53. 54(52).

54. 55(54). 55. 56(55). 56. 57(56). 57. 58(57). 58. 59(58). 59. 60(59).

60. 61(60).

61.

planocompressed, indehiscent, 1-seeded articles and at each isthmus twisted through ca. 90° .......................................................... 7. Cedrelinga Fruit other than as above, usually shorter, if segmented much narrower and not twisted between seeds ............................................................ 51 Pods massive, either pendulous or less often ascending, borne on a stoutly claviform or abruptly broad-capitate, pitted axis, in outline oblong or broad-linear, laterally compressed and low-corrugate lengthwise, contracted at base into a stipe 2–8 cm long ..................................... 17. Parkia Pods sessile or shortly stipitate (to 15 mm long), then unlike the preceding in all other respects ....................................................................... 52 Seed coat without a pleurogram .............................................................. 53 Seed coat with a pleurogram ................................................................... 54 Fruits borne on the trunk or on previous year’s and older branches, far below the leaves ............................................................................ 26. Zygia Fruits borne on present year’s branches, axillary to coeval or lately fallen leaves ............................................................................. 14. Macrosamanea Pods coiled into a ring or a compressed spiral, indehiscent, the sutures immersed, the valves when ripe (or dried) of thick ligneous or fibrous and pulpy consistency, the cavity divided by false septa into 1-seeded locules ............................................................................... 10. Enterolobium Pods either straight or coiled, but if coiled then readily dehiscent, and the cavity 1-locular ..................................................................................... 55 Leaflets palmately veined from pulvinule ....................................... 3. Albizia Leaflets pinnately veined or 1-veined ..................................................... 56 Pod straight, papery, inertly dehiscent through both sutures, the valves continuous .................................................................... 22. Pseudosamanea Pod either straight or falcate, either indehiscent, follicular, or lomentiform, the valves coriaceous, ligneous, or pulpy .................................. 57 Fruits indehiscent, plumply biconvex or subcylindric, neither follicular nor lomentiform ...................................................................... 23. Samanea Fruits strongly compressed, either follicular or lomentiform ................ 58 Dehiscence lomentiform, the valves cracking transversely between seeds ........................................................................................... 11. Hydrochorea Dehiscence opening along one or both sutures, not lomentiform .......... 59 Pods shortly stipitate (stipe 5–15 mm long) ............................... 12. Leucaena Pods sessile or subsessile, the stipe < 5 mm long ................................... 60 Pods dehiscent along both sutures, the sutures commonly sinuous or the seminiferous one deeply recessed between seeds, the valves orange-red internally; dehiscence elastic, through both sutures, the valves recurved and twisted ................................................................................ 1. Abarema Pods dehiscent along a single suture, neither elastically dehiscent nor orange-red internally .............................................................................. 61 Pods linear, the sutures either straight or shallowly constricted between seeds, the stiffly coriaceous valves dehiscent along the semniferous suture; seeds discoid, sharply marginate but scarcely winged, testa thin ......................................................................................... 4. Anadenanthera Pods oblong, the sutures straight and dilated to frame the almost flat valves; dehiscence through the ventral suture, the valves opening like a book; seeds narrowly oblong, testa hard .................................. 5. Balizia

586

M IMOSACEAE

1. ABAREMA Pittier, Bol. Minist. R.R. E.E. 10–12: 56, 1927; reprinted in Arb. Legum., part 1, Trab. Mus. Comercial Venezuela 2: 86. 1927. Pithecellobium sect. Abaremotemo Benth., Trans. Linn. Soc. London 30: 581. 1875. Jupunba Britton & Rose, N. Amer. Fl. 23: 24. 1928. Klugiodendron Britton & Killip, Ann. New York Acad. Sci. 35: 125. 1936. by Rupert C. Barneby, James W. Grimes, and Paul E. Berry Unarmed trees or bushy treelets, commonly evergreen. Stipules small and caducous or obsolete. Leaves bipinnate, the pinnae 1–6 pairs (in the flora area), the longer ones with 1–13 pairs of leaflets, these of median or ample size; petioles charged with a cupular or wart-like nectary. Inflorescence composed of capitula, sometimes spikes outside the flora area, these pedunculate and either solitary or paired in the axil of coeval leaves. Flowers either sessile or short-pedicellate, the one terminating each capitulum usually longer and thicker than the peripheral ones and precociously expanding, its staminal tube also longer. Calyx campanulate; corolla tubular or narrowly funnelform. Stamens many; filaments united into a tube; anthers lacking a terminal gland. Pods linear compressed, recurved into a ring, or falcate, or sometimes nearly straight, the sutures commonly sinuous or the seminiferous one deeply recessed between seeds, the chartaceous, coriaceous, or woody valves orange-red internally; dehiscence elastic, through both sutures, the valves recurved and twisted. Seeds exarillate, the testa opaque, translucent, or partly white, without pleurogram (in the flora area); embryo gray-blue. Neotropics (most diverse in Greater Antilles, Colombia, Venezuela, the Amazon basin, and southeastern Brazil); 46 species, 12 in Venezuela, all in the flora area. Abarema resembles Albizia in dimorphic flowers, but differs in dehiscence of the pod and in the gray-blue or partly white seeds often lacking a pleurogram. From Pithecellobium s. str. it differs in lack of stipular spines and in the lack of fleshy aril invaginating the seed. Key to the Species of Abarema 1. 1. 2(1).

2.

3(2).

3.

4(1). 4.

Leaflets of all pinnae exactly 1 pair .......................................................... 2 Leaflets of distal pinnae 2–13 pairs .......................................................... 4 Leaflets ovate-elliptic, long-caudate, bright green on both surfaces even when dried; filaments of terminal flower erect; pod recurved into a ring, much over 4 cm long ............................................................... A. laeta Leaflets elliptic-obovate, obtuse, noticably bicolored, upper surface lustrous dark green, lower surface pallid and dull; free part of filaments in the heteromorphic terminal flower proximally thickened and recurved; pod almost straight and only 3–4 cm long ............................................ 3 Leaflets bicolored, lustrous on upper surface, pallid dull on lower, broadly rounded at apex; corolla of peripheral flowers of the capitulum 3– 6.5 mm long, not more than twice as long as calyx ............. A. leucophylla Leaflets concolorous, lustrous on both surfaces, shortly bluntly acuminate; corolla of peripheral flowers 11.5–15.5 mm long, 3–4.5 times as long as calyx ................................................................................... A. levelii Leaflets of longer pinnae at least 6 pairs .................................................. 5 Leaflets of longer pinnae 2–5(6)pairs ........................................................ 8

Abarema 587

5(4).

Young branchlets and dorsal face of leaflets softly villosulous with erect hairs ..................................................................................... A. barbouriana 5. Young branchlets and dorsal face of leaflets smooth or strigulose with short, appressed hairs ........................................................................... 6 6(5). Calyx of peripheral flowers 4–9 mm long and their corolla 8–13 mm long; tassel of androecium rose-purple .......................................................... 7 6. Calyx of peripheral flowers not over 2.5 mm long and their corolla not over 7 mm long; tassel of androecium white ............................ A. jupunba 7(6). Longer leaf stalks (6–)8–23 cm and pinnae (3)4–6 pairs; rachis of longer pinnae (7–)8–14(–20) cm; below 200 m in Amazonas state .. A. floribunda 7. Longer leaf stalks 5–7 cm and pinnae 3–4 pairs; rachis of longer pinnae 4– 7 cm; 1100 m and above in Bolívar state ............................... A. ferruginea 8(4). First petiolar nectary funnelform-campanulate (deeply cup-shaped) and 10–13 × 8–12 mm ................................................................. A. adenophora 8. First petiolar nectary either cup- or disk-shaped, mounded, or immersed in the petiolar groove, but < 2 mm tall ................................................. 9 9(8). Petiolar nectary obesely wart-like, with punctiform orifice much narrower than the body; corolla glabrescent or minutely puberulent externally ........................................................................................ A. microcalyx 9. Petiolar nectary cup- or disk-shaped, with broad open cavity; corolla, at least its lobes, silky externally ............................................................ 10 10(9). Calyx of peripheral flowers 1.8–2.5 mm long; dorsal face of leaflets minutely puberulent ................................................................................ 11 10. Calyx of peripheral flowers 3–7 mm long; dorsal face of leaflets smooth or softly pubescent with erect tawny hairs ............................................. 12 11(10). Foliar nectaries cupular, the largest one 2–3 mm diameter and 0.8–1 mm tall; flowers minutely puberulent ............................................... A. acreana 11. Foliar nectaries convex and pored, smaller than above; flowers densely silky .......................................................................................... A. junpunba 12(10). Dorsal face of leaflets smooth; pod nearly straight ....... A. longipedunculata 12. Dorsal face of leaflets softly pubescent with erect tawny hairs; pod unknown .......................................................................................... A. villifera Abarema acreana (J.F. Macbr.) L. Rico, Novon 9: 555. 1999. —Pithecellobium acreana J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 51. 1943, “Pithecolobium.” —Hydrochorea? acreana (J.F. Macbr.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 33. 1996. Tree 15–30 m tall; pinnae 4–6 pairs, the interpinnal segments 2–5.5 cm long, each pinna-pair with a nectary, the one near or above the midpetiole sessile and cupular-elliptic and 2–3 mm diameter, 0.8–1 mm tall, distal ones progressively smaller; leaflets obtusely rhombic-ovate from an unequal, broadly rounded base, the largest ones 3.5–6 × 1.5–3.7 cm; petiole proper 4–6.5 cm long. Nonflooded evergreen lowland forests, 100– 300 m; Bolívar (Reserva Forestal La Para-

gua, Río Toro), Amazonas (east of Guanábano [about halfway between San Carlos and Santa Lucia]). Nicaragua, Costa Rica, Brazil. The Amazonas collection is based on a sterile specimen, Berry & Aymard 7365 (MO, PORT), and the Bolívar specimens are similarly based on nonflowering specimens cited by Barneby in his 1996 monograph of Hydrochorea. Abarema adenophora (Ducke) Barneby & J.W. Grimes, Mem. New York. Bot. Gard. 74: 74. 1996. —Pithecellobium adenophorum Ducke, Arq. Inst. Biol. Veg. 4: 5. 1938, “Pithecolobium.” Tree 12–30 m tall; pinnae of larger leaves 2–5 pairs and leaflets 16–40(–48) per leaf, the distalmost leaflets 5–11 × 3.5–7.5 cm,

588

M IMOSACEAE

asymmetrically obovate or rhombic-obovate; petiole proper 2.5–10 cm long, with a stipitate or sessile, funnel-shaped nectary 8–13 × 6–12 mm situated at insertion of first pinnapair, sometimes a smaller or squatter tubular nectary present at tip of some pinna rachises. Nonflooded evergreen lowland forests, 100–200 m; Amazonas (south of Caño Marimajare near San Carlos de Río Negro). Nicaragua, Pacific coastal area of Colombia, Amazonian Peru and Brazil. Abarema barbouriana (Standl.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 70. 1996. —Pithecellobium barbourianum Standl., Contr. Arnold Arbor. 5: 74. 1933. Discontinuously dispersed from Panama to Guyana, Suriname, French Guiana, and northern Amazonian Brazil; 2 varieties, 1 in Venezuela. A. barbouriana var. arenaria (Ducke) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 72. 1996. —Pithecellobium arenarium Ducke, Arq. Inst. Biol. Veg. 2: 37. 1935. —Guamito (Amazonas), Meseyek (Bolívar). Shrub or treelet 2–7(–9) m tall; pinnae of larger leaves 3–5 pairs and leaflets of longer pinnae (7)8–11 pairs, either pilosulous overall on dorsal face, or glabrate except for pilosulous midrib, or (locally) glabrous, the larger ones 9–21 × 5–10 mm. Bush islands on flooded or nonflooded savannas, rocky torrent banks, meadows at summits of tepuis, sandy lowland savannas, (100–)1200–2000 m; common and widespread in Bolívar and Amazonas. Northern Brazil (Amazonas: Rio Negro). This variety is part of a polymorphic complex. The bark yields saponins used in laundry. Abarema ferruginea (Benth.) Pittier, 3rd. Conf. Interamer. Agric. Caracas 360. 1945. —Pithecellobium ferrugineum Benth., London J. Bot. 3: 216. 1844. Treelet 1.5–8 m tall; leaflets 8–10 pairs, 2–3 × 1.2–2 cm; filaments of androecium said to be deep red when fresh. Tepui escarpments, 500–1200 m; Bolívar (Roraimatepui). Brazil (Serra do Sol).

Abarema floribunda (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 48. 1996. —Pithecellobium floribundum Spruce ex Benth., Trans. Linn. Soc. London 30: 584. 1875. —Marupa. Tree 10–20 m tall. Lowland riparian forests, 100–200 m; Amazonas (Río Casiquiare). Northern inter-Andean Colombia, Peru, western and central Brazil. The single collection from the flora area agrees closely with the Brazilian type. Abarema jupunba (Willd.) Britton & Killip, Ann. New York Acad. Sci. 35: 126. 1936. —Acacia jupunba Willd., Sp. Pl. 4: 1067. 1806. —Pithecellobium jupunba (Willd.) Urb., Symb. Antill. 2: 257. 1900. Tree commonly 5–35(–40) m tall (in uplands sometimes fertile when only 1.5–3 m tall). Lesser Antilles, tropical South America; 2 varieties, both in the flora area. Key to the Varieties of A. jupunba 1. Pinnae of larger leaves 3–7 pairs and leaflets of longer pinnae 7–12 pairs; terminal leaflets of larger leaves (13–)16–35 × 10–20 mm ...................... var. jupunba 1. Pinnae of larger leaves 1–3 pairs and leaflets of longer pinnae 3–6 pairs; terminal leaflets of larger leaves 35–70 mm long ................................... var. trapezifolia A. jupunba var. jupunba Evergreen lowland to lower montane forests, rocky riverbanks, 100–400 m; Amazonas (upper Río Negro, Río Mawarinuma). Zulia; widespread over the range of the whole species except where displaced by var. trapezifolia. A. jupunba var. trapezifolia (Vahl) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 69. 1996. —Mimosa trapezifolia Vahl, Eclog. Amer. 3: 47. 1807. —Pithecellobium trapezifolium (Vahl) Benth., London J. Bot. 2: 142. 1844. —Abarema trapezifolia (Vahl) Pittier, Bol. Minist. R.R. E.E. 10–12: 56, 1927; reprinted in Arb. Legum., part. 1, Trab. Mus. Comercial Venezuela. 2: 86. 1927. —Jupunba trapezifolia (Vahl) Moldenke,

Abarema 589

Bull. Torrey Bot. Club 59: 155. 1932. —Guamo moñtanero, Maripo, Samán. Evergreen lowland to montane forests and shrublands, tepui slopes, (100–)200– 1200 m; widespread in Bolívar and Amazonas. Coastal Cordillera; Trinidad, Guyana, Suriname. This variety is the commonest Abarema throughout the flora area. On the Gran Sabana an undescribed form of this variety differs in having only 1(2) pairs of pinnae per leaf. This resembles the Brazilian A. piresii Barneby & J.W. Grimes, but its pod is that of var. trapezifolia, with outer margin obscurely undulate, not deeply impressed between seeds. Abarema laeta (Benth..) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 79. 1996. —Pithecellobium laetum Benth., London J. Bot. 3: 203. 1844. —Inga laeta (Benth.) Poepp., Nov. Gen. Sp. Pl. 3: 80. 1845. —Klugiodendron laetum (Benth.) Britton & Killip, Ann. New York Acad. Sci. 35: 125. 1936. Shrub or slender treelet 1–5(–6) m tall; leaflets bright green, reticulately veined; flowers vespertine; filaments white, fading to orange. Nonflooded evergreen lowland to lower montane forests, 100–300 m; Amazonas (Río Negro). Central and northern Venezuela; Colombia, French Guiana, Suriname, Ecuador, Peru, Brazil.

chartaceous, lustrous on upper surface, pallid and dull on lower surface; pod short, scarcely curved. Shrublands on white sand, montane forests on tepui slopes, 100–900 m; Amazonas (upper Río Negro, Cerro Huachamacari, Sierra de la Neblina). Brazil. Abarema levelii (R.S. Cowan) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 106. 1996. —Pithecellobium levelii R.S. Cowan, Mem. New York Bot. Gard. 10(4): 69, fig. 48. 1961. Tree 8–10 m; leaves amply 4-foliolate; inflorescences compact terminal, efoliate panicles; flowers white. Riparian forests, 100–200 m; northwestern Amazonas (banks of Río Orinoco and immediately affluent creeks near the mouth of Río Atabapo). Central Amazonian Brazil, expected in Colombia adjacent to the flora area. Abarema longipedunculata (H.S. Irwin) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 52. 1996. —Pithecellobium longipedunculatum H.S. Irwin, Acta Bot. Venez. 2: 223. 1967. Shrub or small tree 2–20 m tall; leaflets 3–5 pairs, ovate, coriaceous, lustrous on upper surface, pallid on lower surface, the larger ones 4–6.5 × 2–4 cm. Montane forests, along streams of tepui summits, 1400–1800 m; Bolívar (Auyán-tepui, Cerro Jaua and Cerro Sarisariñama). Endemic. ◆Fig. 496.

Abarema leucophylla (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 76. 1996. —Pithecellobium leucophyllum Spruce ex Benth., Trans. Linn. Soc. London 30: 581. 1875. —Marepillo. Southwestern Venezuela; eastern Colombia, northwestern Brazil; 2 varieties, 1 in Venezuela. The second variety, var. vaupesensis, is known only from Colombia. It has been found just across the Río Orinoco from San Fernando de Atabapo, Amazonas state, and might eventually be found in the flora area somewhere in that general vicinity.

Abarema microcalyx (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 63. 1996. —Pithecellobium microcalyx Spruce ex Benth., Trans. Linn. Soc. London 30: 582. 1875. Colombia, Peru, Brazil; 3 varieties, 1 in Venezuela.

A. leucophylla var. leucophylla Tree (4–)5–21 m; pinnae of each leaf 1(2) pairs and each pinna bifoliolate; leaflets

Abarema villifera (Ducke) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 108. 1996. —Pithecellobium villi-

A. microcalyx var. microcalyx Tree 5–10 m tall; petiolar nectary obese, opening by a terminal pore; pods black, 11– 20 mm wide, bent into a ring. Nonflooded riparian forests, Río Negro caatinga, 100–300 m; Amazonas (Río Guainía, Río Negro, upper Río Orinoco). Colombia.

590

M IMOSACEAE

Fig. 496. Abarema longipedunculata

ferum Ducke, Arq. Inst. Biol. Veg. 4: 5. 1938. Tree 6–20 m tall. Lowland riparian forests (along black-water rivers), montane forests, 100–1300 m; Bolívar (Sierra Pakaraima), Amazonas (Caño Chola south of San Carlos

de Río Negro, Río Guainía). Suriname, Brazil (Amazonas: upper Rio Negro). This species resembles Abarema jupunba var. trapezifolia except for densely villous dorsal face of leaflets. The bark is mashed for soapy water (in Bolívar).

2. ACACIA Mill., Gard. Dict. Abr. ed. 4. 1754. Vachellia Wight & Arn., Prodr. Fl. Ind. Orient. 272. 1839 [1834]. Farnesia Gasp., Descr. Nuov. Gen. Leg. 1836, non Fabr. 1763. Poponax Raf., Sylva Tellur. 118. 1838. Senegalia Raf., Sylva Tellur. 119. 1838. Manganaroa Speg., Physis (Buenos Aires) 6: 312. 1923. Acaciella Britton & Rose, N. Amer. Fl. 23: 96. 1928. by Lourdes Cárdenas de Guevara and Giovanna De Martino Trees, shrubs, or lianas, spiny or not. Leaves bipinnate or reduced to simple phyllodia; pinnae in few to many pairs; stipules generally small and caducous, sometimes modified into spines; pinnules generally small and numerous; gland usually present on the petiole and frequently also on the rachis. Inflorescence either a cylindric spike or axillary, head-like glomerules that are solitary or fasciculate and sometimes forming panicles at the tip of the branches. Flowers small, numerous, yellow,

Acacia 591

cream, or white. Calyx tubular or campanulate, 4- or 5-dentate; corolla tubular, funnelform or campanulate, 4- or 5-lobed. Stamens numerous, exserted, free or slightly joined basally; anthers small, sometimes with an apical gland; intrastaminal disk sometimes present; pollen in polyads. Ovary sessile or stipitate. Legume compressed, subcylindric or cylindric, 2-valved or tardily dehiscent to indehiscent, straight or curved, rarely articulate. Seeds with pleurogram. Pantropics and subtropics, most diverse in Australia; ca. 1200 species, ca. 15 in Venezuela, 8 of these in the flora area. Key to the Species of Acacia 1. 1. 2(1).

2.

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

Plant with stipules transformed into spines; flowers yellow ................... 2 Plant with foliaceous stipules; flowers white, cream, or yellow ............... 3 Pinnae in ≤ 7 pairs; lower surface of pinnules with evident secondary veins; corolla < 2.5 mm long; stamens > 4.5 mm long; fruit cylindric, light brown, with a whitish line along the dorsal suture ..... A. farnesiana Pinnae in 8–17 pairs; lower surface of pinnules with secondary veins not evident; corolla > 2.5 mm long; stamens < 3 mm long; fruit subcylindric or cylindric, uniformly light brown ................................... A. macracantha Flowers in elongated, cylindric spikes ...................................................... 4 Flowers in head-like glomerules ................................................................ 5 Spikes (including the peduncle) ≥ 14.5 cm long; calyx ≥ 3 mm long; corolla ≥ 5 mm long ...................................................................... A. alemquerensis Spikes (including the peduncle) ≤ 8.5 cm long; calyx < 2 mm long; corolla ≤ 3.5 mm long .......................................................................... A. articulata Stipules persistent, basally cordate, usually ≥ 10 × 10 mm; floral rachis ca. 7 mm long; calyx ≥ 2 mm long; stamens ≥ 9 mm long .... A. tamarindifolia Stipules caducous, basally oblique or obliquely angled, ≤ 7 × 1.5 mm; floral rachis ≤ 2 mm long; calyx < 2 mm long; stamens ≤ 7 mm long ............ 6 Pinnae in 8–12 pairs; pinnules in 13–30 pairs, ≥ 8 × 2 mm ...... A. glomerosa Pinnae in 10–35 pairs; pinnules in 33–70 pairs, ≤ 4.5 × 1 mm ................ 7 Pinnules falcate, middle vein excentric (almost bordering the interior margin); anther (in bud) with apical gland ........................... A. paniculata Pinnules linear, middle vein subcentral; anther without gland ................................................................................................. A. podadenia

Acacia alemquerensis Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 380. 1909. —Manganaroa alemquerensis (Huber) Speg., Physis (Buenos Aires) 6: 313. 1923. Acacia alvaroi Cárdenas & De Martino, Ernstia 56: 10. 1989. Shrub or liana 10–12 m tall; flowers fragrant. Disturbed areas, 50–300 m; northern Bolívar. Colombia, Brazil, Bolivia. Acacia articulata Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 73. 1922. —Manga-

naroa articulata (Ducke) Speg., Physis (Buenos Aires) 6: 313. 1923. Tree or liana to 16 m tall. Riparian forests, ca. 100 m; Bolívar (vicinity of Río Cuyuní). Apure, Barinas, Guárico, Portuguesa, Zulia; Brazil. ◆Fig. 497. Acacia farnesiana (L.) Willd., Sp. Pl. 4: 1083. 1806. —Mimosa farnesiana L., Sp. Pl. 521. 1753. —Vachellia farnesiana (L.) Wight & Arn., Prodr. Fl. Ind. Orient. 272. 1834. —Poponax farnesiana (L.) Raf., Sylva Tellur. 118. 1838. —Cují cabro, Yaque.

592

M IMOSACEAE

Fig. 497. Acacia articulata

Fig. 498. Acacia tamarindifolia

Acacia 593

Fig. 499. Acacia farnesiana

Fig. 500. Acacia macracantha

594

M IMOSACEAE

Fig. 501. Acacia glomerosa

Shrub or tree 3–5 m tall; flowers fragrant. Deciduous forests, disturbed areas, 50–300 m; northern Bolívar. Aragua, Carabobo, Cojedes, Distrito Federal, Guárico, Falcón, Lara, Mérida, Nueva Esparta, Portuguesa, Táchira, Trujillo, Zulia; widespread in the Neotropics from Mexico to Argentina. ◆Fig. 499.

Acacia glomerosa Benth., London J. Bot. 1: 521. 1842. —Senegalia glomerosa (Benth.) Britton & Rose, N. Amer. Fl. 23: 116. 1928. —Chigüire, Güigüire sabanero. Spiny tree 4–25 m; flowers fragrant. Deciduous and disturbed forests, 100–400 m; northern and central Bolívar (near Upata).

Albizia 595

Anzoátegui, Aragua, Barinas, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Guatemala, Honduras, El Salvador, Costa Rica, Colombia, Peru, Brazil. ◆Fig. 501. Acacia macracantha Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1080. 1806. —Mimosa macracantha (Humb. & Bonpl. ex Willd.) Poir. in Lam., Encycl. suppl. 1: 78. 1810. —Poponax macracantha (Humb. & Bonpl. ex Willd.) Killip, Caribbean Forest. 9: 248. 1948. —Cují. Acacia flexuosa Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1082. 1806. —Mimosa flexuosa (Humb. & Bonpl. ex Willd.) Poir., Encycl. suppl. 1: 78. 1810. —Poponax flexuosa (Humb. & Bonpl. ex Willd.) Britton & Rose, Ann. New York Acad. Sci. 35: 139. 1936. Mimosa lutea Mill., Gard. Dict. ed. 8, 18. 1768. —Acacia lutea (Mill.) Britton, Bull. Torrey Bot. Club 16: 327. 1889, non Leavenw. 1824. —Poponax lutea (Mill.) Britton & Rose, N. Amer. Fl. 23: 90. 1928. Poponax canescens Britton, Ann. New York Acad. Sci. 35: 138. 1936. Tree or shrub 3–7 m tall; stem with unpleasant odor. Deciduous forests, savannas, and shrublands, 50–400 m; northern Bolívar. Widespread elsewhere in Venezuela; Mexico, Panama, West Indies, Colombia, French Guiana. ◆Fig. 500.

Acacia macracantha is used to build fences. Acacia paniculata Willd., Sp. Pl. 4: 1074. 1806. —Mimosa paniculata (Willd.) Poir. in Lam., Encycl. suppl. 1: 74. 1810. —Senegalia paniculata (Willd.) Killip, Trop. Woods 63: 6. 1940. Tree, shrub, or liana 4–5 m tall. Riparian and disturbed forests, 100–300 m; northeastern Bolívar. Apure, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Monagas, Zulia; Colombia, Ecuador, Peru, Brazil, Bolivia. Acacia podadenia (Britton & Killip) L. Cárdenas, Revista Fac. Agron. (Maracay) 7(3): 135. 1974. —Senegalia podadenia Britton & Killip, Phytologia 1: 24. 1933. —Bejuco jala pa’trás. Small tree or climber to 5 m tall. Disturbed forests, 100–500 m; western and northeastern Bolívar. Colombia, Peru, Brazil. Acacia tamarindifolia (L.) Willd., Sp. Pl. 4: 1092. 1806. —Mimosa tamarindifolia L., Sp. Pl. 523. 1753. —Senegalia tamarindifolia (L.) Britton & Rose, N. Amer. Fl. 23: 120. 1928. Tree or shrub 3–6 m tall. Semideciduous forests, deciduous scrub 50–400 m; northern Bolívar. Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Miranda, Monagas, Nueva Esparta, Sucre, Zulia; Lesser Antilles, Colombia. ◆Fig. 498.

3. ALBIZIA Durazz., Mag. Tosc. 3: 11. 1772. Arthrosamanea Britton & Rose, Ann. New York Acad. Sci. 35: 128. 1936. by Rupert C. Barneby and James W. Grimes Unarmed trees of sympodial growth. Leaves bipinnate, at least one nectary on each petiole; stipules small, caducous. Inflorescences of capitula either terminally pseudoracemose-paniculate or borne in specialized efoliate pseudoracemes axillary to primary cauline leaves, the flowers of each capitulum dimorphic, at least the androecium of the terminal one modified. Flowers all small (< 7 mm long in American species). Androecium whitish, < 20 mm long in American species; stamens more than twice as many as the divisions of the perianth; filaments united into a tube. Pods either sessile or shortly stipitate, in profile linear or broad-linear, compressed or planocompressed, the valves either papery and inertly dehiscent through the sutures, or leathery and segmented between seeds. Seeds with hard testa and pleurogram. Circumtropics; over 100 species, 7 in Venezuela, 3 of these in the flora area.

596

M IMOSACEAE

The genus in its full circumtropical dispersal is polymorphic in form of inflorescence, in form, texture, and dehiscence of fruit, and in other characters. All species found in the flora area are referable to the American sect. Arthrosamanea (Britton & Rose) Barneby & J.W. Grimes. Albizia lebbeck (L.) Benth. is an Asian species that is widely cultivated in northern Bolívar state as a street tree, particularly in cities like Ciudad Bolívar and Ciudad Guayana. This species is characterized by its white flowers and persistent fruits, the latter oblong (13–32 × 2.5–5 cm), light tan, and with papery valves that are bullately dilated over each seed. The fruits are very tardily dehiscent, often only after they fall off the trees, and while on the tree the seeds often become detached from the placenta, which makes the fruits rattle characteristically in strong winds. Key to the Species of Albizia 1.

1.

2(1).

2.

Inflorescence composed of leafless pseudoracemes of capitula, each arising from the axil of a mature coeval leaf; pinnae of larger leaves 2–5 pairs and the leaflets of larger pinnae 9–12 pairs, the larger leaflets to 15–17 × 6–10 mm; upper surface of leaflets finely but prominulously (3)4(5)-veined, the primary veins branched; pods ca. 6–13 × 0.8–1.2 cm, torulose, septiferous between seeds but not or very tardily dehiscent ............................................................................................ A. subdimidiata Inflorescence paniculate, terminating branches of the current year; pinnae of larger leaves (5)6–12 pairs and the leaflets of larger pinnae 15– 20 pairs, the larger leaflets to 5–13 × 1.5–5 mm; upper surface of leaflets either veinless or 1-veined; pods 20–25 mm wide ......................... 2 Central flower of each capitulum greatly differentiated, with staminal tube exserted from corolla; calyx and corolla glabrous or minutely ciliolate, not silky; larger leaflets ca. 5–7 × 1.5–2.5 mm; pod lomentiform, when ripe articulate between seeds .................................... A. glabripetala Central flower of each capitulum not greatly differentiated; calyx and corolla gray-hispidulous; larger leaflets ca. 9–13 × 3–5 mm; pod continuous, with straight margins, resembling that of some acacias ................................................................................................. A. barinensis

Albizia barinensis L. Cárdenas, Ernstia 21: 5. 1983. Tree 7–20 m tall. Seasonally dry woodland, sometimes surviving as a pasture shade tree, ca. 200–300 m; northern Bolívar (near Represa Guri). Northern Venezuela; southwestern Guyana. ◆Fig. 502. Albizia glabripetala (H.S. Irwin) G.P. Lewis & P.E. Owen, Legumes Ilha de Maracá 42. 1989. —Pithecellobium glabripetalum H.S. Irwin, Mem. New York Bot. Gard. 15: 109. 1966. Tree 5–40 m tall; inflorescence of small paniculate capitula; flowers greenish; filaments white. Savannas, gallery forests, 100–

400 m; northeastern Bolívar (80 km S Caicara, Río Orinoco), Amazonas (La Esmeralda). Guyana, Brazil (Roraima). Albizia subdimidiata (Splitg.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 232. 1996. —Acacia subdimidiata Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 112. 1842. —Hueso de pescado blanco. Pithecellobium multiflorum auct. non Acacia multiflora H.B.K. 1824: sensu Benth., London J. Bot. 3: 240. 1844. Tree 5–30 m tall; inflorescence of small capitula disposed in leafless axillary pseudoracemes; pod compressed-moniliform, 10–16

Anadenanthera 597

Fig. 502. Albizia barinensis

× 0.7–1.2 cm. Nonflooded evergreen lowland forests, 50–300 m; uncommon in southern Delta Amacuro (Caño Joba-Suburu east of Caño Sacupana, Sabuca), eastern Bolívar (El

Palmar, San Pedro de las dos Bocas). Guyana, Suriname, French Guiana, and Amazonian Peru and Brazil.

4. ANADENANTHERA Speg., Physis (Buenos Aires) 6: 313. 1923. by Rupert C. Barneby Unarmed trees. Leaves elaborately bipinnate; stipules setiform, caducous; petiole with a shield-shaped or shallow dish-shaped nectary. Inflorescence of subglobose capitula borne in fascicles or on brachyblasts either shortly below the foliage, in leaf axils, or in exserted pseudoracemes; peduncle with a small cup-shaped involucre. Flowers whitish, 5-merous. Calyx campanulate; corolla lobes weakly

598

M IMOSACEAE

united below middle. Stamens 10; filaments free; anthers ovate-oblong, the connective either charged or not with a caducous gland. Ovary subsessile, glabrous. Pods linear, planocompressed, the sutures either straight or shallowly constricted between seeds, the stiffly coriaceous valves dehiscent along the semniferous suture to release the seeds. Seeds discoid, sharply marginate but scarcely winged; testa thin, glossy brown, with a small delicate pleurogram; endosperm lacking. Neotropics; 2 species, 1 in Venezuela. The second species, Anadenanthera macrocarpa (Benth.) Brenan, is found only south of the Amazon River.

Fig. 503. Anadenanthera peregrina var. peregrina

Anadenanthera peregrina (L.) Speg., Physis (Buenos Aires) 6: 313. 1923. —Mimosa peregrina L., Sp. Pl. 520. 1753. —Piptadenia peregrina (L.) Benth., J. Bot. (Hooker) 4: 340. 1941. —Niopa peregrina (L.) Britton & Rose, Addisonia 12: 37. 1927. —Yopo. Inga niopo Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1027. 1806. —Piptadenia niopo (Humb. & Bonpl. ex Willd.) Spruce, Notes. Bot. Amaz. 2: 426. 1908. Widespread in Neotropics; 2 varieties, 1 in Venezuela. This is the source of a powerful hallucinogenic snuff called yopo.

Balizia 599

A. peregrina var. peregrina Tree 10–25 m tall, the trunk to 0.2–0.4 m diameter; bark rugulose-verrucose; pinnae of larger leaves 12–30 pairs, leaflets of longer pinnae 35–80 pairs, linear, acute, 1-veined, the largest 3–8 × 0.5–1.5 mm; pods commonly 10–30 × 1–2.5(–3) cm, the sutures often undulately constricted, the valves dull, the epicarp eventually exfoliating in scurfy scales. Semideciduous forests, forest patches in savannas, gallery forest margins, shrub-

lands, and disturbed areas, 50–800 m; Bolívar (El Tigre, near La Paragua, Maripa, Túriba-San Pablo road, between San Pedro de las dos Bocas), Amazonas (near Puerto Ayacucho, Río Manapiare basin, Río Ocamo near Raudal Arata). Widespread elsewhere in Venezuela (e.g., Apure, Barinas, Guárico, Lara, Miranda); West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Paraguay, some of this distribution most probably due to dispersal by man. ◆Fig. 503.

5. BALIZIA Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 34. 1996. by Rupert C. Barneby Trees, resembling Albizia in most respects, but different in indeterminate fertile branches with umbelliform capitula axillary to coeval leaves (not terminally pseudoracemose nor assembled into efoliate pseudoracemes), in pinnate (not palmate) venation of leaflets, and (in the flora area) in tardily dehiscent, transversely fibrous fruit. Pubescence of young growth brownish. Stipules linear-ligulate, caducous; petiolar nectaries present; pinnae of larger leaves 6–10(–14) pairs and leaflets of longer pinnae 17–29 pairs, the larger leaflets 6–14 mm long. Capitula 20–40-flowered, the flowers heteromorphic, the pedicel of small peripheral ones 4–7.5 mm, the 1–3 central flowers subsessile and larger. Filaments 12–22, the tube included. Pods sessile, oblong in profile, 7–13 × 1.8–3.2 cm, planocompressed, the flat valves framed by dilated sutural rim, composed of thin blackish exocarp, coarsely transverse-fibrous mesocarp, and crustaceous endocarp; dehiscence follicular. Seeds narrowly oblong, testa hard; pleurogram narrow. Neotropics; 3 species, 1 in Venezuela.

Fig. 504. Balizia pedicellaris

600

M IMOSACEAE

Balizia pedicellaris (DC.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 37. 1996. —Inga pedicellaris DC., Prodr. 2: 441. 1825. —Pithecellobium pedicellare (DC.) Benth., London J. Bot. 3: 219. 1844. —Samanea pedicellaris (DC.) Killip ex Record, Trop. Woods 63: 4. 1940. —Macrosamanea pedicellaris (DC.) Kleinhoonte in Pulle, Fl. Suri-

name 2(2): 329. 1940. —Hueso de pescado, Kairaimundek (Pemón). Tree 20–45 m tall; flowers pink; filaments white. Evergreen lowland to lower montane forests, 100–800 m; scattered in Delta Amacuro, Bolívar, and Amazonas. Scattered in central Venezuela; widespread nearly throughout the Amazon basin, disjunct to Atlantic forests of southeastern Brazil. ◆Fig. 504.

6. CALLIANDRA Benth., J. Bot. (Hooker) 2: 138. 1840, nom. cons. by Paul E. Berry and Rupert C. Barneby Trees, shrubs, and subshrubs, sometimes functionally herbaceous from perennial roots or rhizome. Leaves bipinnate, rarely simply pinnate; petiolar nectaries lacking; pinnae in 1–many pairs; leaflets opposite or rarely subopposite basally, 4– numerous, oblique, midvein excentric; stipules herbaceous or subcoriaceous, deciduous or persistent, rarely spinescent (not in the flora area). Inflorescence composed of capitula or condensed racemes or umbels. Flowers sessile to rarely long-pedicellate, few to numerous, often dimorphic in the capitulum, 5(occasionally 3, 4, or 6)-merous. Stamens numerous, long-exserted, monadelphous, the tube included or exserted; filaments united into a tube 1/3–2 times the length of the corolla, white or bright red to purple, often pigmented only apically; anthers dorsifixed; pollen shed in 8-grained polyads. Ovary sessile to short-stipitate, tapering into the style; stigma dilated or scarcely so, the stigmatic surface convex, shallowly cupulate, or obscurely penicellate; ovules mostly 8, sometimes fewer, or rarely to 11. Pods narrowly oblanceolate or linear-oblanceolate in profile, straight or slightly falcate, the valves plane or low-convex, stiffly coriaceous or woody, the margin thickened; valves elastically dehiscent from the apex. Seeds discoid or compressed-ovoid or -rhomboid, the testa hard, the pleurogram either U-shaped or lacking. Southwestern U.S.A. south to Uruguay, warm-temperate Argentina, and northern Chile; ca. 135 species, 18 in Venezuela, 13 of these in the flora area. This treatment is based on the monograph of Calliandra by R. C. Barneby (Memoirs of the New York Botanical Garden 74(3): 1–223. 1998). Key to the Species of Calliandra 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(2). 5.

Leaves with pinnae always in 1 pair ......................................................... 2 Leaves, at least the lower ones, with pinnae in 2–many pairs ................ 7 Leaflets generally > 20, < 2 cm long .......................................................... 3 Leaflets 4–8, generally much longer than 2 cm long ................................ 5 Leaflets in 2.5–5 pairs, each of the terminal pair measuring 2.5–10.5 × 1.2–3.6 cm ............................................................................... C. guildingii Leaflets in 3–13 pairs, or the longest leaflets < 3 cm long ....................... 4 Leaflets broadly linear, 2.5–6 times as long as wide, 2.5–4.5 mm wide ...................................................................................................... C. riparia Leaflets rhombic or rhombic-ovate, 1.5–2.3 times as long as wide, 2– 15 mm wide ........................................................................ C. surinamensis Distal pair of leaflets small, the largest leaflets 1.5–4 cm long .... C. tergemina Distal pair of leaflets larger, the largest blades 4–12 cm long ................. 6

Calliandra 601

6(5).

Leaflets narrowly or broadly but inequilaterally elliptic or oblance-elliptic, the distal pair 4–10 × 1–3.2 cm; pods stiffly ascending; seeds 7.5–9 × 5.5–7 mm .................................................................................... C. coriacea 6. Leaflets semi-ovate or inequilaterally ovate or lance-elliptic, the distal pair 5.5–16 × 2–6(7) cm; pods plaiotropic or geotropic; seeds 11–22 × 7– 12 mm ........................................................................................ C. trinervia 7(1). Primary vein of leaflets immediately posterior to midrib incurved-ascending through 1/4–3/4 of blade, or, in some very small leaflets, barely visible, the primary venation thus either palmate or simple ................... 8 7. Primary vein of leaflets immediately posterior to midrib produced, parallel to midrib, almost or quite to blade’s apex, 3-veined, its whole length; peduncles arising from primary leaf axils toward apex of stem, the inflorescence, by suppression of distal leaves, pseudoracemose ........... 10 8(7). Capitula sessile or almost so, the peduncle < 3 mm long or obsolete; leaflets acute at apex, the longest ones 4–7(–9) mm long; pod 6–8 mm wide ..................................................................................................... C. cruegeri 8. Capitula clearly pedunculate, the peduncle (4–)15–60 mm long; leaflets either longer, or obtuse at apex; pod 10–23 mm wide .......................... 9 9(8). Capitula with 5–9(10) flowers; seed coat with pleurogram .... C. glomerulata 9. Capitula with (9)10–28 flowers; seed coat lacking pleurogram .......... C. laxa 10(7). Perianth completely glabrous; androecium 6- or 7-merous .............. C. rigida 10. Perianth, at least the corolla, white-silky externally; androecium (10–) 12–57-merous ....................................................................................... 11 11(10). Larger leaflets 13–22 × 2.4–3.6 mm ................................... C. pakaraimensis 11. Larger leaflets 6–12 × 1–2.5 mm ............................................................. 12 12(11). Primary and secondary leaf axes mostly straight, ascending; leaflets of longer pinnae in 21–52 pairs; androecium 12–20-merous; no granular trichomes .................................................................................. C. tsugoides 12. Primary and secondary leaf axes widely spreading and recurved; leaflets of longer pinnae in 46–80 pairs; androecium mostly 36–57-merous; some granular trichomes mixed with plain gray ones in the inflorescence ...................................................................................... C. vaupesiana Calliandra coriacea (Humb. & Bonpl. ex Willd.) Benth., London J. Bot. 3: 95. 1844. —Inga coriacea Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1010. 1806. Small tree 2–10 m tall. Riparian forests along black-water rivers, 100–300 m; Amazonas (Río Casiquiare, Río Siapa near base of Cerro Aracamuni, Río Ucata, base of Sierra de la Neblina). Belize, Panama, Colombia, Guyana, Suriname, Brazil. ◆Fig. 507.

level to 300 m; Bolívar (Río Orinoco), Amazonas (near Puerto Ayacucho). Apure, Falcón, Guárico, Nueva Esparta, Sucre; Trinidad, Guyana, Brazil (Roraima). ◆Fig. 508.

Calliandra cruegeri Griseb., Fl. Brit. W.I. 224. 1860, “Cruegerii.” Calliandra affinis Pittier, Bol. Minist. R.R. E.E. no. 10–12, 51. 1927, reprinted in Arb. Legum., part 1, Trab. Mus. Comercial Venezuela 2: 81. 1927. Shrub or small tree to 6 m tall. Deciduous forests, savannas, granitic outcrops, near sea

C. glomerulata var. glomerulata Savanna thickets, deciduous forests, granitic outcrops, 50–500 m; Bolívar (lower Río Caroní, Serranía de Imataca), Amazonas (Coromoto). Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Monagas, Sucre, Táchira, Yaracuy, Zulia; Colombia (Norte de Santander), Guyana, Brazil (Roraima). ◆Fig. 512.

Calliandra glomerulata H. Karst., Fl. Columb. 2: 5, t. 103. 1862. Shrub or tree 2–10 m tall. Colombia, Venezuela, Guyana, Brazil; 2 varieties, both in Venezuela, 1 of these in the flora area.

602

M IMOSACEAE

Calliandra guildingii Benth., London J. Bot. 3: 96. 1844. Tree 3–12 m tall. Riparian forests, 100– 200 m; Amazonas (Río Casiquiare). Aragua, Distrito Federal, Miranda, Monagas, Sucre, Táchira, Yaracuy; Lesser Antilles (cultivated?), Colombia, Trinidad, Ecuador, Peru. Calliandra laxa (Willd.) Benth., Trans. Linn. Soc. London 30: 551. 1875. —Acacia laxa Willd., Sp. Pl. 4: 1069. 1805. Large shrub or small tree 5–20 m tall. Panama, Colombia, Venezuela, Guyana, Brazil; 3 varieties, all in the flora area.

Rocky stream banks, savanna margins, 300–500 m; Bolívar (Río Acanán, Río Asa 62 km southeast of La Paragua, Río Urimán above Salto Acarima, Río Kavac near base of Auyán-tepui). Endemic. Calliandra pakaraimensis R.S. Cowan, Mem. New York Bot. Gard. 10(1): 142. 1958. Calliandra resupina R.S. Cowan, Mem. New York Bot. Gard. 10(4): 65, fig. 45. 1961. Shrub 1–7 m tall. Montane savanna, xeromorphic woodlands, 500–1500 m; Bolívar (Gran Sabana). Guyana. ◆Fig. 514.

Key to the Varieties of C. laxa 1. Petiole of well-developed leaves ca. twice as long as the first interpinnal segment of leaf stalk; larger leaflets 4–6.5 mm long ................................ var. urimana 1. Petiole of well-developed leaves < 1.5 times as long as the first interpinnal segment of leaf stalk; larger leaflets 8– 21 × 2–8 ............................................. 2 2. Longer peduncles 6–20 mm long; leaflets of longer pinnae in 18–30 pairs .............. .............................................. var. laxa 2. Longer peduncles 20–65 mm long; leaflets of longer pinnae in 8–18 pairs ................ ..................................... var. stipulacea C. laxa var. laxa Calliandra trijugata Schery, Fieldiana, Bot. 28: 256. 1952. Savannas, rocky plains, 50–300 m; Bolívar (Cuidad Bolívar to Guasipati and El Miamo, Serranía de Imataca). Anzoátegui, Aragua, Distrito Federal, Miranda, Monagas, Nueva Esparta, Sucre, Trujillo; Panama, Colombia. C. laxa var. stipulacea (Benth.) Barneby, Mem. New York Bot. Gard. 74(3): 33. 1998. —Calliandra stipulacea Benth., J. Bot. (Hooker) 2: 137. 1840. Forest-savanna ecotone, riparian forests, 100–1000 m; widespread in Bolívar, Amazonas (slopes of Cerro Duida, between Culebra and Cerro Duida, Puerto Ayacucho, near Río Coro-Coro, Río Ocamo, base of Sierra de la Neblina). Colombia, Guyana, Brazil. ◆Fig. 511. C. laxa var. urimana Barneby, Mem. New York Bot. Gard. 74(3): 33. 1998.

Calliandra rigida Benth., London J. Bot. 5: 103. 1848. Erect shrub 1–3 m tall. Upland savannas, gallery forests on sandy soil, forest edges, 200–1200 m; Bolívar (base of La Escalera, Salto Kamá in Río Aponguao basin, Río Kamoirán, Río Karaurín). Guyana. ◆Fig. 505. Calliandra riparia Pittier, Bol. Minist. RR. EE. no. 8/9, reprinted in Arb. Arbust. Venez. 6–8: 80. 1927. Shrub 1.5–6 m tall. Semideciduous forests, rocky river banks, 50–200 m; northern Bolívar. Venezuelan Andes and Coastal Cordillera, Falcón, Zulia; West Indies, Panama, Colombia, Guyana, widely cultivated elsewhere. Calliandra surinamensis Benth., London J. Bot. 3: 105. 1844. Calliandra tenuiflora Benth., Trans. Linn. Soc. London 30: 547. 1875. Small tree 1.5–8 m tall. Shrublands, rocky riverbanks, 50–1200 m; Bolívar (base of Apacará-tepui, Gran Sabana), Amazonas (Isla Sebastián in Río Casiquiare, slopes of Sierra de la Neblina). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 510. Calliandra tergemina (L.) Benth., London J. Bot. 3: 96. 1844. —Mimosa tergemina L., Sp. Pl. 517. 1753. Calliandra caripensis Willd., Sp. Pl. 4: 1009. 1806. Shrub 1–4 m tall. Savanna margins, deciduous forests, 50–200 m; Bolívar (Guasipati to El Callao). Lara east to Sucre; Mexico, Central America, Colombia, Guyana, Suri-

Calliandra 603

Fig. 505. Calliandra rigida

Fig. 507. Calliandra coriacea

Fig. 506. Calliandra trinervia var. trinervia

604

M IMOSACEAE

Fig. 508. Calliandra cruegeri

Fig. 509. Calliandra vaupesiana var. oligandra

Fig. 510. Calliandra surinamensis

Calliandra 605

Fig. 511. Calliandra laxa var. stipulacea

Fig. 512. Calliandra glomerulata var. glomerulata

606

M IMOSACEAE

Fig. 513. Calliandra tsugoides

Fig. 514. Calliandra pakaraimensis

Cedrelinga 607

name, French Brazil.

Guiana,

Ecuador,

Peru,

Calliandra trinervia Benth., London J. Bot. 3: 94. 1844. Shrub or tree 3–10(–30) m tall. Mexico, Central America, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; 7 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of C. trinervia 1. Young stems and leaf axes with straight, vertically erect hairs to 0.8–2 mm long ..................................... var. pilosifolia 1. Young stems and leaf axes either glabrous or pilosulous with hairs < 0.5 mm long ....................................... var. trinervia C. trinervia var. trinervia Small spreading tree or shrub 3–8 m tall. Riparian forests, 50–200 m; Amazonas (Río Casiquiare, near San Fernando de Atabapo). Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 506. C. trinervia var. pilosifolia (R.S. Cowan) Barneby, Mem. New York Bot. Gard. 74(3): 116. 1988. —Calliandra pilosifolia

R.S. Cowan, Mem. New York Bot. Gard. 10(1): 142. 1958. —Hoja de venado. Small tree 3–5 m tall. Riparian forests, river banks, 100–200 m; Amazonas (western base of Cerro Sipapo, Río Casiquiare, Río Yatúa, near San Carlos de Río Negro). Brazil. Calliandra tsugoides R.S. Cowan, Mem. New York Bot. Gard. 10(1): 143. 1958. Shrub 1–10 m tall; leaflets ventrally glossy. Tepui shrublands, rocky lowland black-water river banks, 100–1500 m; Bolívar (Cerro Guaiquinima, Uei-tepui), Amazonas (Cerro Autana, Cerro Sipapo, Cerro Yapacana, and associated rivers). Brazil (Amazonas). ◆Fig. 513. Calliandra vaupesiana R.S. Cowan, Bot. Mus. Leafl. 18: 142, t. 29d–f. 1958. Shrub 0.5–2 m tall. Colombia (Vaupés), Venezuela; 2 varieties, 1 in Venezuela. C. vaupesiana var. oligandra Barneby, Mem. New York Bot. Gard. 74(3): 193. 1998. White-sand savannas and shrub islands, 100–200 m; Amazonas (Río Atabapo, Río Guayapo, Río Orinoco 10 km above mouth of Ventuari). Colombia (Vaupés). ◆Fig. 509.

7. CEDRELINGA Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 70. 1922. by Rupert C. Barneby and James W. Grimes Potentially gigantic emergent forest trees, unarmed. Leaves amply bipinnate; a depressed petiolar nectary between or below the first of 2 or 3 pairs of pinnae; leaflets (2)3 or 4 pairs, glabrous, lustrous and reticulate on upper surface, dull on lower surface, the larger ones 6–11 × 3–5.5 cm. Inflorescence of small capitula fasciculate at nodes of terminal panicle. Flowers isomorphic. Calyx campanulate; corolla 5merous, its lobes longer than the tube. Filaments numerous, proximally united to corolla and connate into a tube a little longer than the corolla, the anthers without terminal gland. Pods pendulous, lomentiform, the 2–6 monospermous articles chartaceous, flat, twisted through ± 90° (so resembling links of a chain). Seeds large, flattened, with membranous testa and lacking pleurogram. Southeastern Colombia, southern Venezuela, Suriname, French Guiana, eastern Ecuador, eastern Peru, Brazil (Acre, Amazonas, Mato Grosso, Pará); 1 species. Cedrelinga cateniformis (Ducke) Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 70. 1922. —Piptadenia catenaeformis Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 17, pls. 5, 6. 1915. —Pithecellobium catenaeformis (Ducke) L. Cárdenas, Revista Fac. Agron. (Maracay) 7(3): 124. 1974. —Mure, Cachicama, Guaura.

Tree 25–70 m tall; flowers small greenish; pods anemochorous. Nonflooded evergreen lowland forests, 100–300 m; Bolívar (Rio Canaracuni), Amazonas (northern base of Sierra de la Neblina, Maroa to Yavita, San Carlos de Río Negro to Solano). Other distribution as in genus. ◆Fig. 515.

608

M IMOSACEAE

Fig. 515. Cedrelinga cateniformis

Chloroleucon 609

8. CHLOROLEUCON (Benth.) Britton & Rose, N. Amer. Fl. 23: 36. 1928. —Pithecellobium sect. Chloroleucon Benth., Trans. Linn. Soc. London 30: 597. 1875. —Chloroleucon (Benth.) Britton & Rose ex Record, Trop. Woods 10: 24. 1927, without description, “Chlorleucum.” by Rupert C. Barneby and James W. Grimes Deciduous trees with extremely dense wood, armed at some or all nodes of stem with 1 or 2 thorns (modified branchlets) or sometimes unarmed. Leaves bipinnate; stipules membranous, caducous, but each branchlet terminating in a perule of striate scales; petiole charged with a cupular nectary; leaflets of small or median size. Inflorescence of pedunculate capitula arising singly or geminate in the axil of coetaneous leaves. Flowers sessile, 5-merous polyandrous, isomorphic or the terminal one of each capitulum slightly larger. Calyx campanulate; corolla tubular. Filaments united into a tube; anthers lacking terminal gland. Pods linear compressed, either gently incurved or circinnate, tardily inertly dehiscent. Seeds with hard testa and a pleurogram on each face; aril none. Dry regions from tropical Mexico and the Antilles to Argentina; 10 species, 2 in Venezuela, both in the flora area. Key to the Species of Chloroleucon 1.

1.

Pod circinnate, forming about 2 complete turns; pinnae of larger leaves 5 or 6 pairs and their leaflets to 25–30 pairs; leaflets linear, commonly < 8 × 2 mm ............................................................................ C. eurycyclum Pod almost straight or falcate, not twisted; pinnae of larger leaves 2 or 3 pairs and their leaflets 3 or 4 pairs; leaflets obovate, 15–22 × 10– 15 mm ...................................................................................... C. mangense

Fig. 516. Chloroleucon mangense var. tetrazyx

610

M IMOSACEAE

Chloroleucon eurycyclum Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 140. 1996. Tree 12–24 m tall; trunk to 45 cm DBH; pod coiled into a flattened spiral, the valves 11–13 mm wide. Semideciduous forests, 200– 300 m; northern Bolívar (Reserva Forestal Paragua). Endemic. Chloroleucon mangense (Jacq.) Britton & Rose, N. Amer. Fl. 23: 38. 1928. —Mimosa mangensis Jacq., Enum. Syst. Pl.

34. 1760. Greater Antilles, Colombia, northwestern Venezuela, northeastern Bolivia; 6 varieties, 3 in Venezuela, 1 of these in the flora area. C. mangense var. tetrazyx Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 157. 1996. Small tree ca. 2 m tall. Semideciduous scrub on hillsides, 100–300 m; central Bolívar (Río Caroní basin to Altiplanicie de Nuria). Endemic. ◆Fig. 516.

9. ENTADA Adans., Fam. Pl. 2: 318. 1763, nom. cons. Entadopsis Britton & Rose, N. Amer. Fl. 23: 191. 1928. by Rupert C. Barneby Unarmed (in flora area), potentially scandent shrubs of lowland, often riparian forests, pubescent with simple hairs or glabrate, eglandular. Leaves bipinnate, the leaflets in opposite pairs, petiolar nectaries lacking. Inflorescence an ample terminal, usually 1-sided panicle of spiciform racemes. Flowers 5-merous 10-androus. Calyx campanulate, short-toothed; petals erect, valvate in bud, free, glabrous, ca. 2–3 mm long. Filaments free, less than twice as long as perianth; anther-sacs oblong, introrse, the connective sometimes much thickened, always tipped with a (caducous) gland. Pod very large, narrow-oblong, planocompressed in flora area species, ca. 25– 45 × 5.5–8 cm, the thin exocarp exfoliating at maturity and the stiffly papery mesocarp separating from the persistent replum and breaking into 1-sided, individually indehiscent articles 3–6 times wider than long. Seeds exarillate, compressed, with pleurogram; endosperm absent. Americas, Africa; 30 species, 3 in Venezuela, 2 of these in the flora area. Entada is a polymorphic group; the two species in the flora area form the segregate genus Entadopsis, as defined in the foregoing description, and differ from genuine Entada s. str. in lack of foliar tendrils, craspedial dehiscence of the pod, and relatively small seeds with pleurogram. Key to the Species of Entada 1.

1.

Pinnae of largest leaves 4–8 pairs; leaflets of longest pinnae 12–20 pairs; connective of anthers fleshy, as broad as anther-sacs or almost so ................................................................................................ E. polyphylla Pinnae of largest leaves 3 or 4(5) pairs; leaflets of longest pinnae 6–9(10) pairs; connective of anthers narrow, inconspicuous ........... E. polystachya

Entada polyphylla Benth., J. Bot. (Hooker) 2: 133. 1840. —Entadopsis polyphylla (Benth.) Britton, N. Amer. Fl. 23: 191. 1928. Shrub, potentially vining, to 10 m tall, sometimes independently arborescent, with long compound, horizontal or pendulous inflorescences composed of vertically ascending flower spikes; flowers whitish, disagree-

ably strong-scented. Evergeen lowland forests, either seasonally flooded or not, 100– 200 m; Amazonas (along Caño Yapacana and Río Casiquiare). Puerto Rico, Guyana, Suriname, French Guiana, interruptedly widespread over the northern half of the Amazon basin, from interior Ecuador and Peru to northeastern Brazil. ◆Fig. 517.

Entada 611

Entada polystachya (L.) DC., Prodr. 2: 425. 1825. —Mimosa polystachia L., Sp. Pl. 520. 1753. —Entadopsis polystachya (L.) Britton, N. Amer. Fl. 23: 191. 1928. Mimosa bipinnata Aubl., Hist. Pl. Guiane 946. 1775. Potentially scandent shrub or tree to 10 m tall or more; flowers heavily fragrant, white

turning yellowish or reddish brown, attractive to ants. Evergreen lowland forests, usually along river banks, 50–100 m; northern Bolívar (along and near Río Orinoco, downstream from the mouth of Río Parhueña). Western Mexico, Central America, Hispaniola, Puerto Rico, Lesser Antilles, tropical South America.

Fig. 517. Entada polyphylla

612

M IMOSACEAE

10. ENTEROLOBIUM Mart., Flora 20(2) Beibl.: 117. 1837. by Rupert C. Barneby and James W. Grimes Unarmed trees, often of large size, closely resembling Albizia in foliage, petiolar nectaries, inflorescence, and individual flowers, but different in the characteristic fruits. Pods compressed but plump, incurved into a full or nearly full circle, indehiscent, the mesocarp of valves sweet and fleshy but woody in age. Seeds transverse, arranged in either 1 or 2 ranks of 1-seeded compartments, the testa hard, with pleurogram. Southern Mexico and the Antilles to Argentina; 10 species, 3 in Venezuela, 2 of these in the flora area. Enterolobium cyclocarpum Griseb., which is frequent in the Río Orinoco lowlands of central Venezuela, has not yet been found in the flora area, but is to be expected there. It has acuminate, lanceolate leaflets 2–3.5 mm wide and the periphery of the pod is undulately impressed between seeds. Also, Enterolobium barnebianum A.L. Mesquita & M.F. Silva, which is known from areas close to the southern limit of the flora area in northern Amazonas and Roraima states in Brazil, is likely to be

Fig. 518. Enterolobium barinense

Hydrochorea 613

found in the flora area. It differs from E. barinense in its lance-oblong leaflets with a simple and strongly displaced midvein. Key to the Species of Enterolobium 1. 1.

Pinnae of larger leaves 3–8(9) pairs; leaflets lanceolate or oblong, 10–20 × 2.5–6.5 mm; pod diameter 7–10 cm ......................................... E. barinense Pinnae of larger leaves 11–30 pairs; leaflets linear, 2–10 × 0.4–1 mm; pod diameter 3–6.5 cm ............................................................. E. schomburgkii

Enterolobium barinense L. Cárdenas & H. Rodr., Ernstia 21: 1. 1983. —Caro blanco. Tree 8–20 m tall or more. Lower montane and semideciduous forests, 100–300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Represa Guri), Amazonas (Río Ocamo). Barinas, Mérida. ◆Fig. 518. Enterolobium schomburgkii (Benth.) Benth., Trans. Linn. Soc. London 30:

599. 1875. —Pithecellobium schomburgkii Benth., London J. Bot. 3: 219. 1844. —Curariua grande, Guaranaramuji (Yanomami), Ududu (Yekwana). Tree 10–50 m tall. Nonflooded evergreen lowland forests, 100–200 m; Bolívar (lower Río Caroní), Amazonas (Culebra, Río Orinoco, Río Padamo). Southern Mexico, widely but discontinuously dispersed in tropical South America, southeastern Brazil.

11. HYDROCHOREA Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 23. 1996. Pithecellobium sect. Samanea ser. Corymbosa Benth., London J. Bot. 3: 221. 1844. by Rupert C. Barneby and James W. Grimes Unarmed trees, in most respects resembling Albizia and Balizia, differing from the first and resembling the second in inflorescence architecture, differing from Albizia further in pinnate venation of leaflets, and from Balizia further in lomentiform fruits. Pubescence of young growth gray-brown. Stipules linear-subulate, caducous. Petiolar nectaries either sessile or stipitate. Pinnae 1–7 pairs per leaf and leaflets of longer pinnae 3–35 pairs, the longest leaflets 1.2–6 cm. Capitula arising from leaf axils, the flowers heteromorphic, the peripheral ones slenderly pedicellate, the 1–4 innermost (sub)sessile, larger. Filaments including tubes mostly 10–24. Ovary truncate at apex. Pods sessile or almost so, in profile linear or broad-linear, nearly straight, ca. 5–12 × 0.75–1.9 cm, laterally compressed but low-convex over each seed, at maturity lomentiform, the coriaceous or subligneous valves breaking horizontally into square or oblong, 1-seeded articles. Seed coat hard, areolate; pleurogram complete. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 4 species, 3 in Venezuela, all in the flora area. Key to the Species of Hydrochorea 1. 1. 2(1).

Leaflets of longer pinnae 14–35 pairs; pods 7.5–9 mm wide ... H. gonggrijpii Leaflets of longer pinnae 3–11 pairs; pods (9–)10–19 mm wide ............... 2 Pinnae of larger leaves 2–7 pairs, nearly always 3 pairs or more; leaflets of longer pinnae 4–11(–14) pairs and the larger ones 1–3 cm long ................................................................................................ H. corymbosa

614

2.

M IMOSACEAE

Pinnae of larger leaves mostly 1 pair (randomly 2 pairs only in var. scheryi) local on upper Río Orinoco; leaflets of longer pinnae (2)3– 6 pairs and the larger ones ca. 3–6 cm long .......................... H. marginata

Hydrochorea corymbosa (Rich.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 27. 1996. —Mimosa corymbosa Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. —Pithecellobium corymbosum (Rich.) Benth., London J. Bot. 3: 221. 1844. —Samanea corymbosa (Rich.) Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 55. 1926. —Arthrosamanea corymbosa (Rich.) Kleinhoonte in Pulle, Fl. Suriname 2(2): 327. 1940. —Cathormion corymbosum (Rich.) Burkart, Darwiniana 13: 446. 1964. —Albizia corymbosa (Rich.) G.P. Lewis & P.E. Owen, Legumes Ilha de Maracá 42. 1989. —Hueso de pescado rebalsero. Tree (3–)5–20 m tall. Seasonally flooded forests, along rocky river banks, 50–400 m; frequent in central and western Bolívar (Río Parguaza) and southern Amazonas (Río Baría, Río Siapa). Widely dispersed almost throughout the Amazon basin and Guyana, Suriname, and French Guiana. ◆Fig. 519.

Hydrochorea gonggrijpii (Kleinhoonte) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 25. 1996. —Pithecellobium gonggrijpii Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 44. 1925. —Arthrosamanea gonggrijpii (Kleinhoonte) Kleinhoonte in Pulle, Fl. Suriname 2(2): 238. 1940. Tree or bushy treelet 1.5–20 m tall. Flooded riparian forests, rocky river banks or creek beds, 100–500(–1200) m; southern Bolívar (Gran Sabana, Río Acanán), southern Amazonas. Guyana, Suriname. The typical form, uncommon in the flora area, has gray-puberulent flowers and slender pods with sutures scarcely 0.5 mm wide. Commoner in the foothills of tepuis of the Gran Sabana is a form with glabrous flowers and sutures of the pod to 1.5 mm wide. Yet another form, known only from southern Amazonas, has an atypically broad pod to 11 mm wide with slender sutures.

Fig. 519. Hydrochorea corymbosa

Inga 615

Hydrochorea marginata (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 29. 1996 —Pithecellobium marginatum Spruce ex Benth., Trans. Linn. Soc. London 30: 586. 1875. Arborescent shrub, precociously flowering when 2–3 m tall, in age attaining ca. 25 m tall, closely resembling H. corymbosa except for fewer pinnae and leaflets, and for larger flowers. Venezuela, Brazil; 3 varieties, 2 in Venezuela, both in the flora area. The third variety, marginata, is endemic to Amazonian Brazil. Key to the Varieties of H. marginata

H. marginata var. panurensis (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 32. 1996. —Pithecellobium panurense Spruce ex Benth., Trans. Linn. Soc. London 30: 586. 1875. —Arthrosamanea panurensis (Spruce ex Benth.) Pittier, 3rd Conf. Interamer. Agric. Caracas 359. 1945. Tree mostly 3–10 m tall. Seasonally flooded forests along black-water streams, 100–300 m; western Amazonas. Adjacent Colombia and Brazil. A leaf of Colella 2161 (NY) from Río Casiquiare, Amazonas, has two pairs of pinnae, suggesting passage to var. scheryi.

1. Pedicel of lower peripheral flowers 6.5–13 mm; pinnae consistently one pair in all leaves; upper Río Negro and north to mouth of Río Atabapo, below lat. 4° N ................................... var. panurensis 1. Pedicel of lower peripheral flowers 4–5 mm; pinnae of random upper leaves 2 pairs; local on upper Río Orinoco between lat. 5° and 6° N ..... var. scheryi

H. marginata var. scheryi Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 32. 1996. —Pithecellobium bijugatum Schery, Fieldiana, Bot. 28: 258. 1952. Treelet to 7 m tall. Margins of gallery forests, 50–200 m; Amazonas (Río Orinoco basin between Sanariapo and Puerto Ayacucho). Endemic.

12. INGA Mill., Gard. Dict. Abr. ed. 4. 1754. Affonsea A. St.-Hil., Voy. Distr. Diam. 1: 385. 1833. Ingaria Raf., Sylva Tellur. 119. 1838. Feuilleea Kuntze, Revis. Gen. Pl. 1: 182. 1891, pro parte, non Fevillea L. 1737.

by Lourdes Cárdenas and Paul E. Berry Unarmed trees or shrubs. Branches generally pubescent when young, frequently lenticellate. Leaves once-pinnate, always even-pinnate; leaflets opposite, 1– many pairs, glabrous to pubescent; veins frequently more prominent on the upper surface; rachis winged, marginate, or terete; gland sessile or stipitate, present between each pair of leaflets, rarely obsolete; stipules mostly small and caducous, occasionally larger or persistent. Inflorescences axillary or terminal, solitary or of fascicles, spikes, racemes, pseudopanicles, corymbs, capitula, or umbels; floral bracts often conspicuous and persistent, occasionally partly fused to form an involucre. Flowers sessile, subsessile, or pedicellate, whitish, greenish yellow, or yellowish. Calyx tubular, turbinate, or campanulate, usually 5-dentate, or short-lobed, sometimes sub-bilabiate, frequently pubescent; corolla tubular, obconical, or funnelshaped, (4)5(6)-lobed, usually pubescent. Stamens ca. 20–350, basally connate, the tube exserted or included; anthers without glands; pollen in polyads. Ovary sessile or stipitate, ovules 10–32; style filiform, frequently longer than stamens; stigma discoid or capitellate. Legume oblong, straight, falcate, twisted or spiral, flattened, 4angled, subcylindric, or cylindric, generally with thickened margins, indehiscent or late-dehiscent, glabrous or pubescent. Seeds fleshy, thin-walled, the testa developing a thick white sugary pulp (sarcotesta).

616

M IMOSACEAE

Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay, Chile, Argentina; ca. 300 species, 56 in Venezuela, 39 of these in the flora area. The species delimitations used in this treatment follow those of T. D. Pennington (The Genus Inga. Royal Botanic Gardens, Kew. 1997). Key to the Species of Inga 1. 1.

Inflorescences umbelliform or simple-capitate ......................................... 2 Inflorescences not umbelliform, consisting of spikes, racemes, pseudopanicles, corymbs, or capitula arranged in a non-umbelliform way .............................................................................................................. 11 2(1). Calyx inflated, striate, 16–30 mm long; staminal tube 43–53 mm long ........................................................................................................ I. inflata 2. Calyx neither inflated nor striate, < 10 mm long; staminal tube shorter ... 3 3(2). Inflorescence umbellate (pedicels mostly ≥ 3 mm long) ........................... 4 3. Inflorescence capitate (pedicels ≤ 2 mm long) ........................................... 8 4(3). Foliar nectary shortly stalked (to 1 mm tall), ± cylindrical in side view with unexpanded head ca. 0.5 mm diameter ...................... I. heterophylla 4. Foliar nectary sessile or slightly raised, cup-shaped, shield-shaped, orbicular, or urn-shaped ............................................................................ 5 5(4). Leaf rachis unwinged .................................................................... I. paraensis 5. Leaf rachis winged ..................................................................................... 6 6(5). Pedicels 1–5 mm long ................................................................... I. umbratica 6. Pedicels > 7 mm long ................................................................................. 7 7(6). Leaf rachis 1–4 mm wide, foliar nectaries cup-shaped, 0.75–1.5 mm diameter; inflorescence glabrous ..................................................... I. sertulifera 7. Leaf rachis 3–12 mm wide, foliar nectaries flat or disk-shaped, 1.5– 2.5 mm diameter; inflorescence puberulous ......................... I. umbellifera 8(3). Leaf rachis winged ..................................................................................... 9 8. Leaf rachis unwinged ............................................................................... 10 9(8). Leaflets (3–)5(6) pairs; peduncle 2.5–4.5 cm long, pedicels ca. 0.5 mm long; corolla ca. 5 mm long; fruit 10–30 cm long, flat or only slightly swollen around the seeds ........................................................ I. gracilifolia 9. Leaflets 1–4 pairs; peduncle 0.3–1.2 cm long, pedicels 0.5–2 mm long; corolla 3–5 mm long; fruit 8–16 cm long, strongly swollen around the seeds and ± moniliform ............................................................ I. lateriflora 10(8). Leaflets in 2 pairs; largest leaflets 12–15 cm long ............................ I. bijuga 10. Leaflets in (3–)5(6) pairs; largest leaflets 2–7 cm long .............. I. gracilifolia 11(1). Calyx 1–2(–3) mm long; corolla glabrous, scarcely pubescent, or puberulous ....................................................................................................... 12 11. Calyx > 3 mm long; corolla tomentose, silky-villose, strigose or woolly, rarely glabrous ..................................................................................... 20 12(11). Leaves with 1 pair of leaflets .................................................................. I. ulei 12. Leaves with 2 or more pairs of leaflets ................................................... 13 13(12). Inflorescences elongate, mostly > 7 cm long; floral rachis usually longer than peduncle ....................................................................................... 14 13. Inflorescences short, ≤ 6 cm long; floral rachis usually shorter than peduncle ................................................................................................... 16

Inga 617

14(13). Leaflets coriaceous; foliar rachis not winged or somewhat narrowly winged .......................................................................................... I. laurina 14. Leaflets membranous or subcoriaceous; foliar rachis clearly winged .... 15 15(14). Leaflets 3–4 times as long as broad, with 5–9 pairs of secondary veins; flowers sessile ......................................................................... I. marginata 15. Leaflets slender, 4–6 times as long as broad, with 10–12 pairs of secondary veins; flowers with pedicels 1.5–2 mm long ...................... I. salicifolia 16(13). Foliar rachis entirely or partly winged ................................................... 17 16. Foliar rachis not winged .......................................................................... 18 17(16). Leaf rachis winged only part way below each leaflet node; fruits 2.5– 3.5 cm wide ................................................................................ I. bourgonii 17. Leaf rachis winged from base to apex of each leaflet internode; fruits 1.5– 2.3 wide ................................................................................... I. auristellae 18(16). Leaves with 2 pairs of leaflets ........................................................... I. huberi 18. Leaves with 3–6 pairs of leaflets ............................................................. 19 19(18). Largest leaflet mostly 6–10 cm long; peduncle 0.4–2 cm long; corolla 2.5– 4.5 mm long; fruit 1.5–2 cm wide ...................................................... I. alba 19. Largest leaflet 11–19 cm long; peduncle 1–6 cm long; corolla 5–7.5 mm long; fruit 2–4 cm wide ............................................................. I. pezizifera 20(11). Fruit cylindric or subcylindric when mature, margins conspicuously veined, wider than faces; faces overlapped partially or completely by the margins; leaves with 3 or more leaflet pairs ................................ 21 20. Fruit flattened when mature, margins usually not sulcate, narrower than the sides or equal, not overlapping them or almost so; leaves with 1 or more leaflet pairs ................................................................................. 23 21(20). Gland between each leaflet pair concave, transversely compressed, mouth-shaped, wider in the middle than in the extremes, 2–3 mm diameter; fruit (30–)50–100(–200) cm long, 2–5 cm diameter, deeply furrowed in cross section; flowers sessile ........................................... I. edulis 21. Gland between each leaflet pair shield-shaped or cup-shaped, 0.75–2 mm diameter; fruit 5–30 × 1.3–2.5 × 1.5–2.5 cm, cylindrical and furrowed or somewhat flattened, partly covered by the strongly expanded margins; flowers sessile or pedicellate ............................................................... 22 22(21). Inflorescence a raceme or corymb, the flowers pedicellate; distal leaflets 10–23 × 4.5–14 cm; calyx tubular-campanulate, 4–6 mm diameter; fruit cylindrical, deeply furrowed ....................................................... I. ingoides 22. Inflorescence a spike or shortly-pedicellate raceme; distal leaflets 4.8– 13 × 1.5–5.5(–7) cm; calyx tubular to funnel-shaped, 2–4 mm diameter; fruit cylindrical or quadrangular ...................................................... I. vera 23(20). Foliar rachis unwinged or marginate; gland between each pair of leaflets sessile, never stipitate ......................................................................... 24 23. Foliar rachis winged; gland between each pair of leaflets sessile or stipitate ....................................................................................................... 30 24(23). Leaves with conspicuous, cordate-reniform, persistent stipules 1.1–3.3 × 0.7–3.8 cm ................................................................................. I. stipularis 24. Leaves with inconpicuous stipules, or if conspicuous, then elliptical, falciform, or spatula-shaped, deciduous or persistent .............................. 25 25(24). Branches lenticellate-warty; corolla glabrous except on the lobes; leaflets glabrous ....................................................................................... I. capitata

618

M IMOSACEAE

25. 26(25). 26. 27(26).

27. 28(27). 28. 29(28).

29. 30(23).

30.

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

Branches lenticellate; corolla pubescent; leaflets glabrous or more often pubescent ............................................................................................. 26 Corolla densely woolly, the trichomes forming waves towards the lobes ................................................................................................. I. rubiginosa Corolla tomentose, silky-villose, or strigose, the trichomes not undulate .............................................................................................................. 27 Leaflets glabrous except the main and secondary veins, which are somewhat puberulous; fruit 8–15(–20) × 1.4–3.1 × 0.7–1(–1.5) cm, flat when young but becoming nearly cylindrical at maturity ..................... I. nobilis Leaflets pubescent; fruit (12–)15–40 × 2–4 × 0.7–1.2 cm, mostly flat when mature .................................................................................................. 28 Leaves with 2(3) pairs of leaflets .............................................. I. leiocalycina Leaves with (3)4 or more pairs of leaflets ............................................... 29 Indumentum of lower leaflet surface with curly trichomes, foliar nectary 1–1.5 mm diameter, calyx 5–12 mm long, often striate; fruit glabrous ................................................................................................... I. multijuga Indumentum of lower leaflet surface sericeous-strigose, foliar nectary 1.5– 3.5 mm diameter, calyx 3.5–6.5 mm long; fruit velutinous ... I. thibaudiana Plants usually with stiff trichomes; fruits with faces wider than margins; foliar rachis winged; leaves with 3 or more leaflet pairs; leaflets pubescent ....................................................................................................... 31 Plants without stiff trichomes; fruits with faces wider than margins or equal (except sometimes I. sapindoides, and then fruit with subequal or equal faces and margins); foliar rachis winged, marginate, or unwinged; leaves with 1 or more pairs of leaflets; leaflets glabrous or pubescent .................................................................................................. 35 Calyx tube 3–9 mm long; corolla tube 16–30 mm long ........................... 32 Calyx tube 9–27 mm long; corolla tube 30–60 mm long, densely villose .............................................................................................................. 33 Foliar nectary subsessile or shortly stalked, cup-shaped, not sunken in the rachis; corolla tube 16–28 mm long ............................... I. cayennensis Foliar nectary partly sunken in the leaf rachis, pulvinate, with a small aperture; corolla tube 27–30 mm long ............................... I. lomatophylla Floral bracts persistent and reflexed; mature fruit puberulent and scarcely strigose ................................................................... I. macrophylla Floral bracts caducous and not reflexed; mature fruits hispid .............. 34 Nectary gland between each pair of leaflets stipitate; fruit 15–35 × 4–5 × 0.9–1 cm, straight to spirally twisted ......................................... I. fastuosa Nectary gland between each pair of leaflets sessile or somewhat raised; fruit (only young ones known) 10–12 × 1.7–2 × 0.5 cm, flat .... I. micradenia Fruit 3–4 cm wide, strongly curved or coiled; inflorescence ramiflorous; leaflets in 4 or 5 pairs ............................................................... I. melinonis Fruit 1.6–4 cm wide, ± straight; inflorescence axillary; leaflets in 1–6 pairs .............................................................................................................. 36 Flowers yellow, calyx closed in bud and splitting irregularly; leaflets in 2 or 3 pairs ........................................................................................... 37 Flowers white; calyx open in bud or splitting regularly; leaflets in (1)2– 6 pairs ................................................................................................... 38

Inga 619

37(36). Leaflets in 2 or 3 pairs, largest ones 9.5–15.5 × 2.9–5.1 cm; young shoots and leaves subglabrous; fruit 2.3–2.5 cm wide, prominently transverse striate ..................................................................................... I. neblinensis 37. Leaflets in 2 pairs, largest ones 12–21 × 6.5–11.5 cm; young shoots and leaves usually pubescent; fruit (2–)3–4 cm wide, usually without transverse striations............................................................................. I. pilosula 38(36). Leaf rachis narrowly winged, < 5 mm wide ............................................ 39 38. Leaf rachis more broadly winged, 5–15 mm wide .................................. 40 39(38). Leaflets in 2 or 3(4) pairs, largest leaflets 11–22 × 3.2–10 cm; petiolar portion of leaf rachis winged ......................................................... I. stenoptera 39. Leaflets in 4–6 pairs, largest leaflets 7–17.5 × 2.5–8.5 cm; petiolar portion of leaf rachis usually unwinged ........................................... I. thibaudiana 40(38). Leaflets in (2)3(4) pairs; foliar rachis 6–17 cm long; petiole 0.8–1.1 cm long ............................................................................................... I. sapindoides 40. Leaflets in 1–3 pairs; foliar rachis 2.5– 7.5 cm long; petiole 2–7 cm long ................................................................................................... I. splendens Inga alba (Sw.) Willd., Sp. Pl. 4: 1013. 1806. —Mimosa alba Sw., Prodr. 85. 1788, non Vahl 1807. —Guamito, Guamito montañero, Guamo, Guamo colorado, Kuarikyek (Arekuna). Inga carachensis Pittier, Bol. Minist. RR. EE. no. 8–12, reprinted in Arb. Arbust. Venez. 9/10: 262. 1929. Tree 10–35 m tall; leaflets (3)4, 5(6) pairs, rachis unwinged, foliar nectaries sessile or shortly stalked, disk-shaped or cup-shaped, 1–2.5 mm diameter; inflorescence a congested spike, axillary, often clustered on short leafless axillary shoots to 3–4 cm long, 1–4 per axil, peduncle 0.4–2 cm long, floral rachis 5–8 mm long; fruits 6–25 × 1.4–3 × 0.4–0.9 cm, convex, straight or curved, glabrous, margins 1.5–2(–5) mm thick, slightly raised. Evergreen lowland to montane forests, secondary forests, granitic outcrops, and riparian areas, 100–1300 m; Delta Amacuro (Río Cuyubini, Río Toro, Santa Catalina, Serranía de Imataca), widely scattered in Bolívar, Amazonas (San Juan de Manapiare, Isla Ratón, Río Mawarinuma, Yavita). Apure, Guárico, Táchira, Trujillo, Zulia; Mexico, Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 521. The scraping of the interior layer of the bark, mixed with water produces a soapy, brick-red fluid that is used by Arekuna Amerindians to treat scabies, eczema, and fungal infections.

Inga auristellae Harms, Notizbl. Königl. Bot. Gart. Berlin 6: 298. 1915. Tree or shrub 3–8 m tall; leaflets 2 or 3 pairs, rachis winged from base to apex of each leaflet internode, foliar nectaries sessile or shortly stalked, cup-shaped, 1–1.25 mm diameter; inflorescence axillary, a congested spike, 2 or 3 per axil, clustered near the shoot apex, peduncle 1–2 cm long, floral rachis 7–10 mm long, pedicels 0–0.5 mm long; fruits 7–15 × 1.5–2.3 × 0.5–0.9 cm, flat to convex, straight or slightly curved, margins 2–4 mm thick, slightly raised. Swampy riparian forests of black-water rivers, 100– 200 m; Amazonas (base of Sierra de la Neblina, Río Mawarinuma). French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. Inga bijuga Schery, Fieldiana, Bot. 28: 257. 1952. Tree 7–8 m tall; leaflets 2 pairs, rachis unwinged, foliar nectaries disk-shaped, 1.5– 2 mm diameter, absent between distal pair of leaflets; inflorescence axillary, capitate, solitary, peduncle 5–10 mm long, pedicels ca. 1 mm long; fruit unknown. Tepui summit forest, ca. 2200 m; Bolívar (Sororopán-tepui). Endemic. Inga bourgonii (Aubl.) DC., Prodr. 2: 434. 1825. —Mimosa bourgonii Aubl., Hist. Pl. Guiane 941, t. 358. 1775. Tree 7–15 m tall; leaflets (2)3(4) pairs, rachis winged only part way below each leaflet

620

M IMOSACEAE

node, foliar nectaries sessile or slightly stalked, cup-shaped, 1–1.5 mm diameter; inflorescence axillary, a congested spike, 2 or 3 per axil, peduncle 1–4 cm long, floral rachis 1–3 cm long; fruits 15–17 × 2.5–3.5 × 0.6–1 cm, flat when young then becoming convex, straight, faces with transverse venation, margins 1–2.5 mm thick, not or slightly raised. Evergreen lowland and seasonally flooded forests, along rivers and streams, near sea level to 200 m; Delta Amacuro (Curiapo, Misión San Francisco de Guayo, Río Cuyubini), Amazonas (base of Sierra de la Neblina, Raudal de los Guaharibos on the upper Río Orinoco, Río Yureba). Monagas; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 523. Inga capitata Desv., J. Bot. Agric. 3: 71. 1814. —Guama comestible. Tree 6–14 m tall; leaflets 2 or 3 pairs, terminal pair 8–25 × 3–10 cm, rachis terete, foliar nectaries flat or disk-shaped, often slightly sunken in the rachis, 1–2.5 mm diameter; inflorescence axillary, a congested spike, 1–3 per axil, peduncle 3–11 cm long, floral rachis 0.5–3.5 cm long; fruit 6–25 × 2–5 × 0.5–1.5 cm, flat to convex, straight or curved, faces smooth to densely lenticellate, glabrous, margins 2–10 mm thick and slightly raised in young fruit. Evergreen lowland to lower montane forests, riparian forests, 50–600 m; Bolívar (33 km south of El Dorado, near Cerro Abismo, El Tigre, Río Caura at mouth of Río Nichare, Serranía de los Pijiguaos), Amazonas (San Carlos de Río Negro). Carabobo, Miranda, Yaracuy; Costa Rica, Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 524. Inga cayennensis Sagot ex Benth., Trans. Linn. Soc. London 30: 626. 1875. -–Feuilleea cayennensis (Sagot ex Benth.) Kuntze, Revis. Gen. Pl. 1: 187. 1891. Inga dysantha Benth., Trans. Linn. Soc. London 30: 626. 1875. Tree 3–15 m tall; young growth golden-tomentose to villose; leaflets 3–6(–8) pairs, rachis winged, foliar nectaries shortly stalked or subsessile, head cup-shaped, 0.5–1 mm diameter; inflorescence a lax or congested spike or raceme, peduncle 1–3 cm long, floral rachis 2.5–6 cm long, pedicels 0–8 mm long;

fruits 15–20 × 1.7–4.2 × 0.6–0.8 cm, straight or slightly curved, faces smooth, densely hispid to villose, margins 4–6 mm thick, slightly raised. Evergreen lowland and riparian forests, 100–300 m; Bolívar (vicinity of Corozal), Amazonas (Rio Cataniapo). Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Inga edulis Mart., Flora 20(2) Beibl.: 113. 1837. —Feuilleea edulis (Mart.) Kuntze, Revis. Gen. Pl. 1: 187. 1891. —Guama, Guamo, Guamo bejuco, Guamo liso, Guamo rabo de mono, Jaguayak (Curripaco), Maipá (Arekuna), Wajuna (Yekwana). Mimosa inga Vell., Fl. Flumin. Icon. 11: pl. 3. 1827 [1831], as Mimosa ynga, non L. 1753. —Inga inga (Vell.) J. Moore, Bernice P. Bishop Mus. Bull. 10(19): 6. 1934, as Inga ynga, non Britton 1918. Inga vera H.B.K., Nov. Gen. Sp. [quarto ed.] 6: 289. 1823 [1824], non Willd. 1806. Tree 5–30 m tall; leaflets 4–6 pairs, the rachis broadly winged, foliar nectaries sessile, 2–3 mm diameter, aperture transversely compressed to reniform; inflorescence a congested or sometimes lax spike, axillary, up to 6 per axil; peduncle 1–5 cm long, floral rachis 1–4.5 cm long; corolla tube 9–20 mm long, sericeous; fruit 30–100(–200) × 2–5 cm, cylindrical, straight or spirally twisted, faces completely covered by expanded margins (deeply furrowed). Evergreen lowland and riparian forests, most often associated with towns or small farm plots through cultivation, also secondary forests, 50–700 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia, Argentina. ◆Fig. 526. Inga edulis is commonly cultivated for the edible pulp surrounding the seeds. It is widely cultivated outside its native range in both Central and South America. Inga fastuosa (Jacq.) Willd., Sp. Pl. 4: 1014. 1806. —Mimosa fastuosa Jacq., Fragm. Bot. 15, pl. 10. 1809 [1801]. —Guamo, Guamo peludo. Inga venosa Griseb. ex Benth., Trans. Linn. Soc. London 30: 623. 1875.

Inga 621

Inga guaremalensis Pittier, Bol. Comerc. Industr. Venez. no. 13, reprinted in Arb. Arbust. Venez. 1: 5. 1921. Tree 5–20 m tall; leaflets 3 or 4 pairs, setose, rachis winged (8–10 mm wide) and setose-ciliate, foliar nectaries stalked (to 1.5 mm long), the head shallowly cup-shaped, ca. 0.5 mm diameter; inflorescence a lax spike, axillary, solitary or paired, peduncle 4–5.5 cm long, flora rachis 2–3 cm long, bracts 1–1.3 cm long; calyx tube 19–25 mm long, corolla tube 35–45 mm long; fruit 15– 35 × 4–5 × 0.9–1 cm, flat, straight or spirally twisted, faces smooth but covered by dense reddish-brown stiff hairs, margins ca. 8 mm thick, not raised. Evergreen to semideciduous lowland forests, secondary forests, near sea level to 300 m; Delta Amacuro (Tucupita, near Los Castillos, Río Orocoima), northeastern Bolívar (Represa Guri, south of El Dorado, San Pedro de las dos Bocas), Amazonas (lower Río Orinoco). Venezuelan Coastal Cordillera, Táchira; Puerto Rico, Colombia, Trinidad, Guyana, French Guiana, Suriname. ◆Fig. 527. Inga gracilifolia Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 52. 1922. —Guamo monero. Tree 12–25 m tall; leaflets (3–)5(6) pairs, rachis channeled (seemingly very narrowly winged), foliar nectaries sessile or stalked, cup-shaped, ca. 0.5 mm diameter; inflorescence subcapitate, axillary on new shoots, solitary, erect, peduncles 2.5–4.5 cm long, slender, pedicels ca. 0.5 mm long; fruit 10–30 × 1.5–2 × 0.4–0.6 cm, flat to convex, straight, slightly constricted between the seeds, faces with faint reticulate venation, margins slightly raised, glabrous. Evergreen lowland forests, riparian forests, 100–600 m; Delta Amacuro (Serranía de Imataca near Río Cuyubini), Bolívar (Altiplanicie de Nuria), Amazonas (Río Cuao, Río Mawarinuma). Guyana, Peru, Brazil. Inga heterophylla Willd., Sp. Pl. 4: 1020. 1806. —Feuilleea heterophylla (Willd.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. —Mimosa parae Poir. in Lam., Encycl. suppl. 1: 44. 1810 (based on Inga heterophylla Willd.). Inga umbellata G. Don, Gen. Hist. 2: 391. 1832.

Inga protracta Steud., Flora 26: 758. 1843. Tree to 5–20 m tall; leaflets 1 or 2(–4) pairs, rachis narrowly winged, foliar nectaries stalked (to 1 mm tall), the head 0.5–1 mm diameter, not expanded; inflorescence a congested raceme or umbel, axillary, solitary or paired, peduncle 1.2–5 cm long, floral rachis 1.5–5 mm long, pedicels 3.5–8 mm long; fruit 5–15 (–20) × 1–1.5(–2.3) × 1–1.2 cm, flat when immature but becoming swollen at maturity and strongly constricted between the seeds, straight, often with undulate margin, glabrous. Evergreen lowland to lower montane and riparian forests, 100–1000 m; Bolívar (Corozal 6 km from Maniapure towards Caicara, Salto Aponguao in the Gran Sabana), Amazonas (Salto Yureba in lower Río Ventuari basin, San Carlos de Río Negro, Yavita). Monagas, Sucre; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆ Fig. 529. Inga huberi Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 49. 1992. —Guamo negro. Tree 10–30 m tall; leaflets 2 pairs, rachis semiterete, foliar nectaries sessile or shortstalked, cup-shaped, 1–1.5 mm diameter; inflorescence capitate, mostly borne below the leaves, usually several borne on short axillary shoots; fruit 8–25 × 3–5 × 1–1.5 cm, flat, straight, faces obliquely transversely ribbed, the ribs often verrucose, puberulent. Evergreen lowland forest, 200–300 m; Delta Amacuro (Río Toro). Guyana, Suriname, French Guiana, Brazil. Inga inflata Ducke, Bol. Técn. Inst. Agron. N. 2: 3. 1944. Tree 4–10 m tall; leaflets 2 pairs, rachis terete, foliar nectaries sessile, disk-shaped, ca. 2 mm diameter; inflorescence umbellate, axillary, solitary or paired, peduncle 1–6 cm long, bracts 5–10 mm long, pedicels 2–9 mm long; calyx tube 13–28 mm long inflated, striate; corolla tube 23–32 mm long, lobes 7– 8 mm long, glabrous; fruit 9–10 × 2.5 × 0.5 cm, flat, straight, faces with conspicuous transverse venation, margins ca. 5 mm thick, raised, glabrous. Seasonally flooded riparian forests of black-water rivers, 100–200 m; Amazonas (near San Carlos de Río Negro, western base of Sierra de la Neblina). Amazonian Brazil. ◆Fig. 530.

622

M IMOSACEAE

Inga ingoides (Rich.) Willd., Sp. Pl. 4: 1012. 1806. —Mimosa ingoides Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. —Feuilleea ingoides (Rich.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. —Guamo, Guamo blanco, Guamo liso, Guamo chivo, Guamo rebalsero, Moya (Panare). Tree 5–25 m tall; leaflets 3–5 pairs, rachis winged, foliar nectaries sessile, flat, 1–2 mm diameter; inflorescence a raceme, axillary, up to 4 per axil, peduncle 1.5–8 cm long, floral rachis 1.5–5 cm long, pedicels 3–10 mm long; calyx tube 6–12 mm long, puberulent, corolla tube 12–16 mm long, sericeous; fruit 25–50 × 1–1.7 cm, cylindrical, straight, faces almost completely covered by the expanded margins (furrowed in cross-section), puberulous. Usually along streams in forests, savanna borders, or semideciduous forests, near sea level to 400 m; Delta Amacuro (widespread), northeastern Bolívar (Caño Colorado, Reserva Forestal El Caura, Tres Moriches), Amazonas (Tencua). Coastal Cordillera, Cojedes, Guárico, Monagas, Lara, Sucre, Zulia; Lesser Antilles, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 531. Inga lateriflora Miq., Linnaea 19: 131. 1847. —Guama yaragua, Guamillo, Guamo, Guamo cinta, Karayek, Pilón, Yaragua (Baniva). Tree 20–30 m tall, sometimes smaller; leaflets 1–4 pairs, rachis winged (wing 2–3 mm wide), glabrous, foliar nectaries sessile, cup-shaped, 1–2 mm diameter; inflorescence subumbellate-capitate, clustered on small woody protuberances or on short leafless shoots, peduncle 0.3–1.2 cm long, puberulous, pedicels 0.5–2 mm long; fruit 8–16 × 1.3–1.5(–2.5) × 1 cm, initially flat, becoming swollen around the seeds and moniliform, margins ca. 2 mm thick, not raised, glabrous. Evergreen lowland forests, forest-savanna ecotones, edge of Mauritia palm swamps, 100–1300 m; Delta Amacuro (Río Toro), northeastern Bolívar (Altiplanicie de Nuria, Maripa), Amazonas (near San Carlos de Río Negro). Monagas; Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Inga laurina (Sw.) Willd., Sp. Pl. 4: 1018. 1806. —Mimosa laurina Sw., Prodr. 85. 1788. —Feuilleea laurina (Sw.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. —Abira

(Arekuna), Abira-yek (Arekuna), Guamo, Guamo camburito, Guamo liso, Guamo negro, Guamo pata de morrocoy, Guamo rebalsero, Kamadak (Arekuna), Pata de morrocoy. Mimosa fagifolia L., Sp. Pl. 516. 1753. —Inga fagifolia (L.) Willd. ex Benth., Trans. Linn. Soc. London. 30: 607. 1875. —Feuilleea fagifolia (L.) Kuntze, Revis. Gen. Pl. 1: 187. 1891. Mimosa fagifolia Jacq., Select. Stirp. Amer. Hist. 264, t. 164. 1763, non L. 1753. Tree 5–20 m tall; leaflets 2 or 3(4) pairs, rachis channeled but generally not winged, foliar nectaries shortly stalked or sessile, cup-shaped, 0.5–1 mm long; inflorescence a loose spike, axillary or arranged on short leafless shoots, 1–3 per axil, peduncle 0.5– 3(–5) cm long, floral rachis 3–10(–16) cm long; fruit 4.5–20 × 1.7–3.2 × 0.4–1 cm, flat to convex, straight or slightly curved, margins slightly raised or not, 2–3 mm thick, glabrous. Evergreen lowland to lower montane and riparian forests, sandy shrublands, savannas, 100–600 m; Delta Amacuro (Río Toro), Bolívar (from area of Canaima northwards). Barinas, Táchira, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 528. Inga leiocalycina Benth., London J. Bot. 4: 598. 1845. —Guama, Guamo, Guamo caraota. Inga multiflora Benth., London J. Bot. 4: 598. 1845. Tree 12–30 m tall; leaflets 2(3) pairs, rachis terete, foliar nectaries sessile to shortly stalked, cup-shaped, 1–2 mm diameter; inflorescence a congested spike, axillary or on short leafless shoots, 1–4 per axil, peduncle 3–8 cm long, floral rachis 0.7–2(–4) cm long, flowers sessile; fruit 20–40 × 2–4 × 0.7–1 cm, flat to convex, straight or slightly curved, faces swollen and sometimes verrucose over the seeds, with fine transverse venation, margins 2–3 mm thick, slightly raised, glabrous. Evergreen lowland to lower montane and riparian forests, near sea level to 900 m; Delta Amacuro (Caño Araguao and Caño Arature, Río Toro, Serranía de Imataca, east of Caño Sacupana, Caño Joba-Suburu), Bolívar (Cerro Uroi in upper Río Cuyuni basin, San Ignacio de Yuruaní), Amazonas (between

Inga 623

La Esmeralda and Mavaca, Río Ocamo). Southern Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Inga lomatophylla (Benth.) Pittier, Contr. U.S. Natl. Herb. 18: 195. 1916. —Inga speciosa var. lomatophylla Benth., Trans. Linn. Soc. London 30: 626. 1875. —Inga amazonica var. lomatophylla (Benth.) L. Cárdenas, Ernstia 51: 1. 1989, nom. superfl. —Guama, Guamo. Inga amazonica L. Cárdenas, Ernstia 51: 1. 1989. —Inga speciosa Benth., Trans. Linn. Soc. London 30: 620. 1875, non M. Martens & Galeotti 1843. Inga speciosa var. bracteifera Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 17. 1925. —Inga amazonica var. bracteifera (Ducke) L. Cárdenas, Ernstia 51: 1. 1989. Inga speciosa var. membranacea Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 59. 1922. —Inga amazonica var. membranacea (Ducke) L. Cárdenas, Ernstia 51: 1. 1989. Tree 8–15 m tall; leaflets 2–4 pairs, rachis winged, tomentose, foliar nectaries sessile, sunken in the rachis, pulvinate, ca. 1 mm diameter; inflorescence a congested spike, axillary, solitary or paired, peduncle 0.5–1.2 cm long, floral rachis 1.5–4 cm long; fruit 10–25 × 2.5–3.5 × 0.4–0.8 cm, flat, straight, faces smooth but sparsely pilose, margins ca. 3 mm thick, slightly raised. Evergreen lowland to lower montane and riparian forests, 100– 500 m; Amazonas (base of Cerro Duida above Culebra, Mavaca, near San Carlos de Río Negro, near San Fernando de Atabapo, San Pedro del Orinoco, Santa Bárbara del Orinoco). Colombia, French Guiana, Brazil. Inga macrophylla Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1015. 1806, non Hook. 1858, nec Billb. ex Beurl. 1856. —Mimosa macrophylla (Humb. & Bonpl. ex Willd.) Poir. in Lam., Encycl. suppl. 1: 42. 1810. —Cahuabari (Baniva), Guama de araguato, Guamo. Inga brachyptera Benth., London J. Bot. 4: 610. 1845. Inga bracteosa Benth., London J. Bot. 4: 609. 1845. Inga calocephala Poepp., Nov. Gen. Sp. Pl. 3: 78. 1845. Tree 4–6 m tall; leaflets (2)3 or 4 pairs, ra-

chis winged (up to 2 cm wide), foliar nectaries sessile or stalked, cup-shaped or unexpanded, 1–2.5 mm diameter; inflorescence axillary, a congested spike, solitary or paired, peduncle 2–8 cm long, floral rachis 1.8–6 cm long, bracts 1–3 cm long, persistent; calyx tube 15–27 mm long, corolla tube 30–60 mm long, villose; fruit 20–45 × 2.5–4.5 × 0.7–1.7 cm, quadrangular, straight or slightly curved, margins 0.7–1.7 mm thick, winged (to 6 mm), hirsute to glabrous. Riparian forests along black-water rivers, 100–200 m; Amazonas (Caño San Miguel, Río Guainía, San Carlos de Rio Negro to Solano). Colombia, Trinidad, Guyana, French Guiana, Brazil, Ecuador, Peru, Bolivia. ◆Fig. 532. Inga marginata Willd., Sp. Pl. 4: 1015. 1806. —Feuilleea marginata (Willd.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. —Guamita yaragua. Tree 6–20 m tall; leaflets 2 or 3 pairs, rachis winged (to 8 mm wide), foliar nectaries sessile, cup-shaped or disk-shaped, 1–1.5 mm diameter; inflorescence axillary, spicate, solitary or clustered, peduncle 0.6–3(–4) cm long, floral rachis 3.5–11.5 cm long, bracts 0.5–3 mm long, caducous, flowers sessile or with pedicels 0.5–1 mm long; fruit 7–12.5 × 0.9–1.6 × 0.6–0.9 cm, convex, straight or slightly curved, swollen around the seeds and constricted between them, margins ca. 3 mm thick, slightly raised or not, glabrous. Río Negro caatinga, evergreen lowland forests, 100–200 m; Amazonas (near San Carlos de Río Negro). Coastal Range, Zulia; Mexico, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Inga melinonis Sagot, Ann. Sci. Nat. Bot. sér. 6, 13: 335. 1882. ––Guamo balbino. Inga cyclocarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 14. 1925. Tree 8–20 m tall; leaflets 4 or 5 pairs, rachis marginate or narrowly winged below the nodes, foliar nectaries sessile, cup-shaped, 1.75–2 mm diameter; inflorescence a congested spike, mostly ramiflorous, peduncle 1–4 cm long, floral rachis 1–2 cm long; fruit 11–23 × 3–4 × 1.5 cm, flat, curved or coiled, base strongly asymmetrical, lenticellate with age, margins 6–10 mm thick, slightly raised or not, glabrous. Evergreen lowland forests, 100–300 m; Delta Amacuro (Río Toro

624

M IMOSACEAE

in Serranía Imataca). Guyana, French Guiana, Peru, Brazil. Inga micradenia Spruce ex Benth., Trans. Linn. Soc. London 30: 620. 1875. Inga negrensis Spruce ex Benth., Trans. Linn. Soc. London 30: 621. 1875. —Inga disticha var. negrensis (Spruce ex Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 6: 13. 1933. Shrub or tree 5–10 m tall; leaflets 3–5 pairs, hispid-pilose, rachis winged (to 7 mm wide), foliar nectaries shortly stalked or sessile, cup-shaped, 0.75–1.75 mm diameter; inflorescence axillary, a congested spike, solitary or paired, peduncle 1.5–4 cm long, floral rachis 0.5–1.5 cm long; calyx tube 9–11 mm long, corolla tube 21–24 mm long; fruit 10– 12 × 1.7–2 × 0.4 cm, flat, straight, faces smooth, margins ca. 3 mm thick, slightly raised, densely hispid-tomentose. Seasonally flooded riparian forests of black-water rivers, 100–500 m; Bolívar (Rio Tirica), Amazonas (widespread in southwestern part of state, Salto Yureba on lower Río Ventuari). Brazil (Amazonas). ◆Fig. 525. Inga multijuga Benth., Trans. Linn. Soc. London 30: 615. 1875. —Mimosa multijuga (Benth.) Kuntze, Rev. Gen. 1: 188. 1891. Tree 10–25 m tall; leaflets 4–10 pairs, with crinkly hairs, rachis terete, foliar nectaries sessile, 1–1.5 mm diameter, aperture circular or laterally compressed; inflorescence a congested spike, axillary, solitary or clustered, peduncle 3–5.5 cm long, with crinkly hairs, floral rachis 1.5–4 cm long; fruit 12–26 × 1.5–3.5 × 0.3–0.8 cm, flat, straight or curved, margins 4–6 mm thick, slightly raised and slightly ribbed, glabrous. Southern Mexico, Central America, Colombia, Venezuela, Ecuador; 3 subspecies, 1 in Venezuela. I. multijuga subsp. multijuga Leaflets (4)5–10 pairs, foliar nectaries disk- to cup-shaped, aperture circular; calyx 6–10 mm long, corolla 1.5–3 mm long; fruit 2.5–3.5 cm wide. Montane forests, 1200– 1400 m; Bolívar (along Río Karuay at base of Ptari-tepui). Táchira; Honduras, Costa Rica, Panama, Ecuador. Inga neblinensis L. Cárdenas & De Martino, Ann. Missouri Bot. Gard. 76: 1179. 1989.

Shrub or tree 3–8 m tall; leaflets 2 or 3 pairs, rachis winged (5–9 mm wide), foliar nectaries sessile, disk-shaped to cup-shaped, ca. 2 mm diameter; inflorescence a congested spike, axillary, solitary or paired, peduncle 1.8–2.3 cm long, glabrous, flowers yellow; fruit 8–13 × 2.3–2.5 × 0.5 cm, flat, straight or curved, faces with fine raised transverse striations, margins ca. 4 mm thick, slightly raised, glabrous. Seasonally flooded riparian forests, lower montane tepui slope forests, 100–1100 m; Amazonas (Río Mawarinuma, base and slopes of Sierra de la Neblina). Endemic. ◆Fig. 522. Inga neblinensis is closely related to I. pilosula and may prove upon further study to be a form of that species. Inga nobilis Willd., Enum. Pl. 1047. 1809. Southern Mexico, Central America, Puerto Rico, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, both in Venezuela, 1 of these in the flora area. The second subspecies, Inga nobilis subsp. quaternata (Poepp.) T.D. Penn., occurs in the Venezuelan Andes and Coastal Cordillera. It differs in having a capitate or umbellate inflorescence and pedicellate flowers. I. nobilis subsp. nobilis —Awenadu (Yekwana), Guamo, Guamo caraota, Komik (Arekuna), Poté (Arekuna). Tree or shrub 3–15 m tall; leaflets(3)4 or 5 pairs, rachis terete, tomentose to pubescent, foliar nectaries sessile, disk-shaped, 1–2.5 mm diameter, sometimes absent; inflorescence a short congested spike, axillary, solitary or paired, peduncle 0.5–8 cm long, flowers sessile or shortly pedicellate; fruit 8–15(– 20) × 1.4–3.1 × 0.7–1(–1.5 cm), flat when young becoming convex to almost cylindrical when ripe, straight or slightly curved, faces smooth or with faint transverse venation, margins 2–4 mm thick, not or only slightly raised, glabrous to puberulent, the pulp edible. Evergreen lowland and riparian forests, near sea level to 300 m; Delta Amacuro (Río Amacuro near border with Guyana), Bolívar (Serranía de Imataca), Amazonas (Río Casiquiare, Río Matacuni, Río Padamo, Río Pamoni in Casiquiare basin, western base of Sierra de la Neblina). Apure, Táchira, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 533.

Inga 625

Inga paraensis Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 12. 1925. Tree 10–30 m tall; leaflets 2 or 3 pairs, rachis semiterete, foliar nectaries sessile, diskshaped, 1–2 mm diameter; inflorescence umbellate, axillary or clustered on short axillary shoots, peduncle 2.6–8.5 cm long, pedicels 3–6 mm long; fruit 14–25(–35) × 2.5–3.5 × 0.9–1.7, flat, straight or slightly curved, slightly constricted between the seeds, faces slightly transversely striate, margin sometimes undulate, not raised, 1–3 mm thick, glabrous. Lower montane forests, 500–700 m; Bolívar (El Abismo). Guyana, French Guiana, Brazil. Inga pezizifera Benth., London J. Bot. 4: 587. 1845. —Feuilleea pezizifera (Benth.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. —Guamo, Guama pompeya, Guamo rabo de mono. Tree 8–30 m tall; leaflets (3)4–6 pairs, rachis terete, puberulous, foliar nectaries sessile or shortly stalked, cup-shaped, 1.5–3 mm diameter; inflorescence axillary clusters of congested racemes, peduncles 1–6 cm long, floral rachis 0.8–2(–3.5) cm long, pedicels 0.5–1 mm long; fruit 15–21 × 2–4 × 1–2 cm, flat to convex, straight or curved, asymmetrical, faces with faint to prominent transverse of oblique venation and often with strongly verrucose protuberances over the seeds, margins 5–6 mm thick, slightly raised or not, glabrous, pulp edible. Evergreen lowland to montane forests, riparian forests, 50–1100 m; Delta Amacuro (near Los Castillos, Río Toro), Bolívar (Altiplanicie de Nuria, La Escalera), Amazonas (slopes of Cerro Marahuaka, Río Guianía, uppermost Río Orinoco, Sierra de la Neblina). Aragua; Costa Rica, Panama, Colombia, Guayana, Suriname, French Guiana, Ecuador, Brazil. Inga pilosula (Rich.) J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 41. 1943. —Mimosa pilosula Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. —Feuilleea pilosula (Rich.) Kuntze, Revis. Gen. Pl. 1: 186. 1891. —Guama, Guamita, Guamo, Guamo amarillo, Guamo montañero, Jonanae (Guahibo), Pate (Curripaco). Mimosa lucida Vahl, Eclog. Amer. 3: 31. 1807.

Inga nitida Willd., Sp. Pl. 4: 1014. 1806. —Mimosa nitida (Willd.) Poir. in Lam., Encycl. suppl. 1: 41. 1810. Inga pilosiuscula Desv., J. Bot. Agric. 3: 71. 1814. Inga setifera DC., Prodr. 2: 432. 1825. —Feuilleea setifera (DC.) Kuntze, Revis. Gen. Pl. 1: 186. 1891. Inga affinis Steud., Flora 26: 758. 1843. Inga macrophylla Hook., Bot. Mag. t. 5075. 1858, non Humb. & Bonpl. 1806. Tree or small shrub 3–8(–18) m tall; leaflets 2 pairs, terminal pair 12–21 × 6.5–11.5 cm, rachis winged (4–9 mm wide), foliar nectaries sessile, disk-shaped or cup-shaped, 2– 3 mm diameter; inflorescence axillary, a congested spike, solitary or paired, peduncle 3–8 cm long, floral rachis 1–2.5 cm long, pedicels 0–1 mm long; fruit 8.5–20 × (2–)3–4 × 0.5– 0.6 cm, flat, straight, faces with faint to prominent transverse venation, margins 4–6 mm thick, slightly raised, longitudinally ribbed, glabrous. Riparian forests, Mauritia palm swamps, Río Negro caatinga, secondary vegetation, 100–1200 m; Bolívar (Cerro Ceitá near Santa Elena de Uairén, Gran Sabana, Río Nichare basin, near Santa María de Erebato), Amazonas (northwestern corner, lower Río Ventuari, San Carlos de Río Negro). Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 536. Inga rubiginosa (Rich.) DC., Prodr. 2: 434. 1825. —Mimosa rubiginosa Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. —Guamo, Guamo terciopelo. Tree 10–22 m tall; leaflets 3–5 pairs, velutinous, rachis terete, foliar nectaries sessile, disk-shaped, 1.5–2 mm diameter; inflorescence a congested spike or raceme, axillary and 1–3 per axil, peduncle 0.8–4 cm long, floral rachis 2–6 cm long, pedicels 0–1 mm long; corolla 20–34 mm long, undulatevillose; fruit 15–30 × 2.5–3.2 × 0.7–1 cm, flat to quadrangular, straight or curved, faces with faint transverse venation, margins 1– 1.2 mm thick, raised, longitudinally ridged, shortly velutinous. Evergreen lowland to montane forests, 100–1200 m; Delta Amacuro (Río Toro, Serranía Imataca), northeastern Bolívar (Río Toro), Amazonas (Puerto Ayacucho to Gavilán). Panama, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 535.

626

M IMOSACEAE

Inga salicifolia T.D. Penn., The Genus Inga 188. 1997. Small tree; leaflets 2 or 3 pairs, rachis narrowly winged, foliar nectaries sessile, shallowly cup-shaped, ca. 1 mm diameter; inflorescence a congested raceme, axillary, solitary, peduncle 2–2.2 cm long, floral rachis 4– 5 cm long, pedicels 1.5–2 mm long; fruit unknown. Lower montane forests, 100–600 m; Amazonas (expected). Brazil (known only from the type collection on the lower slopes of Serra da Neblina). Inga sapindoides Willd., Sp. Pl. 4: 1012. 1806. —Mimosa sapindoides (Willd.) Poir. in Lam., Encycl. suppl. 1: 40. 1810. —Feuilleea sapindoides (Willd.) Kuntze, Revis. Gen. Pl. 1: 189. 1891. —Guamo morrocoyero, Guamo rastrojero. Inga lindeniana Benth., London J. Bot. 4: 608. 1845. —Feuilleea lindeniana (Benth.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. Inga caracasana Pittier, Bol. Minist. RR. EE. no. 8–12, reprinted in Arb. Arbust. Venez. 9/10: 107. 1929. Inga grandifolia Pittier, Bol. Minist. RR. EE. no. 8–12, reprinted in Arb. Arbust. Venez. 9/10: 108. 1929. Inga camuriensis Pittier, Bol. Minist. R.R. E.E. no. 8–12, reprinted in Arb. Arbust. Venez. 9/10: 110. 1929. Inga antioquensis Britton & Killip, Ann. New York Acad. Sci. 35: 118. 1936. Tree 10–20 m tall; leaflets (2)3(4) pairs, rachis winged (wings to 15 mm wide), rarely terete, pubescent, foliar nectaries shortly stalked, head flat or cup-shaped, 1–2 mm diameter; inflorescence a congested spike, axillary, solitary or clustered, peduncle 1–5.3 cm long, floral rachis 1.8–5 cm long, bracts 7–16 mm long; fruit 10–24 × 2–4.5 × 1–3 cm, flat to quadrangular, straight or slightly curved, margins winged (to 1 cm wide) and longitudinally ribbed, glabrous. Semideciduous and evergreen lowland forests, especially secondary vegetation, 100–300 m; Delta Amacuro (Río Toro, Serranía de Imataca), Bolívar (near El Palmar). Aragua, Barinas, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, Colombia, Trinidad, Ecuador, Peru. ◆Fig. 537. Inga sertulifera DC., Prodr. 2: 436. 1825.

Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, both in Venezuela, 1 of these in the flora area. The second subspecies, Inga sertulifera subsp. leptopus (Benth.) T.D. Penn., is known from Sucre state in northern Venezuela and differs in its longer fruit, chartaceous leaflets, and narrowly attenuate leaflet bases. I. sertulifera subsp. sertulifera. —Abira (Arekuna), Abira-yek (Arekuna), Guamo. Mimosa coriacea Pers., Syn. Pl. 2: 262. 1806. —Inga coriacea (Pers.) Desv., J. Bot. Agric. 3: 71. 1814, non Willd. 1806. Tree 5–15 m tall; leaflets 2(3) pairs, coriaceous, rachis terete to winged (to 4 mm wide), glabrous, foliar nectaries sessile or shortly stalked, cup-shaped, 0.75–1.5 mm diameter; inflorescence umbellate, axillary, solitary or paired, peduncle 2.5–7.5 cm long, pedicels 7–20 mm long; fruit 4.5–10(–12.5) × 1.8–2.8 × 0.5–1 cm, flat to convex, or subcylindric at maturity, straight or curved, margins not or only slightly raised, faintly transversely striate, glabrous, pulp edible. Lowland to lower montane riparian forests, 200–700 m; Bolívar (Gran Sabana, Kamarata to Urimán), Amazonas (Cañon Grande of Sierra de la Neblina). Guyana, Suriname, French Guiana, Peru, Brazil. Inga splendens Willd., Sp. Pl. 4: 1017. 1806. —Mimosa splendens (Willd.) Poir. in Lam., Encycl. suppl. 1: 43. 1810. —Feuilleea splendens (Willd.) Kuntze, Revis. Gen. Pl. 1: 189. 1891. —Guamo, Guama, Guamo blanco, Guamo cambur, Guamo liso, Guamo pata de morrocoy, Guamo verde. Inga floribunda Benth., J. Bot. (Hooker) 2: 143. 1840. Tree 5–30 m tall; leaflets 2 or 3 pairs, rachis winged (to 10 mm wide), foliar nectaries sessile, disk-shaped, 1.5–2.5 mm wide; inflorescence a congested spike, clustered around the shoot apex or on short leafless shoots, 1– several per axil, peduncle 2–3.1 cm long, puberulous, floral rachis 1–4 cm long, flowers sessile; fruit 15–25 × 2.5–3.7 × 0.8–1.2 cm, flat, straight or curved, margins 4–10 mm wide, not or only slightly raised, glabrous, pulp edible. Evergreen lowland forests, seasonally flooded riparian forests,

Inga 627

river banks, forest-savanna ecotones, 100– 500 m; Delta Amacuro (Río Toro), Bolívar (Río Caura), Amazonas (base of Sierra de la Neblina). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 539.

nally ribbed, velutinous to subglabrous with age. Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 subspecies, 1 in Venezuela.

Inga stenoptera Benth., J. Bot. (Hooker) 2: 143. 1840. Tree 15–20(–30) m tall; leaflets 2 or 3(4) pairs, rachis winged (3–5 mm wide), foliar nectaries sessile, flat to cup-shaped, 1.5–3 mm diameter; inflorescence a congested spike, axillary or clustered near the shoot apex, 1–4 per axil, peduncle 1.5–6 cm long, floral rachis 0.5–2 cm long; fruit 9.5–30 × 2.1–2.5 × 0.6–1 cm, flat to convex, straight, faces slightly swollen over the seeds, with fine transverse venation, margins ca. 5 mm thick, slightly raised, glabrous. Seasonally flooded riparian forests, gallery forests, near sea level to 400 m; Delta Amacuro (Tucupita), Bolívar (mouth of Río Maruanta), Amazonas (Río Siapa). Apure, Barinas, Guárico; Colombia, Ecuador, Peru, Brazil.

I. thibaudiana subsp. thibaudiana. —Cudami (Yekwana), Curanbo-yek (Arekuna), Guama, Guamita, Guamo, Guamo negro. Inga gladiata Desv., Ann. Sci. Nat. (Paris) 9: 427. 1826. Inga tenuiflora Salzm. ex Benth., London J. Bot. 4: 596. 1845. Inga macradenia Mart. ex Benth., Trans. Linn. Soc. London 30: 615. 1875, nom. nud. Tree 6–15 m tall; leaf rachis terete; indumentum of young growth and inflorescence puberulous, lower leaflet surface sericeousstrigose to subglabrous; fruit pulp edible. Secondary forests, forest gaps, 100–1300 m; widespread in Bolívar and Amazonas. Coastal Cordillera, Apure, Táchira; Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 520.

Inga stipularis DC., Prod. 2: 435. 1825. —Guamita. Tree 6–15 m tall; leaflets (2)3(4) pairs, rachis terete, foliar nectaries sessile, cupshaped, 0.75–2 mm wide, stipules foliaceous, suborbicular to ovate, persistent, 1.1–3.3 × 0.7–3.8 cm; inflorescence a congested spike or subcapitate, axillary, 1–4 in each axil, peduncle 2–7 cm long, glabrous, floral rachis 0.4–5 cm long; fruit 10–22 × 1.5–2 × 0.4–0.8 cm, flat, straight, constricted between the seeds at maturity, faces smooth, margins ca. 2 mm thick, slightly raised, glabrous, pulp edible. Evergreen lowland to montane forests, riparian forests and riverbanks, secondary forests, 50–1300 m; Bolívar (Río Orinoco in northwest corner of state), Amazonas (Sierra Parima). Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 541. Inga thibaudiana DC., Prodr. 2: 434. 1825. Tree to 25 m tall; leaflets 4–6 pairs, rachis terete to winged, foliar nectaries sessile, disk-shaped or cup-shaped, 1.5–3.5 mm diameter; inflorescence a congested spike, axillary, 2–5 per axil, peduncle 1–4.5 cm long, floral rachis 1–3.5(–5) cm long; fruit 8–30 × 1.8–2.5(–3) × 0.4–0.9 cm, flat, straight or slightly curved, faces smooth, margins 2.5–5 mm thick, slightly raised, finely longitudi-

Inga ulei Harms., Verh. Bot. Vereins. Prov. Brandenburg 48: 161. 1907. Inga racemiflora Ducke, Arq. Inst. Biol. Veg. 4: 4. 1938. Shrub or tree 3–12 m tall; leaflets 1 pair, petiole narrowly winged (to 2.5 mm wide), foliar nectary sessile, cup-shaped or diskshaped, 1–2 mm diameter; inflorescence racemose, clustered near the shoot apex, peduncle 1.9–5.5 cm long, floral rachis 0.7–2 cm long, pedicels 1–2 mm long; fruit 7–14.3 × 2–3.4 × 0.5 cm, flat, becoming convex and slightly swollen around the seeds, slightly curved, base sometimes asymmetrical, margins raised, 2–3 mm thick, glabrous. Seasonally flooded forests along black-water rivers, 100–200 m; Amazonas (Caño San Miguel, Río Guainía, San Carlos de Río Negro). Colombia, Brazil. ◆Fig. 538. Inga umbellifera (Vahl) Steud. ex DC., Prodr. 2: 432. 1825. —Mimosa umbellifera Vahl, Eclog. Amer. 3: 30. 1807. —Feuilleea umbellifera (Vahl) Kuntze, Revis. Gen. Pl. 1: 189. 1891. —Guamita, Huama, Tumba potro, Yaragua. Inga sciandion Steud., Flora 26: 758.

628

M IMOSACEAE

1843. —Feuilleea sciadon (Steud.) Kuntze, Revis. Gen. Pl. 1: 189. 1891. Inga myriantha Poepp., Nov. Gen. Sp. Pl. 3: 77, t. 289. 1845. —Feuilleea myriantha (Poepp.) Kuntze, Revis. Gen. Pl. 1: 188. 1891. Inga lawrenceana Britton & Killip, Phytologia 1: 23. 1933. Tree or shrub 3–15 m tall; leaflets (1)2(3) pairs, rachis winged (to 6 mm wide), the wings widest near the distal part of the internodes, foliar nectaries sessile, flat or diskshaped, 1.5–2.5 mm diameter; inflorescence umbellate, axillary, solitary or along short axillary shoots to 6 cm long, peduncles 1.2–5 cm long, pedicels 7–20 mm long; fruit 5–15 × 2.1–3 × 1–1.4 cm, flat when immature then becoming convex, straight or curved, margins raised when young, rounded when mature, 5–7 mm thick, glabrous. Evergreen lowland to lower montane forests, riparian and secondary forests, 50–900 m, Delta Amacuro (Río Amacuro near Guyana border), Bolívar (Altiplanicie de Nuria, between El Dorado and La Escalera, Macizo del Chimantá [Apacará-tepui]), Amazonas (Río Cataniapo, lower Río Ventuari, western base of Sierra de la Neblina). Sucre; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 534. Inga umbratica Poepp., Nov. Gen. Sp. Pl. 3: 77. 1845. —Feuilleea umbratica (Poepp.) Kuntze, Revis. Gen. Pl. 1: 189. 1891. Tree 8–20 m tall; leaflets 2 or 3 pairs, rachis winged toward the apex of the leaflet internodes, foliar nectaries sessile, flat or diskshaped, 2–3 mm diameter; inflorescence umbellate, axillary or clustered on woody protuberances below the leaves, peduncle 3– 12 cm long, densely puberulous, pedicels 0.75–1.5 mm long; fruit 8–20 × 2–2.7 × 0.5– 0.8 cm, flat, straight, margins usually narrowly winged (3–4 mm), occasionally unwinged, puberulous on the margins, pulp edible. Evergreen lowland forests, 100–300 m, Bolívar (Santa María de Erebato). French Guiana, Ecuador, Peru, Brazil, Bolivia. Inga vera Willd., Sp. Pl. 4: 1010. 1806. Mimosa inga L., Sp. Pl. 516. 1753, non Vell. 1831. Tree 8–30 m tall; leaflets 4–8 pairs, rachis winged (to 12 mm wide), pubescent to tomen-

tose, foliar nectaries sessile or rarely shortly stalked, flat or shallowly cup-shaped, 0.75– 1.75 mm diameter, aperture usually circular, rarely laterally compressed; inflorescence axillary, a congested or lax spike or rarely a raceme, solitary or paired, peduncle 1–5 cm long, floral rachis 1.5–5 cm long, pedicels 0– 1(–5) mm long; fruit 5–15(–20) × 1.3–2.5 × 1.5–2.5 cm, cylindrical or quadrangular, straight or slightly curved, faces 0.5–1.5 cm wide, partly covered by the strongly expanded margins, margins longitudinally ribbed, pubescent to tomentose. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Uruguay, Argentina; 3 subspecies, 2 in Venezuela, both in the flora area. Key to the Subspecies of I. vera 1. Calyx usually 7.5–9.5 mm long; ratio of corolla length to calyx length 1.8–2.3:1 ...................................... subsp. affinis 1. Calyx mostly 11–14 mm long; ratio of corolla length to calyx length 1.4–1.5:1 .......................................... subsp. vera I. vera subsp. affinis. —Alowata lua (Hohontu), Guama, Guamo chivo, Wajunuma (Yekwana). Evergreen lowland to lower montane forests, riparian and secondary forests, 100–400 m; widespread in lowland areas of Bolívar and Amazonas. Widespread elsewhere in Venezuela; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay, Uruguay, Argentina. ◆Fig. 540. I. vera subsp. vera. —Guamo. Inga spuria Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1011. 1806. —Mimosa spuria (Humb. & Bonpl. ex Willd.) Poir. in Lam., Encycl. suppl. 1: 40. 1810. Inga riparia Pittier, Bol. Minist. RR. EE. no. 8–12, reprinted in Arb. Arbust. Venez. 9/10: 268. 1929. Evergreen lowland and riparian forests, 50–200 m; Delta Amacuro (Serranía de Imataca), northernmost Bolívar. Coastal Cordillera, Apure, Barinas, Zulia; Mexico, Central America, Greater Antilles, Colombia, Ecuador.

Inga 629

Fig. 520. Inga thibaudiana subsp. thibaudiana

Fig. 521. Inga alba

630

M IMOSACEAE

Fig. 522. Inga neblinensis

Fig. 523. Inga bourgonii

Inga 631

Fig. 524. Inga capitata

Fig. 525. Inga micradenia

632

M IMOSACEAE

Fig. 526. Inga edulis

Inga 633

Fig. 527. Inga fastuosa

Fig. 528. Inga laurina

634

M IMOSACEAE

Fig. 529. Inga heterophylla

Fig. 530. Inga inflata

Inga 635

Fig. 531. Inga ingoides

636

M IMOSACEAE

Fig. 532. Inga macrophylla

Inga 637

Fig. 533. Inga nobilis subsp. nobilis

Fig. 534. Inga umbellifera

638

M IMOSACEAE

Fig. 535. Inga rubiginosa

Fig. 536. Inga pilosula

Inga 639

Fig. 537. Inga sapindoides

Fig. 538. Inga ulei

640

M IMOSACEAE

Fig. 539. Inga splendens

Fig. 540. Inga vera subsp. affinis

Leucaena 641

Fig. 541. Inga stipularis

13. LEUCAENA Benth., J. Bot. (Hooker) 4: 416. 1842. by Paul E. Berry Unarmed, small to medium-sized trees 2–20 m tall. Leaves bipinnate, paripinnate, with 1 or rarely 2 sessile or stipitate nectaries at the apex of the petiole, 1–many nectaries at the base of the terminal and subterminal pairs of pinnae; (1)2– many pinnae; leaflets few to numerous, short-petiolate or sessile; stipules ovate, subulate, or long-pointed at the apex, with asymmetric wings at the base, persistent or tardily deciduous. Inflorescence capitate, pedunculate, the peduncle with an involucel of united bracts at the distal end, the capitula in fascicles of 1–several in leaf axils on 0–2-branched shoots. Flowers mostly bisexual, 5-merous, subtended by

642

M IMOSACEAE

peltate bracts. Calyx united for 2/3 its length, the free lobes valvate in bud. Petals 5, 1-veined, usually free. Stamens 10, usually in 2 distinct ranks; anthers ovate to oblong, dorsifixed to nearly basifixed, usually sparsely to densely pilose, but a few species glabrous (not in the flora area). Ovary sessile or subsessile. Fruits short-stipitate, pendulous, linear to oblong, compressed, in most species opening simultaneously along both sutures. Seeds circular, ovate, or rhomboidal in outline, compressed, with a U-shaped pleurogram on both sides. Southern U.S.A. (Texas), Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru, and one species widely cultivated or naturalized throughout the tropics worldwide; 22 species, 2 in Venezuela, 1 of these in the flora area. This treatment is largely based on the monograph of Leucaena by C. Hughes (Syst. Bot. Monogr. 55: 1–244. 1998). Leucaena trichodes (Jacq.) Benth. is a common, native shrub in semideciduous regions of coastal Venezuela. Leucaena leucocephala (Lam.) de Wit, Taxon 10: 53: 1961. —Mimosa leucocephala Lam., Encycl. 1: 12. 1783. —Acacia leucocephala (Lam.) Link, Enum. Hort. Berol. Alt. 2: 444. 1822. Small to medium-sized tree 3–15 m tall; leaves 10–25 × 5–16 cm, pinnae 4–9 pairs, leaflets 13–21 pairs per pinna, each 9–16 × 2–5 mm, nearly sessile and strongly asymmetric; petioles 13–24 mm long; capitula 12– 21 mm diameter, in fascicles of (1)2–6 in leaf

axils, peduncles 15–20 mm long; fruiting pods on stipes 4–14 mm long; seeds 8–18 per pod, ovoid. Likely native to southern Mexico or northern Central America, now widely naturalized and domesticated in tropical and subtropical regions of the world; 3 subspecies, 1 in Venezuela. L. leucocephala subsp. glabrata (Rose) S. Zárate, Phytologia 63: 305. 1987. —Leucaena glabrata Rose, Contr. U.S. Natl.

Fig. 542. Leucaena leucocephala subsp. glabrata

Macrosamanea 643

Herb. 5: 140. 1897. Semideciduous forests, disturbed areas, 50–100 m; Bolívar (Ciudad Bolívar, Puerto Ordaz). Widely scattered in northern Ven-

ezuela; probably native to Mexico, widely cultivated and naturalized in Mexico and Central America, as well as tropical areas worldwide. ◆Fig. 542.

14. MACROSAMANEA Britton & Rose, Ann. New York Acad. Sci. 35: 131. 1936. Pithecellobium sect. Samanea ser. Coriaceae Benth., Trans. Linn. Soc. London 30: 589. 1875. Tribe Ingeae genus D I.C. Nielsen in Polhill & P.H. Raven, Adv. Leg. Syst. 1: 190. 1981. by Rupert C. Barneby and James W. Grimes Trees of moderate size and shrubs, mostly of riparian woodland and moist savanna, the branches sometimes sarmentose or vining. Leaves bipinnate; stipules caducous or obsolete; pinnae 1–10 pairs; leaflets 2–20 pairs, lanceolate or rhombic-oblong, sometimes falcate, the leaf stalks charged at base and between pulvinules of some pinna-pairs with depressed nectaries. Flowers sessile or rarely short-pedicellate, disposed in capitula or dense short spikes either on axillary peduncles or sometimes shortly corymbose-paniculate, the floral bracts all or at least the lowest ventrally with a concave nectary. Calyx tubular; corolla narrowly tubular-funnelform, densely puberulent externally. Stamens numerous; filaments white, united into a tube as long as or somewhat longer than the corolla. Pods few, subsessile, broadly linear or oblong in profile, inertly dehiscent through the seminiferous suture but scarcely opening. Seeds compressed but turgid, with dry fragile testa and no pleurogram, the cotyledons hard. Tropical South America (most endemic to basins of Río Orinoco and Río Amazon); ca. 15 species, 6 in Venezuela, all in the flora area. Key to the Species of Macrosamanea 1. 1. 2(1). 2. 3(1). 3. 4(3).

4.

5(3).

5.

Pinnae either 1 or alternatively 1 and 2 pairs per leaf ............................ 2 Pinnae of each leaf 3–10 pairs ................................................................... 3 Pinnae 2 pairs, 1 pair in some leaves; leaflets (12–)14–24 pairs and the larger ones 15–32 × 3.5–7 cm; calyx 8–16 mm long ................. M. discolor Pinnae 1 pair in all leaves; leaflets 2–4 pairs and the larger ones 30–65 × (6–)8–35 mm; calyx 6–7 mm long ........................................ M. simabifolia Leaflets coriaceous, obtuse at apex ........................................................... 4 Leaflets chartaceous, acuminate and falcate at apex ............................... 5 Primary leaf stalk 4.5–18 cm; larger leaflets 17–25 × 6–10 mm; pedicel absent; calyx 6.5–12 mm; corolla not over 2.5 cm; androecium 4–6 cm long; pods 2.5–3.5 cm wide ................................................... M. pubiramea Primary leaf stalk 20–50 cm; larger leaflets to 45–60 × 18–27 mm; pedicel 4–10 mm; calyx 12–19 mm long; corolla 3.5–5 cm; androecium 7–8.5 cm long; pods 6–7 cm wide ........................................................ M. amplissima Primary leaf stalk 10–14 cm long; pinnae about 4 pairs and their leaflets to 10–14 pairs; calyx 10–11 mm long; expanded corolla unknown .......................................................................................... M. consanguinea Primary leaf stalk 13–29 cm long; pinnae to 7–10 pairs and their leaflets to 15–20 pairs; calyx 15–25 mm long; corolla 4–5 cm long ....... M. spruceana

644

M IMOSACEAE

Macrosamanea amplissima (Ducke) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 198. 1996. —Pithecellobium amplissimum Ducke, Arq. Inst. Biol. Veg. 4: 6. 1938. Tree ± 5 m tall, with scandent branches. Seasonally flooded riparian forests, 100–300 m; Amazonas (Caño Curumoni, Caño Morocoto, black-water tributaries of Río Casiquiare, Río Guainía, San Carlos de Río Negro). Colombia (Río Vaupés), Brazil (Amazonas: Rio Negro). Macrosamanea consanguinea (R.S. Cowan) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 189. 1996. —Pithecellobium consanguineum R.S. Cowan, Mem. New York Bot. Gard. 10(4): 68. 1961. Tree 4–6 m tall, sometimes semiscandent. Seasonally flooded riparian forests along black-water rivers, 100–200 m; Amazonas (Limoncito on the left bank of Caño San Miguel). Adjacent Colombia and Brazil. Macrosamanea discolor (Humb. & Bonpl. ex Willd.) Britton & Rose, Ann. New York Acad. Sci. 35: 131. 1936. —Inga discolor Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1023. 1806. —Pithecellobium discolor (Humb. & Bonpl. ex Willd.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3): 63. 1943. Inga adiantifolia Kunth, Mimoses 66, t. 21. 1819 [1820]. —Pithecellobium adiantifolium (Kunth) Benth., London J. Bot. 3: 218. 1844. —Samanea adiantifolia (Kunth) Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 55. 1926. Macrosamanea aquatica Pittier, Bol. Soc. Venez. Ci. Nat. 11: 15. 1947. —Pithecellobium aquaticum (Pittier) R.S. Cowan, Mem. New York Bot. Gard. 10(4): 67. 1961. Pithecellobium subaquaticum Schery, Fieldiana, Bot. 28: 259. 1952. Pithecellobium arenicola R.S. Cowan, Mem. New York Bot. Gard. 10(4): 67. 1961. Shrub or treelet 1–5 m tall. Bush islands on white-sand savannas, Mauritia palm swamps, and regularly flooded riparian or gallery forests, 100–200 m; Amazonas (Caño Ucata, basin of Río Casiquiare and Río Gui-

anía, upper Río Orinoco). Brazil (along the Rio Negro to its confluence with the Amazon). Macrosamanea pubiramea (Steud.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 194. 1996. —Inga pubiramea Steud., Flora 26: 759. 1843. Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 varieties, 1 in Venezuela. M. pubiramea var. lindsaeifolia (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 197. 1996. —Pithecellobium lindseifolium Spruce ex Benth., Trans. Linn. Soc. London 30: 590. 1875. Weak tree or vine 2–7 m tall. Riparian forests, 100–200 m; Amazonas (base of Cerro Sipapo, Río Cataniapo, Santa Lucía on Río Negro). Apure; Brazil (scattered in central Amazon basin). Macrosamanea simabifolia (Spruce ex Benth.) Pittier, 3rd Conf. Interamer. Agric. Caracas 358. 1945. —Pithecellobium simabifolium Spruce ex Benth., Trans. Linn. Soc. London 30: 589. 1875. —Samanea simabifolia (Spruce ex Benth.) Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 55. 1925. Shrubby treelet 2–4(–8) m; leaflet width and pod size polymorphic. Seasonally flooded savannas, shrublands on sandy soil or open shrubby patches on granitic outcrops, 100– 200 m; Bolívar (mouth of Río Caroní), Amazonas (Río Baría, frequent on upper Río Guainía and Río Orinoco). Amazonian Colombia and Brazil. ◆Fig. 543. The record from Bolívar is a single collection of a somewhat aberrant, perhaps varietally distinct form distinguished by subobsolete leaf stalks. Macrosamanea spruceana (Benth.) Killip, Trop. Woods 63: 6. 1940. —Pithecellobium spruceanum Benth., Trans. Linn. Soc. London 30: 590. 1875. Tree, sometimes sarmentose, 2–16 m tall. Riparian forests, 50–100 m; Delta Amacuro (along Caño Aragua between mouth of Caño Jajene and Isla Mono Burojo). Scattered through upper Amazonian Colombia, Peru, and Brazil.

Mimosa 645

Fig. 543. Macrosamanea simabifolia

15. MIMOSA L., Sp. Pl. 516. 1753; Gen. Pl. ed. 5: 233. 1754. Schrankia Willd., Sp. Pl. 4: 888, 1041. 1806. Morongia Britton, Mem. Torrey Bot. Club 5: 191. 1894. by Rupert C. Barneby Trees, shrubs, vines, subshrubs, and herbs, a few of the latter monocarpic, either armed or not with epidermal prickles, most but not all lacking petiolar nectaries. Indumentum composed in various measure and random combination of: (a) minute puberulence, (b) short plain hairs, (c) short-clavate or granular, discolored but nonsecretory trichomes, (d) filiform setulae or tapering, basally enlarged and at times laterally attached (seemingly spurred) setae, these sometimes backwardly produced into the epidermis, (e) scabrous or plumulose setae that become stelliform by shortening of the primary axis, (f) gland-tipped setae or sessile glands, and (g) in one group with petiolar nectaries peltate-squamiform trichomes (exceptionally quite glabrous). Leaves often sensitive to touch, bipinnate; pinnae 1–20; leaflets 1–many pairs, all opposite in ours, generally ample when few and smaller with increase in number. Inflorescence composed either of spikes (subracemose) or capitula either pedunculate in leaf axils, terminally paniculate-pseudoracemose, or fasciculate on brachyblasts, no involucre on peduncle; flowers 3–5(6)-merous, either all haplo- or all diplostemonous and bisexual, or some proximal ones staminate; calyx either campanulate and short-toothed or truncate, or lobed (the lobes sometimes

646

M IMOSACEAE

paleaceous-pappiform), or subobsolete; corolla sympetalous. Filaments free or shortly connate into a tube or stemonozone, commonly pink or white, in one group yellow; anthers ovate or orbicular, the dilated connective not crowned with a caducous gland. Pod a craspedium, the ripe valves separating from the persistent sutures (replum) and usually breaking transversely into 2–20 1-seeded articles, rarely falling in one piece. Seeds plumply ovoid or discoid, with pleurogram and endosperm. Neotropics, some temperate American, Paleotropics (with endemic radiation in Madagascar); ca. 475 species, 39 in Venezuela, 30 of these in the flora area. The genus Mimosa contains all isostemonous Mimosaceae of the flora area that lack petiolar nectaries with the exceptions of Anadenanthera peregrina, of which the peduncles bear a small involucre below the capitulum, and the aquatic Neptunia oleracea. Our few mimosas with petiolar nectaries differ from Piptadenia either in capitulate (not spicate) units of inflorescence, or if these are spicate in haplostemonous flowers; and further from all other Mimosaceae in peltate trichomes of the hypophyllum and from all isostemonous Mimosaceae in the dilated anther-connective that lacks a terminal gland. The craspedial dehiscence of the fruit occurs in the flora area in no other genus except Entada, which is easily known by a charateristic 1-sided compound inflorescence, the presence of anther-glands, and the immense size of the craspedia. Key to the Species of Mimosa 1.

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

5(1). 5. 6(5). 6. 7(6). 7. 8(5).

Leaf petioles with a nectary, this either cupular-patelliform and widely openly recessed, or mounded (verruciform) and small-pored; hypophyllum of leaflets and valves of pod charged with pallid or reddish scales or glanular trichomes .................................................................. 2 Leaf petioles without nectary; no scale-like trichomes ............................. 5 Flowers arranged in a spike; stamens 5 .................................. M. myriadenia Flowers arranged in a capitellate head; stamens 10 ................................ 3 Leaflets of all pinnae exactly 1 pair, ample, 4.5–12 × 3–7.5 cm .............................................................................................. M. colombiana Leaflets of distal pinnae 3 or more pairs, variable in size but mostly smaller than in the preceding ............................................................... 4 Petiolar nectary cupular; pod 7–8 mm wide; leaflets of distal pinnae 3 or 4 pairs .................................................................................... M. guanchezii Petiolar nectary mounded, small-pored, the orifice much narrower than the verruciform body; pod 17–33 mm wide; leaflets variable, but seldom < 4–9 pairs in distal pinnae ............................................. M. rufescens Flowers spicate, the spike (excluding filaments) much > 2 times as long as diameter ................................................................................................. 6 Flowers capitellate, the capitula (excluding filaments) globose or plumply ovoid-ellipsoid, < 2 times as long as diameter ...................................... 8 Stamens as many as corolla lobes; craspedium 1–2 cm wide .......... M. invisa Stamens 2 times as many as corolla lobes; craspedium 4–9 mm wide .... 7 Corolla glabrous or thinly puberulent with simple hairs; leaflets eglandular .................................................................................. M. arenosa Corolla externally densely stellate-pubescent; leaflets glandular on lower surface ............................................................................... M. schomburgkii Stamens 6, 8, or 10, 2 times as many as corolla lobes .............................. 9

Mimosa 647

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

12.

13(12). 13. 14(13). 14. 15(14). 15. 16(15). 16. 17(14). 17.

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

19. 20(8). 20. 21(20).

Stamens 4 or 5, as many as corolla lobes ................................................ 20 Stems with gland-tipped setae (in addition to other types of trichomes) .............................................................................................................. 10 Stems variably pubescent but eglandular ............................................... 11 Lobes of corolla striately 5–several-veined ................................ M. somnians Lobes or corolla weakly 1-veined .............................................. M. orthocarpa Flowers 3-merous; stamens 6 ................................................... M. piscatorum Flowers 4- or 5-merous; stamens 8 or 10 ................................................ 12 Stipules linear-setiform, 1-veined; craspedium linear, obtusely quadrangular, the replum about as broad as the valves and these separating from replum in 1 piece; corolla 5-merous .......................... M. quadrivalvis Stipules triangular or ovate, 3–many-veined (the veins sometimes not apparent dorsally); craspedium linear, planocompressed, the valves much broader than the replum, breaking up into 1-seeded articles; corolla nearly always 4-merous ....................................................................... 13 Setae of stems plumose; corolla lobes 7–11-veined ............... M. surumuënsis Setae of stems either smooth or minutely scaberulous; corolla lobes 1– 3-veined ................................................................................................ 14 Setae of stem attached laterally above base, in profile appearing basally spurred ................................................................................................. 15 Setae of stem either symmetrically or obliquely basifixed, not produced backward from point of attachment .................................................... 17 Pinnae of larger leaves 2–4 pairs; local in western Bolívar state ...................................................................................... M. brachycarpoides Pinnae of larger leaves (7–)9–30 pairs; widely dispersed in the flora area .............................................................................................................. 16 Capitula without filaments 2.5–4.5 mm diameter; craspedia 3–5 mm wide ........................................................................................... M. microcephala Capitula without filaments 6–7 mm diameter; craspedia 13–16 mm wide ............................................................................................. M. orinocoënsis Shrubs (1–)1.5–5 m, sometimes sarmentose; craspedia 8–15 mm wide, at least 12-seeded; pinnae of larger leaves mostly 8–14 pairs ............... 18 Monocarpic herbs, sometimes weakly suffrutescent in age but not over 1.2 m, usually much less; craspedium 3–8 mm wide, 1–5-seeded; pinnae mostly 2–7 pairs ............................................................................ 19 Stipules firm, densely appressed-setose dorsally and imperceptibly venose externally ......................................................................... M. pellita Stipules chartaceous, glabrous dorsally and striately 10–20-veined ........................................................................................................ M. tarda Interpinnal segments of leaf rachis with a pair of stiff straight horizontal aculei; craspedium in profile obliquely obovate, 7–11 × 6–8 mm .................................................................................................. M. dormiens Leaf rachis unarmed; craspedium in profile elliptic-oblong, 7–15 × 3–4 mm ............................................................................................... M. camporum Lower surface of leaflets densely gray-pubescent with stellate trichomes; cultivated and potentially naturalized tree ............................ M. scabrella Lower surface of leaflets pubescent with simple hairs or setae, or glabrous; native shrubs, subshrubs and herbs ........................................ 21 Pinnae at least 4 pairs per leaf and well spaced along rachis; leaflet mar-

648

M IMOSACEAE

21.

22(21).

22. 23(22).

23. 24(23).

24.

25(22). 25. 26(25). 26. 27(25). 27. 28(27). 28. 29(28). 29.

30(28). 30.

gins lacking basally dilated or setiform cilia; craspedium 7–14 × 1–2 cm, 10–18-seeded ................................................................................. M. invisa Pinnae often 1 pair, if 2 or 3(4) pairs then crowded near tip of leaf stalk, appearing subpalmate; leaflets ciliate with basally dilated trichomes; craspedium much shorter and narrower, mostly 2–5-seeded ............ 22 Leaflets of single pair of pinnae exactly 2 pairs, the distal pair much the largest, the anterior leaflet of the proximal pair much smaller than its fellow or obsolescent, the whole leaf therefore 6(8)-foliolate ............. 23 Leaflets of distal (or only) pair of pinnae > 2 pairs ................................. 25 Stems and leaf stalks weakly armed, the internodes ordinarily with only 1– 3 or randomly a few more aculei, the leaf stalks often unarmed; distal leaflets obovate, broadest above mid-blade; calyx minute ......... M. debilis Stems and leaf stalks serially armed on ribs with rows of scattered aculei; distal leaflets elliptic, mostly broadest near or below middle ............ 24 Calyx pappiform, 0.7–1.6 mm long, irregulary cleft nearly to base into pale straw-colored setae; capitula prior to anthesis concealed by enmeshed bracteal setae; craspedium 4–7 mm wide .................. M. sensitiva Calyx minute, 0.25–0.4 mm long, the rim glabrous or minutely ciliolate; capitula prior to anthesis not concealed by enmeshed bracteal setae; craspedium 8–13 mm wide .................................................... M. velloziana Stems and leaf stalks serially armed on ribs with rows of scattered recurved aculei; pinnae exactly 1 pair .................................................... 26 Stems armed at or below each node with 2 or 3 aculei, or unarmed, and leaf stalks unarmed or with 1 or 2 random aculei .............................. 27 Leaflets of each pinna 8–16 pairs and the longest ones ca. 1–2.5 cm long .......................................................................................... M. schrankioides Leaflets of each pinna 3–6 pairs and the longest ones 2–3.5 cm long ........................................................................................................ M. casta Pinnae of all leaves exactly 1 pair; leaflets 1-veined ........... M. xanthocentra Pinnae 2–4 pairs, at least in leaves subtending peduncles; leaflets 4- or 5-veined from pulvinule ....................................................................... 28 Calyx lobes well-developed, paleaceous or pappiform, pallid, variably decompound into setiform divisions ........................................................ 29 Calyx minute or vestigial ......................................................................... 30 Stems unarmed; calyx pappiform, the lobes divided to base; pinnae subpalmate ......................................................................................... M. huberi Stems almost always armed at or immediately below nodes with 1–3 weak aculei; calyx paleaceous, the divisions of the lobes coalescent at base; pinnae separated along rachis by short spaces ................ M. hirsutissima Stipules 1-veined; pinnae 3–5 pairs and leaflets of longer pinnae ca. 30– 50 pairs ................................................................................. M. polydactyla Stipules 5–13-veined; pinnae (1)2 or 3 pairs and leaflets of longer pinnae 11–30(–35) pairs ........................................................................... M. pudica

Mimosa arenosa (Willd.) Poir. in Lam., Encycl. suppl. 1: 66. 1810. —Acacia arenosa Willd., Sp. Pl. 4: 1060. 1806. Mimosa xantholasia Benth., London J. Bot. 5: 88. 1846. Mimosa fasciculata var. ernestiana

Kuntze, Revis. Gen. Pl. 1: 198. 1891. Southern Mexico, Central America, Lesser Antilles, Colombia, Venezuela, Brazil; 2 varieties, both in Venezuela, 1 of these in the flora area. The other variety, Mimosa arenosa var.

Mimosa 649

leiocarpa (DC.) Barneby, is found erratically from southern Mexico to northern Colombia and northwestern Venezuela. M. arenosa var. arenosa Arborescent shrub to 6 m tall, often randomly armed on internodes but some flowering branches unarmed; pinnae of larger leaves 4–10; leaflets of longer pinnae 3–6.5 mm, in 12–30 pairs; inflorescence spicate; flowers white; craspedium stipitate, the body ca. 3–6 × 0.4–0.6 cm, 7–10-seeded. Brush woodlands, seasonally dry communities, sometimes aggressive in disturbed places, 50–500 m; Bolívar (Altiplanicie de Nuria). North-central Venezuela; interruptedly widespread north to Martinique and south to southeastern Brazil. Mimosa brachycarpoides Barneby, Mem. New York Bot. Gard. 65: 471. 1991. Stiffly branched shrub ca. 1.5 m tall, the stems densely strigose with basally spurred setae; stipules 3–7-veined; pinnae 2–4 pairs; leaflets 5–7 mm, to 26–36 pairs. Granitic outcrops (inselbergs), ca. 100 m; northwestern Bolívar (ca. 30 km downstream from Puerto Ayacucho). Endemic. Mimosa brachycarpoides is related to and resembles M. microcephala, but differs in fewer pinnae and in a campanulate calyx nearly 1 mm long. Mimosa camporum Benth., J. Bot. (Hooker) 2: 130. 1840. Mimosa flavescens Splitg., Tijdschr. Natuurl. Gesch. Physiol. 9: 110. 1842. Mimosa pusilla Benth., Bot. Voy. Sulphur 90. 1844. Mimosa flaviseta Benth., London J. Bot. 5: 90. 1846. Mimosa martensis Britton & Killip, Ann. New York Acad. Sci. 35: 152. 1936. Monocarpic herb, very variable in stature (2–120 cm tall), the stems hispid; stipules striately 12–25-veined; pinnae mostly 2–6 pairs; leaflets (5–)12–35 pairs. Open and disturbed, usually moist places, on savannas, along roadside ditches (depauperate in dry soils), 50–500 m; Delta Amacuro, northern Bolívar (Río Orinoco), Amazonas (upper Río Negro). Scattered over much of Venezuela; widely distributed from southern Mexico and Lesser Antilles to Amazonian Brazil and Bolivia.

Mimosa casta L., Sp. Pl. 518. 1753. —Paranovillo. Weakly suffrutescent vine 2–4 m tall, prickly throughout, with mostly axillary capitula. Pinnae 1 pair; leaflets 4 or 5(6) pairs, lance-oblong-elliptic, 2–3.5 cm. Craspedium oblong, plano-compressed, 2–4 × 1–1.4 cm, 3–5(6)-seeded, the replum stiffly setose. Scrambling in thickets and hedges and forming diffuse tangles in open places in lowland scrub-woodland, sometimes in wet pastures, 50–200 m; scattered in Delta Amacuro. Infrequent in northeastern Venezuela; seemingly native to Windward Islands, Trinidad-Tobago, and lower Río Orinoco and Rio Amazonas, but found in recent times on Puerto Rico and Jamaica, and at isolated stations along the Panama Canal, in central Colombia, and northwestern Brazil. Mimosa colombiana Britton & Killip, Ann. New York Acad. Sci. 35(3): 153. 1936. —Mimosa bauhiniaefolia H. Karst., Fl. Columb. 2: 65, pl. 133. 1862 [1863], non Salisb. 1796. —Zarzaparilla (Meta). Potentially high-climbing vine forming tangled thickets in forest openings and in hedges; nectary near base of petioles patelliform; pinnae 2 pairs and leaflets of each pinna 1 pair, the blades 4.5–12 × 3–7.5 cm; capitula small, white, paniculate, fragrant; craspedium 7–13 × 1.5–2.7 cm. Gallery forests, riparian woodlands, disturbed places, 50–200 m; northwestern Bolívar (Río Parguaza), northern Amazonas (Isla Ratón, near Puerto Ayacucho, between road Puerto Ayacucho-Sanariapo, Río Cataniapo) (confluence of Río Meta and Río Orinoco). Apure; scattered in northeastern Colombia. Mimosa debilis Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1029. 1806. Mimosa adhaerens H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 249. 1823 [1824]. Mimosa notata Steud., Flora 26: 758. 1843. Mimosa hostmannii Benth., London J. Bot. 5: 84. 1846. Mimosa debilis var. (?)panamensis Benth., Trans. Linn. Soc. London 30: 391. 1875. —Mimosa panamensis (Benth.) Standl., Contr. U.S. Natl. Herb. 18: 104. 1916. Costa Rica, Panama, tropical South America (most diverse in Brazil south of the Amazon basin); 5 varieties, 1 in Venezuela.

650

M IMOSACEAE

M. debilis var. debilis Weakly suffrutescent herb, prematurely flowering and then appearing monocarpic, weakly prickly; pinnae 1 pair and leaflets of each leaf 8, the distal pairs obovate or broadly oblanceolate 2–6.5 cm; craspedia radiating from receptacle in a usually dense cluster, 10–18 × 3–5 mm, the replum coarsely hispid, the valves varying from glabrous to setulose. Lowland savannas, weedy pastures, disturbed woodland, sometimes in fencerows, 100–500 m; northern Bolívar. Panama to Paraguay and Argentina. Mimosa dormiens Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1035. 1806. Mimosa humilis Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1035. 1806. Mimosa intermedia Kunth, Mimoses 16, t. 6. 1819. Monocarpic, in age sometimes basally indurated herb 15–60 cm tall, armed on internodes and on interpinnal segments of leaf stalks with straight horizontal aculei 3–6 mm, and with a similar ascending spicule arising from between each pair of pinnae; stipules ovate, 5–9-veined; pinnae 4–7 and leaflets 10–17 pairs, the latter to 4–8.5 mm long; craspedium in profile asymmetrically obovate, mostly 2-seeded, the distal segment of valves broader. Muddy shores, wet savannas, openings of flooded riparian forests, ca. 100–200 m; Delta Amacuro (near Tucupita), northwestern Bolívar. Venezuelan llanos; discontinuously distributed from southern Mexico to northern Colombia and central and eastern Amazonian Brazil. Mimosa guanchezii Barneby, Mem. New York Bot. Gard. 65: 39. 1991. Vine resembling M. colombiana in habit, armature and inflorescence, but pinnae mostly 3 or 4, and leaflets 3 or 4 pairs, the larger distal leaflets 16–28 × 11–18 mm; craspedium ca. 45–60 × 7–8 mm. Edges of forests, 50–100 m; Amazonas (near Puerto Ayacucho). Endemic. Mimosa hirsutissima Mart., Flora 21(2) Beibl.: 55. 1838. Mimosa tomentosa Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1033. 1806, non Rottling 1803. Functionally herbaceous from woody roots, the diffuse stems and foliage densely

loosely pubescent, the stems weakly armed at most nodes with 1–3 small aculei; pinnae in leaves subtending peduncles 2 or 3 pairs, several-veined on lower surface; capitula excluding filaments 6–8 mm diameter; calyx lustrously paleaceous, at least 1 mm long; craspedium ca. 10–25 × 4–6 mm, 2–4-seeded. Savannas, Mauritia palm swamps, 50–200 m; northwestern Bolívar (Caicara del Orinoco, Maripa), Amazonas (Río Orinoco and immediate affluents from the Río Manapiare basin). Adjacent Venezuela; northeastern Colombia, northeastern Brazil, common and polymorphic on the Brazilian Planalto and in adjoining Paraguay and Argentina. ◆Fig. 545. Mimosa huberi Barneby, Mem. New York Bot. Gard. 65: 632. 1991. Unarmed suffrutescent herb or clambering subshrub 0.5–2 m tall, the stems hirsute with scaberulous setae, the foliage densely subappressed-silky; pinnae of larger leaves 2–4 pairs, subpalmate near apex of leaf stalk, the leaflets of distal pair ca. 20–30 pairs and to 1–2 cm long; capitula ovoid-ellipsoid, excluding filaments 4.5–5 diameter; calyx pappiform, the tube ca. 0.2 mm, fringed with fine setulae 0.6–1 mm; craspedium ca. 15–20 × 4 mm, 3–5-seeded. Bush islands in savannas, exposed lajas, 50–200 m; along and close to the Río Orinoco in Bolívar (261 km SW of Caicara del Orinoco) and Amazonas (near Puerto Ayacucho). Endemic. Mimosa huberi is to be expected across the river in Colombia. Mimosa invisa Mart. ex Colla, Herb. Pedem. 2: 255. 1834, non Mart. 1837. Interruptedly widespread from Colombia and Venezuela south to southern Brazil and Paraguay; 2 subspecies and 4 varieties, 1 subspecies and 2 varieties in Venezuela, 1 variety in the flora area. M. invisa subsp. spiciflora var. spiciflora (H. Karst.) Barneby, Mem. New York Bot. Gard. 65: 307. 1991. —Mimosa spiciflora H. Karst., Fl. Columb. 2: 61, t. 131. 1862 [1863]. Mimosa brevispica Harms, Notizbl. Königl. Bot. Gart. Berlin 6: 303. 1915. Weakly fruticose and to 2–5 m tall, the stems and leaf stalks serially armed with recurved prickles; pinnae of larger leaves 6–16

Mimosa 651

and leaflets ca. 25–35 pairs, the latter not over 8 × 2 mm; flowers loosely capitate or shortly spicate, 5-merous, 5-androus; craspedium broad-linear planocompressed ca. 1– 2 cm wide, 10–18-seeded. Climbing in thickets and trailing in open rocky places, 100– 1400 m; Bolívar (expected in the vicinity of Urimán). Northern Venezuela; northern Colombia, Guyana, Brazil (Roraima: Serra Pacaraima). The combination of haplostemonous flowers, lack of basally dilated leaflet-cilia, and broad multiovulate pod is unique in the flora area. Mimosa microcephala Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1039. 1806. Shrub or subshrub 1–5 m tall, either armed or not and either erect or sarmentose over and through shrubs and smaller trees; stems pubescent with basally spurred setae; inflorescence of small capitula either pseudoracemose or paniculate; pinnae of larger leaves (7–)10–30 and leaflets to 20–60 pairs; craspedium linear, 3–5 mm wide, 3–10(–16)seeded. Savannas, outcrops, rocky river banks, 200–1200 m; Bolívar (Caicara del Orinoco, Río Guayaraca), Amazonas (Puerto Ayacucho, Río Cataniapo). Interruptedly dispersed in Guyana, Suriname, French Guiana, southeastern Colombia, southern Venezuela, and Amazonian Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of M. microcephala 1. Plants armed; corolla lobes 1-veined; filaments white ........... subsp. cataractae 1. Plants unarmed; corolla lobes 1- or 3veined; filaments either pink or white ........................... subsp. microcephala M.

microcephala subsp. cataractae (Ducke) Barneby, Mem. New York Bot. Gard. 65: 468. 1991. —Mimosa cataractae Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 75. 1922. Colombia, Venezuela, Brazil; 2 varieties, 1 in Venezuela. M. microcephala subsp. cataractae var. lumaria Barneby, Mem. New York Bot. Gard. 65: 469. 1991. Rocky ridges and outcrops, 200–500 m; Bolívar (Altiplanicie de Nuria, Río Caroní

and Río Orinoco), Amazonas (Río Sipapo). Colombia (Guainía). M. microcephala subsp. microcephala Colombia, Venezuela; 3 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of M. microcephala subsp. microcephala 1. Corolla lobes 1-veined ...... var. communis 1. Corolla lobes 3-veined .. var. microcephala M. microcephala subsp. microcephala var. communis Barneby, Mem. New York Bot. Gard. 65: 469. 1991. Savannas, outcrops, rocky streambanks, lajas, 100–1200 m; western and northeastern Bolívar (southwest of Caicara, Río Guayaraca), northwestern Amazonas (Río Guianía and upper Río Orinoco). Adjacent Colombia. M. microcephala subsp. microcephala var. microcephala Mimosa microcephala Humb. & Bonpl. ex Willd., s. str. Savanna thickets, laja outcrops, 50–200 m; locally plentiful in northwestern Amazonas (Río Orinoco between Sanariapo and Cerro San Borja along lower Río Suapure). Apure; Colombia (Vichada). Mimosa myriadenia (Benth.) Benth., Trans. Linn. Soc. London 30: 408. 1875. —Entada myriadenia Benth., J. Bot. (Hooker) 2: 133. 1840. —Raspa. Acacia paniculiflora Steud., Flora 26: 760. 1843. Scrambling shrub or potentially highclimbing, prickly vine with a cupular nectary on leaf stalks and peltate trichomes on hypophyllum of leaflets and on valves of pod; inflorescence paniculate, of spike-like racemes; flowers white, haplostemonous, fragrant. Forests, gallery margins, 50–500 m; common in southern Delta Amacuro and western half of Bolívar, Amazonas (upper Río Orinoco). Widespread in Venezuela; Costa Rica, Pacific Colombia, Guyana, Suriname, French Guiana, Amazon basin. Mimosa myriadenia is highly variable in number of pinnae and in number and size of leaflets. In the flora area, the pinnae are 9– 18 pairs and the leaflets of longer ones are

652

M IMOSACEAE

16–42 pairs and to 3.5–9.5 mm long. The more common form has relatively large leaflets (to 5–9.5 × 1.5–3 mm) and a less common form has smaller, more crowded leaflets (to 3–6.5 × 0.6–1.4 mm). Mimosa orinocoënsis Barneby, Mem. New York Bot. Gard. 65: 436. 1991. Riparian shrub, closely resembling the common M. pellita, but different in glabrate venulose stipules, basally spurred cauline setae, and minute calyx; craspedium 7–10 × 1.3–1.7 cm, 12–18-seeded, the valves hispid with stout, basally dilated, vertically erect setae, breaking up into water-borne articles 4.5–8 mm long, each sealed at both ends by a septum 0.6–1 mm wide. River banks, near sea level to 200 m; Delta Amacuro (lower delta region), northwestern Amazonas. Northeastern Colombia, Brazil (Pará). Mimosa orthocarpa Spruce ex Benth., Trans. Linn. Soc. London 30: 437. 1875. Herbaceous or weakly suffruticose, to 1.5 m tall, resembling M. somnians in armament and gland-tipped setae, but corolla lobes 1-veined; pinnae 4–9 and leaflets ca. 15–22 pairs, none over 7 mm long; craspedium stipitate, the body 16–32 × 4–5 mm, viscid-setulose. River banks, seasonally wet savannas, forest margins, 50–300 m; Bolívar (Río Orinoco southwest of Ciudad Bolívar), Amazonas (Río Orinoco northeast of Isla Ratón). Venezuelan llanos; discontinuously dispersed in southern Mexico, Central America, Lesser Antilles, Colombia, and northern and Amazonian Brazil. ◆Fig. 547. The closely related Mimosa corynadenia Britton & Rose differs in fewer (3–5) pairs of pinnae and pod 8–11 mm wide. It is known from flooded lowlands in the middle Río Orinoco basin, but so far only from well north of the river; it may be looked for in the northwestern corner of the flora area. Mimosa pellita Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1037. 1805. —Arana de gato negro. Mimosa pigra auct. non L. 1755. Mimosa asperata auct. non L. 1759. Tropical lowland Mexico and Cuba south to warm temperate Argentina, Africa, southeastern Asia; 3 varieties, 2 in Venezuela, 1 of these in the flora area.

M. pellita var. pellita Potentially arborescent shrub 1–5 m tall, either free-standing, or when crowded either forming thickets or sarmentose, armed on stems and leaf stalks with either straight or recurved aculei and between each pair of pinnae with an ascending spicule 1.5–15 mm long, the peduncles axillary or in early anthesis pseudo-racemose; stipules firm, dorsally strigose-silky; pinnae of larger leaves 7–14 and leaflets of longer pinnae 25–45 pairs, the largest leaflets 5–12 mm long; craspedium 4–12 × 0.8–15 cm, mostly 14–20seeded, the valves densely hispid, breaking up into articles sealed at each end by a septum. Stream banks and shores, along ditches, seasonally flooded savannas, edges of gallery forests, low elevations, near sea level to 300 m; Delta Amacuro (mouth of Caño Güiniquina), Bolívar (Río Caura 5–20 km south of Salto Pará, Río Cuchivero, San Félix), northern Amazonas. Common in the Venezuelan Llanos; range of the species. ◆Fig. 548. Mimosa piscatorum Barneby, Mem. New York Bot. Gard. 65: 286. 1991. Weak, inconspicuous, weedy subshrub, randomly armed on stem-angles and leaf stalks with recurved aculei, puberulent but lacking setae; leaf stalks 11–26 mm, bearing at apex a spicule and 1 pair of pinnae, the leaflets 4–9(–11) pairs, to 3.5–8 × 1.4–2 mm; flowers in axillary pedunculate capitula, trimerous, hexandrous; craspedia linear-oblong, 25–42 × 3.5–4.5 mm, (5)6–10-seeded, the replum randomly aculeate, the valves papery pilosulous. Stony savannas, fallow fields, shores, open matorrales, near sea level to 1300 m; Bolívar (La Encrucijada). Lara, Nueva Esparta; eastern Brazil (Pernambuco, Bahia). Mimosa piscatorum resembles forms of M. pudica with conjugate pinnae, but its flowers are trimerous and diplostemonous. Mimosa polydactyla Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1033. 1806. Suffrutescent, ascending or scrambling, weedy, opportunistic herb mostly 0.5–1.6(–2) m tall, armed at and below each node with 2 or 3 aculei, thinly strigose or hispid, the leaflets setose-ciliate; leaf stalks 2–7 cm bearing near apex 3–5 closely approximate pairs of pinnae, the leaflets of longer ones 30–55

Mimosa 653

pairs and < 1 cm long; craspedia radiating from receptacle in a dense cluster, in profile 9–15 × 3.5–4.5 mm, 3- or 4-seeded, the replum always and the valves often hispid. Disturbed forests, brush-woodlands, thickets, pastures, ditch banks, 100–300 m; Delta Amacuro (Serranía de Imataca), central Bolívar, Amazonas (Río Negro). Discontinuously widespread from Costa Rica and Lesser Antilles to southeastern Brazil. Mimosa pudica L., Sp. Pl. 518. 1753. Weakly frutescent, precociously maturing (then sometimes monocarpic), diffuse or scrambling, weedy herb mostly < 1 m, armed at and below each node with 1–3 aculei, variably puberulent or hispid, the foliage very sensitive; leaf stalks 2–5.5 cm, the pinnae either 1 or 2(3), hairs all inserted near tip of leaf stalk, the leaflets 11–27 pairs, to 5–13 mm long; capitula ellipsoid, without filaments 3.5–5 mm diameter; calyx minute; craspedia in profile undulately narrow-oblong 8–15 × 3.5–4.5 mm, commonly 4-seeded, the replum hispid but the valves mostly glabrous. Disturbed and waste places, near sea level to 1200 m. American in origin but now circumtropical; 4 varieties, 2 in Venezuela, both in the flora area. Key to the Varieties of M. pudica 1. Corolla puberulous distally .................... ...................................... var. tetrandra 1. Corolla glabrous ................ var. unijuga M. pudica var. tetrandra (Humb. & Bonpl. ex Willd.) DC., Prod. 2: 426. 1825. —Mimosa tetrandra Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1032. 1806. Mimosa hispidula H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 252. 1823 [1824]. Scattered in Bolívar and Amazonas. Native from Mexico and Cuba south to southeastern Brazil, weedy in Old World. ◆Fig. 546. M. pudica var. unijuga (Walp. & Duchass.) Griseb., Abh. Königl. Ges. Wiss. Göttingen 7: 211. 1857. —Mimosa unijuga Walp. & Duchass., Linnaea 23: 744. 1850. Northwestern Bolívar, eastern Amazonas (Río Casiquiare, upper Río Orinoco). Common in northern Venezuela; scattered in Pantropics.

This is the prevalent circum-Caribbean variety. Mimosa quadrivalvis L., Sp. Pl. 522. 1753. Central and eastern U.S.A., Mexico, Central America, Cuba, Puerto Rico, Martinique, South America to northern Argentina; 16 varieties, 1 in Venezuela. M. quadrivalvis var. leptocarpa (DC.) Barneby, Mem. New York Bot. Gard. 65: 298. 1991. —Schrankia leptocarpa DC., Prod. 2: 443. 1825. —Leptoglottis leptocarpa (DC.) Standl., J. Wash. Acad. Sci. 15: 458. 1925. Diffuse, weakly suffrutescent herb with 4angulate, serially aculeate stems; capitula small, globose, axillary, borne on short prickly peduncles; pinnae 2–4 pairs; flowers mostly 5-merous, diplostemonous; craspedium bluntly 4-angular, with replum 2–3.5 mm wide and valves 1–3 mm wide, these separating from the replum in 1 piece. Savannas and disturbed lowland habitats, near sea level to 200 m; along and near the Río Orinoco in Delta Amacuro, Bolívar, and northwestern Amazonas. Well distributed over northern and central Venezuela; widespread over much of tropical South America and extending feebly north to southern Mexico and Martinique. ◆Fig. 544. The widespread and opportunistically weedy Mimosa diplotricha C. Wright (M. invisa Mart., non Mart. ex Colla; Schrankia brachycarpa Benth.) resembles M. quadrivalvis in habit and armament, but has 4–8 rather than 2–4 pairs of pinnae per leaf, a 4merous perianth, and a compressed craspedium with narrow replum and articulate valves. It has a nearly coextensive neotropical dispersal, but is inexplicably rare in Venezuela. It is known from Portuguesa and should be looked for south of the Río Orinoco in the flora area. Mimosa rufescens Benth., Trans. Linn. Soc. London 30: 418. 1875. Vine armed on stem and leaf stalks with many small retrorse prickles, the hypophyllum of leaflets and the broad papery craspedia sprinkled with small peltate trichomes; inflorescence a terminal panicle of small white fragrant capitula. Lowland forests, climbing into the canopy to 25 m, or

654

M IMOSACEAE

forming dense tangles in forest openings, cleared woodlands, in hedges, 100–400 m. Widespread almost throughout the Amazon basin and north to the lower Río Orinoco basin; 2 varieties, both in the flora area. Key to the Varieties of M. rufescens 1. Pinnae of larger leaves 3 pairs and their leaflets only 2 or 3 pairs; highly localized .................................. var. amnis-nigri 1. Pinnae of larger leaves 4–10 pairs and leaflets of distal pinnae 4–9(–11) pairs; range of the species ..... var. rufescens M. rufescens var. amnis-nigri Barneby, Mem. New York Bot. Gard. 65: 42. 1991. —Mimosa micracantha Benth., J. Bot. (Hooker) 2: 131. 1840, non Poir. 1817, “microcantha.” Amazonas (northern base of Sierra de la Neblina). Brazil (Amazonas: middle Río Negro). This is perhaps no more than a minor variant of the relatively common and variable Mimosa rufescens var. rufescens. M. rufescens var. rufescens Mimosa micrantha var. plurijuga Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 583. 1932. Delta Amacuro (expected), widespread in Bolívar and Amazonas. Distribution as in species. Mimosa scabrella Benth., J. Bot. (Hooker) 4: 387. 1841. Unarmed, broad-crowned tree of rapid growth, to 20 m tall and in girth to ca. 40 cm, the young growth scabrous with densely plumulose setules; leaflets gray-stellate beneath; pinnae 3–9 and leaflets to ca. 15–30 pairs, the leaflets 3.5–9 mm long; capitula globose, the flowers 4-merous, 4-androus, the corolla yellow-silky externally, the filaments sulfur-yellow; craspedium 15–42 × 6– 8 mm, densely scabrous-setulose. Montane forests, ca. 1100 m; Bolívar (Sierra de Lema). Brazil. Mimosa scabrella is a native of mountainous southeastern Brazil where it is associated with Araucaria and forms associations called bracaatingal. It is grown in Old and New World tropics for ornament, fuel, pulp, and charcoal. In Venezuela it is cul-

tivated in Distrito Federal and probably elsewhere, and occasionally escapes cultivation. Mimosa schomburgkii Benth., J. Bot. (Hooker) 2: 133. 1840. Broad-crowned tree 5–12(–15) m tall, trunk to ca. 40 cm diameter; flowering stems unarmed, the young branchlets pubescent with medusoid setules; foliage aromatic, the leaflets charged beneath with minute glands; inflorescence a panicle of spikes; pinnae to 7–13 and leaflets to 20–26 pairs; flowers either 4- or 5-merous, diplostemonous; perianth densely silky-tomentulose overall with intermeshed stellae; craspedium 6–9 mm wide, 4–12-seeded. Bush-islands of seasonally wet savannas, margins of gallery forests, 100–300 m; northern and central Bolívar (Río Cuyuní). Scattered in the lower Río Orinoco basin and around the southeastern periphery of the Guayana Highland in Venezuela; Honduras, Nicaragua, Guyana, Brazil (northern Roraima and cultivated in parks and gardens). Mimosa schrankioides Benth., London J. Bot. 5: 86. 1846. Prickly vine in all respects like M. casta except for number and size of leaflets, these 8–16 pairs, broad-linear, to 11–26 × 2–6.5 mm, very variable in density and quality of pubescence, either puberulent, or strigose, or glabrate; craspedia 18–40 × 7–12 mm, 3- or 4-seeded, setose overall but between setae either glabrous or densely puberulent. Riverbanks, seasonally wet savannas, sometimes weedy in pastures and along roadside ditches, 50–300 m; Bolívar, Amazonas. Discontinuously widespread in Guatemala, Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 2 varieties, both in the flora area. Key to the Varieties of M. schrankioides 1. Calyx membranous 0.1–0.4 mm, the rim not ciliolate or only minutely so; flower buds not concealed by setae .................... ..................................... var. sagotiana 1. Calyx paleaceous-pappiform ca. 1–1.2 mm, the lobes deeply decompound into setiform divisions; flower-buds concealed by a mesh of bracteal setae .......................... ............................... var. schrankioides

Mimosa 655

M. schrankioides var. sagotiana (Benth.) Barneby, Mem. New York Bot. Gard. 65: 529. 1991. —Mimosa sagotiana Benth., Trans. Linn. Soc. London 30: 394. 1875. Mimosa santanderensis Britton & Rose ex Britton & Killip, Ann. New York Acad. Sci. 35: 151. 1936. Amazonas (Puerto Ayacucho). Apure; dispersed throughout the range of the species. M. schrankioides var. schrankioides Northern Bolívar (Cerro San Borja along lower Río Suapure). Middle and lower Río Orinoco in Venezuela; Guyana, Brazil (northern Roraima). Mimosa sensitiva L., Sp. Pl. 518. 1753. Diffusely tangled or vine-like, softly frutescent, very prickly subshrubs to 2 m tall but often smaller, the stems and leaf stalks armed with files of cat’s-claw aculei; pinnae 1 pair and leaflets of each 2 pairs, the larger distal pair dimidiately ovate or falcately lance-elliptic 2.5–6 cm long; capitula globose, mostly axillary; calyx pappiform 0.7–1.6 mm; craspedia in profile oblong-elliptic 13– 23 × 4–7 mm, setose especially along replum. Thickets, seasonally dry woodlands, savannas, in the lowlands becoming aggressively weedy in disturbed places and on roadsides, 50–200 m; Bolívar (along and near Río Orinoco south of Ciudad Bolívar), adjacent Amazonas. Coastal Cordillera, Llanos; common in eastern Brazil between the mouth of the Amazon and Bahia and interruptedly north (perhaps adventive with cattle). Mimosa somnians Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1036. 1806. Mimosa palpitans Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1036. 1806. Mimosa somniculosa H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 257. 1823 [1824]. Mimosa tobagensis Urb., Repert. Spec. Nov. Regni Veg. 15: 307. 1918. Potentially fruticose to 3 m tall but often weakly so, extremely variable in habit, foliage, and indumentum, armed or not, but with characteristically firm, striately veined corolla lobes; craspedium usually stipitate, commonly 3–9 cm long and 7–16-seeded. Mexico, Central America, South America to northern Argentina, adventive in Jamaica; 4 subspecies, 1 in Venezuela.

M. somnians subsp. somnians Armed and ± glandular. Range coextensive with the species; 2 varieties, both in the flora area. Key to the Varieties of M. somnians subsp. somnians 1. Leaf stalks of longer leaves subfiliform 0.25–0.35 mm diameter; rachis of longer pinnae 9–14 mm, bearing 13–19 pairs of leaflets .......................... var. deminuta 1. Leaf stalks of longer leaves > 0.35 mm diameter; rachis of longer pinnae 15–55 mm, bearing 20–50 pairs of leaflets ...................................... var. somnians M. somnians subsp. somnians var. deminuta Barneby, Mem. New York Bot. Gard. 65: 454. 1991. On and about granitic outcrops, 100–200 m; Bolívar (along Río Orinoco from Puerto Ayacucho downstream to mouth of Río Caura), adjacent northern Amazonas. Colombia (Vichada). M. somnians subsp. somnians var. somnians. —Arestín. Savannas, brush-woodland, open hillsides, becoming weedy in lowland pastures, 100–300 m; northeastern Bolívar (east from lower Río Caroní basin). Mexico, Central America, South America to eastern Brazil and Argentina. Mimosa surumuënsis Harms, Notizbl. Königl. Bot. Gart. Berlin 6: 304. 1915. Shrub 0.5–4 m tall, randomly remotely aculeate, remarkable for the indumentum of plumulose setae combined with diplostemonous flowers; pinnae to 5–11 and leaflets to 25–36 pairs, the largest leaflets ca. 2.5–4.5 mm long; inflorescence a panicle of small capitula; craspedium 2.5–3 mm wide, 3–6seeded. Outcrops in savannas, rock ledges, 300–600 m; Bolívar (northern slope of Cerro Bolívar). Guyana, Brazil (northern Roraima: Serra Pacaraima). Mimosa tarda Barneby, Mem. New York Bot. Gard. 65: 434. 1991. Shrub 2–3 m tall. River banks, 200–300 m; Amazonas (Río Ocamo). Interruptedly dispersed in subequatorial latitudes in Central America and South America.

656

M IMOSACEAE

Mimosa tarda closely resembles M. pellita and M. orinocoësis, except as noted in the key to species. Mimosa velloziana Mart., Flora 22(1) Beibl.: 9. 1839. —Mimosa viva Vell., Fl. Flumin. Icon. 11: pl. 33. 1827 [1831]. Mimosa jiramenensis H. Karst., Fl. Columb. 2: 59, t. 130. 1862 [1863]. —Mimosa velloziana var. jiramenensis (H. Karst.) Benth., Trans. Linn. Soc. London 30: 390. 1875. Essentially like M. sensitiva in habit, foliage, and ecology, different in minute calyx 0.25–0.45 mm long and in broad pod, 20–42 × 8–13 mm. Secondary lowland and ruderal habitats, 50–100 m; Amazonas (Raudal de Atures). Maracaibo basin, Llanos; widespread from southern Mexico to northern Argentina. Mimosa velloziana is a weedy mimosa of doubtful native range.

Mimosa xanthocentra Mart., Flora 21(2) Beibl. 4–5: 50. 1838. Herbaceous or suffruticose, armed at and close below nodes or unarmed; pinnae of all leaves 1 pair and the many small leaflets forwardly imbricated, 1-veined; capitula ovoid, gray-puberulent; flowers 4-merous, 4androus; calyx paleaceous-pappiform, the lobes decompound into capillary divisions. Widespread over much of South America east of the Andes; 4 subspecies and 8 varieties, none yet found in the flora area. M. xanthocentra subsp. xanthocentra var. xanthocentra Armed. Grassland and brush communities, to be expected in savannas south of Río Orinoco in northwestern Bolívar. Common in the Maracaibo and upper Río Orinoco basins east to Guárico; Colombia, southeastern Brazil, northwestern Argentina.

Fig. 544. Mimosa quadrivalvis var. leptocarpa Fig. 545. Mimosa hirsutissima

Mimosa 657

Fig. 546. Mimosa pudica var. tetrandra

Fig. 548. Mimosa pellita var. pellita

Fig. 547. Mimosa orthocarpa

658

M IMOSACEAE

16. NEPTUNIA Lour., Fl. Cochinch. 2: 653. 1790. by Rupert C. Barneby Herbs, pubescent with simple hairs, or glabrescent. Leaves bipinnate, the leaflets in opposite pairs; stipules ± dilated and cordate, commonly striate-veined; petiolar nectary present or not. Inflorescence axillary, the peduncle 2-bracteate. Flowers densely spicate or capitate, dimorphic, the upper ones bisexual, the lower staminate or (in ours) sterile, with petaloid yellow filaments. Perianth 5-merous, the campanulate tube shortly toothed, the petals nearly or quite free, valvate in bud. Stamens either 10 or 5, the anther-connective gland-tipped or not. Pods stipitate, narrowly oblong in profile, gently curved downward, compressed but turgid over each seed, the valves inertly dehiscent downward from apex. Seeds compressed, exarillate, with hard testa, a pleurogram on each face, and thin endosperm adherent to the testa. Circumtropical and warm-temperate areas; 11 species, 2 in Venezuela, both in the flora area. Neptunia is a genus of sunny, moist or swampy lowland habitats, one species floating in still waters, four native in the New World. The leaves are very sensitive. Key to the Species of Neptunia 1.

1.

Floating herb with markedly spongy-inflated internodes and many fine adventitious roots at each node; no petiolar nectary; anther-connective glandless; seeds 4–8 per pod ...................................................... N. oleracea Erect herb of wet ground or standing water, the stems then somewhat spongy but not adventitiously rooting; petiolar nectary near insertion of first pair of pinnae; connective of fertile anthers gland-tipped (in bud); seeds 8–20 per pod ................................................................ N. plena

Fig. 549. Neptunia oleracea

Parkia 659

Neptunia oleracea Lour., Fl. Cochinch. 654. 1790. Mimosa natans L. f., Suppl. Pl. 439. 1781. —Neptunia natans (L. f.) Druce, Bot. Soc. Exch. Club Brit. Isles 4: 637. 1916 [1917], non W. Theob. 1883. Mimosa prostrata Lam., Encycl. 1: 10. 1783, nom. illeg. —Neptunia prostrata (Lam.) Baill., Bull. Mens. Soc. Linn. Paris 1: 356. 1883. Floating herb with 1 leaf and 1 peduncle arising from each node, the leaf consisting of usually 2 or 3 pairs of pinnae, these composed of < 20 pairs of small leaflets. Pools, river backwaters, at low elevations; Bolívar (expected south of Río Orinoco). Scattered in Venezuelan Llanos;

scattered in Neotropics, Africa, and Malesia. ◆ Fig. 549. Neptunia plena (L.) Benth., J. Bot. (Hooker) 4: 355. 1842. —Mimosa plena L., Sp. Pl. 519. 1753. Neptunia surinamensis Steud., Flora 26: 759. 1843. Usually erect perennial herb from orange taproot; larger leaves consisting of 2–4 pairs of pinnae, these composed of 20–35 pairs of leaflets. Moist or wet ground, sometimes in shallow water, lowlands; northern Delta Amacuro, Bolívar (Ciudad Bolívar). Rather common in the Llanos; widely dispersed in the Neotropics, also India, where thought to be introduced.

17. PARKIA R. Br. in Denham & Clapperton, Narr. Travels Africa app. 289. 1826. by Helen C. F. Hopkins Trees to 40 m tall or more, or rarely shrubs; spines absent. Leaves opposite, alternate, or in clusters, bipinnate; petiole and rachises usually nectariferous. Pinnae 2–55 pairs. Leaflets sessile, to 100 or more pairs. Compound inflorescence axis to 5 m long, branched or not. Peduncles to 1 m or more long, pendent or erect. Capitula capitate, spherical, globose, clavate, or biglobose, from 1 × 1 to 20 × 8 cm. Flowers in a single capitulum either all fertile (bisexual or if staminate, with the gynoecium reduced or absent), or of 2 kinds (fertile ones at base of capitula; modified, structurally hermaphrodite but functionally sterile, nectariferous ones around the apex), or of 3 kinds (fertile ones in distal ball; modified, structurally staminate and nectariferous ones below; and neuter ones, often bearing elongated staminodia, at base of capitulum). Bracts obdeltate-spathulate, slightly longer than calyx. Calyx long-tubular or funnel-shaped, gamosepalous, the 5 lobes usually imbricate, 2 larger and 3 smaller, or rarely induplicate and ± equal in size; corolla longer than calyx, the 5 lobes narrowly spathulate or lingulate, free at apex and either connate in middle and free at base, or free in middle and connate at base. Stamens 10, all fertile, shortly exserted; filaments connate below into a tube, to which the corolla may also be adnate; anthers, at least in New World species, usually bearing a small apical gland. Ovary shortly stipitate; style exserted as far as anthers or beyond; stigma terminal, forming a pore at the tip. Infructescence pendent or erect; receptacle swollen, claviform, bearing 1–several pods and rhomboidal scars where flowers have aborted. Pods stipitate, coriaceous, ligneous or rarely fleshy, dehiscent or not, sometimes velutinous, linear, strap-shaped, or oblong in outline, sometimes falcate, submoniliform, terete, or broadly crescent-shaped, sometimes twisted or curled, sometimes containing mealy pulp or gum. Seeds numerous, in 1 or 2 rows, discoid, ellipsoid, or rarely otherwise; testa usually thick and hard, bearing a horseshoeshaped line (pleurogram) on either face, or sometimes thin and pleurogram lacking. Pantropics; ca. 35 species, 9 in Venezuela, all of these in the flora area. Huber 10332 (NY and VEN, from Bolívar state, savanna borders of the Río Trueno west of Chiguao, 480 m) may represent an additional new taxon, but more complete specimens are required.

660

M IMOSACEAE

Key to the Species of Parkia 1.

1.

2(1).

2.

3(2).

3.

4(3).

4.

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

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

Leaflets with numerous erect, white or yellowish hairs, especially on the margins and underside; pods velutinous, the valves 11–16 × 4–5 cm plus stipe of 2–3 cm long; tree ca. 9 m tall ........................... P. barnebyana Leaflets glabrous or rarely sparsely hairy on margin; pods glabrous, or if velutinous then valves 20–45 × 3–6(–8) cm, plus stipe of 2–13 cm long; tree to 40 m tall ...................................................................................... 2 Capitula spherical or oblate both in bud and at anthesis, lacking basal fringe and constricted ring of nectar-secreting flowers; mature pods dehiscent though sometimes tardily so ................................................. 3 Capitula clavate in bud and biglobose at anthesis, having a broad basal fringe of neuter flowers with far exserted staminodia, an apical ball of fertile flowers, and a constricted ring of modified staminate, nectar-secreting flowers in between; mature pods indehiscent .......................... 5 Peduncles to 50 cm long, flexible, pendent; capitula oblate with basal flowers fertile and apical flowers modified-hermaphrodite and nectariferous; pods glabrous, dehiscent along thickened adaxial suture only .............................................................................................. P. pendula Peduncles to 4 cm long, rigid, ± erect; capitula spherical, composed entirely of fertile flowers; pods velutinous, dehiscent along both sutures ................................................................................................................ 4 Main axes of compound inflorescence unbranched, arising proximally to the leaves; pinnae (17–)30–45(–55) pairs; leaflets 40–110 pairs per pinna; pods 18–50 × 4.7–6 cm; capitula red ............................... P. velutina Main axes of compound inflorescence much-branched, arising distally to the leaves; pinnae 9–20 pairs; leaflets 30–55 pairs per pinna; pods 17– 36 × 3–4.7 cm; capitula yellow ........................................................... P. ulei Peduncles opposite or subopposite, or several inserted together in opposite or subopposite groups; leaves opposite .................................. P. nitida Peduncles alternate and usually inserted singly; leaves alternate ......... 6 Valves of pod somewhat spongy or corky with air between layers, glabrous, not or scarcely corrugated over seeds; seasonally flooded forests ...................................................................................................... P. discolor Valves of pod coriaceous, not spongy or corky, glabrous or velutinous, usually corrugated over seeds; nonflooded forests ..................................... 7 Leaflets weakly or strongly sigmoid; capitula bicolored with red fringe and yellow distal ball ............................................................. P. panurensis Leaflets oblong, never sigmoid; capitula ± unicolored, either all reddish or all yellowish ........................................................................................... 8 Largest leaflet on the leaf 27–39 × 5.5–10 mm; capitula usually with red dish fringe and pink distal ball; pods glabrous ........................ P. igneiflora Largest leaflet on the leaf 13–20 × 3–4.5 mm; capitula with cream fringe and yellow distal ball; pods velutinous ..................................... P. truncata

Parkia barnebyana H.C. Hopkins, Kew Bull. 55: 133. 2000. Small tree to 9 m tall; leaflets oblong, 3–5 × ca. 1–1.5 mm; compound inflorescence axis

short and erect; capitula and flowers unknown. Savannas bordering forests, riparian forests, ca. 100 m; Amazonas (Cucurital near Caname). Endemic.

Parkia 661

Parkia discolor Spruce ex Benth., Trans. Linn. Soc. London 30: 363. 1875. —Cacani-caritsapi, Casabe de murciélago. Parkia auriculata Spruce ex Benth., Trans. Linn. Soc. London 30: 363. 1875. Tree to 15 m tall or more, branching low, often with an irregular, straggly, open crown and long tortuous branches; leaflets much darker on upper surface, ± oblong to sigmoid, 10–26 × 2.5–7 mm; capitula ca. 4.5–5.5(–6.5) cm long, with deep red fringe and pink-red distal ball. Common on white sand in forests seasonally flooded by black-water rivers, 50– 200 m; Amazonas (basins of Río Atabapo, Río Casiquiare, Río Negro, and upper Río Orinoco). Amazonian Colombia and Brazil. ◆Fig. 550. Parkia igneiflora Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 472. 1932. Tree to 18 m tall, with small rounded crown and usually very upwardly extending, ± erect compound inflorescence axes; capitula ca. 6.5–8.5 cm long. Low- to mediumheight forests on white sand (caatinga woodland), swampy ground near black-water streams but not flooded forests, 100–200 m; Amazonas (Río Guainía, Río Negro). Amazonian Colombia, Peru, and Brazil. Parkia nitida Miq., Stirp. Surinam. Select. 7. 1850 [1851]. —Caro blanco, Caro montañero, Hueso de pescado, Makumik. Parkia oppositifolia Spruce ex Benth., Trans. Linn. Soc. London 30: 363. 1875. Medium or large tree to 40 m tall, the bole sometimes buttressed, the bark often smelling of garlic, onions, or methyl salicylate, and the crown sometimes appearing silverygrey from the white, waxy coating on leaflets; leaflets oblong, 10–25 × 2.5–6.5 mm; capitula ca. 4.5–8 cm long, with cream fringe and yellow distal ball. Nonflooded forests, old secondary forests, 100–1300 m; Delta Amacuro (El Palmar, near Bolívar border), Bolívar (southwest of El Manteco, Macizo del Chimantá [Acopán-tepui]), Amazonas (Sierra de la Neblina). Southern Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. Parkia panurensis Benth. ex H.C. Hopkins, Brittonia 34: 348. 1982. —Casabe murciélago, Clavellina.

Parkia pectinata auct. non Benth. 1875: sensu Benth., Trans. Linn. Soc. London 30: 362. 1875, pro parte. Small to large tree 15–35 m tall; compound inflorescence axes extending shortly beyond crown, ± erect; leaflets 7–13 × 1.5–3 mm; capitula ca. 4.5–7 cm long. Nonflooded evergreen lowland forests on sandy soils (caatinga forests and caatinga woodlands), sometimes near streams, 100–300 m; Amazonas (Piedra Arauicaua, Reserva Forestal El Sipapo, Río Pacimoni-Río Yatúa, San Carlos de Río Negro, San Carlos to Solano road). Amazonian Colombia, Ecuador, Peru, Brazil. Parkia pendula (Willd.) Benth. ex Walp., Repert. Bot. Syst. 5: 577. 1846. —Inga pendula Willd., Sp. Pl. 4: 1025. 1806. —Aruro (Yekwana), Caro, Cascarón, Day’ e’mon, Deyi, Dividive, Pega-pego, Tanyeechemen, Warada-kuma-tare, Zarcillo. Large tree to ca. 40 m tall, bole ca. 1 m dbh, crown broad, flattened, layered, or umbrella-shaped with capitula or pods pendent below; leaflets linear, 3–6 × 0.5–1 mm; capitula bright red; pods with amber-colored, sticky gum along thickened suture on dehiscence. Evergreen lowland forests, 50–500 m; widely distributed in Bolívar and Amazonas. Táchira, Zulia; Honduras, Nicaragua, Costa Rica, Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador and Peru, Amazonian and coastal Brazil. ◆Fig. 551. Parkia truncata R.S. Cowan, Mem. New York Bot. Gard. 9: 336. 1957. Tree to 20 m tall; capitula ca. 5–6 cm long. Forests on granitic outcrops, ca. 200 m; Amazonas (Rio Cunucunuma). Endemic. Parkia ulei (Harms) Kuhlm., Arch. Jard. Bot. Rio de Janeiro 4: 356, t. 30. 1925. —Leucaena ulei Harms, Verh. Bot. Vereins Prov. Brandenburg 48: 162. 1906 [1907]. Tree 3–50 m tall; leaflets oblong, 3–10 × 0.5–2 mm; capitula 1–1.8 mm diameter. Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 2 varieties, 1 in Venezuela. P. ulei var. surinamensis Kleinhoonte, Recueil Trav. Bot. Néerl. 30: 169. 1933. —Caro, Clavelino, Clavellina.

662

M IMOSACEAE

Fig. 550. Parkia discolor

Parkia 663

Fig. 551. Parkia pendula

664

M IMOSACEAE

Large tree to 35(–50) m tall, sometimes with plank buttresses to 2 m tall and a large rounded crown. Evergreen lowland forests, occasionally riparian forests, near sea level to 100 m; Bolívar (El Palmar). Monagas; Guyana, Suriname, French Guiana, eastern Amazonian Brazil. Parkia velutina Benoist, Notul. Syst. (Paris) 3: 271. 1914. —Uaira tombepe.

Large tree to 40 m tall, with buttresses to 2 m tall, crown with clusters of leaves borne on thickened velutinous twigs; leaflets linear or narrowly sigmoid, 5–11.5 × 1–2 mm; capitula ca. 2.5–3.5 cm diameter. Evergreen lowland forests, often near streams, 400–500 m; Bolívar (upper Río Caroní). Colombia west of the Andes, French Guiana, Amazonian Ecuador, Peru, and Brazil.

Fig. 552. Pentaclethra macroloba

Piptadenia 665

18. PENTACLETHRA Benth., J. Bot. (Hooker) 2: 127. 1840. by Rupert C. Barneby Unarmed trees with amply bipinnate leaves, thinly pilosulous, eglandular. Leaf stalks lacking nectary, the narrow crowded leaflets linear-falcate, glabrous or almost so; stipules lanceolate, deciduous. Inflorescence a terminal, efoliate or proximally few-foliate panicle of stout elongate, densely many-flowered spikes. Flowers 5-merous. Calyx small, campanulate, shortly lobed, the lobes imbricate in bud; corolla reddish purple, gamopetalous for at least half its length, valvate in bud. Stamens white or yellowish, dimorphic, 5 antesepalous fertile, their anthers shortly exserted from corolla and the connective tipped with a bilobed gland, 5 antepetalous, much longer, staminodal. Pod ascending and outwardly falcate, in profile oblanceolate, the valves woody, coarsely striate lengthwise, elastically dehiscent from apex downward through both sutures and coiled into a ring. Seeds ca. 3–5, large, plumply compressed-obovoid, the brown testa lacking pleurogram; endosperm absent. Central America, Lesser Antilles, Colombia, Venezuela, northern Amazonian Brazil, Africa; 2 species, 1 in Venezuela. Pentaclethra macroloba (Willd.) Kuntze, Revis. Gen. Pl. 1: 201. 1891. —Acacia macroloba Willd., Sp. Pl. 4: 1060. 1806. —Abarkasa-dek (Arekuna), Arabakasayek (Arekuna), Behebehedu (Warao), Bihibihidu (Warao), Clavellino, Clavellín, Clavellina. Pentaclethra filamentosa Benth., J. Bot. (Hooker) 2: 127. 1840. Tree 7–30 m tall, with trunk to 30 cm diameter; leaves 20–40 cm long, the 11–17

pairs of pinnae opposite or alternate, the leaflets of longer pinnae ca. 35–45 opposite pairs, proximally auriculate, acute; spikes including short peduncle 10–25 cm, without filaments ca. 1.5 cm diameter; pod 18–31 × (3–)3.5–5 cm, glabrous, dark brown. Evergreen lowland to lower montane forests, 50– 500 m; frequent and locally plentiful in southern Delta Amacuro and central and northern Bolívar. Distribution as in genus, except Africa. ◆Fig. 552.

19. PIPTADENIA Benth., J. Bot. (Hooker) 2: 135. 1840. Piptadenia sect. Pityrocarpa Benth., J. Bot. (Hooker) 4: 339. 1841. —Pityrocarpa (Benth.) Britton & Rose, N. Amer. Fl. 23: 190. 1923. by Rupert C. Barneby Trees, some with several trunks and bushy habit, shrubs, and scrambling vines, commonly armed on stems and leaf stalks with scattered recurved prickles, rarely armed only at stem nodes with a pair of stout prickles, rarely unarmed except for spinescent stipules, but 2 of ours completely unarmed, almost all puberulent or pilosulous eglandular; a cupular, patelliform or mounded nectary on petioles at or below first pair of pinnae. Leaves variable in numbers of pinnae and leaflets, these either many and small or few and ample. Inflorescence of narrow spikes axillary to coeval leaves or terminally paniculate. Flowers 5-merous, 10-androus. Calyx campanulate, shortly toothed; petals either free or connate into a tube. Filaments free or almost so; anthers parallel, the narrow connective tipped with a caducous gland. Ovary stipitate. Pod commonly resembling that of many American Acaciae, in profile linear-oblong or oblong-elliptic, straight, narrowed at base into a stipe, the sutures either straight or deeply constricted between seeds, the body planocom-

666

M IMOSACEAE

pressed, the papery or subcoriaceous valves finally separating inertly along one or both sutures. Seeds subdiscoid, the testa not winged, bearing a small pleurogram; endosperm present. Neotropics to Argentina (most numerous in eastern and Amazonian Brazil); ca. 20 species, 6 in Venezuela, 4 of these in the flora area. Piptadenia is heterogeneous and is of disputed definition. Key to the Species of Piptadenia 1.

1.

2(1). 2.

3(1).

3.

Plant wholly unarmed; peduncle of spikes charged below the first flower with a lobed involucre, this early dry and circumscissile; sutures of pod undulately constricted between seeds, the ripe valves becoming reddish brown and furfuraceous, dull (section Pityrocarpa) ..................... 2 Plant armed either at nodes or on internodes; peduncle of spikes without an involucre, sometimes with a single small bract; sutures of pod (not known in P. imatacae) straight, and ripe valves lustrous, venulose (section Piptadenia) ..................................................................................... 3 Leaflets of distal pinnae (3)4–7(8) pairs, the larger blades (30–)35–70 × 13–18 mm; tall trees of lowland forests ................................. P. leucoxylon Leaflets of distal pinnae 9–13 pairs, the blades relatively small, the largest 9–21 × 4–11 mm; shrubby trees of savannas and scrub ............................................................................................. P. moniliformis Long forest vines, armed on stems and leaf stalks with scattered prickles; young growth tawny-pilosulous; leaf nectary mounded, with small terminal pore; pinnae to ca. 3 and leaflets of distal ones 4 or 5 pairs, the largest leaflets obovate and to 20–33 mm; corolla scarcely 2 mm long, not over 2–3 times as long as calyx, the essentially free segments villosulous dorsally .................................................................... P. imatacae Free-standing trees, armed at random nodes with a pair of stout prickles, elsewhere unarmed; young growth finely gray-puberulent or glabrate; leaf nectary depressed-patelliform, longer than wide; pinnae to 8–12 and leaflets to 25–45 pairs, the leaflets linear-oblong, < 7 mm long; corolla 4–6 mm long, at least 4 times as long as calyx, the glabrous segments united into a tube .................................................................... P. viridiflora

Piptadenia imatacae Barneby, Brittonia 38: 227, fig. 1C. 1986. —Arespín. Vine to 30 m high; flowers greenish white or ochroleucous, the concolorous filaments exserted only ca. 1 mm; pod unknown. Lower montane forests, 300–500 m; northeastern Bolívar (Serranía de Imataca). Endemic.

Piptadenia moniliformis Benth., J. Bot. (Hooker) 4: 339. 1841. Small shrubby tree 1–6 m tall forming thickets by suckering. Colonizing savannas, seasonally dry thickets, 50–100 m; Bolívar (San Félix, Puerto Ordaz). Anzoátegui, Sucre; eastern Brazil (Pará southeast to Bahia).

Piptadenia leucoxylon Barneby & J.W. Grimes, Brittonia 36: 236, fig. 1. 1984. —Palo blanco, Clavellino. Tree 10–30 m tall, trunk to 40 cm diameter. Semideciduous forests, 50–300 m; northeastern Bolívar (islands in Lago Guri, El Paraíso camo northeast of Upata in Altiplanice de Nuria). Anzoátegui, Monagas; Trinidad, southern Guyana. ◆Fig. 553.

Piptadenia viridiflora (Kunth) Benth., J. Bot. (Hooker) 4: 337. 1841. —Acacia viridiflora Kunth, Mimoses 81, t. 25. 1819 [1821]. —Pityrocarpa viridiflora (Kunth) Brenan, Kew Bull. 10: 177. 1955. —Tiamo güire (in Guárico). Piptadenia biuncifera Benth., J. Bot. (Hooker) 4: 337. 1841. Piptadenia rubescens Pittier, Bol. Minist.

Pithecellobium 667

Fig. 553. Piptadenia leucoxylon

RR. EE. no. 8/9, reprinted in Arb. Arbust. Venez. 6–8: 84. 1927. Piptadenia speciosa Britton & Killip, Ann. New York Acad. Sci. 35: 155. 1936. Tree 6–20 m tall, often several-trunked; corolla greenish white, sometimes rubescent in age; pod excluding stipe (6–)8–12(–15) × (1.5–)1.8–2.2(–2.5) cm, the valves reticulate in age. Semideciduous forests, matorral, and physiognomically similar, seasonally dry

communities, 50–600 m; northeastern Bolívar (lower Río Caura). Venezuelan Llanos; discontinuously dispersed from southern Mexico (Chiapas) south through interAndean Colombia to eastern Bolivia and northwestern Argentina, eastern Brazil. Piptadenia viridiflora is anomalous in the genus by its elongate tubular corolla, longexserted filaments, and armature of paired infranodal prickles.

20. PITHECELLOBIUM Mart., Flora 20(2) Beibl.: 114. 1837, nom. cons., s. str. Pithecellobium sect. Unguis-cati Benth., Trans. Linn. Soc. London 30: 571. 1875, “Pithecolobium.” by Rupert C. Barneby and James W. Grimes Trees and subarborescent shrubs, mostly xerophytic and deciduous, armed with stipular spines (but flowering branchlets of some unarmed). Leaves bipinnate, the pinnae 1 or 2 (in ours 1) pairs, the leaflets of each pinna 1 pair, in ours relatively large (5–12 cm). Inflorescence of spikes or (ours) capitula. Flowers isomorphic or the terminal one larger. Calyx campanulate; corolla tubular. Filaments united into a tube, the anthers without terminal gland. Pods linear, compressed but turgid, either decurved, or twisted, or coiled, the chartaceous or fleshy valves separating in dehiscence but remaining attached to receptacle, red or orange internally. Seeds invested in lower half by a fleshy aril.

668

M IMOSACEAE

Fig. 554. Pithecellobium roseum

U.S.A. (Florida), Mexico, Neotropics (most diversity), poorly represented south of the Equator, a few species native or naturalized in Paleotropics; ca. 20 species, 5 in Venezuela, 2 of these in the flora area. Key to the Species of Pithecellobium 1.

1.

Peduncles at most nodes of pseudoracemose inflorescence 2–4 (solitary only at random nodes where subtended by a rudimentary leaf); tassel of androecium red or red-purple .................................................. P. roseum Peduncles at all or at most nodes of pseudoracemose inflorescence solitary; tassel of androecium white or ochroleucous ................ P. unguis-cati

Pithecellobium roseum (Vahl) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(2): 21. 1997. —Mimosa rosea Vahl, Eclog. Amer. 3: 33, t. 25. 1807. —Inga rosea (Vahl) Steud., Nomencl. Bot. 1: 431. 1821. —Taguapira. Inga forfex Kunth, Mimoses 51, t. 16. 1819 [1820]. —Pithecellobium forfex (Kunth) Benth., London J. Bot. 3: 199. 1844. Inga pubescens DC., Prod. 2: 437. 1825. —Pithecellobium pubescens (DC.) Benth., J. Bot. (Hooker) 2: 141. 1840. Pithecellobium guaricense Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 51. 1926. Pithecellobium parviflorum Pittier, Bol. Soc. Venez. Ci. Nat. 5: 305. 1939.

Tree of bushy outline; seeds lustrous black with white aril. Disturbed matorrales, xeromorphic woodlands, dry hills, sometimes in seasonally flooded savannas, 50–400 m; Bolívar (near Ciudad Bolívar and Upata). Scattered in northern Venezuela; discontinuously dispersed in Tobago, Colombia, Guyana, and northern Brazil. ◆Fig. 554. Pithecellobium unguis-cati (L.) Benth., London J. Bot. 3: 200. 1844. —Mimosa unguis-cati L., Sp. P. 517. 1753. —Inga unguis-cati (L.) Willd., Sp. Pl. 4(2): 1006. 1806. —Zygia unguis-cati (L.) Sudw., U.S.D.A. Div. Forest. Bull. 14: 248. 1897. Inga microphylla Humb. & Bonpl. ex

Pseudopiptadenia 669

Willd., Sp. Pl. 4: 1004. 1806. —Pithecellobium microphyllum (Humb. & Bonpl. ex Willd.) Benth., London J. Bot. 3: 200. 1844. Pithecellobium microchlamys Pittier, Bol. Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 48. 1926. Pithecellobium pulchellum Pittier, Bol. Soc. Venez. Ci. Nat. 4: 82. 1937, non Pittier 1922. —Pithecellobium larensis

L. Cárdenas, Revista Fac. Agron. (Maracay) 7: 123. 1974. Arborescent, drought-deciduous shrub, flowering after rains. Semideciduous forests, 200–300 m; northern Bolívar (islands in Lago Guri). Anzoátegui, Falcón, Nueva Esparta, Sucre, Táchira; U.S.A. (Florida), Pacific southern Mexico, Central America, West Indies, south sparingly to the Orinoco basin.

21. PSEUDOPIPTADENIA Rauschert, Taxon 31: 559. 1982. Newtonia sect. Neonewtonia Burkart in Reitz, Fl. Ilustr. Catarin. LEGU/I: 285. 1979, s. str. by Rupert C. Barneby Unarmed trees (in flora area), puberulent, eglandular, with amply bipinnate leaves and a linear-elliptic nectary sessile in the groove of the petiole. Inflorescence of axillary spikes. Flowers small, whitish or ochroleucous, 5-merous, 10-androus.

Fig. 555. Pseudopiptadenia suaveolens

670

M IMOSACEAE

Calyx shallowly campanulate; corolla lobes erect. Anthers parallel, the narrow connective tipped with an early caducous gland. Ovary stipitate, pilosulous. Pods pendulous, linear in profile, planocompressed, the subligneous valves splitting when ripe along the ventral suture and narrowly gaping to release the seeds. Seed planocompressed, the thin lustrous testa produced around the margin as a narrow wing, lacking pleurogram; endosperm absent. Costa Rica, Venezuela to Bolivia and southeastern Brazil; 9 species, 2 in Venezuela, 1 of these in the flora area. Pseudopiptadenia suaveolens (Miq.) J.W. Grimes, Brittonia, 45: 27. 1993. —Piptadenia suaveolens Miq., Linnaea 18: 589. 1844. —Newtonia suaveolens (Miq.) Brenan, Kew Bull. 1955. 182. 1955. —Chirima (Arekuna), Yigüire, Hueso de pescado (applied to a variety of mimosoid trees). Tree 30–50 m tall, with trunk above the buttressed base to 60 cm diameter; leaves of about 7–12 pairs of pinnae of which the

longer have 25–40 pairs of opposite leaflets to 5–9 mm, lustrous above and dull or tawny beneath; pods (10–)20–40 × 1–1.7 cm; flowers fragrant. Nonflooded evergreen lowland forests, 50–200 m; Delta Amacuro (Río Toro), northwestern Bolívar (Altiplanicie de Nuria, Río Tabaro in Río Nichare basin, Serranía de Imataca), Amazonas (San Carlos de Río Negro). Coastal Cordillera; frequent in Guyana, Suriname, French Guiana, and lower Amazonian Brazil. ◆Fig. 555.

Fig. 556. Pseudosamanea guachapele

Samanea 671

22. PSEUDOSAMANEA Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 54. 1930. by Rupert C. Barneby and James W. Grimes Deciduous or semideciduous unarmed trees. Leaflets relatively few and large; stipules lanceolate, caducous; pinnae of larger leaves 3–6 pairs; leaflets of longer pinnae 5–8 pairs, the longer leaflets ca. 2.5–5 cm, pinnately veined. Inflorescence composed of umbelliform capitula axillary to coeval leaves. Flowers of each capitulum heteromorphic, the peripheral ones slenderly pedicellate, the terminal one sessile and much larger. Androecium 16–30-merous; filaments united into a tube; anthers eglandular. Pod sessile or shortly stipitate, broad-linear, planocompressed, straight or almost so, dehiscent through ventral suture or subindehiscent. Seeds transverse; testa crustaceous; pleurogram U-shaped. Southeast Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru; 2 species, 1 in Venezuela. Pseudosamanea guachapele (H.B.K.) Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 54. 1930. —Acacia guachapele H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 281. 1823 [1824]. —Lysiloma guachapele (H. B.K.) Benth., Trans. Linn. Soc. London 30: 533. 1875. —Pithecellobium guachapele (H.B.K.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 13(3.1): 54. 1943. —Albizia guachapele (H.B.K.) Dugand, Phytologia 13: 389. 1966. —Guacha, Masaguaro, Samanigua, Tabaca. Samanea samanigua Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 54. 1926. —Pithecellobium samanigua (Pittier) J.F. Macbr., Candollea 6: 4. 1934.

Pithecellobium longepedatum Pittier, Contr. U.S. Natl. Herb. 20: 464. 1922. —Albizia longepedata (Pittier) Britton & Rose, Trop. Woods 11: 14. 1927. Tree to 21 m tall, flowering shortly before or with the young leaves of the year. Gallery forests, disturbed matorrales, seasonally flooded forests, 50–100 m; Bolívar (Puerto Ordaz). Frequent in northern and western Venezuela, Apure; widespread in Central America, Colombia, Ecuador, northwestern Peru, sometimes cultivated. ◆Fig. 556. The “Samanigua” (Pseudosamanea guachapele) at anthesis resembles Samanea saman, but differs in longer pedicels of peripheral flowers (11–22 mm, not less than 5 mm long).

23. SAMANEA (Benth.) Merr., J. Wash. Acad. Sci. 6: 46. 1916. —Pithecellobium sect. Samanea Benth., Trans. Linn. Soc. London 30: 585. 1875. Pithecellobium sect. Samanea ser. Carnosae Benth., Trans. Linn. Soc. London 30: 587. 1875. by Rupert C. Barneby and James W. Grimes Semideciduous trees, some attaining vast size. Leaves bipinnate, with relatively few and large leaflets folding forward at dusk or after shock; stipules herbaceous, early deciduous; pinnae of larger leaves 3–6 pairs; leaflets of longer pinnae 3– 8 pairs, pinnately veined. Capitula axillary to coeval leaves, umbelliform, the flowers heteromorphic, the peripheral ones at least shortly pedicellate, the terminal one stouter, sessile. Calyx of peripheral flowers vase-shaped, 4.5–11 mm long; corolla 7– 14.5 mm long. Stamens 16–36, united into a tube shorter than corolla, the tassel pink or reddish. Pods subsessile, broad-linear, straight or nearly so, either compressed but plump or biconvex, indehiscent, the exocarp thin, the mesocarp pulpy, the endocarp crustaceous-lignescent, narrowly septiferous between seeds. Seeds released by weathering or predators, or excreted in cattle dung; testa hard, areolate (with a pleurogram).

672

M IMOSACEAE

Fig. 557. Samanea saman

Central America, West Indies, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia, Argentina, Paraguay, cultivated and naturalized in Mexico, West Indies, and Old World; 3 species, 2 in Venezuela, both in the flora area. Key to the Species of Samanea 1.

1.

Leaflets of longer pinnae 3–5 pairs, usually puberulent on upper surface; one petiolar nectary at base of leaf stalk; pod strongly biconvex, densely minutely velutinous; native in Bolívar state ............. S. inopinata Leaflets of longer pinnae 6–8(9) pairs, glabrous, lustrous on upper surface; petiolar nectaries between first and furthest pinna pairs, never at base of leaf stalk; interfloral bracts spatulate, with linear claw and rhombic blade 1–2.3 mm wide; pod glabrous, the broad faces of the valves depressed and nearly plane; common in northern and central Venezuela, to be expected, at least cultivated, in the flora area .................... S. saman

Stryphnodendron 673

Samanea inopinata (Harms) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 123. 1996. —Serianthes inopinata Harms, Notizbl. Bot. Gart. Berlin-Dahlem 11: 55. 1930. —Pithecellobium inopinatum (Harms) Ducke, Mem. Inst. Oswaldo Cruz 51: 426. 1953. —Albizia inopinata (Harms) G.P. Lewis, Leg. Bahia 162. 1987. Pithecellobium nuriense H.S. Irwin, Mem. New York Bot. Gard. 15: 107. 1966, “nuriensis.” Broad-crowned tree 4–11 m tall. Evergreen lowland and lower montane forests, 100–400 m; Bolívar (Altiplanicie de Nuria, lower Río Caroní and Río Paragua basins). Guyana, Brazil (discontinuously dispersed in Atlantic area from Alagoas to Bahia, cultivated in Rio de Janeiro). Samanea inopinata has long been confused with S. saman, but is consistently different in the position of the leaf nectary and in the plumply convex (in the flora area velutinous) pod.

Samanea saman (Jacq.) Merr., J. Wash. Acad. Sci. 6: 47. 1916. —Mimosa saman Jacq., Fragm. Bot. 15, t. 9. 1809 [1800]. —Pithecellobium saman (Jacq.) Benth., London J. Bot. 3: 216. 1844. —Samán. Broad-crowned tree to 45 m tall; trunk diameter to 2 m. Not yet collected from the flora area, but its presence as a cultivated or escaped tree is highly probable. ◆Fig. 557. Samanea saman is native in Central America and South America, and is adapted to either wet or seasonally dry lowland climates. The Samán is widely cultivated as a shade tree in city streets, parks, gardens, and plantations in the Neotropics and also in eastern Asia. Although frequent in northern and central Venezuela, there is no present evidence of its occurrence south of the Río Orinoco. The pulpy pods provide nutritive forage for cattle and goats. The sensitive leaves close at nightfall and during rains.

24. STRYPHNODENDRON Mart., Flora 20(2) Beibl.: 117. 1837.

by Elêna Maria de Lamare Ochionni Prostrate subshrubs, shrubs, or small to large trees. Leaves often alternate, rarely subopposite, bipinnate, small, with wart-like gland near the base of the petiole; leaflets 2–23, 0.4–12 cm long. Inflorescence of spikes, these solitary or forming groups that may be fasciculate, paniculate, or racemose; bracts always geminate; bracteoles usually shell-like (conchiform). Flowers bisexual, rarely staminate, 5merous, 2–4 mm long. Calyx usually campanulate, glabrous to villous; petals connate at base and forming a tube for 1/2–3/4 of their length, laciniate, the laciniae usually acute. Stamens 10, free or shortly fused at base; anthers elongate with a small, round, caducous, apical gland. Ovary superior, rarely short-stipitate, unicarpellate, unilocular, many-ovulate, usually elongate, glabrous to densely villous. Fruit a legume, tardily dehiscent, elongate, subturgid or compressed, mostly ferruginous, glabrous or pubescent, septate between the seeds. Seeds usually ellipsoid, disposed transversely inside the legume, to 6 mm long. Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay; 29 species, 5 in Venezuela, all of these in the flora area. Key to the Species of Stryphnodendron 1. 1. 2(1).

Leaflets ca. 30 × 15 mm; spikes arranged in well-branched panicles ........................................................................................... S. polystachyum Leaflets smaller than above; spikes solitary or clustered, but never in brached panicles .................................................................................... 2 Lower surface of leaves with tufts of hairs present along the lower half of midrib ..................................................................................................... 3

674

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

M IMOSACEAE

Lower surface of leaves without tufts of hairs along the lower half of midrib ........................................................................................................... 4 Shrub or small tree 4–8 m tall; leaves with 3 or 4 pairs of opposite pinnae; leaflets 10–16 × 7–10 mm .............................................................. S. levelii Large tree to 35 m tall; leaves with 7–10 pairs of subopposite pinnae; leaflets ca. 10 mm long ......................................................... S. microstachyum Leaflets 8–10 pairs, alternate, oblong-rectangular, 8–15 mm long; calyx and corolla glabrous ................................................................ S. guianense Leaflets 10–20 pairs, alternate or subopposite, linear, 4–7 mm long; calyx and corolla pubescent ...................................................... S. pulcherrimum

Stryphnodendron guianense (Aubl.) Benth., Trans. Linn. Soc. London 30: 374. 1875. —Mimosa guianensis Aubl., Hist. Pl. Guiane 938, t. 357. 1775. —Acacia guianensis (Aubl.) Willd., Sp. Pl. 4: 1061. 1806. —Folianthera guianensis (Aubl.) Raf., Sylva Tellur. 120. 1838. —Piptadenia guianensis (Aubl.) Benth., J. Bot. (Hooker) 4: 335. 1842. Venezuela, Guyana, French Guiana, Brazil, Peru, Bolivia; 2 subspecies, 1 in Venezuela. S. guianense subsp. guianense. —Brusquillo, Cañafistolillo, Masaguaro, Vainefá. Tree to 15(–25) m tall. Margins of gallery forests, disturbed forests, 100–400 m; Bolívar (Altiplanicie de Nuria, La Paragua). Guyana, French Guiana, Brazil. Stryphnodendron levelii R.S. Cowan, Mem. New York Bot. Gard. 10(1): 144. 1958. Shrub or small tree 4–8 m tall; flowers yellow. Riparian forests, 100–200 m; Amazonas (Caño Guazuriapana in lower Río Atabapo). Endemic. Stryphnodendron microstachyum Poepp., Nov. Gen. Sp. Pl. 3: 82. 1845. —Caro, Clavelino, Josefina. Stryphnodendron inaequale Benth., Trans. Linn. Soc. London 30: 374. 1875. Stryphnodendron purpureum Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 16. 1915. Large tree to 35 m tall. Evergreen and semideciduous lowland to lower montane forests, riparian forests, 50–900 m; Delta Amacuro (Serranía de Imataca), Bolívar

(scattered in northern part), Amazonas (Caño Cupaven por el Río Orinoco frente a San Fernando de Atabapo). Costa Rica, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 559. Stryphnodendron polystachyum (Miq.) Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 416. 1925. —Piptadenia polystachya Miq., Linnaea 18: 590. 1844. —Chincharrón, Cobija, Masaguaro. Piptadenia tocantina Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 33. 1925. Tree 15–25 m tall; sap red. Evergreen lowland and lower montane forests, 100–600 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, near El Dorado, Serranía de Imataca). Guyana, Suriname, Brazil. ◆Fig. 560. Stryphnodendron pulcherrimum (Willd.) Hochr., Bull. New York Bot. Gard. 6: 274. 1910. —Acacia pulcherrima Willd., Sp. Pl. 4: 1061. 1806. —Mimosa pulcherrima (Willd.) Poir. in Lam., Encycl. suppl. 1: 66. 1810 [1811]. Stryphnodendron floribundum Benth., J. Bot. (Hooker) 4: 343. 1842. —Stryphnodendron guianense f. floribundum (Benth.) Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 250. 1925. Stryphnodendron angustum Benth., Trans. Linn. Soc. London 30: 375. 1875. Stryphnodendron melinonis Sagot, Ann. Sci. Nat. Bot. sér. 6, 13: 322. 1882. Large tree to 30 m tall; sap red. Lower montane forests, 200–800 m; Bolívar (Cerro Camarón near base of Cerro Guaiquinima, northeast of Cerro Marutaní, upper Río Caroní). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 558.

Stryphnodendron 675

Fig. 558. Stryphnodendron pulcherrimum

Fig. 559. Stryphnodendron microstachyum

676

M IMOSACEAE

Fig. 560. Stryphnodendron polystachyum

25. ZAPOTECA H. Hernández, Ann. Missouri Bot. Gard. 73: 757. 1986. by David Brunner and Enrique Forero Unarmed erect or scandent shrubs; branches terete or 4-angled. Leaves bipinnate; pinnae in 1–many pairs; leaflets opposite, sessile, 4–many; stipules generally persistent, foliaceous. Inflorescence capitate, axillary and/or pseudopaniculate. Flowers sessile, (4)5(6)-merous. Calyx cup-shaped, dentate or denticulate; corolla campanulate or funnelform. Stamens ca. 30–60, the filaments long-exserted, white to pink or bright red apically; anthers dorsifixed, eglandular; pollen united into 16grained polyads. Ovary short-stipitate; ovules ca. 10–15; style filiform, stigma cup-

Zapoteca 677

shaped. Pods linear to oblong, membranous to coriaceous, compressed intraseminally, margins thickened; valves dehiscing elastically from the apex. Seeds ovoid to rhomboid, rarely ellipsoid, nonarillate, usually with an irregular or regular pleurogram. Southwestern U.S.A., Mexico, Central America, West Indies, South America; 17 species, 4 in Venezuela, 2 of these in the flora area. Zapoteca was described as a segregate genus from Calliandra based primarily on palynological and cytological data, Zapoteca having 16-grained polyads and x=13 whereas Calliandra has 8-grained polyads and x=8 or 11. Key to the Species of Zapoteca 1. 1.

Leaflets broadly obovate, rarely only oblong-obovate, 3–10 pairs per pinna ............................................................................................ Z. formosa Leaflets oblong, the terminal pairs somewhat obovate-oblong, generally 10 or more pairs per pinna ................................................. Z. portoricensis

Zapoteca formosa (Kunth) H. Hernández, Ann. Missouri Bot. Gard. 73: 757. 1986. —Acacia formosa Kunth, Mimoses 102, t. 32. 1819 [1822]. —Calliandra formosa (Kunth) Benth., London J. Bot. 3: 98. 1844. Shrub or small tree to 5 m tall. Southwestern U.S.A. (Arizona), Mexico, Central America, West Indies, Colombia, Venezuela, Brazil, Bolivia, Paraguay, Argentina; 6 subspecies, 1 in Venezuela.

Fig. 561. Zapoteca formosa subsp. formosa

Z. formosa subsp. formosa Calliandra marginata Griseb. ex R.O. Williams, Fl. Trinidad 1: 299. 1931. Semideciduous forests, disturbed areas, 50–300 m; Bolívar (Puerto Ordaz, islands in Lago Guri, headwaters of Río Saca, Tumeremo). Widespread along northern coast of Venezuela and dry areas of the Andes; Mexico, Central America, West Indies, Colombia, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 561.

678

M IMOSACEAE

Fig. 562. Zapoteca portoricensis subsp. flavida

Zapoteca portoricensis (Jacq.) H. Hernández, Ann. Missouri Bot. Gard. 73: 758. 1986. —Mimosa portoricensis Jacq., Collectanea 4: 143. 1790 [1791]. —Calliandra portoricensis (Jacq.) Benth., London J. Bot. 3: 99. 1844. Erect or scandent shrubs 3–7 m tall. U.S.A., eastern Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 subspecies, 1 in Venezuela.

Z. portoricensis subsp. flavida (Urb.) H. Hernández, Ann. Missouri Bot. Gard. 76: 825. 1989. —Calliandra flavida Urb., Ark. Bot. 24A(4): 4. 1931. Lowland riparian forests, near sea level to 100 m; Delta Amacuro (Río Cuyubini), northern Bolívar. Carabobo, Yaracuy; southeastern Mexico, Central America, Grenada, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 562.

26. ZYGIA P. Browne, Civ. Nat. Hist. Jamaica 279. 1756. Pithecellobium sect. Caulanthon Benth., London J. Bot. 3: 197. 1844, “Pithecolobium.” Marmaroxylon Killip, Trop. Woods 63: 3. 1940. by Rupert C. Barneby, James W. Grimes, and Paul E. Berry Unarmed, cauliflorous trees and shrubs. Leaf stalks either well-developed (Marmaroxylon group), with several pairs of pinnae and these with numerous small leaflets, or short (Zygia, s. str.) with a single pair of pinnae and few pairs of ample

Zygia 679

leaflets, in either case charged with a concave nectary; stipules small, caducous or obsolete, or sometimes small, striate and subpersistent. Inflorescence arising from previous year’s branchlets or from the trunk of earlier years, consisting of either spikes or capitula, always leafless, simply pedunculate or sometimes shortly racemose-paniculate, the sessile or subsessile flowers isomorphic. Calyx either campanulate or shallowly concave; corolla tubular or narrowly funnelform, commonly striate-veined. Filaments numerous, connate into an included or shortly exserted tube, the anthers without terminal gland. Pods linear or oblong in profile, either straight, or simply curved, or rarely twisted, laterally compressed but either flat or somewhat turgid (exceptionally cylindric), the sutures either thickened or not, the valves coriaceous or woody, the cavity ordinarily continuous; dehiscence (not seen in several) tardy, inelastic, or follicular, or through both sutures. Seeds compressed, with thin testa and no pleurogram, exarillate. Tropical lowland Mexico and Greater Antilles to northern Argentina; about 60 species, 15 in Venezuela, 11 of these in the flora area. This is the only genus of Ingeae in the flora area with both cauliflory and bipinnate leaves. Key to the Species of Zygia 1. 1. 2(1). 2.

3(2).

3.

4(3).

4.

5(1).

5.

Pinnae of each leaf exactly 1 pair, borne at apex of a petiole < 1 cm long, this sometimes reduced to the primary pulvinus of the leaf ................ 2 Pinnae of each leaf 2–8 pairs disposed along a well-developed primary rachis ......................................................................................................... 5 Leaflets of each pinna solitary, the entire leaf therefore bifoliolate, and the rachis of each pinna either obsolete or under 1 cm long ..... Z. unifoliolata Leaflets of each pinna either one pair or more often more, frequently of uneven number (3, 5, 7); rachis of each pinna much longer than 1 cm ................................................................................................................ 3 Corolla not more than 3 times as long as the deeply campanulate calyx, this 2–4.3 mm long, and the perianth glabrous overall; leaflets of each pinna 5–9; pod woody when ripe, commonly decurved or coiled, (16–) 20–30 mm wide ....................................................................... Z. inaequalis Corolla at least 3 times as long as the shortly campanulate or shallow dish-shaped calyx, this 0.5–2 mm long; perianth often (but not consistently) puberulent or hispidulous, overall or in part; leaflets of each pinna 2–5(–7) pods various in texture and width ................................. 4 Secondary veins arising from midrib of leaflets of very unequal caliber, 1 or 2 on proximal side of blade longer and stronger than succeeding ones and produced well beyond midblade .................................. Z. latifolia Secondary veins arising from midrib of leaflets of nearly equal caliber, the lowest on proximal side of blade neither stronger nor longer than more distal ones nor extending beyond midblade ........................... Z. cataractae Capitula in fascicles of 2–4 at all nodes of the pseudoraceme, each fascicle subtended by a simple bract; flowers small, the corolla 3.2–4.3 mm long; pod 3.5–7 × 0.7–1 cm ........................................................Z. racemosa Capitula solitary or seemingly fasciculate, if the latter, then the unit not subtended by a simple bract; flowers larger, the corolla 6–19 mm long; pods longer or wider, or both ................................................................. 6

680

M IMOSACEAE

6(5).

Posterior basal angle of leaflets obtuse; pod smaller and arcuate and twisted or else not known ...................................................................... 7 6. Posterior basal angle of leaflets triangular acute; pod almost straight, 12– 30 × 1.2–2 cm ......................................................................................... 8 7(6). Leaflets of larger pinnae 12–18 pairs and the largest of them ca. 15– 30 mm long; pod not seen; localized in Bolívar ............................ Z. collina 7. Leaflets of larger pinnae 7–10 pairs and the largest of them ca. 30–60 × 8– 18 mm; pod ca. 7–10 × 0.7–1 cm, arcuate or twisted, the valves tomentulose with short bronze hairs; commoner than the preceding .................................................................................................. Z. ramiflora 8(6). Stipules triangular, 0.5–2 mm long, veinless or faintly 1-veined; capitula evidently pseudoracemose, the primary axis ± extended ........ Z. basijuga 8. Stipules lanceolate or linear-lanceolate,3–19 mm long, striately (1)3–12veined; capitula solitary or seemingly fasciculate ................................ 9 9(8). Pinnae 8 or 9 pairs and the largest leaflets 7–10 mm long; pod plurilocular, the cavity interrupted between seeds by complete septa .... Z. potaroënsis 9. Pinnae 4–6 pairs and the largest leaflets 11–32 mm long; pod unilocular, lacking interseminal septa .................................................................. 10 10(9). Larger leaflets 11–18 × 3–5 mm, obtuse-mucronulate; valves of pod densely pilosulous .................................................................... Z. claviflora 10. Larger leaflets 18–32 × 5–9 mm, acute-mucronate; valves of pod glabrous ................................................................................................... Z. palustris Zygia basijuga (Ducke) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 68. 1997. —Pithecellobium basijugum Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 122. 1930, “Pithecolobium.” —Macrosamanea basijuga (Ducke) Dugand, Mutisia 9: 6. 1952. Tree 5–20 m tall; petiole reduced to a pulvinus. Evergreen lowland forests, 50–200 m; Bolívar (lower Río Caura). Colombia, Peru, Brazil. ◆Fig. 563. Zygia cataractae (H.B.K.) L. Rico, Kew Bull. 46: 496. 1991. —Inga cataractae H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 297. 1823 [1824]. —Guamo de rano, Clavellino. Inga glomerata DC., Prod. 2: 438. 1825. —Pithecellobium glomeratum (DC.) Benth., London J. Bot. 3: 213. 1844. —Zygia glomerata (DC.) Pittier, Bol. Minist. R.R. E.E. 10–12: 1927, reprinted in Trab. Mus. Com. Venez. 2: 70. 1927. Pithecellobium divaricatum Benth., London J. Bot. 3: 213. 1844. —Zygia divaricata (Benth.) Pittier, Bol. Minist. R.R. E.E. 10–12: 1927; reprinted in Trab.

Mus. Com. Venez. 2: 69. 1927. Slender tree 4–10 m tall. Seasonally flooded forests, sandy river banks, near sea level to 500 m; widely scattered in Delta Amacuro, Bolívar, and Amazonas. Apure, Guárico; widespread in Amazon basin and South American countries north of Brazil. ◆Figs. 566 and 568. Zygia cataractae is almost everywhere as frequent as Z. latifolia var. communis, which differs in asymmetric venation of the leaflets and ordinarily in narrower pods (9–15 versus 14–26 mm wide). Zygia claviflora (Spruce ex Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 65. 1997. —Pithecellobium claviflorum Spruce ex Benth., Trans. Linn. Soc. London 30: 596. 1875, “Pithecolobium.” —Abarema claviflora (Spruce ex Benth.) Kleinhoonte in Pulle, Fl. Suriname 2(2): 320. 1939. —Marmaroxylon claviflorum (Benth.) L. Rico, Kew Bull. 46: 515. 1991. —Arepillo hoja fina. Bushy treelet 2–8 m tall. Riparian forests, low forests on hills, 100–300 m; Amazonas (Río Negro, upper Río Orinoco, Río Cunucu-

Zygia 681

numa, San Carlos de Río Negro, San Pedro de Cataniapo, Yavita to Maroa). Apure; Guyana, Suriname, French Guiana, central and western Amazon basin. ◆Fig. 564. Zygia collina (Sandwith) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 80. 1997. —Pithecellobium collinum Sandwith, Kew Bull. 1948: 315. 1948. —Marmaroxylon collinum (Sandwith) L. Rico, Kew Bull. 46: 516. 1991. —Espina de pescado. Tree of medium or large size, generally 10 m tall or more. Evergreen lowland forests, 50–200 m; Delta Amacuro (Los Castillos de Guayana), Bolívar (near Upata), Amazonas (Río Sipapo). Guyana. ◆Fig. 565. Zygia inaequalis (Humb. & Bonpl. ex Willd.) Pittier, Bol. Minist. R.R. E.E. 10– 12: 1927; reprinted in Trab. Mus. Com. Venez. 2: 69. 1927. —Inga inaequalis Humb. & Bonpl. ex Willd., Sp. Pl. 4: 1019. 1806. —Pithecellobium inaequale (Humb. & Bonpl. ex Willd.) Benth., Trans. Linn. Soc. London 30: 596. 1875. Pithecellobium pilosulum Pittier, Contr. U.S. Natl. Herb. 20: 466. 1922, “Pithecollobium.” Shrub or weak treelet 2–8 m tall; epicarp of ripe fruit often cracks into short oblique slits. Riparian forests, 50–200 m; Bolívar (lower Río Orinoco basin), Amazonas (along Río Orinoco, Río Negro and Río Casiquiare basins). Apure, Zulia; Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 567. Zygia latifolia (L.) Fawc. & Rendle, Fl. Jamaica 4: 150. 1920. —Mimosa latifolia L., Syst. Nat. ed. 10: 1310. 1759. —Inga latifolia (L.) Willd., Sp. Pl. 4: 1020. 1806. —Pithecellobium latifolium (L.) Benth., London J. Bot. 3: 214. 1844. —Calliandra latifolia (L.) Griseb., Fl. Brit. W. I. 225. 1864. Shrub or tree (2–)3–15 m tall; flowers either white or pink, capitate or incipiently spicate. A polymorphic species of wide dispersal in the Neotropics (southern Mexico and Antilles, to southeastern Brazil and northeastern Bolivia); 6 varieties, 3 in Venezuela, all in the flora area.

Key to the Varieties of Z. latifolia 1. Tube of corolla pilosulous or subappressedpuberulent ..................... var. lasiopus 1. Tube of corolla glabrous, only the lobes puberulent or ciliolate ...................... 2 2. Larger leaflets typically (4–)5–12 × 2– 5 cm; pod 11–19(–20) cm wide; abundant throughout lowland flora area .................................... var. communis 2. Larger leaflets (9–)11–20 × 3.5–8 cm; pod 20–33 mm wide; not yet found in the flora area but closely approaching it in Río Orinoco delta ........... var. latifolia Z. latifolia var. communis Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 119. 1997. —?Inga cauliflora Willd., Sp. Pl. 4: 1021. 1806. —Mimosa cauliflora (Willd.) Poir. in Lam., Encycl. suppl. 1: 45. 1810. —Pithecellobium cauliflorum (Willd.) Mart., Flora 20(2) Beibl. 8: 116. 1837, “Pithecolobium.” Seasonally flooded lowland forests, swampy shores, stream banks, from near sea level ascending in gallery forests to 600 m, common and widespread in Delta Amacuro, Bolívar, and Amazonas; scattered in northern and western Venezuela; Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Braail, Bolivia, sometimes associated with the ecologically similar Z. cataractae, from which it differs in distinctive leaflet venation, as described in the key to varieties. Z. latifolia var. lasiopus (Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 120. 1997. —Pithecellobium lasiopus Benth., J. Bot. (Hooker) 2: 141. 1840. Inga ramiflora Steud., Flora 26(1): 759. 1843, non G. Don 1832, nec Pithecellobium ramiflorum Benth. 1844. Seasonally flooded forests, river banks, and understories of gallery woodlands, below 200 m; common in Delta Amacuro, Bolívar, and Amazonas. Guyana, Suriname, French Guiana, eastern Peru, Brazil (Amazon basin south to Mato Grosso). ◆Fig. 569.

682

M IMOSACEAE

Fig. 563. Zygia basijuga

Fig. 564. Zygia claviflora

Zygia 683

Fig. 565. Zygia collina

Fig. 566. Zygia cataractae

684

M IMOSACEAE

Fig. 567. Zygia inaequalis

Fig. 568. Zygia cataractae

Zygia 685

Fig. 569. Zygia latifolia var. lasiopus

Fig. 570. Zygia unifoliolata

686

M IMOSACEAE

Z. latifolia var. latifolia Seasonally flooded forests, river banks, near sea level to 100 m; Delta Amacuro (lower delta region). Southern Mexico, Central America, West Indies including Trinidad, Guyana, French guiana, Brazil (Amapá).

corolla 3.2–4.5 mm long. Evergreen lowland forests, 100–300 m; Bolívar (western edge of state near border with Amazonas), (Amazonas (upper Río Ventuari). Apure; Colombia (Amazonas), Guyana, French Guiana, Suriname, Peru (Loreto), Brazil.

Zygia palustris Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 66. 1997. Small tree 2–4 m tall. Lowland riparian forests (along black-water rivers), 100–200 m; Amazonas (between Río Baria and Río Mawarinuma, uppermost Río Yatua). Guyana (Kaieteur Plateau). Zygia potaroënsis Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 68. 1997. Small tree 2–6 m tall. Riparian forests, 100–300 m; Bolívar (upper Río Caura). Guyana (Kaieteur Plateau).

Zygia ramiflora (Benth.) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 76. 1997. —Pithecellobium ramiflorum Benth., London J. Bot. 3: 215. 1844. —Marmaroxylon ramiflorum (Benth.) L. Rico, Kew Bull. 46: 518. 1991. Pithecellobium dinizii Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 66. 1922. Treelet or slender tree 3–15 m tall; narrow tomentulose pod and sigmoid-rhombic leaflets are characteristic. Evergreen lowland forests, granitic outcrops along creeks, 100–300; Amazonas (lower Río Ventuari). Scattered through central and western Amazonian Peru and Brazil.

Zygia racemosa (Ducke) Barneby & J.W. Grimes, Mem. New York Bot. Gard. 74(1): 71. 1997. —Pithecellobium racemosum Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 272a, Corrigenda. 1922, “Pithecolobium,” a legitimate substitute for Pithecolobium racemiflorum Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 14, pl. 4. 1915, non Pithecolobium racemiflorum Donn. Sm. 1913. Trees 6–30 m tall; capitula in bracteate fascicles on an extended pseudoracemose axis; leaflets emarginate, flowers small, the

Zygia unifoliolata (Benth.) Pittier, 3rd Conf. Interamer. Agric. Caracas 259. 1945. —Pithecellobium unifoliolatum Benth., London J. Bot. 3: 212. 1844. Treelet, in habit and habitat resembling Z. cataractae, but differing especially in the simplified leaves. Riparian forests, understories in seasonally flooded forests, 100–200 m; northeastern Bolívar (upper and middle Río Orinoco), Amazonas (upper Río Negro, Río Casiquiare and tributaries). Apure, Guárico, Monagas; Panama, Colombia, Peru, Brazil. ◆Fig. 570.

MOLLUGINACEAE by Julian A. Steyermark Annual herbs, rarely shrubs. Leaves verticillate, opposite, or alternate, simple, not succulent, entire; exstipulate or stipules deciduous. Inflorescence of axillary cymes, the cymes often loose and open, or flowers solitary. Flowers actinomorphic, small, usually with a single perianth of (4)5 distinct segments. Petaloid staminodes small or absent, rarely connate-tubular (outside the flora area). Stamens (2)3–5 or more; anthers 2-locular. Ovary superior, (2)3–5-locular, rarely more, but 3–5 in the flora area; ovules numerous in each locule, campylotropous, placentation axile; styles

Glinus 687

usually distinct (solitary in Glinus). Fruit a capsule, loculicidally dehiscent and 3–5valvate in the flora area. Seeds few or numerous, reniform, with a small aril (strophiole) at the hilum, or strophiole absent; true endosperm lacking. Mainly tropics and subtropics, especially Africa; ca. 13 genera and 100 species; 2 genera and 2 species in the flora area. The family Molluginaceae has previously been included by many authors in the Aizoaceae. It is separated from the Aizoaceae by the hypogynous flowers, distinct perianth segments, and nonfleshy habit. In addition, the Molluginaceae possess anthocyanins but lack betalains, whereas the Aizoaceae have betalains but lack anthocyanins. Key to the Genera of Molluginaceae 1. 1.

Plants densely pubescent; leaves obovate, oblanceolate, or spatulate; flowers sessile or barely pedicellate; seeds strophiolate ............. 1. Glinus Plants glabrous or nearly so; leaves chiefly linear; flowers pedicellate; seeds exstrophiolate .................................................................... 2. Mollugo

1. GLINUS L., Sp. Pl. 563. 1753. Prostrate to ascending, much-branched plants, often forming mats. Leaves in unequal verticils. Flowers glomerate in leaf axils, inconspicuous. Stamens 3–5 or more. Ovary 3–5-locular; style solitary, short; stigmas 3–5. Capsules 3–5-valvate, loculicidally dehiscent. Seeds strophiolate. Tropics and subtropics of both hemispheres; ca. 10 species, 1 in Venezuela.

Fig. 571. Glinus radiatus

688

M OLLUGINACEAE

Glinus radiatus (Ruiz & Pav.) Rohrb. in Mart., Fl. Bras. 14(2): 238, t. 55, fig. l. 1872. —Mollugo radiata Ruiz & Pav., Fl. Peruv. 1: 48. 1798. Usually prostrate annual herb with stellate hairs; flowers yellowish. Sand bars and mud flats, usually along larger rivers, some-

times a weed in cotton fields, and in cut-over gallery forests, near sea level to 400 m; scattered in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Barinas, Cojedes, Monagas, Portuguesa, Sucre; southwestern U.S.A. (Texas), Mexico, Central America, Greater Antilles, South America. ◆Fig. 571.

2. MOLLUGO L., Sp. Pl. 89. 1753. Herbaceous annuals or perennials, usually much-branched. Leaves verticillate, pseudoverticillate, or basal, simple; stipules deciduous; blades entire, linear to obovate. Flowers minute. Stamens 3–10. Ovary 3–5-locular. Capsules 3–5-valvate, loculicidally dehiscent. Seeds exstrophiolate, usually numerous. Widely distributed in U.S.A., Mexico, Central America, West Indies, South America, and West Africa; ca. 15 species, 3 in Venezuela, 1 of these in the flora area. Mollugo verticillata L., Sp. Pl. 89. 1753. Annual herb, dichotomously muchbranched, prostrate or erect; leaves verticillate. Sand bars, rock outcrops, savannas, roadsides, waste ground, near sea level to

200(–400) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; U.S.A., Mexico, Central America, West Indies, South America, West Africa. ◆Fig. 572.

Fig. 572. Mollugo verticillata

Mollinedia 689

MONIMIACEAE by Paul E. Berry and Ariane Luna Peixoto Evergreen trees or shrubs, rarely semiscandent. Leaves simple, decussate or in whorls of 3 or 4, petiolate, exstipulate, glabrous or pubescent with simple or stellate, hairs; margins entire or serrate to dentate. Inflorescence axillary or cauliflorous, cymose or fasciculate. Flowers radially or bilaterally symmetric, unisexual or rarely bisexual. Receptacle well developed, subglobose to cup-shaped, with 3–many tepal lobes, these either sepaloid, petaloid, or connate into a calyptra. Staminate flowers with few to numerous stamens irregularly disposed in the floral cup, usually without basal appendages but sometimes with paired basal nectary glands on the filaments; anthers dehiscent longitudinally, transversely, or circumscissilely. Pistillate flowers with 1–many carpels, free or fused to the receptacle wall; stigmas and/or styles free or joined by mucilage plugs; ovules solitary, pendulous. Fruiting receptacle repand, with few to many sessile or stipitate drupes, or the drupes enclosed in a fleshy receptacle and exposed at maturity by the splitting of the receptacle; mesocarp fleshy, rarely half covered by an orange stylar aril; endocarp bony; endosperm abundant, oily; embryo small, straight. Tropics and subtropics, rarely in temperate regions; ca. 22 genera and nearly 200 species, 1 genus and 3 species in the flora area. This treatment follows the more limited circumscription of Monimiaceae employed by S. Renner in the treatment in Flora of Ecuador (Monimiaceae. Flora of Ecuador. Opera Botanica 59: 100–116. 1997). Siparuna, which has often been treated under a broader concept of Monimiaceae, will be treated in the our flora under the family Siparunaceae as in S. Renner (Siparunaceae. Flora of Ecuador. Opera Botanica 59: 1–99. 1997). 1. MOLLINEDIA Ruiz & Pav., Fl. Peruv. Prodr. 83, pl. 15. 1794. Dioecious scrambling shrubs or small trees, usually aromatic. Leaves decussate or in whorls of 3, membranous to coriaceous, glabrous to pilose, with simple hairs; margins entire or dentate. Cymes axillary, often formed of 3-flowered dichasia. Flowers unisexual, pedicellate and bracteate, the floral cup campanulate to obconical in staminate flowers, urceolate to utriculate in pistillate flowers, usually completely enclosing the stamens or carpels. Staminate flowers with 4 subequal tepal lobes, the 2 inner ones smaller and with a thin, terminal, inflexed appendage; stamens 8–60, irregularly dispersed on the floral cup, sessile or with short filaments; anthers with 2 pollen sacs apically confluent and opening by a single slit. Pistillate flowers with connate tepal lobes dehiscing circumscissilely as a calyptra soon after anthesis; carpels 10–50, free on the receptacle; styles very short; stigmas punctate. Fruiting receptacle repand and persistent, usually densely pubescent, fleshy when fresh and sometimes red or orange; drupes sessile or stipitate, ellipsoid, black and shiny when mature. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 20 species, 3 or 4 in Venezuela, 3 of these in the flora area.

690

M ONIMIACEAE

Key to the Species of Mollinedia 1.

1.

2(1). 2.

Branchlets and cymes densely pubescent; lower surface of leaves with hairs 0.5–2 mm long; pedicels of cymes conspicuously 4-winged in cross section ........................................................................................ M. repanda Branchlets and cymes glabrous or finely pubescent; lower surface of leaves with hairs 0.5 mm long; pedicels of cymes not conspicuously winged in cross section .......................................................................... 2 Leaves narrow, 3.5–5 cm wide, with 7 or 8 pairs of straight and closely spaced secondaries; drupes stalked, the stalks 8–12 mm long ..... M. killipii Leaves broader, > 5 cm wide, with 4–12 pairs of arcuate, loosely spaced secondaries; drupes sessile ............................................................ M. ovata

Mollinedia killipii J.F. Macbr., Candollea 5: 351. 1934. Shrub 3–6 m tall, completely glabrous; leaves narrowly elliptic, 8–14 × 3.5–5 cm, long-acuminate; drupes with stalks 8–12 mm long. Montane forests, 1000–1100 m; Amazonas (Caño Piedra near Cerro Cuao). Colombia, Ecuador, Peru, Brazil. Mollinedia ovata Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 143. 1798. —Yara-yara. Mollinedia laurina Tul., Ann. Sci. Nat. Bot. sér. 4, 3: 43. 1855. Mollinedia ptariensis Steyerm., Fieldiana, Bot. 28: 123. 1951.

Fig. 573. Mollinedia ovata

Mollinedia glabricaulis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 115. 1989. Mollinedia neblinensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 113. 1989. Shrub or tree 3–15 m tall, branchlets and lower surface of leaves finely pubescent to glabrescent; leaf blades broadly ovate, (10–) 15–22(-33) × 6–15 cm, entire or markedly toothed; drupes sessile. Evergreen lowland forests, tree islands in savannas, lower to upper montane forests and tepui slopes, 50– 1900 m; Bolívar (Altiplanicie de Nuria, Cerro Aracamuni, Cerro Venamo, southwest of El

Pakaraimaea 691

Dorado along Río Chicanán slopes of Auyántepui, El Paují, Gran Sabana, near Kavanayén, Macizo del Chimantá [Apacarátepui], base of Perai-tepui, slopes of Ptaritepui, Uaipán-tepui), Amazonas (base of Cerro Duida, Cerro Sipapo, Río Atabapo, Sierra de la Neblina). Barinas, Mérida, Sucre, Táchira, Yaracuy, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 573. Mollinedia repanda Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 142. 1798. Mollinedia yaviensis Lasser & Maguire, Brittonia 7: 79. 1950.

Mollinedia roraimensis Steyerm., Fieldiana, Bot. 28: 236. 1951. Shrub or small tree 3–12 m tall, with conspicuous brown or yellow, tomentose indumentum on branchlets, inflorescence, and lower surface of leaves; blades elliptic, 14–22 × 6.5–11 cm, with 5–7 pairs of arcuate secondaries; pedicels of cymes 4-winged in cross section; drupes sessile. Lower montane to montane forests, especially along tepui bluffs, 900–2200 m; Bolívar (Roraima-tepui), Amazonas (Cerro Coro Coro, Cerro Yaví, Río Coro Coro west of Cerro Yutajé). Colombia, Guyana, Ecuador, Peru, Bolivia.

MONOTACEAE by James S. Miller Trees, usually evergreen, with resinous wood and often with stellate or glandular hairs or peltate scales. Leaves simple, usually coriaceous; stipules well developed, often enclosing the terminal bud, usually caducous. Inflorescences axillary or terminal, paniculate or racemose, ebracteate or with small bracts. Sepals 5, imbricate or less commonly valvate, free or basally connate in a short tube, sometimes adnate to the ovary, usually enlarging and becoming wing-like in fruit. Petals 5, convolute in bud, free or basally connate, usually thick, often pubescent. Stamens (5–)10–numerous, usually 15, in 1–3 whorls; filaments free or basally connate; anthers bithecal, dehiscing longitudinally. Ovary 2–5-locular, superior, the placentation axile; style terminal, entire or shortly lobed; ovules 2–4 per locule. Fruits dry, indehiscent or tardily loculicidally dehiscent, borne with the persistent, enlarged, wing-like calyx lobes. Venezuela, Guyana, tropical Africa; 3 genera and 41 species, 1 species in Venezuela. 1. PAKARAIMAEA Maguire & P. Ashton, Taxon 26: 354. 1977. Trees to 20 m tall, often multistemmed, the branches short-tomentose, later glabrate, with prominent stipule scars. Leaves coriaceous, petiolate, with caducous, triangular stipules. Inflorescence axillary, paniculate, bracteoles small. Sepals 5, imbricate, persistent and accrescent. Petals 5, shorter than the sepals, imbricate. Stamens 40–50, free, in three whorls; the filaments inserted on a short androphore; anthers bithecal, the cells lateral on an enlarged, triangular, apical connective. Ovary 4- or 5-locular; style entire, the stigma capitate; ovules (2–)4 per locule, pendulous. Fruits indehiscent, 4- or 5-locular. Endemic to the Guayana Shield in Venezuela and Guyana; 1 species.

692

M ONOTACEAE

Pakaraimaea dipterocarpacea Maguire & P. Ashton, Taxon 26: 354. 1977. Venezuela, Guyana; 2 subspecies, 1 in Venezuela. P. dipterocarpacea subsp. nitida Maguire & Steyerm., Mem. New York Bot. Gard. 32: 308, fig. 59. 1981.

—Kanayek (Arekuna). Tree to 20 m tall; trunks in clumps of 2– many stems; flowers creamy white. Montane forests on sandy soil, 500–1100 m; Bolívar (Cerro Guaiquinima, Icabarú, upper and lower Río Paragua). Endemic. ◆ Fig. 574.

Fig. 574. Pakaraimaea dipterocarpacea subsp. nitida

M ORACEAE 693

MORACEAE by Cornelis C. Berg Trees, shrubs, or herbs (Dorstenia), usually with milky sap, dioecious, monoecious, androdioecious, or gynodioecious. Leaves alternate, in spirals or distichous; stipules fully amplexicaul to lateral, free or connate; blade basally attached, entire or pinnately incised, with pinnate venation. Inflorescences bisexual or unisexual, pedunculate or sessile, mostly bracteate, racemose, spicate, globose-capitate, with a discoid to cup-shaped receptacle, often involucrate, or with an urceolate receptacle (fig or syconium). Flowers unisexual, free or connate, or adnate to the receptacle. Staminate flowers with 2–4(–7) free or connate tepals or perianth lacking; stamens 1–4, straight or inflexed before anthesis; pistillode present or absent. Pistillate flowers with 3 or 4(–7) free or connate tepals; pistil 1; ovary 1-locular; ovule 1, apically attached; stigmas 2 or 1, filiform, band-shaped (vittiform), or tongue-shaped (linguiform). Fruit an achene or ± drupaceous, or forming a drupaceous structure with the enlarged and fleshy perianth, or forming (1–many-seeded) drupaceous or syncarpous structures. Seed large and without endosperm or small and with endosperm; cotyledons flat or thick, equal or unequal. Pantropics; 37 genera and ca. 1100 species, 14 genera and 62 species in the flora area. Two species of the Asiatic genus Artocarpus (breadfruit, fruto de pan, yaca) are commonly cultivated for their edible fruit and are present in the flora area. Artocarpus altilis (Parkinson) Fosberg has lobed, pinnatifid leaves and ramiflorous fruits, whereas A. heterophyllus Lam. has entire leaves and cauliflorous fruits. Morus alba L. is a small tree with ovate-cordate, serrate leaves that is cultivated locally in the flora area (e.g., Ciudad Bolívar, Mannetta 1, VEN; Sierra de Lema area, ca. 1300 m, Steyermark 104416, MO). The widespread Amazonian Batocarpus amazonica (Ducke) Fosberg might be discovered in the flora area. Key to the Genera of Moraceae 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

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

Herbs ............................................................................................ 4. Dorstenia Trees or shrubs ........................................................................................... 2 Stipules fully amplexicaul, leaving annular scars .................................... 3 Stipules semi-amplexicaul or lateral ......................................................... 9 Inflorescences bisexual .............................................................................. 4 Inflorescences unisexual ............................................................................ 5 Receptacle urceolate and all flowers entirely enclosed (figs); lower surface of leaves with waxy glandular spots at the base of the midrib ..... 5. Ficus Receptacle (sub)globose, covered with peltate bracts and flowers partly exposed; lower surface of leaves lacking waxy glandular spots ................................................................................................. 1. Brosimum Inflorescences not involucrate; all or most bracts peltate ......... 1. Brosimum Inflorescences involucrate; all bracts basally attached ............................ 6 Stipules connate .............................................................................. 2. Castilla Stipules free ............................................................................................... 7

694

M ORACEAE

7(6).

7.

8(7).

8.

9(2). 9. 10(9). 10.

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

Inflorescences sessile; staminate inflorescences with free stamens among concentrically arranged bracts; pistillate inflorescences 1-flowered; stigmas filiform ............................................................... 12. Pseudolmedia Inflorescences pedunculate or subsessile; staminate inflorescences with distinct flowers; pistillate inflorescences with several to many flowers, or if 1-flowered, then the stigmas tongue-shaped ................................ 8 Inner involucral bracts of staminate inflorescences broad, covering all flowers until anthesis; pistillate flowers connate and the marginal ones adnate to the receptacle, free parts of the tepals aculeate to cushion-shaped, hardened in fruit; margin of the blade always entire ............................................................................................. 10. Naucleopsis Inner involucral bracts of staminate inflorescences not covering all flowers, or if partly so, then narrow and elongate; pistillate flowers free or at anthesis basally connate; margin of the blade often dentate .................................................................................................... 11. Perebea Inflorescences bisexual ............................................................................ 10 Inflorescences unisexual .......................................................................... 12 Receptacle cylindrical or turbinate with the staminate flowers at the top of the receptacle; bracts not peltate .............................. 14. Trymatococcus Receptacle globose or discoid, or if turbinate, then (part of) the bracts peltate, the staminate flowers not confined to the top of the receptacle .............................................................................................................. 11 Stamens mostly 4, rarely 3 or 5; pistillode usually well developed ......................................................................................... 6. Helianthostylis Stamens 1 or 2, rarely 3; pistillode absent ................................. 1. Brosimum Inflorescences involucrate, mostly with a discoid to cup-shaped receptacle .............................................................................................................. 13 Inflorescences not involucrate, racemose, spicate, globose-capitate, or 1-flowered ............................................................................................. 14 Leaf blades glabrous or sparsely hairy, the upper surface smooth, usually drying greenish .......................................................................... 9. Maquira Leaf blades distinctly hairy, the upper surface often scabrous and usually drying brownish .................................................................... 7. Helicostylis Plants with staminate inflorescences [always (sub)spicate] .................. 15 Plants with pistillate inflorescences ........................................................ 17 Flowers indistinct (variable numbers of floral parts, tepals, and stamens, mixed with interfloral bracts); stamens < 4 ............................... 3. Clarisia Flowers distinct; stamens 4 ..................................................................... 16 Filaments of the stamens longer than the perianth; spines often present .................................................................................................... 8. Maclura Filaments of the stamens shorter than the perianth; spines absent .................................................................................................... 13. Sorocea Stigmas 2 and unequally long or 1; spines often present on branches in young specimens ........................................................................ 8. Maclura Stigmas 2 and equally long; spines absent ............................................. 18 Inflorescences borne in leaf axils, racemose or subcapitate ......... 13. Sorocea Inflorescences mostly borne on older wood, 1-flowered or capitate ..................................................................................................... 3. Clarisia

Brosimum 695

1. BROSIMUM Sw., Prodr. 12. 1788, nom. cons. Piratinera Aubl., Hist. Pl. Guiane 888. 1775, nom. rejic. Brosimopsis S. Moore, Trans. Linn. Soc. London, Bot. 4: 473. 1895. Trees, monoecious or dioecious. Leaves distichous; stipules fully amplexicaul and connate or semi-amplexicaul and free; blades entire. Inflorescences in the leaf axils, bisexual or unisexual, pedunculate; receptacle (sub)globose, hemispheric, turbinate, or convexly discoid, at first completely covered with peltate bracts. Staminate flowers several to numerous, borne on the surface of the receptacle; stamens 1–4; pistillode absent. Pistillate flowers 1–several, embedded in the receptacle; ovary adnate to the perianth; stigmas 2, filiform to band-shaped. Fruit(s) forming a drupaceous structure with the enlarged, fleshy, and often colored receptacle. Seed large, without endosperm; cotyledons thick, equal or unequal. Mexico, Central America, Cuba, Jamaica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, introduced in Trinidad; 15 species, 7 in Venezuela, all in the flora area. Key to the Species of Brosimum 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5. 6(5). 6.

Stipules semi-amplexicaul and free (subgenus Brosimum) ..................... 2 Stipules fully amplexicaul and connate (subgenus Ferolia) ..................... 4 Midribs on upper surface of leaf blades impressed; stipule venation prominent, subflabellate-furcate .................................................. B. lactescens Midribs on upper surface of leaf blades plane; stipule venation inconspicuous, if visible, then only a single midrib ....................................... 3 Lower surface of leaf blades (sub)glabrous; inflorescences unisexual ................................................................................................ B. alicastrum Lower surface of leaf blades ± hairy; inflorescences bisexual ... B. guianense Areoles of the lower surface of the leaf blades densely white-pubescent ..................................................................................................... B. potabile Areoles of the lower surface of the leaf blades not clearly hairy ................. 5 Epidermis of the petiole flaking off ..................................................... B. utile Epidermis of the petiole not flaking off ..................................................... 6 Stipules (0.5–)1–2.5(–4) cm long; leaf blades usually equal-sided ............. .................................................................................................. B. rubescens Stipules 0.3–0.7 cm long; leaf blades distinctly unequal-sided .................. ................................................................................... B. melanopotamicum

Brosimum alicastrum Sw., Prodr. 12. 1788. Tree to 35 m tall. Mexico, Central America, Cuba, Jamaica, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, introduced in Trinidad; 2 subspecies, 1 in Venezuela. B. alicastrum subsp. bolivarense (Pittier) C.C. Berg, Acta Bot. Neerl. 19: 326. 1970. —Helicostylis bolivarensis Pittier, Contr. U.S. Natl. Herb. 20: 96. 1918. —Brosimum bolivarense (Pittier) Ro-

mero, Mutisia 33: 2. 1970. —Chara, Guaimaro, Misionero negro. Brosimum latifolium Standl., Trop. Woods 42: 26. 1935. Dioecious tree to 35 m tall. Evergreen and deciduous forests, 50–500 m; Bolívar (Altiplanicie de Nuria, El Dorado, Puerto Ordaz, lower Río Caura and Río Paragua, Río Toro, east of Túriba). Widespread elsewhere in Venezuela; Panama, Colombia, Guyana, Ecuador, Peru, Brazil (Acre, Roraima), Bolivia, introduced in Trinidad. ◆Fig. 577.

696

M ORACEAE

Brosimum guianense (Aubl.) Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 337. 1909. —Piratinera guianensis Aubl., Hist. Pl. Guiane 888. 1775. —Æræ æphi (Piaroa), Almidón, Charo blancho, Charo macho, Erenpi, Juacha (Yekwana), Majomo, Manichi, Mato Palo. Brosimum palmarum Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 158. 1937. Brosimum rotundatum Standl., Bull. Torrey Bot. Club. 75: 293. 1948. Monoecious tree to 30 m tall, usually with buttress roots. Semidedicuous to evergreen and riparian forests, secondary vegetation, 50–1000 m; Delta Amacuro (Río Toro), northern Bolívar, Amazonas (widespread). Anzoátegui, Barinas, Mérida, Miranda, Sucre, Zulia; Mexico, Belize, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 576. The fruits of Brosimum guianense are edible. Brosimum lactescens (S. Moore) C.C. Berg, Acta Bot. Neerl. 19: 326. 1970. —Brosimopsis lactescens S. Moore, Trans. Linn. Soc. London, Bot. 4: 473. 1895. —Æphí (Piaroa), Caraashiji (Yanomami), Charo, Charo colorado, Charo macho, Guaímero, Misionero negro, Sajurua (Yekwana), Yuquito. Dioecious tree to 40 m tall. Nonflooded forests, 100–200 m; scattered throughout most of Bolívar and Amazonas. Apure, Barinas, Mérida, Miranda, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 575. The fruits, which are produced in the rainy season, are edible. Brosimum melanopotamicum C.C. Berg, Acta Bot. Neerl. 19: 327. 1970. —Sajurua, Shada (Yekwana), Takuramo. Dioecious tree to 20 m tall. Nonflooded forests, 100–200 m; Amazonas (upper Río Cuao, Río Cunucunuma, Río Mawarinuma, Río Negro, Yavita to Maroa road). Guyana, Brazil (Amazonas). The wood of Brosimum melanopotamicum is used locally to make bows for hunting. Brosimum potabile Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 26. 1922. —Majagua.

Brosimum myristicoides Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 157. 1937. Tree to 30 m tall. Nonflooded forests, 100– 200 m; Amazonas (near San Fernando de Atabapo, Yavita to Maroa road). Colombia, Ecuador, Peru, Brazil. Brosimum rubescens Taub., Bot. Jahrb. Syst. 12(Beibl. 27): 4. 1890. —Cajimán, Fuiyu (Yekwana), Guanacaste, Marima, Palo de Brasil, Wadimashu (Yekwana), Wãrãke duthæ (Piaroa), Wishoguatemosi (Yanomami). Brosimum paraense Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 6: 67. 1910. Brosimum lanciferum Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 24. 1922. Monoecious tree to 40 m tall. Nonflooded forests, 100–200(–1200) m; Bolívar (El Cácaro between Río Caura and Río Paragua, Río Caruay, lower Río Caura), Amazonas (Culebra, La Esmeralda, Río Casiquiare, middle Río Cataniapo, upper Río Manapiare basin, Río Mawarinuma, upper Río Orinoco, Río Padamo, Yavita to Maroa road). Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Branches of Brosimum rubescens are carved and used as handles, the latex is used to plug holes in boats, and the fruits are eaten by capuchin monkeys, parrots, and insects. Brosimum utile (H.B.K.) Pittier, Contr. U.S. Natl. Herb. 20: 102. 1918. —Galactodendrum utile H.B.K., Nov. Gen. Sp. (quarto ed.) 7. 163. 1825. Monoecious tree to 50 m tall. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 7 subspecies, 3 in Venezuela, 2 of these in the flora area. Key to the Subspecies of B. utile 1. Leaf margins repand; indument sparse; inflorescences usually geminate; peduncle 25–35 mm long; perianth none .............................. subsp. longifolium 1. Leaf margins not repand; indument variable; inflorescences usually solitary; peduncle 1–12 mm long; perianth 0.1–0.3 mm tall ................. subsp. ovatifolium

Brosimum 697

Fig. 575. Brosimum lactescens

Fig. 576. Brosimum guianense

Fig. 577. Brosimum alicastrum subsp. bolivarense

698

M ORACEAE

B. utile subsp. longifolium (Ducke) C.C. Berg, Acta Bot. Neerl. 19: 328. 1970. —Brosimum longifolium Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 27. 1922. —Marima. Nonflooded forests, 100–200 m; Amazonas (San Carlos to Solano). Colombia, Ecuador, Peru, Brazil. B. utile subsp. ovatifolium (Ducke) C.C. Berg, Acta Bot. Neerl. 19: 328. 1970. —Brosimum ovatifolium Ducke, Arch.

Jard. Bot. Rio de Janeiro 3: 27. 1922. —Dayacabi (Baré), Marima. Lowland forests, 100–600 m; widely scattered in Bolívar and western and southwestern Amazonas. Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. Piaroa Amerindians strip the bark of Brosimum utile subsp. ovatifolium and pound it into a cloth for carrying loads and for ceremonial outfits; edible caterpillars are also found on the trees. The inner bark of saplings is twisted and used for bow strings.

Fig. 578. Castilla elastica subsp. elastica

Clarisia 699

2. CASTILLA Sessé in Cerv., Gaz. Lit. Mexico (suppl.) 7. 1794. Castilloa auct. non Endl. 1837. Trees, dioecious or monoecious, with self-pruning lateral branches. Leaves distichous on the lateral branches; stipules fully amplexicaul, connate; blades with an entire or dentate margin. Inflorescences on short spurs in the leaf axils, unisexual, involucrate. Staminate inflorescences pedunculate, with a flabellate and bivalvate receptacle (or those accompanying pistillate inflorescences funnel- to cupshaped and entire to 2-lobed) with stamens solitary or in pairs on radial and branched ridges of the receptacle and among free or connate interstaminal “bracts.” Pistillate inflorescences solitary (or accompanied by staminate inflorescences), sessile, with a discoid to cup-shaped receptacle. Flowers several to many, free or connate. Ovary adnate to the perianth; stigmas 2, band-shaped. Fruiting perianth enlarged, fleshy, yellow to dark orange; fruit adnate to the perianth. Seed large, without endosperm; cotyledons thick and equal. Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia, possibly naturalized in other countries; 3 species, 1 in Venezuela. Two species of Castilla yield rubber: C. ulei Warb., which is widely distributed in the Amazon Basin, and C. elastica, which occurs as a native from Mexico to western Ecuador. Members of this genus are also cultivated as shade trees in cacao or coffee plantations. Castilla elastica Sessé in Cerv., Gaz. Lit. Mexico (suppl.) 7. 1794. Tree to 30 m tall, with low buttresses and abundant, white latex. Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, possibly cultivated or naturalized in the flora area; 3 subspecies, 2 cultivated in Venezuela, 1 of these in the flora area.

C. elastica subsp. elastica. —Caucho. Lower montane forests, 500–600 m; Bolívar (Cerro Tomasote northwest of Santa Cruz). Cultivated or possibly naturalized in parts of northern Venezuela; Mexico, Central America, cultivated or naturalized elsewhere. ◆Fig. 578.

3. CLARISIA Ruiz & Pav., Fl. Peruv. Prodr. 128. 1794, nom. cons. Sahagunia Liebm., Mexic. Halvgr. 128: 1850, preprinted from Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 5, 2: 316. 1851. Acanthinophyllum Allemão, Revista Brazil. 1: 210. 1857. Trees or shrubs, dioecious. Leaves distichous; stipules lateral, free; blades entire or pinnately incised. Inflorescences on leafless branchlets or spurs on the older wood, pedunculate, bracteate. Staminate inflorescences spicate; flowers indistinct, tepals 2–7, free or connate; stamens 1–3; pistillode absent. Pistillate inflorescences capitate or 1-flowered; perianth tubular, 4-lobed; ovary adnate to the perianth; stigmas 2, filiform or tongue-shaped. Fruiting perianth enlarged, fleshy, red or orange; fruit adnate to the perianth. Seed large, without endosperm; cotyledons thick and equal. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, all in the flora area. Key to the Species of Clarisia 1. 1.

Margin and/or apex of the blade spinulose, midrib prominent above .................................................................................................... C. ilicifolia Margin and apex of the blade not spinulose, midrib slightly impressed above ....................................................................................................... 2

700

M ORACEAE

Fig. 579. Clarisia ilicifolia

Fig. 580. Clarisia racemosa

Dorstenia 701

2(1). 2.

Inflorescences borne in the leaf axils or on leafless branchlets just below the leaves; uncinate hairs usually present .................................. C. biflora Inflorescences borne on leafless branchlets on the older wood; uncinate hairs absent .............................................................................. C. racemosa

Clarisia biflora Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 255. 1798. Tree to 35 m tall. Riparian forests, 300– 400 m; Amazonas (upper Río Orinoco). Barinas, Mérida, Zulia; Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. Clarisia ilicifolia (Spreng.) Lanj. & G. Rossberg, Recueil Trav. Bot. Néerl. 33: 717. 1936. —Exoecaria ilicifolia Spreng., Neue Entd. 2: 117. 1821. —Acanthinophyllum ilicifolia (Spreng.) W. Burger, Ann. Missouri Bot. Gard. 49: 27. 1962. —Cafecillo, Marima. Acanthinophyllum strepitans Allemão, Revista Brazil. 1: 368. 1858. —Clarisia strepitans (Allemão) Lanj., Recueil Trav. Bot. Néerl. 33: 272. 1936. Clarisia spruceana Lanj., Recueil Trav. Bot. Néerl. 33: 272. 1936. —Acanthino-

phyllum spruceana (Lanj.) W. Burger, Ann. Missouri Bot. Gard. 49: 30. 1962. Shrub or small tree, sometimes to 20 m tall. Lowland forest understory, 50–600 m; widely scattered in Bolívar and Amazonas. Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 579. The fruits of Clarisia ilicifolia are eaten raw or cooked by Piaroa Amerindians. Clarisia racemosa Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 255. 1798. —Billo, Cajimán. Tree to 40 m tall, often with buttress roots. Nonflooded forests, 100–300 m; Delta Amacuro (Río Toro), Bolívar (Río Canaracuni and Río Caura, Serranía de Imataca, near Upata), Amazonas (Isla Ratón). Mexico, Nicaragua, Costa Rica, Panama, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 580.

4. DORSTENIA L., Sp. Pl. 121. 1753. Treelets, shrubs, subshrubs, or herbs (and then usually perennial, rarely annual), monoecious, non-succulent or ± succulent (and with or without tubers), branched or unbranched. Stems mainly supraterraneous to entirely subterraneous, internodes long, both long and short, or only short; latex white, sometimes yellow or watery; indument mostly consisting of unicate hairs. Leaves in spirals (rosulate in plants with stems with short internodes), sometimes (sub)distichous, petiolate; blade basally attached or peltate, entire or pinnately to palmately or pedately lobed to parted; margin usually dentate to crenate; venation pinnate to subpalmate or in peltate leaves almost radiate, brochidodromous to reticulate; stipules subfoliaceous, subulate, 1–many-veined, persistent, sometimes caducous, mostly firmly coriaceous. Inflorescences solitary, in pairs, or several together on minute short-shoots in the leaf axils, pedunculate; receptacle discoid to turbinate to cup-shaped, orbicular, elliptic, quadrangular, (irregularly) lobed to stellate, naviculate, bifurcate or ligulate, green, yellow, purplish, or reddish brown, at least in part, with the bracts on the outer surface of the receptacle ± dispersed or in 1–several, marginal to submarginal rows, sometimes on appendages, or ebracteate and then mostly with the marginal and/or submarginal appendages short and tooth-like, subulate, long and spathulate, or ribbon-shaped; usually more or less protogynous, occasionally unisexual, flowerless border (fringe) between flower-bearing part and the edge of the receptacle narrow to broad, often with minute, globose to club-shaped or conical, hyaline, purplish, or reddish brown hairs on the perianth, among the flowers, and on the fringe. Flowers connate, staminate ones among the pistillate ones or concentrated in or confined to the periphery of the flowering face, pedicellate, the pedicels connate

702

M ORACEAE

and adnate to the perianth of the pistillate flowers; tepals (1)2 or 3(4), (almost) free. Stamens (1)2 or 3(4), epitepalous, inflexed in bud, pistillode sometimes present. Pistillate flowers sessile, perianth tubular, only the upper, entire or 2- or 3-lobed part free from the surrounding flowers, ovary free; stigmas filiform to band-shaped, 2 and then often unequal in length or 1. Fruit a drupe or drupelet, usually dehiscent, often ± stipitate, exocarp white and turgid-fleshy, smooth or tuberculate, usually pushing outward and ejecting the endocarp. Seed large or small, testa thin, with a small, slightly thickened, vascularized part below the hilum; endosperm present or absent; embryo with flat, equal cotelydons and a relatively long radical, or with thick ± unequal cotyledons and a small radicle. Neotropics, Africa, and adjacent parts of Asia; ca. 105 species, 3 in Venezuela, 1 of these in the flora area. Dorstenia brasiliensis Lam., Encycl. 2: 317. 1786. —Tusilla. Dorstenia tubicina Ruiz & Pav., Fl. Peruv. 1: 65. 1798. Dorstenia sabanensis Cuatrec., Bol. Soc. Venez. Ci. Nat. 17: 93. 1956. Small herb to 10 cm tall, with subterranean rhizomatous stem. Savannas, often in rocky places near streams, 50–400 m; widespread in northern Bolívar, Amazonas (near Puerto Ayacucho). Anzoátegui, Carabobo, Guárico, Monagas; Trinidad, Guyana, French Guiana, Peru, southern Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 581. Dorstenia brasiliensis is sometimes used medicinally.

Fig. 581. Dorstenia brasiliensis

5. FICUS L., Sp. Pl. 1059. 1753. Urostigma Gasp., Nov. Gen. Fic. 9. 1844. Pharmacosycea Miq., London J. Bot. 6: 525. 1847. Trees, shrubs, or climbers, often with adventitious aerial roots (in hemiepiphytes and root climbers), monoecious or (gyno)dioecious (at least functionally staminate and pistillate); uncinate hairs rare, waxy glandular spots, usually present on leaves at the base of the midrib, in the axils of the basal or other lateral veins, in the main furcations of the venation on the lower surface, or at the nodes of leafy twigs. Leaves alternate, in spirals or distichous, (sub)opposite or sometimes subverticillate; stipules fully amplexicaul to lateral, mostly free. Inflorescences with an urceolate receptacle (syconium, fig) with a narrow circular or slit-shaped orifice (ostiole), bracts on the peduncle (peduncular bracts), subtending the receptacle, mostly 2 or 3 (basal bracts), on the outer surface of the receptacle (lateral bracts), in

Ficus 703

the orifice of the receptacle (ostiolar bracts), among the flowers (interfloral bracts), and/or subtending (staminate) flowers (bracteoles). Flowers bisexual (with staminate flowers and pistillate flowers with styles of different length) or (functionally) unisexual, either with staminate flowers and (nonseed-producing) pistillate flowers with short styles or with long-styled pistillate flowers and neuter flowers, pronouncedly protogynous. Staminate flowers sessile or pedicellate, with 2–5 (or more) tepals, free to almost fully connate, stamens 1–5, pistillode absent or present. Pistillate flowers sessile or pedicellate, with 3–5 (or more) free to fully connate tepals, ovary free; styles different in length; stigmas 2 and filiform or 1 and filiform or clavate to funnelform. Fruits drupelets or achenes, small. Seed with endosperm, embryo with flat and equal cotyledons. Pantropics; ca. 720 species, 48 native species in Venezuela, 33 of these in the flora area. Numerous species of Ficus have been introduced and are commonly cultivated in Venezuela and the flora area. Among them are F. carica L., the cultivated fig (higo), a shrub or small tree with 3–5-lobed leaves, and F. pumila L., a small-leaved, climbing species often cultivated to cover outside walls. The main introduced tree species found in gardens, parks, and along roads are F. benjamiana L., F. drupacea Thunb., F. elastica Roxb., F. lyrata Warb., F. microcarpa L. f., and F. religiosa L. Ficus gracilis was reported by Pittier (Bol. Soc. Venez. Ci. Nat. 9: 119. 1944, type Williams 15070a). However, this has been redetermined as Ilex divaricata Mart. ex Reiss. (Aquifoliaceae). Key to the Species of Ficus 1. 1.

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

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

Figs solitary in the leaf axils; basal bracts 3; stamens 2; stigmas 2; aerial roots absent and plants always terrestrial (subg. Pharmacosycea) ..... 2 Figs in pairs in the leaf axils or several together on minute spurs also below the leaves; basal bracts 2; stamen 1; stigma usually 1; aerial roots usually present and plants often hemiepiphytic (subg. Urostigma) ................................................................................................................ 6 Epidermis of the petiole flaking off; lower surface of leaf blades and figs often scabrous or scabridulous ................................................... F. maxima Epidermis of the petiole not flaking off; lower surface of leaf blades and figs not scabrous or scabridulous .......................................................... 3 Lateral veins ca. 20–30 pairs .................................................. F. sphenophylla Lateral veins 8–18 pairs ............................................................................ 4 Stipules 1–2.5 cm long, drying brown, often puberulent ............. F. piresiana Stipules 2–13 cm long, drying greenish, usually glabrous ....................... 5 Figs when dry 1–1.8 cm diameter and with a protruding apex; stipules 2–4(–7) cm long; petioles 0.8–2.5 cm long .............................. F. yoponensis Figs when dry 1.5–3(–4) cm diameter and usually without a protruding apex; stipules (3.5–)5–10(–13) cm long; petioles (1–)3–8 cm long ..................................................................................................... F. insipida Figs on spurs, also below the leaves and then often > 2 together ............ 7 Figs mostly confined to the leaf axils, mostly in pairs ............................ 10 Lower surface of leaf blades with dense brown hairs ........... F. albert-smithii Lower surface of leaf blades glabrous or with sparse minute brown hairs ................................................................................................................ 8

704

M ORACEAE

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

9.

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

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

20.

Stipules 2–8 cm long; leaf blades ≥ 20 cm long, narrowly obovate ................................................................................................... F. caballina Stipules 0.3–1(–2) cm long; leaf blades < 16 cm long, or, if to 22 cm long, then elliptic or oblong ............................................................................ 9 Leaf blades usually < 10 cm long, mostly (ob)lanceolate to (sub)obovate; petioles to 1(–1.8) cm long, ca. 1.5 mm thick; figs when dry 0.3–0.4 cm diameter; peduncles to 0.2(–0.3) cm long ................................ F. mathewsii Leaf blades usually 10–20 cm long and mostly elliptic to oblong; petioles ≤ 3 cm long and 2–3 mm thick; figs when dry 0.3–1.2 cm diameter, and/or peduncles 0.2–0.8 cm long ...................................................... F. guianensis Lower surface of leaf blades ± conspicuously hairy, at least at the midrib, or all veins (minutely) puberulent ...................................................... 11 Lower surface of leaf blades glabrous or almost so, at most sparsely and minutely puberulent on the midrib ..................................................... 23 Figs pedunculate (peduncles at least 0.2 cm long if the receptacle is at least 1 cm diameter, or to 0.1 cm long if the receptacle is to 1 cm diameter) ........................................................................................................ 12 Figs sessile or subsessile .......................................................................... 16 Indumentum consisting of short hairs of equal length; stipules puberulent ........................................................................................................ 13 Indumentum consisting of hairs of different lengths; stipules (sub)hirsute .............................................................................................................. 14 Leaf blades 2.5–4 cm wide, apex acuminate to acute; peduncles (0.1–)0.2– 0.4 cm long ...................................................................... F. donnell-smithii Leaf blades (2–)4.5–10.5 cm wide, apex rounded; peduncles 0.5–1.7 cm long ........................................................................................... F. pakkensis Lateral veins (7–)11–16 pairs; figs when dry (1–)1.5–2.5 cm diameter ................................................................................................. F. gomelleira Lateral veins 4–8(–10) pairs; figs when dry 0.8–1.3 cm diameter ......... 15 Petioles 0.5–2(–5) cm long, hirsute to hirtellous .................... F. malacocarpa Petioles (1.5–)2–7 cm long, minutely puberulent ...................... F. hebetifolia Upper surface of leaf blades scabrous ..................................................... 17 Upper surface of leaf blades smooth ....................................................... 19 Lateral veins (8–)10–15 pairs; ostiole surrounded by a 3-lobed rim .................................................................................................. F. matiziana Lateral veins 4–8(–10) pairs; ostiole without a rim or surrounded by an entire rim ............................................................................................. 18 Figs when dry 0.6–0.7 cm diameter, subglabrous ..................... F. dendrocida Figs when dry 0.8–1 cm diameter, usually hirtellous to subhirsute ............................................................................................. F. malacocarpa Ostiole surrounded by a 3-lobed rim ....................................................... 20 Ostiole surrounded by an entire rim or rimless ...................................... 21 Stipules 1–1.8 cm long, striate, often subpersistent; leaf blades with tertiary venation reticulate to scalariform and ± prominent on the lower surface .......................................................................... F. panurensis Stipules 0.3–1(–3) cm long, not striate, caducous; leaf blades with tertiary venation for the greater part scalariform, the lower surface usually plane .............................................................................................. F. trigona

Ficus 705

21(19). Lower surface of leaf blades and figs (minutely) puberulent ................ ......................................................................................... F. donnell-smithii 21. Lower surface of leaf blades and figs more pronouncedly hairy ............ 22 22(21). Figs when dry 0.4–0.6 cm diameter, densely tomentose to sublanate ................................................................................................... F. mollicula 22. Figs when dry 0.8–1.2 cm diameter, hirtellous to subhirsute .................... ............................................................................................. F. malacocarpa 23(10). Figs when dry ≤ 1 cm diameter ............................................................... 24 23. Figs when dry > 1 cm diameter ............................................................... 32 24(23). Figs sessile ..................................................................................... F. paraensis 24. Figs pedunculate (peduncles ≥ 0.1 cm long) ............................................ 25 25(24). Ostiole ± sunken ...................................................................................... 26 25. Ostiole not sunken, flat or slightly prominent ........................................ 27 26(25). Ostiole deeply sunken into the ± crateriform apex of the receptacle; figs when dry 0.4–0.8 cm diameter; peduncles 3–4 mm long; leaf blades usually elliptic to oblong .............................................................. F. pertusa 26. Ostiole slightly impressed; figs 0.3–0.4(–0.5) cm diameter; peduncles 1– 2 mm long; leaf blades lanceolate ....................................... F. schumacheri 27(25). Figs and stipules minutely puberulent .............................. F. donnell-smithii 27. Figs and stipules (sub)glabrous ............................................................... 28 28(27). Bases of leaf blades emarginate, with a small notch slightly wider than the petiole .................................................................................... F. krukovii 28. Bases of leaf blades acute, obtuse, rounded, truncate, or (broadly) subcordate ............................................................................................ 29 29(28). Figs when dry 0.4–1.2 cm diameter; peduncles 0.2–1 cm long .............. 30 29. Figs when dry 0.3–0.4(–0.5) cm diameter; peduncles 0.1–0.2 cm long .............................................................................................................. 31 30(29). Figs when dry 0.4–1 cm diameter; petioles 0.5–2.5(–5) cm long; leaf blades mostly 5–10 cm long ............................................................... F. amazonica 30. Figs when dry (0.8–)1–1.2 cm diameter; petioles (2–)4–10 cm long; leaf blades mostly 10–20 cm long ......................................................... F. eximia 31(29). Leaf blades lanceolate, to 9 cm long; mature figs greenish with red spots .............................................................................................. F. schumacheri 31. Leaf blades elliptic to oblong or (sub)obovate, to 4.5 cm long; mature figs red or pink ............................................................................... F. americana 32(23). Figs pedunculate ...................................................................................... 33 32. Figs sessile or subsessile (peduncles ≤ 0.2 cm long) ............................... 38 33(32). Ostiole surrounded by a triangular (or circular) rim ....................... F. crocata 33. Ostiole not surrounded by a rim .............................................................. 34 34(33). Ostiole sunken into the crateriform apex of the receptacle ................... 35 34. Ostiole not sunken, flat or prominent ..................................................... 36 35(34). Leaf blades mostly ≤ 10 cm long, coriaceous to subcoriaceous; figs when dry 0.4–0.8 cm or 1–1.6 cm diameter .......................................... F. pertusa 35. Leaf blades mostly 10–20 cm long, ± thickly coriaceous; figs when dry 1.6– 2 cm diameter ......................................................................... F. broadwayi 36(34). Basal bracts to 4 mm long; leaf blades acuminate to subcaudate at apex ................................................................................................... F. paraensis 36. Basal bracts 5–10 mm long; leaf blades rounded at apex ....................... 37

706

M ORACEAE

37(36). 37. 38(32). 38. 39(38). 39. 40(38). 40. 41(40). 41. 42(41). 42. 43(41). 43. 44(43). 44.

Leaf blades acute to obtuse at base ............................................ F. obtusifolia Leaf blades cordate (to rounded) at base ................................. F. catappifolia Petioles (2–)4–12(–20) cm long; figs depressed-globose .......................... 39 Petioles to 4 cm long, if longer, then the figs usually ellipsoid and/or with a prominent ostiole ................................................................................. 40 Basal bracts 3–4 mm long; figs when dry 0.8–1.2 cm diameter ... F. paludica Basal bracts 5–8 mm long; figs when dry 1.5–2 cm diameter ........................................................................................... F. nymphaeifolia Ostiole surrounded by a triangular (or circular) rim ....................... F. crocata Ostiole not surrounded by a rim .............................................................. 41 Basal bracts to 4 mm long, and/or leaf blades acuminate to subcaudate at apex ...................................................................................................... 42 Basal bracts 4–8 mm, and/or leaf blades rounded at apex ..................... 43 Ostiole plane; petioles 0.5–1.5 cm long ....................................... F. tepuiensis Ostiole prominent; petioles 1–4.5(–7) cm long ............................. F. paraensis Leaf blades cordate (to rounded) at base ................................. F. catappifolia Leaf blades acute to obtuse at base ......................................................... 44 Petioles 0.5–1.5 cm long; figs (depressed-)globose ...................... F. tepuiensis Petioles (1–)2–4 cm long; figs usually longer than broad .......... F. obtusifolia

Ficus albert-smithii Standl., Lloydia 2: 174. 1939. —Matapalo. Tree to 15 m tall. Riparian forests, forests bordering savannas, upland forests, 100– 1400 m; Delta Amacuro (Río Toro area), Bolívar (Cerro Guaiquinima, Cerro Venado, Gran Sabana), Amazonas (Cerro Sipapo, Piedra Cocuy, Río Pasimoni, Sierra de la Neblina). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 582. Ficus amazonica (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 298. 1867. —Urostigma amazonicum Miq., London J. Bot. 6: 541. 1847. —Higuillo, Matapalo, Mutumutu. Urostigma angustifolium Miq., London J. Bot. 6: 539. 1847. —Ficus angustifolia (Miq.) Miq., Ann. Mus. Bot. LugdunoBatavum 3: 298. 1867. Tree to 20 m tall. Dry forests, riparian forests, swamps, near sea level to 500 m; Delta Amacuro (scattered), northern Bolívar. Widespread elsewhere in Venezuela (except the Llanos); Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 584. Ficus americana Aubl., Hist. Pl. Guiane 952. 1775. —Matapalo. Ficus perforata L., Pl. Surin. 171. 1775. Ficus jacquiniifolia A. Rich. in Sagra,

Hist. Phys. Cuba, Bot. Pl. Vasc. 3: 221. 1850. Tree to 20 m tall. Evergreen lowland to montane forests, 100–2200 m; Bolívar (Auyán-tepui, Gran Sabana, Ilú-tepui), Amazonas (Caño Iguapo, Caño San Antonio, middle Río Pasimoni, lower Río Sipapo). Apure, Guárico, Zulia; Central America, West Indies, Colombia, Guyana, Ecuador, Brazil. ◆Fig. 587. Ficus americana is part of a species complex with F. guianensis and F. mathewsii, both in the flora area, as well as F. maitin Pittier in the coastal mountain and Andean region of Venezuela. Ficus americana is essentially a lowland taxon changing in its features, such as the dimensions of the blade, from the West Indies through Central America to western South America. Material with features similar to that of the Greater Antilles is confined to high altitudes in the flora area. The western South American form (with larger leaves) is represented in Zulia at low altitudes. Ficus broadwayi Urb., Repert. Spec. Nov. Regni Veg. 15: 110. 1917. Ficus mendelsonii Britton, Bull. Torrey Bot. Club 48: 330. 1921 [1922]. Ficus savannarum Standl., Bull. Torrey Bot. Club 75: 298. 1948. Tree to 15 m tall. Evergreen forests, 100–

Ficus 707

900 m; Bolívar (16 km northwest of Kilómetro 88, Santa Elena de Uairén). Anzoátegui, Sucre; Trinidad, Guyana, Suriname, French Guiana, Brazil (Bahia, Roraima).

Ocamo). Apure, Barinas; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Acre, Amapá), Bolivia. ◆Fig. 583.

Ficus caballina Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 13(2): 301. 1937. —Lechero, Mutumutu. Ficus tamatamae Pittier, Bol. Soc. Venez. Ci. Nat. 8: 260. 1943. Tree to 25 m tall. Riparian and evergreen forests, near sea level to 500 m; Delta Amacuro (widespread), northeastern Bolívar, Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 590.

Ficus eximia Schott in Spreng., Syst. Veg. 4(cur. post.): 410. 1827. —Simure. Ficus expansa Pittier, Bol. Soc. Venez. Ci. Nat. 4: 63. 1937. Ficus glandulosa Pittier, Bol. Soc. Venez. Ci. Nat. 4: 70. 1937. Ficus guanarensis Pittier, Bol. Soc. Venez. Ci. Nat. 4: 71. 1937. Ficus turbinata Pittier, Bol. Soc. Venez. Ci. Nat. 4: 61. 1937. Ficus foveolata Pittier, Bol. Soc. Venez. Ci. Nat. 7: 133. 1941. Ficus foveata Pittier, Bol. Soc. Venez. Ci. Nat. 8: 258. 1943. Tree to 15 m tall. Forests, 100–900 m; Delta Amacuro (Serranía de Imataca), Amazonas (Río Asisa, San Carlos de Río Negro, Sierra Parima). Widespread in northern and western Venezuela; Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. In Panama, Ficus eximia gradually merges into F. citrifolia Mill. from the West Indies and Central America. It is distinguished by its smaller leaves and figs. Ficus eximia may eventually be found to be just a subspecies of the rather complex F. citrifolia, perhaps also including F. amazonica as a subspecies.

Ficus catappifolia Kunth & Bouché in A. Braun & Bouché, Index Seminum (Berlin) 14. 1846 [1847]. Tree to 15 m tall. Flooded riparian forests, swamp forests, near sea level to 100 m; Delta Amacuro (Río Cuyubini), Bolívar (south of Altiplanicie de Nuria). Guyana, Suriname, French Guiana, Brazil (Amapá, Pará, Roraima). Ficus crocata (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 297. 1867. —Urostigma crocatum Miq., London J. Bot. 6: 531. 1847. —Matapalo. Tree to 20 m tall. Dry forests, 50–800 m; Bolívar (Ciudad Bolívar, Guasipati, upper Río Túriba). Widespread elsewhere in Venezuela; Mexico, Central America, Greater Antilles, Trinidad, Tobago, Ecuador, Peru, Brazil. Ficus crocata is very closely related to Ficus gomelleira, and some populations have features transitional to the latter. Ficus dendrocida H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 46. 1817. Tree to 20 m tall. Riparian and secondary forests, near sea level to 100 m; Delta Amacuro (Caño Macareo). Widespread elsewhere in Venezuela outside the flora area; Panama, Colombia, Brazil (Pará). Ficus donnell-smithii Standl., Contr. U.S. Natl. Herb. 20: 21. 1917. —Lechero. Tree to 25 m tall. Riparian forests, 100– 600 m; Bolívar (Río Canaracuni, Río Supamo), Amazonas (Caño Colorado 55 km southeast of Puerto Ayacucho, lower Río

Ficus gomelleira Kunth & Bouché in A. Braun & Bouché, Index Seminum (Berlin) 18. 1846 [1847]. —Higuerón, Higuerote, Higuito, Kanwae (Panare). Tree to 25(–40) m tall. Varied forest types, 50–400 m; Delta Amacuro (Caño Atoiba north of Caño Araguao, Caño Güiniquina), northernmost and eastern Bolívar, Amazonas (Isla Ratón, Macuruco to Santa Bárbara, Río Yureba). Anzoátegui, Apure, Miranda, Yaracuy; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 594. Ficus gomelleira is very closely related to F. crocata. These two taxa are sometimes difficult to tell apart and may prove to be distinct only at the subspecific level. Ficus guianensis Desv. ex Ham., Prodr. Pl. Ind. Occid. 62. 1825. —Ducuajo

708

M ORACEAE

(Yekwana), Matapalo, Oí Pharatæ (Piaroa), Tuædu Pharatæ (Piaroa). Ficus corpulenta Pittier, Bol. Soc. Venez. Ci. Nat. 8: 257. 1943. Ficus maroana Pittier, Bol. Soc. Venez. Ci. Nat. 9: 120. 1944. Ficus erratica Standl., Bull. Torrey Bot. Club 75: 295. 1948. Ficus mensalis Standl., Bull. Torrey Bot. Club 75: 297. 1948. Tree to 35 m tall. Varied forest types, especially in savannas and forests bordering savannas, near sea level to 2200 m; Delta Amacuro (scattered), Bolívar (widespread), Amazonas (widespread). Widespread elsewhere in Venezuela; St. Vincent, Grenada, Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Ficus guianensis is very variable with regard to the shape of the blade and dimensions of figs and blade. It is a complex of taxa that includes F. americana and F. mathewsii. High-altitude populations often have thick, coriaceous leaves. Ficus hebetifolia Dugand, Caldasia 1: 50. 1942. Tree to 20 m tall. Lowland forests, 100– 200 m; Amazonas (15 km southeast of San Fernando de Atabapo). Colombia, French Guiana, Brazil. Ficus insipida Willd., Sp. Pl. 4: 1143. 1806. —Higuerón, Higuito, Matapalo. Ficus glabrata H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 47. 1817. Tree to 40 m tall. Widely distributed in Mexico, Central America, Lesser Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 2 subspecies, both in the flora area. Ficus insipida is morphologically variable and probably ecologically diverse as well. Several infraspecific taxa might be (and have been) distinguished, one of them (subsp. scabra) being quite distinct. Key to the Subspecies of F. insipida 1. Leaf blades smooth on the upper surface; leaf blades, petioles, and stipules mostly glabrous .................... subsp. insipida 1. Leaf blades scabrous on the upper surface; lower surface of leaf blades, petioles, and stipules hairy ............... subsp. scabra

F. insipida subsp. insipida Ficus longistipula Pittier, Bol. Soc. Venez. Ci. Nat. 4: 53. 1937. Mostly in riparian forests, 100–200 m; Bolívar (Río Toro). Common elsewhere in Venezuela; Central America, Lesser Antilles, Colombia, Trinidad, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 589. F. insipida subsp. scabra C.C. Berg, Acta Amazonica 14(suppl.): 201. 1986. Open, often riparian forests, 100–400 m; Delta Amacuro (Serranía de Imataca), Bolívar (El Dorado, Isla Anacoco, Serranía de Imataca). Colombia (Meta), Guyana, Suriname, French Guiana, Brazil (Pará). Ficus krukovii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 171. 1937. Tree to 25 m tall. Forests, 100–200 m; Amazonas (Río Mawarinuma). Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 597. Ficus malacocarpa Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 172. 1937. —Matapalo, Palo vieja. Ficus scabrida Pittier, Bol. Soc. Venez. Ci. Nat. 4: 68. 1937. Tree to 10 m tall. Various types of forests, near sea level to 200 m; Delta Amacuro (scattered), northeastern Bolívar, Amazonas (scattered). Aragua, Cojedes, Lara, Mérida, Yaracuy; Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 588. Ficus mathewsii (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 298. 1867. —Urostigma mathewsii Miq., London J. Bot. 6: 549. 1847. —Dukuajo (Yekwana), Phõi pharatæ (Piaroa), Shiigui. Ficus gleasonii Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 170. 1937. Tree to 35 m tall. Forests, 50–1100 m; northwestern Bolívar, Amazonas (widely scattered). Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 593. The dried leaves of the typical form of Ficus mathewsii have a very smooth upper surface of the blade with the secondary veins hardly visible. More eastward in the flora area and Guyana, the secondary (and often also the tertiary) venation is more conspicuous in dried leaves. The latter form is often difficult to distinguish from F. guianensis.

Ficus 709

Ficus matiziana Dugand, Caldasia 4: 116. 1946. Tree to 10 m tall. Forests, 100–200 m; Amazonas (Pamoni, Río Cataniapo, Río Mawarinuma, San Carlos de Río Negro). Colombia, Guyana, Brazil, Bolivia. Ficus maxima Mill., Gard. Dict. ed. 8., Ficus no. 6. 1768. —Matapalo, Mutumutu. Ficus radula Willd., Sp. Pl. 4: 1144. 1806. Ficus parkeri Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 300. 1867. Tree to 25 m tall. Riparian, marshy, or secondary forests, near sea level to 1000 m; widely scattered in Delta Amacuro, northern Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, Greater Antilles, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. Ficus mollicula Pittier, Bol. Soc. Venez. Ci. Nat. 7: 134. 1941. Ficus tigrensis Pittier, Bol. Soc. Venez. Ci. Nat. 7: 135. 1941. Shrub or tree to 6 m tall. Granitic outcrops, 50–200 m; Bolívar (Río Cuchivero, Río Parguaza, Serranía Baraguán), Amazonas (between Puerto Ayacucho and Samariapo). Apure; Colombia (Vichada). ◆Fig. 591. Ficus nymphaeifolia Mill., Gard. Dict. ed. 8, Ficus no. 9. 1768. —Higuerote, Matapalo, Mutumutu. Ficus urbaniana Warb. in Urb., Symb. Antill. 3: 459. 1903. Tree, mostly to 30 m tall, sometimes much taller. Various types of forests, 50–300 m; Delta Amacuro (scattered), northernmost Bolívar, Amazonas, (near Puerto Ayacucho). Widespread elsewhere in Venezuela; Central America, Lesser Antilles, Trinidad, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 596. Ficus obtusifolia H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 49. 1817. Urostigma gardnerianum Miq., London J. Bot. 6: 527. 1847. —Ficus gardneriana (Miq.) Miq., Ann. Mus. Bot. LugdunoBatavum 3: 297. 1867. Urostigma involutum Liebm., Mexic. Halvgr. 132: 1850; preprinted from Kongel. Danske Vidensk. Selsk. Skr.,

Naturvidensk. Math. Afd. ser. 5, 2. 320. 1851. —Ficus involuta (Liebm.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 298. 1867. Ficus pascuorum Pittier, Bol. Soc. Venez. Ci. Nat. 4: 54. 1937. Tree to 25 m tall. Forests, 100–200 m; Amazonas, (Río Manapiare basin). Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. Ficus pakkensis Standl., Bull. Torrey Bot. Club 75: 287. 1948. —Kanwae (Panare), Wajuko (Yekwana). Shrub or tree to 15 m tall. Semideciduous forests, savannas, 100–800 m; Bolívar, (south of Altiplanicie de Nuria, Río Caura, Río Maniapure, Río Túriba). Guyana, Suriname, French Guiana, Brazil. Ficus paludica Standl., Bull. Torrey Bot. Club 75: 298. 1948. —Dan-anatu (Warao). Urostigma leucostictum Miq., London J. Bot. 6: 535. 1847. —Ficus leucosticta (Miq.) Miq., Ann. Mus. Bot. LugdunoBatavum 3: 297. 1867. Tree to 20 m tall. Flooded riparian forests, near sea level to 50 m; Delta Amacuro, (near Curiapo, Río Cuyubini). Guyana, Suriname, French Guiana, Brazil (Amapá, Amazonas). Ficus panurensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 174. 1937. —Dukaja. Ficus maguirei Standl., Bull. Torrey Bot. Club 75: 296. 1948. Tree to 20 m tall. Forests, thickets on granitic outcrops, near sea level to 800 m; Delta Amacuro (Caño Arature), northern Bolívar, Amazonas (Cacurí, near Puerto Ayacucho, Raudal Picure in Río Cunucunuma, near San Fernando de Atabapo). Apure, Cojedes, Lara, Miranda, Táchira, Zulia; Guyana, Suriname, French Guiana, Brazil (Amazonas). Ficus paraensis (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 1867. —Urostigma paraense Miq., London J. Bot. 6: 534. 1847. —Masjrata (Warekena), Matapalo, Sesejudi (Yekwana), Wanawadudu (Yekwana). Ficus hydrophila Pittier, Bol. Soc. Venez. Ci. Nat. 8: 259. 1943.

710

M ORACEAE

Ficus orinocensis Pittier, Bol. Soc. Venez. Ci. Nat. 8: 259. 1943. Ficus arukensis Standl., Bull. Torrey Bot. Club 75: 295. 1948. Tree to 20 m tall. Common, often along rivers, near sea level to 900 m; Delta Amacuro (Caño Araguao, Serranía de Imataca), Bolívar (near Kilómetro 88, Puerto Ordaz, Río Caura, Salto Pará, near Santa María de Erebato, base of Sororopán-tepui), Amazonas (widely scattered). Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 592. Ficus pertusa L. f., Suppl. Pl. 442. 1781. —Matapalo, Mutumutu. Urostigma erythrostictum Miq., London J. Bot. 6: 540. 1847. —Ficus erythrosticta (Miq.) Miq., Ann. Mus. Bot. LugdunoBatavum 3: 298. 1867. Urostigma geminum Miq., London. J. Bot. 6: 547. 1847. —Ficus gemina (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 298. 1867. Ficus palmicida Pittier, Bol. Soc. Venez. Ci. Nat. 4: 69. 1937. Ficus kanukuensis Standl., Lloydia 2: 174. 1939. Tree to 20 m tall. Riparian, montane, and nonflooded lowland forests, granitic domes, near sea level to 900 m; Delta Amacuro (Caño Atoiba), Bolívar (El Palmar, Jabillal on lower Río Caura), Amazonas (Sierra Parima, between Río Baría and Río Mawarinuma). Widespread elsewhere in Venezuela; Mexico, Central America, Jamaica, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 595. Ficus pertusa is represented by two forms: (a) with small figs (0.4–0.8 cm diameter when dry) and (b) with large figs (1–1.6 cm diameter) and often relatively large leaves. Ficus piresiana Vázq. Avila & C.C. Berg, Acta Amazonica 14(suppl.): 207. 1986. Large tree. Riparian and montane forests, 100–1300 m; Amazonas (Río Baría, Sierra de la Neblina). Colombia, French Guiana, Ecuador, Peru, Brazil (Acre, Amapá, Pará), Bolivia. Ficus schumacheri (Liebm.) Griseb., Fl. Brit. W. I. 151. 1859. —Urostigma schumacheri Liebm., Mexic. Halvgr. 140:

1850; preprinted from Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. ser. 5, 2. 328. 1851. —Matapalo, Mutumutu (Warao). Tree to 25 m tall. Riparian and swamp forests, lower montane forests, near sea level to 600 m; Delta Amacuro (widespread), Bolívar (Altiplanicie de Nuria, El Palmar, Río Botanamo, Río Venamo), Amazonas (Raudal Maveri near the mouth of Río Uotamo). Falcón, Sucre; Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 585. Ficus sphenophylla Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 176. 1937. Tree to 15 m tall. Forests, 100–800 m; Bolívar (near Túriba), Amazonas (San Carlos de Río Negro). Colombia, Ecuador, Peru, Brazil, Bolivia. Ficus tepuiensis C.C. Berg & Simonis, Ernstia 6: 8. 1981. Tree to 20 m tall. Montane forests, Bonnettia forests, 1000–1800 m; Bolívar, (Auyán-tepui, Cerro Venamo, Macizo del Chimantá [Abacapá-tepui], Sierra de Maigualida). Endemic. ◆Fig. 586. Ficus trigona L. f., Suppl. Pl. 411. 1781. —Higuillo. Ficus fagifolia (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 299. 1867. Ficus ernstiana Pittier, Bol. Soc. Venez. Ci. Nat. 4: 55. 1937. Ficus plicato-ostiolata Pittier, Bol. Soc. Venez. Ci. Nat. 8: 258. 1943. Tree to 15 m tall. Usually on periodically flooded riverbanks, near sea level to 500 m; Delta Amacuro (Caño Araguao, Caño Arature, La Margarita to Puente Miranda), northeastern Bolívar (base of Guy-tepui near Perai-tepui, 6 km from Maniapure, San Pedro de las dos Bocas), Amazonas (Río Baría, along Río Orinoco, San Carlos de Río Negro). Guárico, Mérida, Táchira, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Ficus yoponensis Desv., Ann. Sci. Nat. Bot. sér. 2, 18: 310. 1842. Tree to 20 m tall. Montane forests, ca. 1000 m; Bolívar (Cerro Uroi in Río Chicanán basin). Widespread elsewhere in Venezuela; Mexico, Central America, Colombia, Trinidad, Tobago, Ecuador, Peru.

Ficus 711

Fig. 582. Ficus albert-smithii

Fig. 583. Ficus donnell-smithii

712

M ORACEAE

Fig. 584. Ficus amazonica

Fig. 585. Ficus schumacheri

Fig. 586. Ficus tepuiensis

Ficus 713

Fig. 587. Ficus americana

Fig. 588. Ficus malacocarpa

Fig. 589. Ficus insipida subsp. insipida

714

M ORACEAE

Fig. 590. Ficus caballina

Fig. 591. Ficus mollicula

Fig. 592. Ficus paraensis

Ficus 715

Fig. 593. Ficus mathewsii

Fig. 594. Ficus gomelleira

716

M ORACEAE

Fig. 595. Ficus pertusa

Fig. 596. Ficus nymphaeifolia

Helianthostylis 717

Fig. 597. Ficus krukovii

Fig. 598. Helianthostylis steyermarkii

6. HELIANTHOSTYLIS Baill., Adansonia 11: 299. 1875. Trees, androdioecious. Leaves distichous; stipules semiamplexicaul, free; blade entire. Inflorescences borne in the leaf axils, bisexual or staminate, globose, hemispherical, or turbinate, pedunculate, bracteate. Staminate flowers (a few in bisexual, numerous in staminate inflorescences) superficial; perianth 3- or 4-lobed to

718

M ORACEAE

3- or 4-fid; stamens (3)4(5); pistillode well developed. Pistillate flower 1, embedded in the receptacle; ovary adnate to the perianth; stigmas 2, filiform. Fruit forming a drupaceous structure with the enlarged, fleshy, yellowish receptacle. Seed large, without endosperm; cotyledons thick and equal. Colombia, Venezuela, Ecuador, Peru, Brazil; 2 species, 1 in Venezuela. Helianthostylis steyermarkii C.C. Berg, Acta Bot. Neerl. 21: 99. 1972. Tree to 15 m tall. Nonflooded forests on white sand, 100–200 m; Amazonas (Río

Casiquiare, road between Solano and San Carlos de Río Negro, between Yavita and Pimichín). Brazil (Amazonas). ◆Fig. 598.

7. HELICOSTYLIS Trécul, Ann. Sci. Nat. Bot. sér. 3, 8: 134. 1847. Trees, dioecious or monoecious, with self-pruning lateral branches. Leaves distichous on the lateral branches; stipules not fully amplexicaul; blade entire or dentate. Inflorescences borne on short spurs in the leaf axils, unisexual, discoidcapitate, involucrate. Staminate inflorescences ≤ 15 together, pedunculate, receptacle (at least before anthesis) with revolute margin; inner involucral bracts not covering all flowers until anthesis; tepals 4, at least basally connate; stamens (2–)4. Pistillate inflorescences solitary or in pairs, sometimes accompanied by 1 or 2 staminate ones, sessile or pedunculate; flowers several to numerous; tepals 4(–6), at least basally connate, the 2 inner ones cohering on the inner surface with entangled thin hairs; ovary almost free; stigmas 2, filiform or band-shaped and then often twisted. Infructescences discoid to subglobose. Fruiting perianth enlarged, fleshy, yellow to orange; fruit basally adnate to the perianth. Seed large, without endosperm; cotyledons thick and equal. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 7 species, 3 in Venezuela, all in the flora area. Key to the Species of Helicostylis 1. 1.

2(1).

2.

Leaf blades scabrous on both surfaces, the acumen short and obtuse; tepals of staminate flower free; stigmas filiform and straight .... H. scabra Leaf blades usually smooth on the upper surface and always on the lower surface, the acumen elongate and subacute; tepals of staminate flower distinctly connate; stigmas band-shaped and usually twisted ............. 2 Leaf blades usually lanceolate, mostly 10–15 cm long, hairs on the lower surface appressed on the main veins; pistillate inflorescences sessile ..................................................................................................... H. elegans Leaf blades usually oblong, mostly 15–25 cm long, hairs on lower surface spreading on the main veins; pistillate inflorescences often pedunculate ....................................................................................... H. tomentosa

Helicostylis elegans (J.F. Macbr.) C.C. Berg, Acta Bot. Neerl. 18: 464. 1969. —Perebea elegans J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 63. 1931. Helicostylis lancifolia Ducke, Bull. Mus. Hist. Nat. (Paris) sér. 2, 4: 721. 1932. Tree to 25 m tall. Nonflooded forests, ca. 100–400 m; Bolívar (upper Río Caura), Ama-

zonas (Yavita to Maroa road). Colombia, Ecuador, Peru, Brazil. Helicostylis scabra (J.F. Macbr.) C.C. Berg, Acta Bot. Neerl. 18: 464. 1969. —Pseudolmedia scabra J.F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 11: 62. 1931. —Lengua de tigre, Palo de morrocoy.

Maclura 719

Fig. 599. Helicostylis tomentosa

Helicostylis asperifolia Ducke, Trop. Woods 31: 11. 1932 Tree to 30 m tall. Nonflooded forests, 100– 200 m; Amazonas (Río Cunucunuma, Río Mawarinuma, San Carlos de Río Negro). Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil. Helicostylis tomentosa (Poepp. & Endl.) Rusby, Mem. Torrey Bot. Club 6: 120. 1896. —Olmedia tomentosa Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 32. 1838. —Aijiadu (Yekwana), Cajimán, Charo macho, Charo peludo, Maskedek (Pemón), Pata de Morrocoy, Shopa (Ya-

nomami), Surja, Yoeru (Piaroa). Olmedia polycephala Pittier, Bol. Soc. Venez. Ci. Nat. 7: 306. 1942. Tree to 30 m tall; latex yellowish. Nonflooded forests, 100–1000 m; Delta Amacuro (Río Toro), Bolívar, (south of El Manteco, Kavanayén, east-northeast of El Palmar, Río Canaracuni, Río Caura, Río Karún, Río Tabaro near Río Nichare, near Upata), Amazonas (widely scattered). Monagas, Táchira, Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 599. The fruit of Helicostylis tomentosa is edible.

8. MACLURA Nutt., Gen. N. Amer. Pl. 2: 233. 1818, nom. cons. Chlorophora Gaudich., Voy. Uranie 509. 1826 [1830]. Climbers, trees, or shrubs (sometimes only in juvenile specimens) armed with straight to curved, reduced branchlets ending in a spinose tip. Leaves chartaceous to subcoriaceous, when dry often brittle, petiole relatively thin; stipules laterally free, mostly very small. Staminate inflorescences globose, (sub)capitate, spicate, or (sub)racemose; stamens straight or inflexed in the bud. Pistillate inflorescences globose-capitate; flowers free or connate; ovary free or the lower part adnate to the perianth, stigmas 2, mostly strongly unequal in length, or one; tepals and interfloral

720

M ORACEAE

bracts mostly accumulating yellow dye in 2 (or more) immersed “glands.” Seeds small to rather large, endosperm present (but scarce) or absent; embryo various, mostly with (rather) thin, folded or plane cotyledons and a long radicle. Pantropics and northern temperate regions; 9 species, 2 in Venezuela, 1 of these in the flora area. Maclura tinctoria (L.) Steud., Nomencl. Bot. ed. 2, 2: 87. 1841 [1840]. —Morus tinctoria L., Sp. Pl. 986. 1753. —Chlorophora tinctoria (L.) Gaudich. ex Benth. & Hook. f., Gen. Pl. 3: 363. 1880. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 2 subspecies, 1 in Venezuela. M. tinctoria subsp. tinctoria. —Mora. Tree to 20 m tall; trunk spiny, with white latex. Semideciduous forests bordering savannas, secondary forests, 50–300 m; Bolívar (Puerto Ordaz, Río Asa at Raudal Cotúa). Widespread elsewhere in Venezuela and the Neotropics south to Argentina. ◆Fig. 600.

Fig. 600. Maclura tinctoria subsp. tinctoria

9. MAQUIRA Aubl., Hist. Pl. Guiane 2(suppl.) 36. 1775. Olmediophaena H. Karst., Bot. Jahrb. Syst. 8: 375. 1887. Olmedioperebea Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 3. 1922. Trees, dioecious or monoecious, with self-pruning lateral branches. Leaves distichous on the lateral branches; stipules fully amplexicaul, free; blade entire. Inflorescences borne on short spurs in the leaf axils, unisexual, discoid-capitate, involucrate. Staminate inflorescences mostly several together, pedunculate; inner involucral bracts not covering all flowers until anthesis; flowers several to numerous; perianth 4-lobed to 4-parted; stamens (3)4; pistillode absent. Pistillate inflorescences mostly solitary, subsessile or pedunculate; flowers many and free, few and connate, or solitary; perianth 4-lobed to 4-fid; ovary adnate to the perianth; stigmas 2, tongue-shaped or filiform. Fruiting perianth enlarged, fleshy, orange; fruit adnate to the perianth. Seed large, without endosperm; cotyledons thick and equal. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 4 species, all in the flora area.

Maquira 721

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

3.

Leaf blades with the tertiary venation predominantly parallel ................. ............................................................................................. M. sclerophylla Leaf blades with the tertiary venation reticulate ..................................... 2 Leaf blades usually > 10 cm long, mostly strongly unequal-sided; peduncle of staminate inflorescence bracteate; stigmas filiform ................. M. coriacea Leaf blades mostly > 10 cm long, usually slightly unequal-sided; peduncle of staminate inflorescence without bracts; stigmas tongue-shaped .... 3 Lower surface of leaf blades mostly slightly scabrous; margins of leaf blades callose; pistillate flowers few and connate, or solitary; staminate inflorescences 0.3–0.8 cm diameter ....................................... M. calophylla Lower surface of leaf blades mostly smooth; margins of leaf blades not or hardly callose; pistillate inflorescences with several to many free flowers; staminate inflorescence 0.8–1.5 cm diameter ........ M. guianensis

Maquira calophylla (Poepp. & Endl.) C.C. Berg, Acta Bot. Neerl. 18: 464. 1969. —Olmedia calophylla Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 32. 1838. —Olmedioperebea calophylla (Poepp. & Endl.) Ducke, Arq. Serv. Florest. 1: 14. 1939. —Marima. Tree to 35 m tall. Periodically flooded forests, 100–200 m; Amazonas (Culebra, near Puerto Ayacucho, Río Mawarinuma, Río Ocamo, Río Putaco, between San Carlos de Río Negro and Solano). Colombia, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 602.

name, French Guiana, Ecuador, Peru, Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of M. guianensis 1. Involucral bracts sparsely whitish-puberulent; peduncles 0–8 mm long; fruiting perianth not or hardly ribbed, mostly subglabrous .......... subsp. costaricana 1. Involucral bracts densely brown-puberulent; peduncles 5–20 mm long; fruiting perianth ribbed, brown-velutinous ......... ............................... subsp. guianensis

Maquira coriacea (H. Karst.) C.C. Berg, Acta Bot. Neerl. 18: 464. 1969. —Pseudolmedia coriacea H. Karst., Fl. Columb. 2: 21, t. 3. 1862. Olmedia obliqua Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 337. 1909. —Olmediophaena obliqua (Huber) Ducke, Arq. Serv. Florest. 1: 12. 1939. Olmedia maxima Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 32. 1922. —Olmediophaena maxima (Ducke) Ducke, Arq. Serv. Florest. 1: 13. 1939. Tree to 30 m tall. Usually in periodically flooded riparian forests, 50–100 m; Bolívar (Ciudad Bolívar, Río Nichare), Amazonas (Capibara). Apure, Barinas, Guárico, Monagas; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 601. Maquira guianensis Aubl., Hist. Pl. Guiane 2(suppl.): 36, t. 389. 1775. Tree to 25 m tall. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suri-

Fig. 601. Maquira coriacea

722

M ORACEAE

Fig. 602. Maquira calophylla

Fig. 603. Maquira guianensis subsp. guianensis

M. guianensis subsp. costaricana (Standl.) C.C. Berg, Novon 6: 235. 1996. —Perebea costaricana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 390. 1937. —Maquira costaricana (Standl.) C.C. Berg, Acta Bot. Neerl. 18: 463. 1969. Tree to 35 m tall. Dry forested slopes, forest edges, 400–700 m; Bolívar (5 km west of Amaruay-tepui, Cerro Abismo, near El Paují, Río Acanán). Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. M. guianensis subsp. guianensis. —Manichi (Yekwana).

Perebea laurifolia Trécul, Ann. Sci. Nat. Bot. sér. 3, 8: 133. 1847. Tree to 25 m tall. Upland riparian forests, 200–1400 m; Bolívar (Río Caura, Río Cuyuní, Sierra de Lema). Zulia; Guyana, Suriname, French Guiana, Brazil (Maranhão, Matto Grosso, Pará). ◆Fig. 603. Maquira sclerophylla (Ducke) C.C. Berg, Acta Bot. Neerl. 18: 463. 1969. —Olmedioperebea sclerophylla Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 34. 1922. Tree to 25 m tall. Nonflooded forests, ca. 100 m; Amazonas (upper Río Orinoco). Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará).

Perebea 723

10. NAUCLEOPSIS Miq. in Mart., Fl. Bras. 4(1): 120. 1853. Ogcodeia Bureau in A. DC., Prodr. 17: 286. 1873. Acanthospaera Warb., Verh. Bot. Vereins Prov. Brandenburg 48: 150. 1907. Palmolmedia Ducke, Arq. Serv. Florest. 1: 20. 1939. Trees, dioecious, with self-pruning lateral branches. Leaves on the lateral branches distichous; stipules fully amplexicaul, free; blades entire, pinnately veined. Inflorescences on minute spurs in the leaf axils, unisexual, discoid-capitate. Staminate inflorescences often as many as 4 together, pedunculate or subsessile, marginal involucral bracts covering the flowers until anthesis; tepals 0–8, often varying in number, mostly 4, free or basally connate; stamens 1–4. Pistillate inflorescences solitary, sessile or subsessile; flowers several to numerous (rarely one), connate and the peripheral ones adnate to the receptacle; tepals (3)4–6(–10), the free parts aculeate to pyramidal to cushion-shaped, often dislocated (not around the style); free parts of the perianth hardened, often spine-like; ovary adnate to the perianth; stigmas 2, filiform, band-shaped, or tongue-shaped. Infructescences discoid to subglobose. Fruits embedded in the pulpy receptacle. Seed large, without endosperm; cotyledons thick and equal. Neotropics; 22 species, 1 in Venezuela. Additional species of Naucleopsis are expected to be reported from the flora area. Naucleopsis oblongifolia (Kuhlm.) Carauta, Albertoa 3: 262. 1994. —Ogcodeia oblongifolia Kuhlm., Anais Reunião SulAmer. Bot. 3: 77. 1939. Brosimum mello-barretoi Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 70.

1940. —Naucleopsis mello-barretoi (Standl.) C.C. Berg, Acta Bot. Neerl. 18: 465. 1969. Tree to 30 m tall. Nonflooded forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Colombia, Ecuador, Peru, Brazil. ◆Fig. 604.

Fig. 604. Naucleopsis oblongifolia

11. PEREBEA Aubl., Hist. Pl. Guiane 953. 1775. Noyera Trécul, Ann. Sci. Nat. Bot. sér. 3, 8: 135. 1847. Trees or shrubs, dioecious or monoecious, with self-pruning lateral branches. Leaves distichous on the lateral branches; stipules fully amplexicaul, free; blades entire, often with a dentate margin. Inflorescences on short spurs in the leaf axils,

724

M ORACEAE

Fig. 605. Perebea guianensis subsp. guianensis

unisexual, discoid-capitate, involucrate. Staminate inflorescences several together, pedunculate; inner involucral bracts not covering all flowers until anthesis; flowers numerous; sepals 4, free or basally connate; stamens 4; pistillode absent. Pistillate inflorescences solitary or accompanied by staminate ones, subsessile or pedunculate; flowers several to numerous, free or basally connate; perianth 4-lobed to 4-fid; ovary free or partly adnate to the perianth; stigmas 2, tongue-shaped or

Pseudolmedia 725

filiform. Fruiting perianth enlarged, fleshy, reddish; fruit free or partly adnate to the perianth. Seed large, without endosperm; cotyledons thick and equal. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 9 species, 3 in Venezuela, 1 of these in the flora area. Perebea guianensis Aubl., Hist. Pl. Guiane 953, t. 361. 1775. Tree to 20 m tall; leaves 20–50 cm long. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 5 subspecies, 1 in Venezuela.

P. guianensis subsp. guianensis Tree 8–14 m tall. Nonflooded forests, 100– 200 m; Amazonas (Río Baría, Río Mawarinuma). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 605.

12. PSEUDOLMEDIA Trécul, Ann. Sci. Nat. Bot. sér. 3, 8: 129. 1847. Trees dioecious, with self-pruning lateral branches. Leaves distichous on the lateral branches; stipules fully amplexicaul, free; blades entire. Inflorescences borne on short spurs in the leaf axils, discoid-capitate, involucrate, sessile. Staminate inflorescences usually 2–4 together; inner involucral bracts covering all flowers until anthesis; perianth absent; stamens free, among concentrically arranged interfloral bracts; pistillode absent. Pistillate inflorescences usually 2 together; perianth tubular, 4-dentate; ovary adnate to the perianth; stigmas 2, filiform. Fruiting perianth enlarged, fleshy, reddish; fruit adnate to the perianth. Seed large, without endosperm; cotyledons thick and equal. Southern Mexico, Central America, Greater Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; 9 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Pseudolmedia 1. 1.

Branchlets and stipules with equally long, appressed hairs; terminal buds slender ....................................................................................... P. laevigata Branchlets and stipules with spreading hairs, distinctly different in length; terminal buds ± swollen ..................................................... P. laevis

Pseudolmedia laevigata Trécul, Ann. Sci. Nat. Bot. sér. 3, 8: 131. 1847. —Catamajaca rebalsera, Fruto de Paloma, Palo de Brasil. Tree to 30 m tall. Evergreen lowland to montane forests, 100–1300 m; Bolívar (Cerro Ichún, Cerro Venamo, Gran Sabana, Río Canaracuni, near Túriba, Uaipán-tepui), Amazonas (scattered). Zulia; Panama, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 606. Pseudolmedia laevis (Ruiz & Pav.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 16. 1931. —Olmedia laevis Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 258. 1798. —Charo, Charo peludo. Pseudolmedia multinervis Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 10: 189. 1927.

Fig. 606. Pseudolmedia laevigata

726

M ORACEAE

Olmedia caurensis Pittier, Bol. Soc. Venez. Ci. Nat. 8: 306. 1943. Tree 5–30 m tall. Nonflooded forests, 100– 400(–900) m; Delta Amacuro (Río Toro), Bolívar (scattered in northern part, Santa María de Erebato), Amazonas (scattered). Apure, Barinas, Monagas, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 607.

Fig. 607. Pseudolmedia laevis

13. SOROCEA A. St.-Hil., Mém. Mus. Hist. Nat. 7: 473. 1821. Pseudosorocea Baill., Adansonia 11: 296. 1875. Paraclarisia Ducke, Arq. Serv. Florest. 1: 2. 1939. Dioecious shrubs or trees. Leaves distichous; stipules lateral, free; blade with an entire or dentate margin. Inflorescences in the leaf axils or just below the leaves, pedunculate, bracteate. Staminate inflorescences spicate to subcapitate; tepals 4, free or basally connate; stamens 4; pistillode absent. Pistillate inflorescences racemose to subcapitate or subumbellate; perianth tubular, entire or 4-lobed, the upper, free part somewhat different in diameter, surface, and indument from the lower part; ovary adnate to the perianth; stigmas 2, tongue- or band-shaped. Fruiting perianth red, turning blackish; fruit adnate to the perianth; rachis and pedicels ± fleshy, red or orange. Seed large, without endosperm; cotyledons very unequal, one minute and flat, the other thick, conduplicate and enclosing the smaller one. Southern Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Curaçao, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 14 species, 3 in Venezuela, all in the flora area. Sorocea guilleminiana Gaudich., widespread in the Amazon basin, may occur in the flora area as well. It has spinulose-denticulate leaves. Key to the Species of Sorocea 1.

1.

Lower surface of leaf blades ± distinctly hairy, usually scabrous to scabridulous; bracts membranous; plants deciduous, with prominent leaf scars .............................................................................................. S. sprucei Lower surface of leaf blades glabrous, or, if hairy, then not scabrous or scabridulous; bracts subcoriaceous; plants evergreen, with plane leaf scars ........................................................................................................ 2

Sorocea 727

2(1).

2.

Stipules 5–12 mm long; petioles (5–)8–16 mm long; staminate inflorescences 2–12 cm long; pistillate inflorescences 3–5 (in fruit to 9.5) cm long; fruiting perianth 1–1.7 cm diameter .................. S. pubivena Stipules 2–4 mm long; petioles 2–7 mm long; staminate inflorescences 1– 3.5(–4.5) cm long; pistillate inflorescences 0.5–1.5 (in fruit to 4.5) cm long; fruiting perianth ca. 0.5–0.8 cm diameter ................... S. muriculata

Sorocea muriculata Miq. in Mart., Fl. Bras. 4(1): 113. 1853. Shrub or small tree. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies (largely allopatric), both in the flora area.

Fig. 608. Sorocea muriculata subsp. uaupensis

Fig. 609. Sorocea pubivena subsp. hirtella

728

M ORACEAE

Key to the Subspecies of S. muriculata

Key to the Subspecies of S. pubivena

1. Tepals of the staminate flower glabrous outside; upper part of the perianth of the pistillate flower distinctly broader than the lower part, the latter subglobose in fruit ....................... subsp. muriculata 1. Tepals of the staminate flower minutely puberulent outside; upper part of the perianth of the pistillate flower almost as wide as the lower part, the latter cylindrical to oblongoid in fruit .............. ................................ subsp. uaupensis

1. Lower surface of leaf blades with dense to rather sparse spreading hairs ................ ..................................... subsp. hirtella 1. Lower surface of leaf blades with sparse appressed hairs on the midrib or (sub) glabrous ................ subsp. oligotricha

S. muriculata subsp. muriculata Sorocea amazonica Miq. in Mart., Fl. Bras. 4(1): 113. 1853. Forest understory, 100–300 m; Bolívar (Serranía de Imataca), scattered in western Amazonas. Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia. S. muriculata subsp. uaupensis (Baill.) C.C. Berg, Proc. Kon. Ned. Akad. Wetensch., C 88: 387. 1985 —Pseudosorocea uaupensis Baill., Adansonia 11: 297. 1875. —Sorocea uaupensis (Baill.) J.F. Macbr., Candollea 5: 348. 1934. —Akwi æ’mi (Piaroa), Pal de brasil, Phõi æ’mi (Piaroa), Phõi tæphi (Piaroa), Sada (Yekwana), Shara (Yekwana), Sharama. Sorocea guayanensis W. Burger, Acta Bot. Neerl. 11: 439. 1962. Forest understory, 100–1300 m; scattered throughout Bolívar and Amazonas. Colombia, Suriname, French Guiana, Brazil. ◆Fig. 608. Woody stems of Sorocea muriculata subsp. uaupensis are used for poles in house building; more flexible parts are used to weave together conical traps for use in the capture of burrowing animals. Sorocea pubivena Hemsl., Biol. Centr. Amer. Bot. 3: 150. 1883. Shrub or tree to 15(–25) m tall. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; 3 subspecies (partly sympatric), 2 in Venezuela, both in the flora area.

S. pubivena subsp. hirtella (Mildbr.) C.C. Berg, Novon 6: 243. 1996. —Sorocea hirtella Mildbr., Notizb. Bot. Gart. Berlin-Dahlem 10: 183. 1927. Sorocea opima J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 64. 1931. Forest understory, 100–200 m; Amazonas (Río Cataniapo, upper Río Cuao, Yavita). Colombia, Ecuador, Peru, Brazil. ◆Fig. 609. S.

pubivena subsp. oligotricha (Akkermans & C.C. Berg) C.C. Berg, Novon 6: 243. 1996. —Sorocea hirtella subsp. oligotricha Akkermans & C.C. Berg, Proc. Kon. Ned. Akad. Wetensch., C 88: 383. 1985. —Charo macho. Forest understory, 100–300 m; Bolívar (Serranía de Imataca), Amazonas (Río Mawarinuma). Aragua, Carabobo; Panama, Colombia, Guyana, Ecuador, Peru, Brazil. Sorocea sprucei (Baill.) J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 16. 1931. —Pseudosorocea sprucei Baill., Adansonia 11: 296. 1875. Shrub or tree to 8 m tall. Rather dry scrub and forests. Curaçao, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina; 3 subspecies, 1 in Venezuela. S. sprucei subsp. sprucei. —Charo, We yó (Panare). Olmedia virgata Pittier, Bol. Soc. Venez. Ci. Nat. 8: 305. 1943. Small tree in relatively dry forests, 100– 200 m; Bolívar (near Maniapure, lower Río Caura). Widespread elsewhere in Venezuela; Curaçao, Colombia, Guyana, Ecuador, Peru, Brazil (Roraima), Bolivia.

14. TRYMATOCOCCUS Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 30. 1838. Lanessania Baill., Adansonia 11: 298. 1875. Trees, monoecious. Leaves distichous; stipules semi-amplexicaul, free; blades entire. Inflorescences borne in the leaf axils, bisexual, pedunculate, with cylindrical to turbinate receptacle, bracteate. Staminate flowers few to many, superficial, on the

M ORINGACEAE 729

Fig. 610. Trymatococcus amazonicus

upper part of the receptacle; perianth 3-lobed to 3-parted; stamens 3; pistillode present. Pistillate flower one, central, embedded in the receptacle; ovary adnate to the perianth; stigmas 2, filiform. Fruit forming a drupaceous structure with the enlarged, fleshy, yellowish receptacle. Seed large, without endosperm; cotyledons thick, equal or unequal. Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil; 2 species, 1 in Venezuela. Trymatococcus amazonicus Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 30, t. 142. 1838. —Eyokoa (Yekwana), Palo de Brasil, Potojimujii (Yanomami), Rutæ (Piaroa). Trymatococcus turbinatus (Baill.) Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 23. 1922. —Lanessania turbinata Baill., Adansonia 11: 298. 1875. Trymatococcus paraensis Ducke, Arch.

Jard. Bot. Rio de Janeiro 3: 22. 1922. Tree to 15 m tall. Nonflooded or sometimes in periodically flooded forests, 100–700 m; Bolívar (near Santa María de Erebato), Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 610. The seeds of Trymatococcus amazonicus are eaten by humans, pacas, and agoutis.

MORINGACEAE by James S. Miller Deciduous trees, the bark with mucilage canals and often gummy. Leaves alternate, twice or three times pinnately compound, the pinnae opposite, estipulate or the stipules reduced to stipitate glands at the base of the petioles and pinnae. Inflorescences axillary, paniculate. Flowers bisexual, zygomorphic, perigynous. Sepals 5, imbricate, unequal, often somewhat petaloid, spreading or reflexed. Petals 5,

730

M ORINGACEAE

Fig. 611. Moringa oleifera

imbricate, the upper two smaller, the lower the largest, all more or less reflexed. Stamens 5, alternating with 5 staminodia; filaments free, inserted on the margin of a nectariferous disk; anthers monothecal, dehiscing longitudinally; pollen (2)3(4) colporate. Ovary borne on a short gynophore, (2)3(4)-carpellate, unilocular, the placentation parietal; style terminal, hollow, the stigma truncate; ovules biseriate, pendulous, anatropous. Fruit capsular, elongate, explosively dehiscent, unilocular and lacking a replum; seeds numerous, 3-winged or rarely lacking wings, the endosperm scant or absent.

Musa 731

Africa through Madagascar and the Middle East to India, cultivated in Neotropics; 1 genus and 13 species, 1 species in the flora area. 1. MORINGA Adans., Fam. Pl. 2: 318. 1763. Characters and distribution the same as the family; 13 species, 1 in Venezuela. Moringa oleifera Lam., Encycl. 1: 398. 1783 [1785]. Tree to 6 m tall; flowers cream with yellow anthers. Secondary vegetation in towns and

cities, 50–200 m; Bolívar (Ciudad Bolívar). Anzoátegui, Falcón; native of India but widely cultivated in tropical areas. ◆Fig. 611.

MUSACEAE by Kay Yatskievych Perennial herbs from a large corm, often tree-like but dying back to the ground after flowering, with lacticifers, glabrous throughout. Pseudostem (false hollow trunk) formed by the closely overlapping and tightly appressed, sheathed bases of the leaves. Leaves spirally arranged, large, with a coarse basal sheath; petioles long, arising from the top of the pseudostem; blades simple, entire but often frayed; midrib prominent, the lateral veins pinnate, parallel, extending nearly to the margin, each curved upward and touching the top of the adjacent one. Inflorescence axis arising from the corm, growing up through the pseudostem and out the top, then bent or drooping (erect), the exserted portion with large, leather-firm, keeled or boat-shaped, spirally arranged bracts, each of which subtends a compact, few-flowered cyme. Flowers epigynous, irregular, functionally unisexual (the plants monoecious), those subtended by the lower bracts pistillate, with reduced, nonfunctional stamens, those subtended by the upper bracts staminate, with a reduced nonfunctional ovary. Tepals 6, all petaloid but unlike, basically in 2 series. Stamens 5(6), sometimes the 6th as a small staminode; filaments slender, distinct; anthers linear, terasporangiate and dithecal, opening by longitudinal slits. Gynoecium of 3 carpels united to form a compound, trilocular, inferior ovary; style terminal, slender, 3lobed; stigma papillate. Fruits fleshy, indehiscent, with a firmer, separable exocarp. Seeds few to numerous (absent in the flora area), not arillate. Native to southern Asia and northern Australia, widely introduced elsewhere in the tropics and subtropics; 2 genera and ca. 35 species, 1 genus and 1 stable hybrid in Venezuela. 1. MUSA L., Sp. Pl. 1043. 1753. All parts of the plant with lacticifers containing a mucilaginous substance. Rhizome sympodial, with branches terminating in aerial shoots, renewal branches arising from near the base of the aerial shoot, lateral branches arise from leaf-opposed buds. Spathes caducous. Free (inner) tepal unlobed. Stamens 5, staminode present or absent.

732

M USACEAE

Southern Asia, northern Australia, widely cultivated throughout the tropics and subtropics worldwide; ca. 35 species, 1 stable hybrid in Venezuela. Musa ×paradisiaca L., Sp. Pl. 2: 1043. 1753. —Cambur, Plátano. Musa sapientum L., Syst. Nat. ed. 10, 2: 1303. 1759. Large erect herb. Widely cultivated

throughout the lowlands and uplands of the flora area, often persisting at abandoned house sites or old cultivation sites. Widely cultivated elsewhere in Venezuela; distribution as in the genus. ◆Fig. 612.

Fig. 612. Musa ×paradisiaca

Myrica 733

Musa ×paradisiaca is a sterile triploid that produces no seeds. It is widely cultivated and is an important food crop for both indigenous groups (especially the Yanomami) as well as the general (criollo) population. There are

many varieties of cultivated bananas, some eaten raw, but more often cooked in various ways (plátano). The leaves are also used to wrap other foods for cooking.

MYRICACEAE by James S. Miller Evergreen or deciduous shrubs or small trees, usually pubescent, often strongly aromatic. Leaves alternate, simple, or less commonly pinnately lobed, stipules absent or present (Comptonia). Inflorescences axillary, dense spikes or aments, bracteate. Flowers small, unisexual, rarely with some bisexual, the plants monoecious or dioecious, each flower subtended by a bract or sometimes a small group of flowers subtended by a single bract. Perianth absent. Stamens 2–8 to numerous, in a single whorl; filaments connate at the base or free; anthers bithecal, dehiscing longitudinally. Ovary 2-carpellate, superior, unilocular; style terminal, bifid; ovule 1, basal,

Fig. 613. Myrica rotundata

734

M YRICACEAE

orthotropous. Fruits drupaceous to nearly dry, often densely covered with waxy glands; seeds lacking endosperm. Cosmopolitan; 2 or 3 genera and ca. 50 species, 1 species in flora area. 1. MYRICA L., Sp. Pl. 1024. 1753. Shrubs or trees, evergreen or deciduous, often aromatic. Leaves alternate, simple, entire or serrate, the lower surface often glandular, estipulate. Inflorescences axillary, spikes or aments, bracteate. Flowers small, unisexual, the plants mostly dioecious, rarely monoecious. Perianth absent. Staminate flowers with 2–20 stamens; filaments basally connate or free; anthers bithecal, dehiscing longitudinally. Fruits drupaceous, the exocarp usually with waxy accretions; seeds lacking evident endosperm. Temperate, subtropical, and high-elevation areas of the tropics of both hemispheres; ca. 50 species, 2 in Venezuela, 1 of these in flora area. Myrica rotundata Steyerm. & Maguire, Mem. New York Bot. Gard. 17: 441. 1967. Shrub or small tree to 4 m tall. Mixed Bonnetia forests, forested slopes, 1900–2200

m; Bolívar (Macizo del Chimantá [Apácaratepui, Toronó-tepui]). Endemic. ◆Fig. 613. The fragrant gland-dotted leaves are characteristic of the genus.

MYRISTICACEAE by William A. Rodrigues, Gerardo A. Aymard C., and Paul E. Berry Evergreen or rarely deciduous trees or shrubs, frequently aromatic, dioecious or rarely monoecious. Leaves alternate, distichous, sometimes pseudowhorled, occasionally with pellucid punctation, exstipulate; blades simple, entire; trichomes dendroid, often stellate, or malpighiaceous (straight hairs attached near the middle). Inflorescences dichasial (mostly in branching of the 1st order), fasciculate, racemose, or paniculate, axillary, rarely terminal or cauliflorous; bracts mostly caducous; bracteoles frequently present. Flowers small, hypogynous, actinomorphic, campanulate, funnel-shaped, or urceolate, seldom scented. Perianth of (2)3(–5) basally fused, often fleshy, valvate tepals. Staminate flowers with 2–40 stamens; filaments partially or entirely fused and forming an androphore; anthers often united with the oblong, roundish, or claviform androphore, rarely free, muticous or shortly apiculate, tetrasporangiate, extrorse, longitudinally dehiscent, or seldom latrorse (dehiscing both longitudinally and laterally). Pistillate flowers 1-carpellate; ovary superior, sessile, or sometimes shortly stipitate; style often absent; stigma simple or bilobed; ovule solitary, subbasal to basal, anatropous, rarely orthotropous or hemiorthotropous, bitegmic. Fruit a fleshy to leathery or woody capsule, generally opening along ventral and dorsal sutures. Seed coat ligneous, mostly covered by a crustaceous to fleshy, laciniate or entire, brightly colored aril; endosperm abundant, frequently ruminate; embryo small. Pantropics and subtropics; 18 genera and ca. 400 species, 4 genera and 27 species in the flora area.

Compsoneura 735

Key to the Genera of Myristicaceae 1.

1. 2(1).

2.

3(2).

3.

Leaves with tertiary veins conspicuously subparallel, prominulous, and somewhat perpendicular to the costa; pericarps very thin, seed coat irregularly black- or purple-spotted .................................... 1. Compsoneura Leaves with more irregular tertiary venation; pericarps generally thick to woody; seed coat not mottled ................................................................. 2 Young branches, petioles, and lower surfaces of leaves usually pubescent, with stellate or dendroid trichomes; flowers ebracteolate; seeds and fruits longitudinally ellipsoid or subglobose; aril deeply laciniate, nearly to base ................................................................................. 4. Virola Young branchlets and leaves generally lacking the above kinds of trichomes; flowers bracteolate; seeds and fruits transversely ellipsoid; aril entire or only apically laciniate ...................................................... 3 Leaves often finely rugose or minutely papillose on both surfaces, variously shaped and mostly acute (but occasionally rounded) at the apex; anthers 3(–6), dorsally connate or free to the base, the connectives inconspicuous or fleshy; inflorescences strigose with usually malpighiaceous trichomes; aril apically laciniate ............................... 2. Iryanthera Leaves usually smooth and shining on upper surface, obovate, mostly rounded and slightly emarginate at the apex, gradually attenuate at the base; anthers 12(–14), dorsally adnate to a conspicuous connective mass; juvenile inflorescences puberulent with multibranched, somewhat lepidote trichomes; aril entire ................................. 3. Osteophloeum

1. COMPSONEURA (A. DC.) Warb., Ber. Deutsch. Bot. Ges. 13(Gen.-Vers.): 83. 1896 [1895]. —Myristica sect. Compsoneura A. DC., Prodr. 14: 199. 1856. Dioecious, glabrous shrubs or small trees, seldom lianas, the sap often reddish. Leaves glabrous, chartaceous to coriaceous, petiolate; costa raised on upper surface, prominent on lower surface, the veins several, irregularly anastomosing near margins, the veinlets subparallel, prominulous on both surfaces, ± perpendicular to the costa, mostly conspicuous. Inflorescences axillary or on leafless branchlets, racemose, fasciculate-racemose, or narrowly paniculate; bracteoles none. Flowers pedicellate. Staminate perianth fleshy or thinly so, sometimes puberulent inside, tepals 3(4 or 5), partially connate; anthers 4–10, oblong, bilocellate, basifixed and free in section Compsoneura or dorsally adnate to a fleshy truncate-obconical connective mass in section Coniostele, the locelli separately adnate and often divergent; filaments fused into a fleshy androphore. Pistillate perianth generally slightly larger and somewhat thicker than the staminate one; ovary subglobose or ellipsoid with a subsessile, peltate or bilobed stigma. Fruits fleshy, ellipsoid, glabrous, dehiscing longitudinally into 2 valves, smooth or obscurely carinate, pedicellate, the pedicels surmounted by the remaining perianth lobes. Pericarp very thin; seed coat black- and purple-spotted; aril brightly colored, entire or subentire; endosperm not ruminate. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia; ca. 12 species, 2 in Venezuela, both in the flora area.

736

M YRISTICACEAE

Key to the Species of Compsoneura 1. 1.

Leaves coriaceous, apex rounded or obtuse; staminate inflorescences < 1.5 cm long, the flowers crowded in rounded clusters .............. C. debilis Leaves chartaceous or thinly coriaceous, often translucent, apex obtusely cuspidate or acuminate; staminate inflorescences > 2 cm long, aggregated in fascicles of 3–15 at ends of lateral branches ................ C. sprucei

Compsoneura debilis (Spruce ex A. DC.) Warb., Ber. Deutsch. Bot. Ges. 13(Gen.Heft): 84. 1896 [1895]. —Myristica debilis Spruce ex A. DC., Prodr. 14: 697. 1857. —Palala debilis (Spruce ex A. DC.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Café de venado, Ceguera, Majagua, Palo aro de atarraya. Shrub, sometimes scandent, 1–6 m tall; stem 1.5–2 cm diameter; leaves 10–16 × 3.5– 8 cm, elliptic or oblong-elliptic, coriaceous, attenuate at base and narrowly decurrent on the petiole, rounded or obtuse at apex, erect on branches when live; inflorescences glabrous throughout, very short and compact, flowers white; anthers free from base or partially connate by inconspicuous connectives; fruits 17–22 × 9–12 mm, orange or orangered, aril red or purplish. Río Negro caatinga, white-sand scrub (bana), 100–200 m; Amazonas (Caño San Miguel, Caño Ucata southeast of Síquita, Cerro El Danto east of San Juan, Río Atacavi, Río Guainía, Río Puruname, Río Sipapo, Río Temi basin, near San Carlos de Río Negro, base of Sierra de la Neblina, road from Yavita to Maroa). Amazonian Colombia (Guianía, Vaupés), Brazil (Amazonas: upper Rio Negro).

Compsoneura sprucei (A. DC.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 143. 1897. —Myristica sprucei A. DC., Prodr. 14: 199. 1856. —Palala sprucei (A. DC.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. Myristica laurifolia Spruce ex A. DC., Prodr. 14: 199. 1856, as synonym. Myristica mexicana Hemsl., Biol. Cent. Amer. Bot. 3: 67. 1882. —Palala mexicana (Hemsl.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. Compsoneura costaricensis Warb., Repert. Spec. Nov. Regni Veg. 1: 71. 1905. Shrub or small tree 3–18 m tall; trunk to 40 cm diameter, the inner bark with a reddish watery exudate; leaves 9–25 × 3.5–15 cm, elliptic or oblong-elliptic, chartaceous or thinly coriaceous, glabrous, acute to attenu-

Fig. 614. Compsoneura sprucei

Iryanthera 737

ate at base, obtusely cuspidate or acuminate at apex; staminate inflorescences glabrous throughout, 2–8 cm long, narrowly paniculate or fasciculate-racemose; flowers aggregated in irregular fascicles at ends of lateral branches, creamy yellow; fruits 1.5–2 × 2–3 cm, orangish; aril bright red, entire, con-

spicuously stipitate at base (stipes ca. 2–5 mm long). Evergreen lowland forests, 100– 200 m; Amazonas (upper Río Guayapo, Río Mawarinuma, near San Carlos de Río Negro). Mexico, Central America, Amazonian Colombia, Suriname, Amazonian Ecuador, Peru, Brazil (Amazonas). ◆Fig. 614.

2. IRYANTHERA (A. DC.) Warb., Ber. Deutsch. Bot. Ges. 13(Gen.-Heft): 84. 1896 [1895]. —Myristica sect. Iryanthera A. DC., Prodr. 14: 201. 1856. Dioecious or sometimes monoecious shrubs or trees, frequently exuding a reddish sap; very young branchlets ferruginous-strigose, the trichomes usually malpighiaceous, soon glabrous. Leaves chartaceous to coriaceous, often finely rugose or minutely papillose on both surfaces, glabrous; the costa prominent on lower surface; venation mostly conspicuously brochidodromous (with a single primary vein, the secondary veins joined together in a series of marginal loops), sometimes camptodromous (with a single primary vein, the secondary veins curved upwards and gradually diminishing distally within the margin, without forming conspicuous marginal loops). Inflorescences 1–3 in leaf axils or on leafless branchlets (pistillate usually cauliflorous or ramiflorous), fasciculate-racemose or narrowly paniculate, minutely strigose or glabrescent, the pistillate often proximally few-branched; fascicles often sessile on the rachis; pedicels distally bracteolate, bracteoles persistent, cupuliform or 1-sided; staminate perianth small, campanulate to cup-shaped, usually fleshy and strigose outside, glabrous inside, 3(4)-lobed; filaments fused into a column; anthers 3 or 4(–6), adnate to the androphore or apically or rarely completely free, obtuse, 2-celled, the connectives inconspicuous or fleshy, rarely glandular; pistillate perianth usually somewhat larger and thicker than the staminate; ovary ellipsoid, conical or cylindric, glabrous; style short or none; stigma inconspicuous. Infructescences entirely glabrous; fruits mostly transversely ellipsoid, occasionally subglobose, coriaceous, 2-valved. Pericarp mostly woody. Seed transversely ellipsoid or subglobose; aril entire or apically inconspicuously laciniate; endosperm not or little ruminate. Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 23 species, 9 in Venezuela, all in the flora area. Key to the Species of Iryanthera 1.

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

3.

Leaf blade ca. 6–9 × 3–4 cm, obovate or obovate-elliptic, rounded or obtuse at apex; veins plane or slightly impressed on the upper surface ...................................................................................................... I. obovata Leaf blade without the above combination of characters ......................... 2 Veins conspicuously raised on both surfaces ............................................. 3 Veins impressed, obscure, or plane on the upper surface ......................... 4 Leaf blade usually elliptic or obovate-elliptic, a few smooth trichomes on both surfaces; veins mostly ≤ 11 per side; perianth cup-shaped; fruit conspicuously angled on both sides, characteristically apiculate at the apex ............................................................................................. I. tricornis Leaf blade oblong or obovate-oblong, the base subcordate, rounded, or obtuse, the apex obtusely cuspidate or short acuminate, finely rugose or

738

4(2).

4.

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

6.

7(5).

7.

8(7).

8.

M YRISTICACEAE

uniformly minutely papillose on both surfaces; veins usually 14– 20 per side; perianth campanulate; fruit not apiculate at the apex .............................................................................................. I. macrophylla Leaf blade thinly coriaceous, smooth on the upper surface; veins often obscure or plane on both surfaces; perianth broadly campanulate ......................................................................................................... I. laevis Leaf blade coriaceous, minutely rugulose or uniformly minutely papillose on both surfaces; veins conspicuously impressed on the upper surface and raised on the lower surface ............................................................ 5 Perianth cup-shaped or rotate at maturity; anthers as long as the androphore or a few shorter .................................................................. 6 Perianth narrowly campanulate; anthers shorter than the androphore . 7 Veins shallowly impressed or irregularly raised on the upper surface and somewhat plane on the lower surface, venation obscurely anastomosing at the margins; flowers covered by dense trichomes, obscuring their surfaces; fruit usually subglobose with pericarp 4–5 mm thick ................................................................................................... I. lancifolia Veins deeply impressed on the upper surface and raised on the lower surface; venation conspicuously anastomosing at margins; flower surfaces not obscured by dense trichomes; fruit transversally ellipsoid with pericarp 0.5–1.5 mm thick ........................................................ I. paraensis Base of the leaf blade usually acute to subattenuate; veins obscurely anastomosing near the margins; pistillate inflorescence cauliflorous .................................................................................................... I. juruensis Base of the leaf blade obtuse to acute; venation conspicuously anastomosing near the margins; pistillate inflorescence ramiflorous or axillary ......................................................................................................... 8 Leaf blade thickly coriaceous, conspicuously papillose on both surfaces, pedicels to ca. 10 mm long, perianth soon glabrous; fruit indistinctly carinate ..................................................................................... I. paradoxa Leaf blade thinly coriaceous, minutely papillose or nearly smooth on both surfaces; pedicels to 5 mm long; perianth outside uniformly minutely strigose; fruit conspicuously carinate ................................... I. hostmannii

Iryanthera hostmannii (Benth.) Warb., Ber. Deutsch. Bot. Ges. 13(Gen.-Heft): 84. 1896 [1895]. —Myristica hostmannii Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 7. 1853, “hostmanni.” —Coloradito, Cuajo, Cuajo montañero, Majagua, Sangrito. Iryanthera leptoclada Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 9: 965. 1926. Iryanthera congestiflora Macbr., Candollea 5: 350. 1934. Tree 8–15 m tall. Evergreen flooded lowland forests to submontane forests, riparian forests, 100–700 m; Bolívar (Cerro Ichún, El Abismo, El Tigre, Icabarú, Macizo del Chimantá [Toronó-tepuí], Río Caura, Río Nichare basin, Río Paragua, Sierra de Lema),

Amazonas (Río Asisa, Río Casiquiare, Río Manapiare, Río Mawarinuma, Río Ocamo, San Carlos de Río Negro, Sierra de la Neblina, Yavita). Zulia; Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonia), Bolivia. Iryanthera juruensis Warb., Verh. Bot. Vereins Prov. Brandenburg 47: 137. 1905. —Cuajo de tierra firme, Cuajo negro, Kuduvi (Yanomami), Majagua, Masarico, Sangrito. Tree 5–16 m tall; infructescences cauliflorous. Evergreen lowland to submontane forests, 100–900 m; Bolívar (El Cácaro between Rio Caura and Rio Paragua, Icabarú, Río Aparamán, Río Caura, Río Chirca near

Iryanthera 739

Urimán, upper Río Paragua, Santa Maria de Erebato), Amazonas (Caño Libreta 35 km southeast of La Esmeralda, Cerro Duida, El Pozo southeast of San Fernando de Atabapo, Río Baría, Río Casiquiare, Río Cunucunuma, Río Manapiare, Río Matacuni, Río Mawarinuma, Río Ocamo, Río Orinoco). Zulia; Panama, Amazonian Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

85. 1896 [1895]. —Myristica macrophylla Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 6. 1853. —Palala macrophylla (Benth.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Cuajo. Tree to 15 m tall with dense ramiflorous fruit clusters. Evergreen lowland forests, 100–200 m; Amazonas (Capibara, Río Sipapo, Río Ventuari, north of Yavita). Amazonian Colombia, Guyana, Peru, Brazil.

Iryanthera laevis Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 9: 965. 1926. —Akodek (Arekuna), Cuajo, Cuajo tierra firme, Dato (Curripaco), Maruetije (Yanomami), Maruetiji (Yanomami), Molenillo, Nopo, Sakuma (Yekwana). Tree 10–35 m tall. Evergreen lowland forests to lower montane forests, 100–700 m; Bolívar (Río Chirca near Urimán, Río Erebato basin, Río Nichare basin), Amazonas (Cerro El Danto near Cerro Yapacana, Reserva Forestal El Sipapo, Río Asisa, Río Caname, Río Negro 20 km south of confluence with Río Casiquiare, Río Orinoco, Río Padamo, Río Ventuari, 43 km northeast of Santa Bárbara del Orinoco, Sierra Tapirapecó, road between Yavita and Maroa). Colombia, Ecuador, Peru, Brazil (Amazonia), Bolivia.

Iryanthera obovata Ducke, J. Wash. Acad. Sci. 26: 221. 1936. —Cuajo, Molenillo, Tapata (Curripaco), Yagua rosorina. Treelet or tree to 30 m tall; leaves 6–9 × 3–4 cm, rounded to bluntly acute at apex. Evergreen lowland forests, white-sand shrublands (bana), tall caatinga forests, 50– 200 m; Amazonas (Cerro El Danto east of Cerro Yapacana, base of Cerro Yapacana, Río Casiquiare, Río Mawarinuma, road between Yavita and Maroa). Brazil (Amazonas: upper Rio Negro).

Iryanthera lancifolia Ducke, J. Wash. Acad. Sci. 26: 217. 1936. —Cacaguaru, Cuajo grande, Cuajo sabanero, Gueguewudu (Yekwana), Guiejudo (Yekwana), Macairicao. Iryanthera porcata A.H. Gentry, Phytologia 48: 233. 1981. Tree to 35 m tall; leaves large, smooth, coriaceous; flowers minute, densely ferruginous; fruits large. Evergreen lowland to montane forests, near sea level to 1000 m; Delta Amacuro (Caño Arature southwest of Atabuina, east of Río Toro, north of Wausa on Río Amacuro), Bolívar (Cerro Venamo, Sierra de Maigualida), Amazonas (San Carlos de Río Negro). Colombia, Peru, Brazil (Amazonas, Pará, Rondônia). The wood of Iryanthera lancifolia is hard and is used for lumber. Iryanthera macrophylla (Benth.) Warb., Ber. Deutsch. Bot. Ges. 13(Gen.-Vers.):

Iryanthera paradoxa (Schwacke) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 160. 1897. —Myristica paradoxa Schwacke, Add. Fl. Bras. pl. 2. 1886. —Cuajo, Wi-wi-hijú-deu (Yekwana). Iryanthera longiflora Ducke, J. Wash. Acad. Sci. 26: 217. 1936. Tree to ca. 15 m tall; leaves coriaceous. Evergreen lowland to lower montane forests, seasonally flooded forests near black-water rivers, 100–300 m; Bolívar (Río Mawela tributary of Río Erebato), Amazonas (Caño Iguapo, La Esmeralda, Río Atacavi, Río Cataniapo, Río Cunucunuma, Río Mawarinuma, San Carlos de Río Negro, San José del Orinoco, Yavita to Maroa). Colombia, Peru, Amazonian Brazil. Iryanthera paraensis Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 5: 358. 1909. —Tabacilla. Iryanthera elongata Huber, Bol. Mus. Goeldi Paraense Hist. Nat. Ethnogr. 6: 68. 1910. Iryanthera sessilis Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 10: 236. 1928. Treelet or tree to 20 m tall; flowers minute. Evergreen lowland forests, 100–500 m; Bolívar (Río Mawela tributary of the Río

740

M YRISTICACEAE

Erebato, Río Oris, Río Tonoro, Santa María de Erebato), Amazonas (La Esmeralda, Río Mawarinuma, San Carlos de Río Negro). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 615.

Iryanthera tricornis Ducke, Trop. Woods 31: 11. 1932. —Guigujuro (Yekwana). Tree 6–12 m tall; leaves elliptic to obovate, rounded to bluntly acute at apex. Evergreen lowland to lower montane forests, ca. 400 m; Bolívar (El Cácaro between Rio Caura and Rio Paragua, southwest of Urimán). Colombia, Peru, Amazonian Brazil.

Fig. 615. Iryanthera paraensis

3. OSTEOPHLOEUM Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 127, 162. 1897. Dioecious trees; branchlets with multibranched trichomes, later glabrous. Leaves petiolate, coriaceous, glabrous at maturity; margins entire and narrowly revolute; costa prominent on lower surface, the veins obscurely anastomosing near margins; veinlets reticulate. Inflorescences axillary, puberulent, short-racemose or

Osteophloeum 741

paniculately branched. Flowers solitary or fasciculate, pedicellate, the pedicels distally bracteolate; staminate perianths funnel-shaped, small, fleshy, glabrous inside, 3-lobed; androphore fused into a fleshy cylindric column; anthers 12–14, 2-celled, dorsally connate into a fleshy connective mass; pistillate perianths few, larger and more fleshy than the staminate ones; ovary 1-carpellate, conical; stigma sessile, oblique. Fruits transversely ellipsoid or subglobose, 2-valved, the pericarp woody. Seeds transversely ellipsoid or suglobose; testa very thick; aril entire. Panama, Colombia, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, Bolivia; 1 species. Osteophloeum platyspermum (Spruce ex A. DC.) Warb., Ber. Deutsch. Bot. Ges. 13: 89. 1896 [1895]. —Osteophloeum platyspermum (Spruce ex A. DC.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 162. 1897. —Myristica platysperma Spruce ex A. DC., Prodr. 14: 695. 1857. —Palala platysperma (Spruce ex A. DC.) Kuntze, Revis. Gen. Pl. 2: 557. 1891. —Azucarito, Cuajo hoja grande, Cumala blanca, Guarodek (Arekuna), Kunuri (Yekwana), Poroi (Yanomami), Prooiji (Yanomami), Sacuma

(Yekwana), Suiguichasto (Yekwana), Tararujuru (Yekwana). Osteophloeum sulcatum Little, Phytologia 18: 404, fig. 6. 1969. Tree 10–45 m tall and to 85 cm diameter; trunk usually erect, with steep rounded buttresses, slash producing abundant ambercolored, watery exudate; crown usually irregular and reduced, the ascending branches and leaves giving the tree an unmistakable character; leaves oblong-obovate or ellipticoblong, gradually attenuate at base, often rounded or slightly emarginate at apex; flowers yellowish; fruits green. Evergreen lowland to lower montane forests, on sandy ultisols, deciduous forests on white sandy soil, tall caatinga forests, 100–900 m; Bolívar (El Cácaro between Rio Caura and Rio Paragua, Río Canaracuni, headwaters of Río Icabarú and Río Hacha, upper Río Paragua, Río Yudi, Uaiparú), Amazonas (35 km southeast of La Esmeralda, Río Casiquiare, Río Cunucunuma, Río Mawarinuma, upper Río Orinoco, Río Padamo, San Carlos de Río Negro, San José del Orinoco, Sierra de la Neblina, near Yavita). Other distribution as in genus. ◆Fig. 616.

Fig. 616. Osteophloeum platyspermum

742

M YRISTICACEAE

4. VIROLA Aubl., Hist. Pl. Guiane 904. 1775. —Myristica sect. Virola (Aubl.) Endl., Enchiridion 419. 1841. Sebophora Necker, Elem. Bot. 2: 188. 1790. Myristica sect. Sychnoneura A. DC., Ann. Sci. Nat. Bot., sér. 4(4): 30. 1855. Dioecious shrubs or trees, the younger parts pubescent with stellate or multibranched hairs, the slash exudate reddish or brownish. Leaves glabrous on upper surface, the lower surface with stellate or multibranched trichomes and often glabrescent, submembranaceous to coriaceous, petiolate, the veinlets usually obscure. Inflorescences paniculate or racemose, the axes usually stellate-pubescent; bracts membranaceous, soon deciduous, bracteoles lacking. Staminate flowers in fascicles of 3–15; perianth campanulate to funnel-shaped, deeply or shallowly 3- or 4-lobed; anthers 2–6, dorsally adnate to the column, at least basally, longer or shorter than androphore. Pistillate flowers solitary or in fascicles of 2–7. Capsules longitudinally ellipsoid or subglobose, 1–40 per infructescence, the woody pericarp dehiscing longitudinally into 2 valves. Seeds with ruminate endosperm; aril usually red, irregularly laciniate. Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Lesser Antilles (Guadeloupe to Grenada), Colombia, Trinidad-Tobago, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 48 species, ca. 18 in Venezuela, 15 of these in the flora area. Key to the Species of Virola 1. 1. 2(1).

2.

3(2).

3.

4(3).

4.

5(1). 5.

Lower surface of leaves tomentose with persistent, stalked or irregularly branched trichomes ............................................................................... 2 Leaves with usually sessile and soon glabrescent trichomes ................... 5 Leaves densely tomentose on lower surface, with few arcuate veins (1 or fewer per cm), venation camptodromous; anthers 2–3 times as long as the androphore ............................................................................. V. sebifera Leaves densely or loosely tomentose on lower surface, with numerous (2 or 3 per cm or more) straight veins, venation conspicuously brochidodromous; anthers shorter than the androphore or subequal ...... 3 Leaf blades bullate, densely tomentose on lower surface; veins 30 or fewer per side; veinlets impressed on upper surface, obscure on lower surface .................................................................................................... V. rugulosa Leaf blades not bullate, loosely tomentose on lower surface; veins usually over 30 per side; veinlets impressed on upper surface and prominulous on lower surface ..................................................................................... 4 Leaf blades chartaceous, veins plane on upper surface, ca. 2 per cm, trichomes not densely overlapping on lower surface, upper surface of young leaves glabrous ........................................................... V. minutiflora Leaf blades coriaceous, veins impressed on upper surface, ca. 1 per cm, trichomes densely overlapping on lower surface, upper surface on young leaves densely furfuraceous but becoming glabrescent with age ........................................................................................................ V. duckei Staminate inflorescences comparatively simple, 1–3-branched ............... 6 Staminate inflorescences broadly paniculate, freely branching, manyflowered ................................................................................................ 10

Virola 743

6(5).

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

9.

10(5).

10.

11(10).

11.

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

13.

14(12).

Leaves 5–12 × 1.5–4 cm, elliptic or obovate-elliptic, essentially glabrous, the apex round or obtuse and often slightly emarginate, 12–16 veins per side ..................................................................................... V. parvifolia Leaves usually larger than above and/or with more secondary veins ..... 7 Leaf blades obtuse at the base; staminate inflorescences once-branched; montane areas over 1000 m elevation .................................................. 8 Leaf blades attenuate or acute at base; staminate inflorescence 1–3branched; lower elevation areas ............................................................ 9 Leaf blades 5–12 × 1.8–4; petioles 3–6 mm long; peduncle ca. 5 cm long, lateral branches 5–8 mm long ............................................. V. steyermarkii Leaf blades 14–25 × 5–9, petioles 6–10 mm long; peduncle 2.5–3.5 cm long, lateral branches 10–20 mm long ............................................ V. sp. A Leaf blades narrowly elliptic, the veins slightly impressed or obscure on the upper surface, the veinlets often obscure; fruit ellipsoid, 2.5–3 × 1.7–2.4 cm, conspicuously carinate on one side and smooth on the other, rounded or short-stipitate at the base, obtuse or minutely apiculate at the apex ........................................................................... V. michelii Leaf blades broadly elliptic, the veins and veinlets usually prominulous on the upper surface; fruits ellipsoid, 1.3–1.9 × 1.1–1.4 cm, faintly carinate, rounded at the base, rounded and apiculate at the apex ....................................................................................................... V. venosa Leaf blades 4–10 × 2–3 cm, totally glabrous except youngest buds, the base attenuate; petiole ca. 10 mm long; inflorescences delicate, 5–10 cm long ................................................................................................... V. sp. B Leaf blades larger than above and usually more persistently pubescent on lower surface, the base of various shapes; inflorescences longer and more robust .......................................................................................... 11 Leaves lanceolate-oblong with margins very often parallel, usually lustrous on upper surface, length:width ratio > 5:1 and apex shortly acute; young staminate inflorescences distally swollen with the flower fascicles enclosed by bracts; fruit subglobose or ellipsoid, 1.3–2.5 × 1.1– 1.8 cm, often apiculate at the apex, slightly or distinctly carinate ............................................................................................ V. surinamensis Leaves oblong-elliptic, ovate, or obovate-elliptic, length:width ratio < 5:1, or, if > 5:1, then the apex gradually attenuate; inflorescences not swollen distally; fruits various ...................................................... 12 Leaves submembranaceous to papyraceous; anthers about twice as long as the androphore. ............................................................................... 13 Leaves coriaceous to thick-coriaceous; anthers < twice as long as the androphore (except longer in V. sebifera) ............................................ 14 Leaves narrowly oblong to oblong-elliptic, usually glabrescent or glaucous on lower surface, the apex gradually attenuate, the base obtuse; fruits globose to subglobose, faintly carinate ...................................... V. elongata Leaves oblong to oblong-elliptic, usually inconspicuously ferruginous-tomentose on lower surface, the apex acute, the base rounded to subcordate; fruits ellipsoid, minutely apiculate at apex, slightly carinate ................................................................................................... V. theiodora Leaves 15–24 × 4–6 cm, coriaceous, oblong-elliptic or narrowly obovate-

744

M YRISTICACEAE

elliptic, the base attenuate or acute, the apex obtusely cuspidate; fruit ellipsoid, 2.5–5 cm long, 1.5–3.5 cm broad, often conspicuously carinate, obtuse or subacute at the apex ........................................... V. pavonis 14. Leaves 15–50 × 6–24 cm, thick-coriaceous, elliptic-oblong, elliptic, or ovate, the base often deeply cordate to rounded; fruit ellipsoid or globose, distinctly smaller than above ..................................................... 15 15(14). Leaves inconspicuously densely puberulent, sparsely dark-punctate and pale (usually sulphur yellow) on the lower surface; androphore much longer than the anthers, thick-fleshy, abruptly narrowed distally ................................................................................................. V. calophylla 15. Leaves puberulent or glabrescent, not punctate on the lower surface; androphore 1/2–1/3 as long as the anthers .................................... V. sebifera Virola calophylla (Spruce) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 231. 1897. —Myristica calophylla Spruce, J. Linn. Soc. 5: 4. 1860. —Palala calophylla (Spruce) Kuntze, Revis. Gen. Pl. 2: 567. 1891. Virola incolor Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 232. 1897. Tree 4–10 m tall; leaves usually large, thick-coriaceous, cordate at the base. Evergreen lowland forests, 100–200 m; Amazonas (Río Temi, near San Carlos de Río Negro, near Solano, road between Yavita and Maroa). Amazonian Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Virola duckei A.C. Sm., Brittonia 2: 487. 1937. —Guayaba de danto. Tree to 20 m tall. Evergreen lowland forests, 100–200 m; Amazonas (Raudal Remo on Río Sipapo, middle Río Baría, 43 km northeast of Santa Bárbara del Orinoco). Colombia, Ecuador, Peru. Virola elongata (Benth.) Warb., Ber. Deutcsh. Bot. Ges. 13: 89. 1895. —Myristica elongata Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 5. 1853. —Palala elongata (Benth.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Aik-yek (Arekuna), Atauashima, Ayudek (Arekuna), Ayuku (Yekwana), Codo rebalsero (Baniva), Cuajo, Cedrillo, Diaru (Warao), Dieru, Kuaho, Masarico, Merecucuri, Reventillo, Trompillo, Yopo-Ayuco (Yekwana). Myristica cuspidata Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 5. 1853. —Palala cuspidata (Spruce ex

Benth.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Virola cuspidata (Spruce ex Benth.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 176. 1897. Myristica punctata Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 6. 1853. —Palala punctata (Spruce ex Benth.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Virola elongata var. punctata (Spruce ex Benth.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 179, t. 5, fig. 1, 2. 1897. Myristica membranacea Poepp. ex A. DC., Prodr. 14: 196. 1856. —Palala membranacea (Poepp. ex A. DC.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Virola cuspidata var. membranacea (Poepp. ex A. DC.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 177. 1897. Myristica uaupensis Spruce ex A. DC., Prodr. 14: 696. 1857. —Palala uaupensis (Spruce ex A. DC.) Kuntze, Revis. Gen. Pl. 2: 568. 1891. Virola elongata var. longicuspis Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 179, t. 5, fig. 1–3. 1897. —Myristica longicuspis Spruce ex Warb., Nova Acta Acad. Caes. Leop.Carol. German. Nat. Cur. 68: 179. 1897, as synonym. Virola elongata var. subcordata Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 180. 1897. Slender tree or treelet 2–12 m tall; leaves narrowly oblong-elliptic, the apex gradually attenuate. Evergreen lowland to lower montane forests, periodically flooded forests,

Virola 745

white-sand shrublands (bana), near sea level to 800 m; Delta Amacuro (Caño Sacupana), Bolívar (Caño Ucata southeast of Síquita, Cerro Ichún, Cerro Marutaní, El Guaniamo, La Esmeralda, Macizo del Chimantá [Abacapá-tepui], middle Río Caura, Río Chirca near Urimán, Río Hacha, Río Karún, Río Nichare, lower Río Siapa, Santa María de Erebato, Serranía Pia-Zoi), Amazonas (Río Atabapo, Río Casiquiare, Río Guainía, upper Río Orinoco, Río Sipapo, near San Carlos de Río Negro). Amazonian Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. The sap of Virola elongata, which is barely observable in the dry season, is used to treat bucal ulcerations and to cure leishmaniasis. It is applied to sores for 3–5 days until a white layer forms. Virola michelii Heckel, Ann. Inst. Bot.Géol. Colon. Marseille 6: 118, fig. 24a–d. 1898. Virola melinonii Benoist, Bull. Mus. Hist. Nat. (Paris) 30: 104. 1924. Tree to 20 m tall; leaves narrowly elliptic, usually dull on upper surface, attenuate or acute at the base; veins and veinlets mostly obscure on upper surface. Evergreen lowland forests, ca. 100 m; Amazonas (near San Carlos de Río Negro). Guyana, Suriname, French Guiana, Amazonian Brazil. Virola minutiflora Ducke, J. Wash. Acad. Sci. 26: 259. 1936. Tree 5–7 m tall with red sap. Evergreen lowland forests, 100–200 m; Amazonas (road between Yavita and Maroa). Amazonian Peru and Brazil. Virola parvifolia Ducke, J. Wash. Acad. Sci. 26: 264. 1936. —Basegua banero. Tree 4–12 m tall; trunk ca. 9 cm diameter; leaves 5–11 × 3–5 cm, elliptic, coriaceous, glabrous, the base obtuse or rounded, the apex obtuse and slightly emarginate; veins 12–16 per side; flowers funnel-shaped, ferruginous-puberulent outside. Río Negro caatinga, ca. 100 m; Amazonas (Río Casiquiare, near San Carlos de Río Negro). Brazil (Amazonas: upper Rio Negro basin). Virola pavonis (A. DC.) A.C. Sm., Brittonia 2: 504. 1937. —Myristica pavonis A. DC.,

Prodr. 14: 697. 1856. —Palala ?pavonis (A. DC.) Kuntze, Revis. Gen. Pl. 2: 567. 1891, “pavonii.” —Myristica venosa var. pavonis (A. DC.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 225, figs. 1 and 2. 1897. —Ashuhua (Yekwana), Cuajo, Cuajo blanco, Cuajo pepa amarilla, Cudo (Baniva), Palo de gallineta, Tani (Yanomami). Virola elliptica A.C. Sm., Bull. Torrey Bot. Club 60: 351. 1933. Tree 15–30 m tall, usually with stilt roots; fruit large (to 5 × 4 cm), ellipsoid or obovoid; sap at first colorless, usually turning white on contact with the air. Evergreen lowland to lower montane forests, white-sand shrublands (bana), 100–400 m; Amazonas (Caño Caname, Caño Ucata southeast of Síquita, Cerro Yapacana, Cuao-Sipapo massif, Río Casiquiare, Río Cunucunuma, Río Jenita in Río Ocamo basin, Río Mawarinuma, upper Río Orinoco, Río Sipapo, Río Temi, San Carlos de Río Negro, Yavita). Colombia, Ecuador, Peru, Brazil (Amazonas, Mato Grosso, Rondônia). Virola rugulosa (Spruce) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 227. 1897. —Myristica rugulosa Spruce, J. Proc. Linn. Soc., Bot. 5: 4. 1861. —Palala rugulosa (Spruce) Kuntze, Revis. Gen. Pl. 2: 567. 1891. Tree to 35 m tall; leaves 20–27 × 7–9.5 cm, oblong, finely coriaceous, somewhat bullate, the base rounded to cordate, the apex cuspidate, often conspicuously revolute at margins; veins 23–27 per side, straight, conspicuously anastomosing near margins. Evergreen lowland forests, usually in seasonally flooded areas, ca. 100 m, Amazonas (near San Carlos de Río Negro). Brazil (Amazonas: upper Rio Negro basin). Virola sebifera Aubl., Hist. Pl. Guiane 904, t. 345, fig. 1–5. 1755. —Myristica sebifera (Aubl.) Sw., Prodr. 96. 1788. —Ayoco (Warekena), Ayuku (Yekwana), Camaticaro, Camaticaro rojo, Carpeto, Cozoiba, Cuajo, Cuajo negro, Cuajo pequeño, Erika-bai-yek (Arekuna), Guachinakua (Yekwana), Ibirkabá (Arekuna), Lancetillo, Masarico, Piassám, Sangrino blanco, Sangrito blanco, Tártago, Yopo (Warekena).

746

M YRISTICACEAE

Myristica mocoa A. DC., Prodr. 14: 195. 1856. —Palala mocoa (A. DC.) Kuntze, Revis. Gen. Pl. 2: 567. 1891. —Virola mocoa (A. DC.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 183. 1897. Myristica sebifera var. cordifolia A. DC., Prodr. 14: 195. 1856. Myristica sebifera var. curvinervia A. DC., Prodr. 14: 195. 1856. Myristica panamensis Hemsl., Biol. Cent.Amer., Bot. 3: 67. 1882. —Virola panamensis (Hemsl.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 185. 1897. Virola boliviensis Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 184, t. 7, 1–5. 1897. Virola peruviana var. tomentosa Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 189. 1897. Virola venezuelensis Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 182. 1897. Virola mycetis Pulle, Recueil Trav. Bot. Néerl. 4(1–2): 125. 1907. Virola warburgii Pittier, Contr. U.S. Natl. Herb. 18: 143, t. 57. 1937. Tree 2–30 m tall; leaf blade coriaceous, the base usually cordate, the lower surface uniformly tomentose, but the mature leaves with some tendency toward glabrescence. Evergreen lowland to montane forests, savannas, 50–1400 m; common and widespread in Delta Amacuro, Bolívar, and Amazonas. Barinas, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Táchira, Trujillo, Yaracuy, Zulia; Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. A hallucinogenic snuff is prepared from the inner bark and is used by some Amerindians in their ceremonial dances to cure fevers and to drive away evil spirits. Virola steyermarkii W.A. Rodrigues, Ann. Missouri Bot. Gard. 76: 1163. 1989. Tree 10–25 m tall. Montane tepui forests, 1400–2000 m; Bolívar (Cerro Sarisariñama), Amazonas (Cerro Parú). Endemic. ◆Fig. 617.

Virola surinamensis (Rol. ex Rottb.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 208. 1897. —Myristica surinamensis Rol. ex Rottb., Descr. Rar. Pl. Surin. 14. 1776. —Palala surinamensis (Rol. ex Rottb.) Kuntze, Revis. Gen. Pl. 2: 568. 1891. —Capau-rit-wareiyek (Arekuna), Cedrillo, Cedro blanco, Cuajo, Cuajo morichalero, Diaru (Warao), Guanabillo, Uéi (Arekuna), Wadasa (Yekwana), Warusey-yek (Arekuna). Myristica sebifera var. longifolia Lam., Encycl. 4: 391. 1797. Virola glaziovii Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 219, t. 6, fig. 1–5. 1897. Myristica fatua auct. non Houtt. 1774(?): sensu Sw., Prodr. 96. 1788. Tree to 40 m tall; trunk with buttresses and some aerial roots. Evergreen lowland to montane forests, abundant especially along riparian forests subject to flooding, near sea level to 400 m; widespread in Delta Amacuro, Bolívar, and northern Amazonas. Apure, Aragua, Barinas, Monagas, Sucre; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. The reddish sap of Virola surinamensis is used to seal wounds. The timber is valuable and is used especially in the plywood industry. Virola theiodora (Spruce ex Benth.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 187. 1897. —Myristica theiodora Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 6. 1853. —Palala theiodora (Spruce ex Benth.) Kuntze, Revis. Gen. Pl. 2: 568. 1891. —Ayuko (Yekwana). Virola rufula Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 181. 1897. Tree 5–12 m tall; trunk ca. 10 cm diameter. Evergreen lowland, usually nonflooded forests, 50–400 m; Bolívar (El Cácaro between Rio Caura and Rio Paragua, Santa María de Erebato), Amazonas (El Pozo southeast of San Fernando de Atabapo, Río Casiquiare, Río Sipapo, San Carlos de Río Negro). Amazonian Colombia, Ecuador, Peru, Brazil, Bolivia.

Virola 747

Virola theiodora is closely related to V. sebifera Aubl. and differs from it by its thinner leaves, less conspicuous pubescence, and shorter anthers. Amerindians produce both a narcotic drug and an arrow poison from the resin of the soft inner bark. Virola venosa (Benth.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 224. 1897. —Myristica venosa Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 3. 1853. —Palala venosa (Benth.) Kuntze, Rev. Gen. Pl. 2: 568. 1891. —Jaico (Curripaco), Palo molenillo. Myristica venosa var. poeppigii A. DC. in Mart., Fl. Bras. 5(1): 118. 1860. —Virola venosa var. poeppigii (A. DC.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 225. 1897. Tree 5–15 m tall; upper leaf surface lustrous, veins obscure on both surfaces. Evergreen lowland forests, 100–200 m; Amazonas

(Cerro El Danto near Cerro Yapacana, upper Río Orinoco). Amazonian Colombia, Brazil. Virola sp. A Tree to 25 m tall; lower surface of leaves very quickly glabrescent; veins on lower surface slightly raised and conspicuously darker than intervening leaf surface. Evergreen montane forests, 1000–1100 m; Amazonas (Caño Piedra 75 km southeast of Puerto Ayacucho). Endemic. Virola sp. A is based on Sanoja et al. 3055 (MO, PORT). Virola sp. B. —Cuajo, Palo carbón, Tirripi (Curripaco). Tree 5–25 m tall; flowers yellowish. White-sand scrub (bana), evergreen lowland forests, 50–200 m; Amazonas (Cerro El Danto east of San Juan, middle Río Atacavi, San Carlos de Río Negro).

Fig. 617. Virola steyermarkii

748

M YRSINACEAE

MYRSINACEAE by John J. Pipoly III and Jon M. Ricketson Trees or shrubs, with dioecious, bisexual, androdioecious, or polygamous individuals. Leaves alternate, simple, exstipulate, sessile or petiolate, with pellucid, orange, light brown, or black, rounded and/or linear punctations. Inflorescences racemose, spicate, glomerulate, umbellate, or paniculate with flowers arranged in any of the aforementioned forms. Flowers actinomorphic, bisexual or functionally unisexual. Calyx (3)4 or 5(6)-merous, cotyliform, cupuliform, or patelliform; aestivation valvate, imbricate, or quincuncial, free or connate at base, punctate and/or punctatelineate, persistent; corolla aestivation valvate, imbricate, quincuncial, or contorted, dialypetalous or gamopetalous, rotate, subrotate, funnelform, urceolate, or campanulate, 3–6-merous, with punctations rounded or linear. Stamen number equal to corolla lobes and opposite them; filaments distinct or connate at base forming a tube, the tube developmentally fused to, free from, or adnate to the corolla tube. Ovary superior, unilocular, with basal or central free placentation; ovules few to numerous, 1–many-seriate; style simple or obsolete. Fruit drupaceous, 1-seeded. Pantropics; 42 genera and > 2200 species, 5 genera and 55 species in the flora area. Species of Myrsinaceae are found primarily in wet environments, such as elfin, cloud, montane, and wet forests, and mangrove swamps. Many species are restricted to riparian forests or forest margins, and their presence is indicative of low disturbance levels. Myrsinaceae provide some ornamental species, and some fruits are edible. Throughout its range, indigenous people and other rural area dwellers use the leaves for fish poison, the roots for toothache, and the wood for knife handles. The family has many taxonomic problems because of the small flowers, frequently unisexual plants, and low numbers of individuals. Key to the Genera of Myrsinaceae 1.

1.

2(1).

2.

Inflorescence simple, umbellate, or fasciculate, shorter than the petioles, sessile or on short, perennating, lignified peduncles girdled by persistent floral bracts (short shoots); calyx and corolla lobes strictly valvate in bud; style obsolete or reduced; inflorescence axis never rufous-glandular-papillate or ferruginous-lepidote ...................................... 4. Myrsine Inflorescence simple or compound (paniculate), the flowers in spikes, umbels, corymbs, or racemes, if simple then longer than the petioles, pedunculate but never on short, perennating, lignified and girdled by persistent floral bracts (short shoots); calyx and corolla lobes quincuncial, imbricate, contorted or rarely valvate in bud, when valvate the inflorescence rufous-glandular-papillate or ferruginous-lepidote ............... 2 Filaments connate basally to form a staminal tube adnate to the corolla tube at least 1/5 its length; corolla tube glandular-granulose within at least at junction of tube and lobes ........................................ 3. Cybianthus Filaments free from each other and free from or variously adnate to corolla tube < 1/5 its length, or filaments basally connate to form a staminal tube not adnate to corolla tube; corolla tube not glandulargranulose within .................................................................................... 3

Ardisia 749

3(2).

3.

4(3). 4.

Calyx and corolla contorted in bud, the corolla also twisted; filaments adnate to corolla; anthers less than 3 times longer than wide; inflorescence rachis translucent when fresh ....................................... 5. Stylogyne Calyx and corolla imbricate or quincuncial in bud, the corolla not twisted; filaments free from corolla; anthers at least 3 times longer than wide; inflorescence rachis opaque when fresh ................................................ 4 Corolla lobes nearly free or connate to 1/5 their length; ovules pluriseriate, the placentation free basal ............................................. 1. Ardisia Corolla lobes connate ca. 1/4 their length; ovules uniseriate, the placentation free central ..................................................................... 2. Ctenardisia

1. ARDISIA Sw. Prodr. 3: 48. 1788, nom. cons., non Gaertn. 1790. Icacorea Aubl., Hist. Pl. Guiane 2(suppl.): 1. 1775, nom. rejic. Ardisia subg. Graphardisia Mez in Engl., Pflanzenr. IV. 236(Heft 9): 59, 78. 1902. —Graphardisia (Mez) Lundell, Phytologia 48: 139. 1981. Auriculardisia Lundell, Phytologia 49: 341. 1981. Oerstedianthus Lundell, Phytologia 48: 141. 1981. Zunilia Lundell, Phytologia 49: 353. 1981. Amatlania Lundell, Wrightia 7: 38. 1982. Valerioanthus Lundell, Wrightia 7: 50. 1982. Bisexual trees or shrubs. Leaves petiolate or sessile. Perianth with imbricate or quincuncial aestivation; sepals and petals weakly connate. Stamens included; filaments free from the corolla; anthers ca. 3 times longer than wide, dehiscent by apical or subapical pores, or by longitudinal slits. Ovary ovoid; ovules few to numerous, pluriseriate; style elongate, normally exserted at anthesis; stigma punctiform. Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, eastern and Southeast Madagascar, Asia, Oceania, Australia; ca. 400–500 species, 2–5 in Venezuela, 1 of these in the flora area. In the Neotropics, many authors have segregated various groups from Ardisia as distinct genera, while in the Paleotropics several genera have been synonymized. Phylogenetic analyses of the Ardisia complex of genera will be necessary to resolve the usually disparate classifications recognized worldwide (see Pipoly, Sida 17(2): 445. 1996). Ardisia guianensis (Aubl.) Mez in Urb., Symb. Antill. 2: 392. 1901. —Icacorea guianensis Aubl., Hist. Pl. Guiane 2(suppl.): 1, pl. 368. 1775. —Tinus guianensis (Aubl.) Kuntze, Revis. Gen. Pl. 2: 973. 1891. Ardisia acuminata Willd., Sp. Pl. 1062. 1797. —Icacorea acuminata (Willd.) Lundell, Phytologia 56: 20. 1984. Ardisia amanuensis Lundell, Amer. Midl. Nat. 29(2): 485. 1943. —Icacorea amanuensis (Lundell) Lundell, Phytologia 49: 346. 1981. Ardisia cookii Lundell, Wrightia 4: 56. 1968. —Icacorea cookii (Lundell) Lundell, Phytologia 49: 348. 1981. Ardisia longicaudata Lundell, Wrightia 4: 60. 1968. —Icacorea longicaudata

Fig. 618. Ardisia guianensis

750

M YRSINACEAE

(Lundell) Lundell, Phytologia 49: 349. 1981. Ardisia acuminifolia Lundell, Wrightia 6: 101. 1980. —Icacorea acuminifolia (Lundell) Lundell, Phytologia 49: 346. 1981. Tree to 5 m tall. Gallery and riparian forests, 100–600 m; Delta Amacuro (Río Amacuro, San Victor), Bolívar (upper Río Caura,

Río Paramichi), Amazonas (Río Casiquiare, upper Río Orinoco, Río Ugueto. Anzoátegui, Barinas, Distrito Federal, Lara, Mérida, Monagas, Sucre, Táchira, Yaracuy, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Acre, Pará, Roraima), Bolivia. ◆Fig. 618.

2. CTENARDISIA Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 179. 1930. Yunckeria Lundell, Wrightia 3: 111. 1964. Bisexual shrubs or small trees, glabrous or nearly so. Branchlets stout, terete. Petioles stout. Blades large, lateral veins slender, punctate; margins mostly entire.

Fig. 619. Ctenardisia stenobotrys

Cybianthus 751

Inflorescences terminal umbellate or corymbose panicles. Flowers usually (4)5merous. Pedicels long, thickened above. Sepals dextrorsely imbricate, rounded, thick and punctate medially; petals dextrorsely imbricate, connate into basal tube, linear-oblong, widest at or near middle, punctate medially. Stamens subequaling petals, attached near base of corolla tube; filaments free, shorter than anthers; anthers large, linear-lanceolate, attenuate to apex, erect, rigid, reddish, dehiscent through apical pores, epunctate. Ovary glabrous; style slender, subequaling the petals; stigma minute, punctiform; placenta obovoid, with 6–8 large, erect, uniseriate ovules. Fruit drupaceous, black at maturity. Seed globose, with basal depression extending into center of endosperm; embryo transverse, elongate. Southern Mexico, Central America, Venezuela, northeastern Brazil; 4 species, 1 in Venezuela. Ctenardisia stenobotrys (Standl.) Lundell & Pipoly, Wrightia 7: 44. 1982. —Ardisia stenobotrys Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 170. 1936. —Ctenardisia stenobotrys (Standl.) Fonnegra &

S.L. Jung, Hoehnea 12: 31. 1985, nom. superfl. Tree to 6 m tall. Seasonally flooded forests, 100–200 m; Amazonas (Río Baria, Río Mawarinuma, Río Negro, upper Río Yaciba). Brazil (Amazonas). ◆Fig. 619.

3. CYBIANTHUS Mart., Nov. Gen. Sp. Pl. 3: 87. 1829 [1831], nom. &. typ. cons. Peckia Vell., Fl. Flumin. 51. 1825 [1829]; Fl. Flumin. Icon. pl. 134, 135. 1827 [1831], nom. rejic. Conomorpha A. DC., Trans. Linn. Soc. London 17: 102. 1834. Weigeltia A. DC., Trans. Linn. Soc. London 17: 102. 1834. Grammadenia Benth., Pl. Hartw. 218. 1839 [1846]. Comomyrsine Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 643. 1876. Grammadenia subg. Cybianthopsis Mez in Urb., Symb. Antill. 2: 425. 1901. —Cybianthopsis (Mez) Lundell, Wrightia 4: 68. 1968. Correlliana D’Arcy, Ann. Missouri Bot. Gard. 60: 442. 1973. Dioecious, monoecious, androdioecious, bisexual, or polygamous shrubs or trees to 15 m tall, terrestrial or epiphytic, monoaxial or polyaxial. Branchlets glabrous, glandular-granulose, dendroid- and stellate-tomentose, furfuraceous- or ferruginous-stipitate-lepidote. Leaves sessile or petiolate, alternate, subopposite, or pseudoverticillate, the venation camptodromous (secondary veins curve toward the margins, but do not form loops) or rarely acrodromous (uniting at the apex of the leaf); petioles obsolete, or, when present, canaliculate or marginate, tapering gradually to the base, abruptly swollen at both ends or only toward the base (here termed pulvinate). Inflorescence staminate, pistillate, bisexual, or polygamous, lateral (axillary), a simple raceme, panicle of racemose or spicate (rarely corymbose) branches, a pleiochasium, or an indeterminate umbel appearing racemose. Flowers 3–6(7)merous. Calyx cotyliform to cupuliform, the lobes imbricate, valvate or aberrantly contorted, basally connate 1/5–2/3 their length, abaxially glabrous, glandulargranulose, ferruginous-stipitate-lepidote, or translucent-lepidote, adaxially glabrous, epunctate or prominently orange-, red-, or black-punctate; corolla rotate, subrotate, cupuliform, or campanulate, rarely funnelform or salverform, the lobes imbricate or valvate, basally connate 1/5–3/4 their length, abaxially glabrous, glandular-granulose, or ferruginous-stipitate-lepidote, adaxially glandular-granulose at least at the junction of the tube and lobe, the margin entire to erose-denticulate, glabrous, glandular-granulose or rufous-glandular-papillate. Stamens and staminodes

752

M YRSINACEAE

adnate to corolla tube at least 2/3 their length; filaments variously connate to form a tube, the staminal tube adnate to the corolla tube or at times developmentally fused with it (thus the stamens appearing epipetalous), bearing fleshy lobes alternate with the apically free portions of the filaments or not; anthers erect or distally curved, ovate, widely ovate, or triangular, basifixed or dorsifixed, dehiscent by apical pores, confluent apical pores (birimose), or by wide or narrow longitudinal slits; staminodes morophologically similar to the stamens but greatly reduced in size, the antherodes at times producing abortive pollen. Pistil obnapiform, ellipsoid, umbonate, or obturbinate; ovary sparsely to densely translucent glandular-lepidote; style glabrous; stigma capitate, capitate-lobate, or punctiform, persistent or early caducous, the placenta free-central, carnose, umbonate or globose; ovules (1)2–5(–7), uni- or biseriate; pistillode conic, lageniform, obturbinate, or irregularly shaped and vestigial, hollow or bearing a sterile placenta, rarely absent. Fruit drupaceous, 1(2)seeded; endosperm translucent, nonstarchy; embryo small. Nicaragua, Costa Rica, Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian and Southeastern Brazil, Bolivia; ca. 170 species, 74 in Venezuela, 38 of these in the flora area. Members of Cybianthus are principally riparian, occurring only in primary forests or rarely in somewhat disturbed ones. Key to the Species of Cybianthus 1.

1. 2(1).

2.

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

Corolla cupuliform to campanulate; anthers longer than wide, usually distally recurved in anthesis, mostly apically acute or minutely apiculate .......................................................................................................... 2 Corolla rotate to subrotate; anthers wider than long, always erect, apically rounded or emarginate .......................................................... 28 Branchlets and calyx ferruginous-stipitate-lepidote; calyx lobe margin glabrous (except in C. apiculatus); abaxial corolla surface mostly glabrous along the margin (subgenus Conomorpha) ................................. 3 Branchlets and calyx stellate and/or dendroid-tomentose; calyx lobe margin glandular-ciliate; abaxial corolla surface always glandulargranulose along the margin (subgenus Laxiflorus) ............................ 24 Leaves mostly < 3 cm long (rarely to 3.5 cm in C. huberi), to 2 cm wide ................................................................................................................ 4 Leaves mostly 3.1–24 × (1.5–)2.1–8.5 cm .................................................. 7 Inflorescence (1)2(3)-flowered; upper surface of leaves smooth; anthers rounded to emarginate apically ............................................................ 5 Inflorescence 4–9-flowered; upper surface of leaves densely and minutely pitted; anthers apiculate to cuspidate apically ..................................... 6 Leaves coriaceous, apex abruptly acuminate; flowers sessile; calyx deeply crateriform, 1.9–2.4 mm long ....................................................... C. huberi Leaves cartilaginous, apex acute; flowers on pedicels 3–4.5 mm long; calyx cotyliform, 0.8–1.2 mm long .............................................. C. julianii Upper surface of leaf blades shiny; flowers fleshy; staminate corolla 2.7– 2.9 mm long, pistillate corolla 1.9–2.2 mm long; stamens 1.6–1.7 mm long, the free portion of the filaments flat, < 0.1 mm long; staminodes 1–1.3 mm long, the free portion of the filaments flat, 0.1–0.2 mm long; stigma capitate, 2-lobed ............................................... C. steyermarkianus

Cybianthus 753

6.

7(3). 7. 8(7). 8. 9(8). 9. 10(9).

10. 11(10).

11.

12(9). 12.

13(12). 13. 14(8). 14. 15(14). 15. 16(14).

16.

17(7).

Upper surface of leaf blades dull; flowers coriaceous; staminate corolla 2– 2.3 mm long; pistillate corolla 2.4–2.8 mm long; stamens 1.4–1.5 mm long, the free portion of the filaments terete, ca. 0.1 mm long; staminodes 1.8–2.1 mm long, the free portion of the filaments terete, 0.2– 0.3 mm long; stigma not capitate, 3- or 4-lobed ..................... C. wurdackii Inflorescence paniculate or spicate ........................................................... 8 Inflorescence racemose ............................................................................ 17 Inflorescence paniculate ............................................................................ 9 Inflorescence spicate ................................................................................ 14 Branchlets 5–12 mm diameter; leaves coriaceous .................................. 10 Branchlets 2–3 mm diameter; leaves membranaceous to chartaceous .............................................................................................................. 12 Branchlets subsucculent, the bark yellowish brown; inflorescence tripinnately paniculate; leaf apices rounded, obtuse, truncate, or emarginate ..................................................................................................... C. quelchii Branchlets stiff, not subsucculent, the bark creamish white; inflorescence a simple panicle, the branches racemose; leaf apices acuminate ...... 11 Branchlets terete, prominently furrowed, moderately appressed-lepidote, without lenticels; upper surface of leaves smooth, the margins flat; pedicels 2–3 mm long; calyx patelliform, coriaceous, the lobes oblate to suborbicular; corollas 3–3.5 mm long, apically cucullate ..... C. plowmanii Branchlets angular, densely lepidote, prominently lenticellate; upper surface of leaves densely and minutely pitted, the margins revolute; pedicels 0.5–1 mm long; calyx cupuliform, fleshy, the lobes ovate-triangular to deltate; corollas 2.3–2.7 mm long, apically flat ............. C. amplus Leaves 5–7 cm wide, apically acute; petioles marginate along entire length; inflorescence 8–15 cm long; pedicels curved upward .............. C. cardonae Leaves 5(–6) mm wide, abruptly acuminate apically; petioles canaliculate, marginate only distally; inflorescence > 8 cm long; pedicels erect .............................................................................................................. 13 Leaf margin revolute; calyx cotyliform, fleshy; corolla fleshy ... C. punctatus Leaf margin flat; calyx subcupuliform, chartaceous; corolla chartaceous ............................................................................................... C. guyanensis Flowers subsessile (pedicels 0.2–0.6 mm long); calyx deeply cupuliform, fleshy, the lobes proximally curved; corolla lobes not keeled ............. 15 Flowers sessile; calyx crateriform, coriaceous, the lobes erect to spreading; corolla lobes prominently keeled .................................................. 16 Upper surface of leaves pustulate; inflorescence (2–)6–16 cm long; corolla lobe apex flat; anthers erect ..................................................... C. lepidotus Upper surface of leaves smooth; inflorescence 0.8–1 cm long; corolla lobe apex cucullate; anthers distally recurved ............................ C. sipapoensis Leaves membranaceous, 6–9.5 × (1.5–)2.2–3.5 cm, the upper surface sawdust-like; petioles 1.2–1.5 cm long; corolla coriaceous; anthers proximally recurved, the apex apiculate, glabrous; pistil sessile ........ C. holstii Leaves coriaceous, 2.5–3.5(–4) × 1.2–1.9 cm, the upper surface smooth; petioles 0.5–1 cm long; corolla fleshy; anthers erect, the apex rounded, glandular-papillate; pistil on a fleshy disk .................................. C. huberi Staminate corolla 2–2.5 mm long; staminate calyx 0.7–1.1 mm long; pistillate calyx 0.8–1.2 mm long .................................................................. 18

754

M YRSINACEAE

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

19.

20(17). 20. 21(20).

21.

22(20).

22.

23(22).

23.

24(2).

Staminate corolla 2.5–3.7 mm long; staminate calyx 1.1–1.5 mm long; pistillate calyx 1.1–1.7 mm long ............................................................... 20 Leaves obovate-spatulate, 4–5 cm long, apices obtuse to rounded; corolla fleshy, glandular-granulose within; fruit 7–8 mm diameter ..... C. breweri Leaves elliptic, 7–20 cm long, apices acuminate; corolla membranaceous or chartaceous; fruit 4–7 mm diameter when dried ........................... 19 Branchlets 2–3 mm diameter, the bark beige; upper surface of leaves smooth, the margins flat; petioles 0.5–2 cm long; pedicels to 1 mm long; corolla campanulate; fertile and sterile anthers strongly recurved dorsally, the apical portion of the sterile anthers tightly twisted ............................................................................................... C. guyanensis Branchlets 3.5–4.5 mm diameter, the bark reddish brown; upper surface of leaves pusticulate, the margins revolute; petioles 2–3 cm long; pedicels 2–3.5 mm long; corolla rotate, the lobes reflexed, perpendicular to the tube in anthesis; fertile and sterile anthers slightly recurved dorsally, the apical portion of the sterile anthers straight ........... C. roraimae Calyx cotyliform; corolla glabrous or glandular-ciliate along the margins; staminal tube conspicuous, chartaceous or fleshy, opaque ................ 21 Calyx cupuliform; corolla glandular-granulose along the margins; staminal tube inconspicuous, membranaceous, hyaline ............................. 22 Leaves membranaceous, long-acuminate apically, the upper surface smooth; petioles (1–)1.3–1.7(–2.3) cm long; pedicels 0.6–1 mm long; calyx without scales, the margin glandular-ciliate; corolla lobe apex cucullate; staminal tube fleshy, costate, without lobes, the free portion of the filaments as long as the anthers; fruit 4–7 mm diameter ......... C. apiculatus Leaves chartaceous to coriaceous, rounded to acute apically, the upper surface pustulate; petioles 0.5–1.1 cm long; calyx sparsely lepidote, the margin glabrous; corolla lobe apex flat; staminal tube chartaceous, terete, bearing lobes to 0.1 mm long alternating with the filaments, the free portion of the filaments shorter than the anthers; fruit 3–3.5 mm diameter .............................................................................. C. agostinianus Upper surface of leaves smooth, the margins flat; petioles 0.1–0.2 cm long; inflorescence tortuous; staminate corolla 3.5–3.7 mm long, pistillate corolla 2.7–3.5 mm long .................................................... C. spathulifolius Upper surface of leaves sawdust-like, the lower surface densely lepidote, the margins revolute; petioles 0.1–0.2 cm long; inflorescence straight; staminate corolla 2.5–3 mm long, pistillate corolla 1.9–3.3 mm long .............................................................................................................. 23 Upper surface of leaves pustulate, the lower surface moderately lepidote, the scales not overlapping; pedicels erect, < 1 mm long; calyx lobes longer than broad; staminate corolla 2.7–3 mm long, pistillate corolla 3–3.3 mm long, the lobes flat; free filaments shorter than the anthers; fruit verruculose, 6–8 mm diameter ........................................ C. maguirei Upper surface of leaves densely and minutely pitted, the lower surface densely lepidote, the scales overlapping; pedicels nodding, 1–3 mm long; calyx lobes broader than long; staminate corolla 2.5–2.7 mm long, pistillate corolla 1.9–2 mm long; free filaments longer than the anthers; fruit smooth, 4.5–5 mm diameter when dried ........... C. crotonoides Leaves coriaceous, the lower surface not black or pellucid-punctate, the

Cybianthus 755

24.

25(24). 25.

26(25).

26.

27(24).

27.

28(1). 28. 29(28). 29. 30(29).

30.

31(29).

31.

upper surface with tertiary veins not prominently raised; staminate flowers with free portion of the filaments > 0.55 mm; pistillate flowers with free portion of the filaments longer than 0.5 mm or longer (staminate and pistillate flowers not known for C. deltatus) ....................... 25 Leaves chartaceous to membranaceous, the lower surface obscurely to densely black or pellucid-punctate, the upper surface with tertiary veins prominently raised; staminate flowers with free portions of filaments 0.3–0.5 mm long; pistillate flowers with free portion of filaments < 0.5 mm long ....................................................................................... 27 Calyx equally divided, the lobes deltate, as broad as long; inflorescence rachis black-punctate; branchlets thin, 2–4 mm diameter ...... C. deltatus Calyx unequally divided, the lobes broadly triangular, broader than long; inflorescence rachis epunctate; branchlets thick, (3.5–)5–12 mm diameter ....................................................................................................... 26 Calyx lobes with scattered translucent lepidote scales along outside margins; anthers and antherodes dorsifixed 1/3 from base; staminate flowers with free portion of filaments 1–1.3 mm long; pistillate flowers with free portion of filaments 0.5–0.6 mm long .................................. C. duidae Calyx lobes without translucent lepidote scales along margins; anthers and antherodes dorsifixed 1/4 from base; staminate flowers with free portion of filaments 0.6–0.9 mm long ...................... C. fulvopulverulentus Inflorescence rachis translucent green; calyx lobes apically obtuse to rounded, the margins erose-dentate; staminal and staminodial tube conspicuous; staminate corolla 1.3–5.5 mm long; pistillate corolla 2.2– 2.6 mm long, the antherodes sessile on staminodial tube; receptacle not enlarged in fruit; densely punctate ....................................... C. reticulatus Inflorescence rachis opaque, straw-colored; calyx lobes acute to acuminate apically, at times minutely erose apically; staminal and staminodial tube inconspicuous; staminate corolla 3.6–4.5 mm long; pistillate corolla 2.7–3.5 mm long, the free portion of the filaments 0.3– 0.5 mm long; receptacle or pedicel or both enlarged in fuit; leaves obscurely punctate ......................................................................... C. spicatus Petioles abruptly swollen basally; anthers wider than long, always erect, apically rounded or emarginate (subgenus Weigeltia) ....................... 29 Petioles not abruptly swollen basally or absent; anthers basifixed, poricidally dehiscent ............................................................................ 34 Leaves narrowly oblong to oblanceolate (3.1–)4–5(–7.5) cm wide .......... 30 Leaves elliptic to obovate, (6.3–)8–10(–14) cm wide ............................... 31 Petiole subterete; lower surface of leaf blades not black-punctate-lineate; staminate pedicels ca. 1 mm long; pistillate pedicels 0.5–0.6 mm long ................................................................................................ C. longifolius Petiole canaliculate to base; lower surface of leaf blades conspicuously black-punctate-lineate; staminate pedicels ca. 1.5–2.5 mm long; pistillate pedicels (0.7–)1–1.5 mm long ..................................... C. surinamensis Leaves membranaceous, the lower surface prominently black-punctate or conspicuously black-punctate-lineate, base acuminate or tapering gradually, decurrent on petiole to base; petiole deeply canaliculate; pistillate pedicels thin, 2–2.5 mm long .................................................... 32 Leaves chartaceous to subcoriaceous or cartilaginous, the lower surface

756

M YRSINACEAE

32(31).

32.

33(31).

33.

34(28).

34.

35(34). 35.

36(35).

36.

obscurely pellucid-punctate, base acute, barely decurrent on petiole; petiole subterete with a narrow, shallow channel barely discernible above; pistillate pedicels obsolete to thicker than long and subobsolete .............................................................................................................. 33 Branchlets terete, not brittle, not semisucculent, with numerous, minute, appressed reddish lepidote scales, inconspicuously black-punctatelineate; lower surface of leaves conspicuously black-punctate-lineate, the quaternary veins not visible; petioles 3–4.5 cm long, not decurrent ........................................................................................... C. multicostatus Branchlets angulate, brittle, semisucculent, with scattered, minute, appressed, brownish-lepidote scales, densely and prominently long blackpunctate-lineate; lower surface of leaves densely and prominently black-punctate, the punctations one per areole, formed by prominently raised quaternary veins; petioles 1.5–2 cm long, decurrent on the branchlet onto rounded ridges on branchlet ............................... C. liesneri Branchlets angulate, with few, rounded angles below decurrent petiole bases; petioles 1.3–1.7 cm long, decurrent onto rounded angles on branchlets; leaves thickly coriaceous to cartilaginous; pistillate calyx lobes deltate, apically acute; staminate calyx lobes linear-lanceolate, apically narrowly acute to attenuate ................................. C. grandifolius Branchlets terete, with numerous, raised narrow longitudinal ridges not corresponding to petiole bases; petioles (1.7–)2–3 cm long, not decurrent on branchlet; leaves chartaceous to subcoriaceous; pistillate calyx lobes very widely ovate to oblate, apically subacute to obtuse; staminate calyx lobes ovate, apically obtuse with a small acumen at tip ....................................................................................................... C. potiaei Leaves apically not mucronate, the petioles present, the margins opaque, the venation camptodromous, the bases attenuate, acute, or cuneate; staminal tube developmentally fused to the corolla tube, the stamens thus appearing epipetalous (subgenus Cybianthus) .......................... 35 Leaves apically mucronate, the petioles absent, the margins scarious, the venation acrodromous, the bases auriculate; staminal tube merely adnate to the corolla tube (subgenus Grammadenia) ............................ 38 Branchlets angulate, (2.5–)3.5–5 mm diameter; leaves pseudoverticillate, dull on both surfaces, the quaternary veins not prominent. .............. 36 Branchlets terete, 1.5–3.5 mm diameter; leaves alternate, nitid on the upper surface and/or the lower surface, the quaternary veins prominently raised ........................................................................................ 37 Flowers nodding; calyx lobes widely ovate, apically rounded, densely and prominently black-punctate; staminate flowers with pedicels 2.1– 2.7 mm long; anthers quadrate, sessile on apex of staminal tube, appearing sessile on corolla lobe, apically truncate, the pores confluent or nearly so; pistillate flowers with pedicels 0.9–1.2 mm long; pistil obturbinate ......................................................................... C. venezuelanus Flowers erect; calyx lobes narrowly ovate, apically obtuse to acute, densely and prominently red- to brown-punctate; staminate flowers with pedicels 2.7–3 mm long; anthers subglobose, apical free portion of filaments 0.8–1 mm long, apically broadly rounded, the pores not confluent; pistillate flowers with pedicels 0.7–0.9(–1) mm long; pistil

Cybianthus 757

obnapiform .................................................................................. C. prieurei 37(35). Leaves apically subacuminate to acuminate; flowers nodding, grayish brown, calyx lobes acuminate to attenuate, the margin erose, short glandular-ciliate ....................................................................... C. resinosus 37. Leaves apically acute; flowers erect, greenish, calyx lobes obtuse, the margin crenulate, long-ciliate ........................................... C. penduliflorus 38(34). Branchlets glandular-papillate apically; pistil ellipsoid; trunk thickened basally (pachycaulous); plants of open savannas ...................... C. lineatus 38. Branchlets glabrous apically; pistil obnapiform; trunk not thickened basally; plants of montane or elfin forests and scrub thickets ............... 39 39(38). Leaf margins minutely crenulate to denticulate; ovary costate; staminal tube fleshy, lobed, the lobes punctate, connectives prominently blackpunctate ventrally and dorsally; inflorescence prominently black-punctate-lineate; endemic to Sierra de la Neblina .............................. C. piresii 39. Leaf margins entire; ovary lobed; staminal tube membranous, subtruncate, epunctate, connectives without punctations ventrally, pellucidpunctate dorsally; inflorescence rachis pellucid-punctate; endemic to Auyán-tepui and Macizo del Chimantá complex ................... C. ptariensis Synopsis of the Subgenera of Cybianthus in the Flora Area Cybianthus subg. Conomorpha (A. DC.) G. Agostini, Acta Biol. Venez. 10: 150. 1980. —Conomorpha A. DC., Trans. Linn. Soc. London 17: 102. 1834. Conomorpha sect. Euconomorpha Miq., Stirp. Surinam. Select. 111. 1850 [1851], pro parte. —Conomorpha subg. Euconomorpha (Miq.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 254. 1902, pro parte. Conomorpha sect. Aconomorpha Miq. in Mart., Fl. Bras. 10: 304. 1856. A subgenus of 44 species found in Nicaragua, Costa Rica, Panama, Lesser Antilles, Trinidad, Tobago, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 20 species in the flora area (Cybianthus agostinianus, C. amplus, C. apiculatus, C. breweri, C. cardonae, C. crotonoides, C. guyanensis, C. holstii, C. huberi, C. julianii, C. lepidotus, C. maguirei, C. plowmanii, C. punctatus, C. quelchii, C. roraimae, C. sipapoensis, C. spathulifolius, C. steyermarkianus, C. wurdackii). Cybianthus subg. Cybianthus Peckia Vell., Fl. Flumin. 51. 1825 [1829]; Fl. Flumin. Icon. pl. 134, 135. 1827 [1831], nom. rejic. Cybianthus sect. Cybianthoides Miq. in Mart., Fl. Bras. 10: 292. 1856. Cybianthus sect. Eucybianthus Miq. in Mart., Fl. Bras. 10: 292. 1856. A subgenus of 50 species found in Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, and Bolivia; 4 species in the flora area (Cybianthus penduliflorus, C. prieurei, C. resinosus, C. venezuelanus). Cybianthus subg. Grammadenia (Benth.) Pipoly, Mem. New York Bot. Gard. 43: 47. 1987. —Grammadenia Benth., Pl. Hartw. 218. 1839 [1846]. Grammadenia subg. Eugrammadenia Mez in Urb., Symb. Antill. 2: 425. 1901. A subgenus of 7 species found in Costa Rica, Panama, West Indies, Colombia,

758

M YRSINACEAE

Venezuela, Guyana, Ecuador, Peru, and Brazil; 3 species in the flora area (Cybianthus lineatus, C. piresii, C. ptariensis). Cybianthus subg. Laxiflorus G. Agostini, Acta Biol. Venez. 10: 144. 1980. Conomorpha sect. Conomorphida Miq., Stirp. Surinam. Select. 111. 1850 [1851], pro parte. Conomorpha subg. Euconomorpha Mez in Engl., Pflanzenr. IV. 236(Heft 9): 254. 1902, pro parte minore. A subgenus of 6 species found in Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 5 species in the flora area (Cybianthus deltatus, C. duidae, C. fulvopulverulentus, C. reticulatus, C. spicatus). Cybianthus subg. Weigeltia (A. DC.) G. Agostini, Acta Biol. Venez. 10: 156. 1980. —Weigeltia A. DC., Trans. Linn. Soc. London 17: 102. 1834. —Cybianthus sect. Weigeltia (A. DC.) Miq. in Mart., Fl. Bras. 10: 299. 1856. A subgenus of 46 species found in Panama, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and Bolivia; 6 species in the flora area (Cybianthus grandifolius, C. liesneri, C. longifolius, C. multicostatus, C. potiaei, C. surinamensis).

Cybianthus Species Entries Cybianthus agostinianus Pipoly, Ernstia 50: 33, fig. 10. 1988. [Subgenus Conomorpha]. Shrub or small tree to 4 m tall. Open, rocky areas on tepui summits, 1500–2700 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. Cybianthus amplus (Mez) G. Agostini, Acta Biol. Venez. 10: 151. 1980. —Conomorpha ampla Mez in Engl., Pflanzenr. IV. 236(Heft 9): 257. 1902. [Subgenus Conomorpha]. Conomorpha macrophylla Mart., Flora 24(2): 20. 1841, [Herb. Fl. Brasil. 260. 1840.], non Cybianthus macrophyllus Miq. & Mart. 1856. Conomorpha utiarityi Hoehne, Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, anexo 6: 64. 1915. Shrub or small tree to 7 m tall. Riparian forests, 100–1100 m; Bolívar (Río Ichún, Río Paragua), Amazonas (Cerro Cariche, Cerro Duida, upper Río Casiquiare, Río Cunucunuma, Río Negro, Río Orinoco, Río Yavita, Sierra de la Neblina). Colombia (Vaupés), Guyana, French Guiana, Peru, Brazil (Amazonas). ◆Fig. 623.

Cybianthus apiculatus (Steyerm.) G. Agostini, Acta Biol. Venez. 10: 168. 1980. —Conomorpha apiculata Steyerm., Fieldiana, Bot. 28: 456. 1953. [Subgenus Conomorpha]. Shrub or small tree to 3 m tall. Evergreen montane forests, 1000–1600 m; Amazonas (Cerro Cuao, Cerro Duida, Cerro Marahuaka, Río Iguapo, Río Ventuari, Sierra de la Neblina). Guyana (Mount Ayanganna). ◆Fig. 625. Cybianthus breweri G. Agostini, Bol. Soc. Venez. Ci. Nat. 22: 384. 1976. [Subgenus Conomorpha]. Shrub or small tree to 3 m tall. Tepui thickets, gallery forests, 1400–2100 m; Bolívar (Cerro Guanacoco, Cerro Jaua). Endemic. Cybianthus cardonae G. Agostini, Bol. Soc. Venez. Ci. Nat. 22: 386. 1976. [Subgenus Conomorpha]. Shrub or tree to 8 m tall. Montane cloud forests, 500–1800 m; Bolívar (Cerro Jaua, upper Río Paragua), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Sierra de la Neblina). Brazil (Amazonas: Serra Aracá).

Cybianthus 759

Cybianthus crotonoides (R.M. Schomb. ex Mez) G. Agostini, Acta Biol. Venez. 10: 153. 1980. —Conomorpha crotonoides R.M. Schomb. ex Mez in Engl., Pflanzenr. IV. 236(Heft 9): 262. 1902. [Subgenus Conomorpha]. Shrub to 4 m tall. On fissures of rocks, associated with Bonnetia roraimae formations, 1800–2500 m; Bolívar (Aparamán-tepui, Auyán-tepui, Carrao-tepui, Kamarkawarai-tepui, Murisipán-tepui, Ptari-tepui, Terekéyurén-tepui). Endemic. ◆Fig. 621. Cybianthus deltatus Pipoly, Brittonia 35: 65. 1983. [Subgenus Laxiflorus]. Shrub to 3 m tall. Granitic outcrops, savanna edges, 100–800 m; Amazonas (summit of Cerro Aracamuni, near Cerro Sipapo, Río Guainía, Río Guasacavi, near Santa Cruz). Endemic. ◆Fig. 622. Cybianthus duidae (Gleason & Moldenke) G. Agostini, Acta Biol. Venez. 10: 145. 1980. —Conomorpha duidae Gleason & Moldenke, Bull. Torrey Bot. Club 58: 445. 1931. [Subgenus Laxiflorus]. Conomorpha ptariensis Steyerm., Fieldiana, Bot. 28: 466. 1953. Shrub or small tree to 3 m tall. Savannas, sandstone formations, 400–1500 m; Bolívar (Auyán-tepui, Ilú-tepui, Luepa to Santa Elena, Murisipán-tepui, lower Río Caroní, Uaipán-tepui), Amazonas (Cerro Duida, Cerro Guaiquinima, Cerro Yutajé, Río Manipiare, Sierra de Magualida). Guyana (Mount Ayanganna). Cybianthus fulvopulverulentus (Mez) G. Agostini, Bol. Soc. Venez. Ci. Nat. 32: 388. 1976. —Conomorpha fulvopulverulenta Mez in Engl., Pflanzenr. IV. 236(Heft 9): 258. 1902. [Subgenus Laxiflorus]. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 subspecies, both in the flora area. Key to the Subspecies of C. fulvopulverulentus 1. Leaves, inflorescences, and calyces densely stellate, dendroid-ferruginous-tomentose, and glandular-granulose; calyx lobes rounded to acute apically; staminate in-

florescence often tortuous; flowers with staminal tube 1.1–1.3 mm long; pistillate flowers with free portion of filaments 0.7–0.9 mm long; restricted to sandstone and quartzite formations of the superimposed Roraima sediments in the Guayana Shield, mostly above 800 m .................. subsp. fulvopulverulentus 1. Leaves, inflorescences, and calyces glabrous or glandular-granulose; calyx lobes acute to acuminate apically; staminate inflorescences never contorted; the flowers with staminal tube 0.8–1.1 mm long; pistillate flowers with free portion of filaments ca. 0.7 mm long; riparian or restricted to wet savannas and lower talus slopes of sandstone formations on the fringe of the Guayana Shield, occurring mostly below 800 m ......................... ......................... subsp. magnoliifolius C. fulvopulverulentus subsp. fulvopulverulentus Conomorpha riparia R.E. Schult., Bot. Mus. Leafl. 15: 72. 1951. Conomorpha leprosa Steyerm., Fieldiana, Bot. 28: 465. 1958. Conomorpha lithophyta R.E. Schult., Bot. Mus. Leafl. 18: 166. 1958. —Cybianthus lithophytus (R.E. Schult.) G. Agostini, Acta Biol. Venez. 10: 145. 1980. Shrub to 10 m tall. Evergreen lowland forests on sandy soils, montane and tepui forests, 100–2200 m; Bolívar (Auyán-tepui, Cerro Akurimá along middle Río Caroní, Cerro Guaiquinima, Cerro Jaua, Cerro Sarisariñama, Ilú-tepui, Macizo del Chimantá [Toronó-tepui]), Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Parú, Río Manipiare, San Carlos de Río Negro, Serranía Yutajé, Sierra de la Neblina). Colombia, Guyana, Suriname, Brazil. ◆Fig. 626. C. fulvopulverulentus subsp. magnoliifolius (Mez) Pipoly, Brittonia 35: 72. 1983. —Conomorpha magnoliifolia Mez in Engl., Pflanzenr. IV. 236(Heft 9): 258. 1902. —Cybianthus magnoliifolius (Mez) G. Agostini, Acta Biol. Venez. 10: 146. 1980. Conomorpha rigida Mez, Repert. Spec. Nov. Regni Veg. 16: 420. 1920. Shrub to 10 m tall. Forest-savanna ecotones, 50–1200 m; Delta Amacuro (Caño

760

M YRSINACEAE

Simoina west of Isla Cocuina), Bolívar (Macizo del Chimantá), Amazonas (upper Río Casiquiare). Guyana, Suriname, French Guiana, Brazil (Amazonas, Matto Grosso). Cybianthus grandifolius (Mez) G. Agostini, Acta Biol. Venez. 10: 159. 1980. —Weigeltia grandifolius Mez in Engl., Pflanzenr. IV. 236(Heft 9): 287. 1902. [Subgenus Weigeltia]. Weigeltia sylvatica Gleason, Contr. New York Bot. Gard. 282: 294. 1926. —Cybianthus sylvaticus (Gleason) G. Agostini, Acta Biol. Venez. 10: 162. 1980. Tree to 8 m tall. Talus forests on steep slopes, ca. 1200 m; Bolívar (Cerro Guaiquinima). Guyana (along Potaro River). Cybianthus guyanensis (A. DC.) Miq. in Mart., Fl. Bras. 10: 298. 1856. —Conomorpha guyanensis A. DC., Ann. Sci. Nat., Bot., sér. 2, 16: 92. 1841. —Peckia guyanensis (A. DC.) Kuntze, Revis. Gen. Pl. 2: 402. 1891. —Conomorpha peruviana var. β guyanensis (A. DC.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 236. 1902. [Subgenus Conomorpha]. Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas, Pará); 3 subspecies, all in the flora area,

staminal tube 0.9–1.1 mm long; apically free portions of the filaments 0.2– 0.4 mm long .......... subsp. guyanensis 2. Petioles canaliculate, 0.5–0.8(–1.5) cm long; staminate peduncle 0.1–0.2 cm long; floral bracts longer than the pedicels, 1.3–1.5 mm long; pedicels 1– 1.2 mm long; corolla carnose, 2.8–3.2 mm long, the lobes ovate; staminal tube 1.2–1.6 mm long; apically free portions of the filaments 0.4–0.6(–0.7) mm long ...................... subsp. pseudoicacoreus C. guyanensis subsp. guyanensis Conomorpha heterantha Benth. ex Miq. in Mart., Fl. Bras. 10: 304. 1856. Shrub or small tree to 5 m tall. Seasonally flooded areas along black-water rivers, 100– 200 m; Amazonas (Río Baría, Río Mawarinuma, Río Negro). Brazil (Amazonas, Pará). C. guyanensis subsp. multipunctatus (A. DC.) Pipoly, Ann. Missouri Bot. Gard. 79: 944, fig. 14. 1992. —Cybianthus multipunctatus A. DC., Ann. Sci. Nat. Bot. sér. 2, 16: 94. 1841. Shrub or small tree to 8 m tall. Mauritia palm swamps, ca. 100 m; Bolívar (between Cerro Gavilán and Río Horeda). Guyana, Suriname, French Guiana, Brazil (Pará).

Key to the Subspecies of C. guyanensis 1. Leaf blades asymmetric; calyx cotyliform; corolla membranaceous, the lobes reflexed-recurved; apically free portions of the filaments longer than the anthers; anthers ovate, apically acute, dehiscent by wide, sublatrorse slits ........................ ........................ subsp. multipunctatus 1. Leaf blades symmetric; calyx subcupuliform; corolla chartaceous or carnose, the lobes erect to spreading; apically free portions of the filaments shorter than the anthers; anthers narrowly triangular or ovate-triangular, apically apiculate, dehiscent by narrow, introrse slits ............................................................... 2 2. Petioles canaliculate and winged, (1–)1.5– 1.9(–2.3) cm long; staminate peduncle 0.2–0.5 cm long; floral bracts shorter than the pedicels, 0.7–0.8 mm long; pedicels 0.5–1 mm long; corolla chartaceous, 2.3–2.6 mm long, the lobes elliptic;

C. guyanensis subsp. pseudoicacoreus (Miq.) Pipoly, Sida 18: 49. 1998. —Ardisia pseudoicacorea Miq. in Mart., Fl. Bras. 10: 284. 1856. —Cybianthus pseudoicacoreus (Miq.) G. Agostini, Acta Biol. Venez. 10: 155. 1980. Shrub or tree to 12 m tall. Riparian forests, 100–600 m; Amazonas (115 km southeast of Puerto Ayacucho, Río Cuao, Serranía Batata in Cuao-Sipapo massif). Guyana, Ecuador, Peru. Cybianthus holstii Pipoly, Ann. Missouri Bot. Gard. 79: 938, fig. 12. 1992. [Subgenus Conomorpha]. Shrub to 2 m tall. Tepui-slope forests, 500–1000 m; Amazonas (Cerro Guanay, Cerro Yutajé, Río Coro Coro). Endemic. Cybianthus huberi Pipoly, Ann. Missouri Bot. Gard. 79: 918, fig. 3. 1992. [Subgenus Conomorpha].

Cybianthus 761

Shrub to 3 m tall. Tepui scrub, ca. 1700 m; Bolívar (Cerro Guanay). Endemic. Cybianthus julianii Pipoly, Ann. Missouri Bot. Gard. 79: 920, fig. 4. 1992. [Subgenus Conomorpha]. Shrub to 1.5 m tall. Boggy areas on tepui summits, 2500–2600 m; Amazonas (Cerro Marahuaka). Endemic. Cybianthus lepidotus (Gleason) G. Agostini, Bol. Soc. Venez. Ci. Nat. 22: 388. 1976. —Conomorpha lepidota Gleason, Bull. Torrey Bot. Club 58: 446. 1931. [Subgenus Conomorpha]. Conomorpha curvivenia Gleason, Bull. Torrey Bot. Club 58: 444. 1931. Conomorpha lepidota f. acutata Steyerm., Fieldiana, Bot. 28: 465. 1953. Shrub or small tree to 6 m tall. Lower to upper montane forests, 600–2300 m; Bolívar (Cerro Jaua, Macizo del Chimantá [Amurítepui, Apacará-tepui]), Amazonas (Caño Iguapo, Cerro Duida, Cerro Parú, Sierra de la Neblina). Guyana, Peru, Brazil (Amazonas: Serra Aracá), Bolivia. ◆Fig. 627. Cybianthus liesneri Pipoly & Ricketson, Sida 18: 1168, fig. 1. 1999. [Subgenus Weigeltia]. Tree to 4 m tall. Evergreen lowland forests, 100–200 m; Amazonas (base of Sierra de la Neblina). Endemic. Cybianthus lineatus (Benth.) Pipoly, Mem. New York Bot. Gard. 43: 64. 1987. —Grammadenia lineata Benth., Pl. Hartw. 218. 1839 [1846]. [Subgenus Grammadenia]. Shrub to 1.5 m tall. Tepui meadows and scrub, 1200–2000 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Jaua, Macizo del Chimantá [Amurí-tepui, Apacará-tepui, Chimantá-tepui], Roraima-tepui), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Cerro Yaví, Cerro Yutajé). Guyana, Peru. ◆Fig. 620. Cybianthus longifolius Miq. in Mart., Fl. Bras. 10. 299. 1856. —Weigeltia longifolia (Miq.) Benth. in Mart., Fl. Bras. 10. 299. 1856. —Peckia longifolia (Miq.) Kuntze, Revis. Gen. Pl. 2: 402. 1891. [Subgenus Weigeltia].

Shrub to 2 m tall. Riparian forests, 100– 500 m; Amazonas (Río Cataniapo, Rio Negro). Colombia, Ecuador, Brazil, Bolivia. Cybianthus maguirei G. Agostini ex Pipoly, Ann. Missouri Bot. Gard. 79: 952, fig. 18. 1992. [Subgenus Conomorpha]. Shrub or small tree to 2 m tall. Tepui bogs and meadows, 1300–2000 m; Amazonas (Sierra de la Neblina). Endemic. Cybianthus multicostatus Miq. in Mart., Fl. Bras. 10: 299. 1856. [Subgenus Weigeltia]. Tree to 15 m tall. Nonflooded forests, 50– 100 m; Amazonas (near Puerto Ayacucho). Brazil. Cybianthus penduliflorus Mart., Nov. Gen. Sp. Pl. 3: 87. 1831 [1829]. —Peckia penduliflora (Mart.) Kuntze, Revis. Gen. Pl. 2: 402. 1891. [Subgenus Cybianthus]. Shrub or tree to 4 m tall. Seasonally flooded forests near black-water rivers, on banks and hummocks, 100–200 m; Amazonas (Río Negro, lower Río Orinoco). Colombia, Ecuador, Peru, Brazil, Bolivia. Cybianthus piresii Pipoly, Mem. New York Bot. Gard. 43: 65. 1987. [Subgenus Grammadenia]. Shrub to 2 m tall. Montane scrub and rock outcrops, 1700–2200 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Cybianthus plowmanii Pipoly, Ann. Missouri Bot. Gard. 79: 925, fig. 7. 1992. [Subgenus Conomorpha]. Shrub or tree to 8 m tall. Tepui scrub, 1700–2000 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra Aracá). Cybianthus potiaei (Mez) G. Agostini, Acta Biol. Venez. 10: 161. 1980. —Weigeltia potiaei Mez in Engl., Pflanzenr. IV. 236(Heft 9): 285. 1902. [Subgenus Weigeltia]. Tree to 5 m tall. Lower montane forests, 200–900 m; Bolívar (Betania on Santa Elena de Uairén to Icabarú road, El Paují, Rio Samay, Sierra de Maigualida). Guyana, Suriname, French Guiana, Brazil.

762

M YRSINACEAE

Cybianthus prieurei A. DC., Ann. Sci. Nat. Bot., sér. 2, 16: 93. 1841. —Peckia prieureii (A. DC.) Kuntze, Revis. Gen. Pl. 2: 402. 1891. [Subgenus Cybianthus]. Cybianthus nitidus Miq. in Mart., Fl. Bras. 10: 295. 1856. Cybianthus subspicatus Benth. ex Miq. in Mart., Fl. Bras. 10: 296. 1856. Cybianthus comatus Mez in Engl., Pflanzenr. IV. 236(Heft 9): 219. 1902 Cybianthus viridiflorus A.C. Sm., Lloydia 2: 202. 1939. Tree to 10 m tall. Evergreen lowland to montane forests, 100–1000 m; Bolívar (Auyán-tepui, between Betania and Santa Elena de Uairén, El Carmen, Kavanayén, middle Río Paragua, middle Río Orinoco east of Pijiguaos), Amazonas (Río Negro). Guyana, Suriname, French Guiana, Brazil (Acre, Amapa, Bahia, Pará). Cybianthus ptariensis (Steyerm.) Pipoly, Mem. New York Bot. Gard. 43: 66. 1987. —Grammadenia ptariensis Steyerm., Fieldiana, Bot. 28: 476. 1953. [Subgenus Grammadenia]. Grammadenia ptariensis subsp. auyantepuiensis G. Agostini, Acta Bot. Venez. 2: 283. 1967 Shrub or small tree to 3 m tall. Tepui scrub, 1600–2500 m; Bolívar (Auyán-tepui, Ilú-tepui, Macizo del Chimantá [Abacapátepui, Amurí-tepui, Apacará-tepui, Chimantá-tepui, Churí-tepui], Ptari-tepui). Guyana. ◆Fig. 634. Cybianthus punctatus (Mez) G. Agostini, Acta Biol. Venez. 10: 155. 1980. —Conomorpha punctata Mez in Engl., Pflanzenr. IV. 236(Heft 9): 260. 1902. [Subgenus Conomorpha]. Conomorpha sessilis A.C. Sm., Bull. Torrey Bot. Club 67: 294. 1940. Shrub or small tree to 3 m tall. Montane forests, 900–1400 m; Bolívar (Auyán-tepui, Gran Sabana, Macizo del Chimantá, Roraima-tepui, trail to Urimán). Guyana. ◆Fig. 629. Cybianthus quelchii (N.E. Br.) G. Agostini, Bol. Soc. Venez. Ci. Nat. 22: 388. 1976. —Ardisia quelchii N.E. Br., Trans. Linn. Soc., Bot. 6: 46. 1901. —Weigeltia quelchii (N.E. Br.) Mez in Engl., Pflanzenr.

IV. 236(Heft 9): 288. 1902. [Subgenus Conomorpha]. Conomorpha depressa Steyerm., Fieldiana, Bot. 28: 458. 1953. Shrub or tree to 5 m tall. Tepui scrub, gallery forests, 1800–2500 m; Bolívar (Aparamán-tepui, Auyán-tepui, Cerro Guaiquinima, Macizo del Chimantá [Acopán-tepui, Churí-tepui, Chimantá-tepui, Toronó-tepui], Ptari-tepui, Roraima-tepui, Serranía Marutaní), Amazonas (Serranía Sipapo, Sierra de la Neblina). Guyana (Mount Ayanganna), Brazil (Amazonas, Roraima). ◆Fig. 633. Cybianthus quelchii is extremely variable throughout its range. Individuals occurring at the edges of bogs often have smaller leaves and larger inflorescences, while those of the gallery forests have larger leaves and more compact inflorescences. Cybianthus resinosus Mez in Engl., Pflanzenr. IV. 236(Heft 9): 219. 1902. [Subgenus Cybianthus]. Tree to 15 m tall. Lowland to upland forests, 100–600(–1300) m; Amazonas (Cerro Huachamacari, Cerro Yutajé, Río Negro). Colombia (Chocó), Ecuador (Napo), Peru (Huánuco, Loreto). Cybianthus reticulatus (Benth. ex Mez) G. Agostini, Acta Biol. Venez. 10: 146. 1980. —Conomorpha reticulata Benth. ex Miq. in Mart., Fl. Bras. 10: 303. 1856. [Subgenus Laxiflorus] Shrub or small tree 1–8 m tall. Whitesand savannas, 100–200(–700) m; Amazonas (Caño Pimichín, Río Casiquiare basin, Río Negro, Río Siapa, Río Temi, San Carlos de Río Negro, Santa Bárbara del Orinoco). Brazil. Cybianthus roraimae (Steyerm.) G. Agostini, Acta Biol. Venez. 10: 155. 1980. —Conomorpha roraimae Steyerm., Fieldiana, Bot. 28: 468. 1953. [Subgenus Conomorpha]. Shrub or small tree to 5 m tall. Montane to upper montane forests, 1100–2600 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Churítepui], Roraima-tepui). Guyana. ◆Fig. 628. Cybianthus sipapoensis Pipoly & G. Agostini, Ernstia 50: 36, fig. 11. 1988. [Subgenus Conomorpha].

Cybianthus 763

Shrub to small tree ca. 2 m tall. Steep slopes of moist forests, 1600–2200 m; Amazonas (Cerro Sipapo). Endemic. Cybianthus spathulifolius G. Agostini ex Pipoly, Ann. Missouri Bot. Gard. 79: 950, fig. 17. 1992. [Subgenus Conomorpha]. Shrub or small tree ca. 2 m tall. Scrub forests, 2000–2200 m; Amazonas (Cerro Sipapo, Río Cuao, Río Orinoco). Endemic. Cybianthus spicatus (H.B.K.) G. Agostini, Acta Biol. Venez. 10: 146. 1980. —Myrsine spicata H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 250. 1818 [1819]. —Conomorpha spicata (H.B.K.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 259. 1902. [Subgenus Laxiflorus]. Walleria laxiflora Mart., Nov. Gen. Sp. Pl. 3: 89. 1829. —Conomorpha laxiflora A. DC., Trans. Linn. Soc. London 17: 102. 1834. Conomorpha laxiflora var. latifolia Miq. in Mart., Fl. Bras. 10: 303. 1856. —Conomorpha latifolia (Miq.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 255. 1902. Conomorpha candolleana Mez in Engl., Pflanzenr. IV. 236(Heft 9): 256. 1902. Conomorpha glaucorubens Mez in Engl., Pflanzenr. IV. 236(Heft 9): 260. 1902. Conomorpha grandiflora Mez in Engl., Pflanzenr. IV. 236(Heft 9): 258. 1902. Conomorpha madeirensis A.C. Sm., J. Arnold Arbor. 20: 300. 1931. Conomorpha gracilis A.C. Sm., Bull. Torrey Bot. Club 67: 295. 1940. Shrub or small tree to 12 m tall. Blackwater or riparian forests, 100–600 m; Bolívar (Cerro Guaiquinima, Laguna de Canaima, Río Acanán, Río Asa, upper Río Caura), Amazonas (Caño San Miguel, Cerro Aracamuni, Cerro Marahuaka, near Chapezón, Río Casiquiare, Río Mawarinuma, Río Ocamo, Río Pasimoní, Río Siapa, San Fernando de Atabapo, near Solano). Apure; Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas, Roraima). ◆Fig. 632. Cybianthus spicatus is a broad-ranging polymorphic species with many semi-isolated populations throughout the Amazon basin and eastern Guayana Shield. These localized populations have produced seemingly distinct ecotypes, resulting in overdescription. Although four basic ecotypes are

discernible, variation within each one is not mutually exclusive and many collections do not respond to any one category (discussed in Pipoly, Brittonia 35: 76. 1983). Additional fieldwork is needed to understand the variation present in this ochlospecies and its ecological life history. Due to the paucity of pistillate flowering specimens, the classification of ecotypes could be oversimplified. More collections are needed for this species. Cybianthus steyermarkianus (G. Agostini) G. Agostini, Acta Biol. Venez. 10: 156. 1980. —Conomorpha steyermarkiana G. Agostini, Acta Bot. Venez. 2: 283. 1967. [Subgenus Conomorpha]. Shrub to 4 m tall. Tepui outcrops, 1900– 2500 m; Bolívar (Apradá-tepui, Macizo del Chimantá [Abacapá-tepui, Acopán-tepui, Apacará-tepui, Chimantá-tepui, Churí-tepui, Toronó-tepui]). Endemic. ◆Fig. 631. Cybianthus surinamensis (Spreng.) G. Agostini, Acta Biol. Venez. 10: 161. 1980. —Salvadora surinamensis Spreng., Syst. Veg. 5(tent. suppl.): 7. 1828. —Peckia surinamensis (Spreng.) Kuntze, Revis. Gen. Pl. 2: 402. 1891. —Weigeltia surinamensis (Spreng.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 289. 1902. [Subgenus Weigeltia]. Tree to 5 m tall. Riparian forests, 50–1200 m; Delta Amacuro (Río Matanaima near Río Amacuro, San Victor on Sierra Imataca), Bolívar (Altiplanicie de Nuria, Cerro Guaiquinima, El Dorado, Macizo del Chimantá [Amurí-tepui], Ptari-tepui, Río Villacoa, Río Zariapo in upper Río Caura basin, Serranía Cerbatana), Amazonas (Cerro Duida, Cerro Marahuaka, Cerro Yutajé, Río Coro Coro, Río Cunucunuma, Río Guayapo, Río Iguapo, Río Yureba). Guyana, Suriname, Brazil (Amapá, Roraima). ◆Fig. 630. Cybianthus venezuelanus Mez in Engl., Pflanzenr. IV. 236(Heft 9): 221. 1902. [Subgenus Cybianthus]. Cybianthus egensis Mez in Engl., Pflanzenr. IV.236 (Heft 9): 222. 1902. Peckia purpurea Rusby, Bull. New York Bot. Gard. 4: 405. 1907. Cybianthus brownii Gleason, Bull. Torrey Bot. Club 53: 293. 1926. Shrub to 3 m tall. Rocky outcrops, lowland

764

M YRSINACEAE

Fig. 620. Cybianthus lineatus

Fig. 621. Cybianthus crotonoides

Fig. 623. Cybianthus amplus

Fig. 622. Cybianthus deltatus

Cybianthus 765

Fig. 624. Cybianthus venezuelanus

Fig. 625. Cybianthus apiculatus

Fig. 626. Cybianthus fulvopulverulentus subsp. fulvopulverulentus

766

M YRSINACEAE

Fig. 627. Cybianthus lepidotus

Fig. 629. Cybianthus punctatus

Fig. 628. Cybianthus roraimae

Cybianthus 767

Fig. 630. Cybianthus surinamensis

Fig. 631. Cybianthus steyermarkianus

Fig. 632. Cybianthus spicatus

768

M YRSINACEAE

Fig. 633. Cybianthus quelchii

to upland forests, 100–1200 m; Bolívar (Río Paragua), Amazonas (Cerro Huachamacari, Cerro Marahuaka, 25 km south of Puerto Ayacucho, Río Baría). Aragua, Carabobo, Cojedes, Distrito Federal, Lara, Táchira, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil (Amazonas, Serra Aracá), Bolivia. ◆Fig. 624.

Fig. 634. Cybianthus ptariensis

Cybianthus wurdackii G. Agostini ex Pipoly, Ann. Missouri Bot. Gard. 79: 922, fig. 6. 1992. [Subgenus Conomorpha]. Treelet to 3 m tall. Scrub forests on tepui summits, 2200–2300 m; Bolívar (Auyán-tepui, Kamarkawarai-tepui, Macizo del Chimantá [Murey-tepui, Sarvén-tepui], Ptaritepui). Endemic.

4. MYRSINE L., Sp. Pl. 196. 1753; Gen. Pl. ed 5: 90. 1754. Rapanea Aubl., Hist. Pl. Guiane 121, t. 46. 1775. Samara Sw., Prodr. 1: 120. 1788, pro parte, non L. 1771. Manglilla A. Juss., Gen. Pl. 151. 1789. Caballeria Ruiz & Pav., Fl. Peruv. Prodr. 1: 141. 1794. Roemeria Thunb., Nov. Gen. Pl. 9: 130. 1798, non Medik. 1792, non Moench 1794. Scleroxylum Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 3: 57. 1809. Normally unisexual, rarely bisexual, monoecious, dioecious, or polygamous shrubs or small trees. Leaves alternate, exstipulate. Inflorescences lateral (axillary), umbellate or fasciculate, sessile or on short, perennating, accrescent peduncles girdled by persistent floral bracts (thus forming short shoots). Flowers 4- or 5(6)merous. Sepals nearly free or united to 1/2 their length, imbricate or valvate, usually ciliate, punctate, persistent; petals nearly free or rarely united to 1/2 their length, usually ciliate, glandular-granulose at least along margin and often throughout

Myrsine 769

within, punctate. Stamens and staminodes similar, subequaling corolla length; filaments free or connate basally to form a tube, the tube with or without sterile appendages alternating with the filaments, and all merely adnate to the corolla tube, or developmentally fused throughout, the anthers or antherodes thus appearing epipetalous; anthers and antherodes similar, ovate or reniform, elliptic or oblong, rarely sagittate, 2-celled, dehiscing by longitudinal slits, or rarely by subterminal pores opening later into wide longitudinal slits. Pistil and pistillode similar, conic, ellipsoid, obturbinate, obnapiform, or variously subglobose; ovary globose, costate or not, glabrous or glabrescent; ovules few, uniseriate, completely immersed in placenta or seated below apical pores in placenta or variously projecting; style obsolete to present, tapering into stigma; stigma morchelliform, liguliform, sinuate to lobate, prismatic and 3(4)-lobed, or rarely conical. Fruit a drupe, with somewhat fleshy exocarp and crusty or leathery endocarp, 1-seeded. Seed occupying cavity, the endosperm horny, ruminate; embryo cylindric, transverse. Pantropics; ca. 300 species, 12 in Venezuela, 10 of these in the flora area. Key to the Species of Myrsine 1.

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

6.

7(5). 7. 8(7).

8.

Branchlets, petioles, and/or midrib of leaf blade generally densely ferruginous to rufous-tomentose, at times early glabrescent, the trichomes uniseriate ................................................................................... M. coriacea Branchlets, petioles, and midrib of leaf blade glabrous or reddish glandular-papillose ............................................................................................ 2 Petioles 0–5 mm long ................................................................................. 3 Petioles 5–20 mm long ............................................................................... 5 Leaf blades (2.3–)3–5 cm wide; petioles thick, 2–3 mm diameter ............................................................................................. M. maguireana Leaf blades 1–2(–2.5) cm wide; petioles thinner, 0–1.5 mm diameter ..... 4 Leaf blades (1.7–)2–3 × 1–1.5 cm, the apex emarginate or retuse; pedicels 1.5–2 mm long ............................................................................ M. minima Leaf blades 4–6.5 × 1.5–3 cm, the apex obtuse or rounded; pedicels 0.5– 1.5 mm long ....................................................................... M. perpauciflora Leaf blades 1–2 cm wide ............................................................................ 6 Leaf blades 2–9 cm wide ............................................................................ 7 Leaf blades 6–7 times longer than wide, 4–15 × 1–2 cm, lorate, the lower surface with few, conspicuous punctate-lineations, 0.5–4 cm long; pedicels 2.5–3.5 mm long .......................................................... M. resinosa Leaf blades 2–3 times longer than wide, 3.5–6 × 1.5–2 cm, oblong to obovate or oblanceolate, the lower surface with many, inconspicuous punctate-lineations, 0.05–0.1 cm long; pedicels 1–1.5 mm long .... M. picturata Pedicels 0–1.4 mm long .............................................................................. 8 Pedicels 1.5–3 mm long .............................................................................. 9 Branchlets, petioles, and midrib of leaf blade glabrous or glandulargranulose; fruits 7.5–8 mm diameter; pedicels 1.1–1.4 mm long ............................................................................................. M. macrocarpa Branchlets, petioles, and midrib of leaf blade reddish-glandular-papillose; fruits 3–3.5 mm diameter; pedicels 0–1 mm long ................... M. pellucida

770

M YRSINACEAE

9(7).

9.

Leaf apices acute, rarely obtuse; lower surface of leaves prominently reddish-punctate and punctate-lineate; gallery and Mora and ClusiaMagnolia forests ........................................................................... M. nitida Leaf apices obtuse to broadly rounded; lower surface of leaves inconspicuously black-punctate to punctate-lineate; cloud forests to lowland savannas ................................................................................ M. guianensis

Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult., Syst. Veg. 4: 511. 1819. —Samara coriacea Sw., Prodr. 1: 32. 1788. —Rapanea coriacea (Sw.) Mez in Urb., Symb. Antill. 2: 428. 1901. Pantropics; 3 subspecies, 2 in Venezuela, both in the flora area.

reticulata Steyerm., Fieldiana, Bot. 28: 477. 1953. Shrub or small tree to 3 m tall. Gallery forests along streams on tepui summits, 2000–2800 m; Bolívar (Ilú-tepui, Macizo del Chimantá [Apacará-tepui, Toronó-tepui], Río Chajura, Roraima-tepui, Sierra de Maigualida, Sororopán-tepui). Endemic.

Key to the Subspecies of M. coriacea 1. Leaf blades 6–13 cm long, inflorescences (3–)5–9(–11)-flowered; caylx lobes longer than wide ................... subsp. coriacea 1. Leaves 1.5–5.5(–5.8) cm long, inflorescences 2(3)-flowered; calyx lobes deltate ................................. subsp. reticulata M. coriacea subsp. coriacea Caballeria ferruginea Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 280. 1798. —Manglilla ferruginea (Ruiz & Pav.) Roem. & Schult., Syst. Veg. 4: 506. 1819. —Myrsine ferruginea (Ruiz & Pav.) Spreng., Syst. Veg. 1: 664. 1825. —Rapanea ferruginea (Ruiz & Pav.) Mez in Urb., Symb. Antill. 2: 429. 1901. Myrsine myricoides Schltdl., Linnaea 1833: 525. 1833. —Rapanea myricoides (Schltdl.) Lundell, Wrightia 3: 109. 1964. Rapanea ambigua Mez in Engl., Pflanzenr. IV. 236(Heft 9): 380. 1902. Shrub or small tree to 5(–30) m tall, 15 (–50) cm diameter. Lower to upper montane forests and tepui scrub, 700–3000 m; Bolívar (Gran Sabana and adjacent areas, Macizo del Chimantá [Apacará-tepui], Ptari-tepui, Río Caroní), Amazonas (Sierra de la Neblina). Anzoátegui, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Sucre, Táchira, Trujillo, Yaracuy; Mexico, Central America, West Indies, South America except Chile. M. coriacea subsp. reticulata (Steyerm.) Pipoly, Novon 1: 210. 1991. —Rapanea

Myrsine guianensis (Aubl.) Kuntze, Revis. Gen. Pl. 2: 402. 1891. —Rapanea guianensis Aubl., Hist. Pl. Guiane 121, t. 46. 1775. Rapanea oblonga Pohl ex Miq. in Mart., Fl. Bras. 10: 308. 1856. Myrsine ovalifolia Miq. in Mart., Fl. Bras. 10: 313. 1856. —Rapanea ovalifolia (Miq.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 391. 1902. Tree to 6(–15) m tall. Evergreen lowland to upper montane forests, riparian forests, upland savannas, near sea level to 1400 (–2800) m; Delta Amacuro (Caño Guiniquina), Bolívar (widespread), Amazonas (Brazo Casiquiare, Cerro Coro Coro, Río Manapiare, Río Matacuni, Río Negro, Río Parucito, San Carlos de Río Negro). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Monagas, Nueva Esparta, Sucre; Guyana, French Guiana, Suriname, Brazil. ◆Fig. 637. Myrsine macrocarpa Pipoly, Novon 1: 207. 1991. Tree to 7 m tall. Margins of gallery forests in highland meadows, 1500–2000 m; Amazonas (Caño Piedra southeast of Cerro Aracapo, Cerro Huachamacari, Río Cunucunuma). Endemic. Myrsine maguireana Pipoly, Novon 1: 204. 1991. Shrub to 1.5(–2) m tall. Montane scrub on quartzite and granite, 1200–2200 m; Amazonas (Río Mawarinuma, Río Yatúa, Sierra

Myrsine 771

Fig. 635. Myrsine nitida

Fig. 636. Myrsine minima

Fig. 637. Myrsine guianensis

772

M YRSINACEAE

de la Neblina, Sierra de Maigualida, Sierra Tapirapecó). Endemic. Myrsine minima (Steyerm.) Pipoly, Novon 1: 210. 1991. —Rapanea minima Steyerm., Fieldiana, Bot. 28: 477. 1953. Shrub 1.5–3 m tall. Scrub forests and exposed areas in tepui meadows, 1900–2800 m; Bolívar (Angasima-tepui, Cerro Jaua, Ilútepui, Kukenán-tepui, Macizo del Chimantá [Abacapá-tepui, Amurí-tepui, Apácara-tepui, Churí-tepui, Murey-tepui], Río Tírica). Guyana, Brazil. ◆Fig. 636. Myrsine nitida (Mez) Pipoly, Novon 1: 210. 1991. —Rapanea nitida Mez, Repert. Spec. Nov. Regni Veg. 16: 424. 1920. Rapanea roraimensis A.C. Sm., Bull. Torrey Bot. Club 67: 296. 1940. —Myrsine roraimensis (A.C. Sm.) Pipoly, Novon 1: 208. 1991. Shrub or tree to 10 m tall. Rocky knolls in savannas, along the margins of montane forests, riparian forests, (800–)1800–2500 m; Bolívar (El Paují, Gran Sabana, Ilú-tepui, Macizo del Chimantá [Agparamán-tepui, Apácara-tepui], Ptari-tepui, Roraima-tepui, Sororopán-tepui). Guyana. ◆Fig. 635. Myrsine pellucida (Ruiz & Pav.) Spreng., Syst. Veg. 1: 664. 1825. —Caballeria pellucida Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 280. 1798. —Manglilla pellucida

(Ruiz & Pav.) Roem. & Schult., Syst. Veg. 4: 506. 1819. —Rapanea pellucida (Ruiz & Pav.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 394. 1902. Rapanea perforata Mez in Engl., Pflanzenr. IV. 236(Heft 9): 395. 1902. Tree to 5(–10) m tall. Lower montane forests, ca. 400 m; Bolívar (Guaniamo). Anzoátegui, Aragua, Barinas, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Táchira, Trujillo; Colombia, Ecuador, Peru, Bolivia. Myrsine perpauciflora Pipoly, Novon 2: 176. 1992. Tree to 4 m tall. Tepui scrub, 1500–1700 m; Amazonas (Sierra de la Neblina). Endemic. Myrsine picturata Pipoly, Novon 1: 204. 1991. Tree to 8 m tall. Montane scrub, 1200– 2500 m; Bolívar (Ilú-tepui), Amazonas (Sierra de la Neblina). Endemic. Myrsine resinosa (A.C. Sm.) Pipoly, Novon 1: 210. 1991. —Rapanea resinosa A.C. Sm., Bull. Torrey Bot. Club 67: 297. 1940. Shrub 1–1.5(–3) m tall. Gallery forests, 400–1000 m; Bolívar (Gran Sabana, Río Aponguao, Río Caruay, Río Karaurín). Guyana.

5. STYLOGYNE A. DC., Ann. Sci. Nat. Bot. sér. 2, 16: 78. 1841. Dioecious, bisexual, androdioecious, or polygamodioecious shrubs or small trees. Branchlets mostly glabrous, rarely with simple hairs or translucent lepidote scales on the buds. Petioles marginate or canaliculate. Blades usually glabrous. Inflorescences corymbose or paniculate with the branches racemose or short-umbellate-corymbose, sessile or subsessile to long-pedunculate, axillary or terminal, rarely axillary and terminal, mostly glabrous, rarely with simple hairs. Flowers corymbose, (4)5-merous, pedicellate. Sepals dextrorsely contorted in bud, free or short-connate basally, punctate and punctate-lineate, generally glabrous, rarely with simple hairs. Petals dextrorsely contorted and doubly twisted in bud, shortconnate, commonly lineate, punctate, and punctate-lineate, mostly glabrous. Stamens usually shorter than the petals; filaments slender, free from each other and adnate to the petals; anthers oblong, subsagittate basally, dorsifixed or basifixed, twisted once or twice at anthesis, usually dehiscent by longitudinal slits, rarely first by apical pores then widening by slits; staminodes similar to stamens but reduced in size, the antherodes devoid of pollen. Pistil obturbinate; ovary ovoid; style long; stigmas punctiform, subequaling or exceeding the stamens; placenta basal; ovules 3–5,

Stylogyne 773

uniseriate. Pistillode lageniform; ovary hollow; style subcapitate, < 1 mm long. Fruit drupaceous, 1-seeded, the endocarp crustaceous or bony. Seed globose or depressed; endosperm corneous, excavate, not ruminate; embryo transverse, elongate. Southwestern Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 40 species; 10 in Venezuela, 5 of these in the flora area. Key to the Species of Stylogyne 1. 1. 2(1). 2. 3(1). 3. 4(3). 4.

Inflorescence peduncle, rachis, and/or pedicels sparsely to densely papillose, of simple to few-celled papillae, often obscure ............................. 2 Inflorescence peduncle, rachis, and/or pedicels glabrous ......................... 3 Anthers narrowly lanceolate, reddish; inflorescences opaque; ovary with scattered papillae apically ............................................................ S. viridis Anthers oblong, whitish yellow; inflorescence translucent pink or pinkishreddish; ovary glabrous apically ........................................... S. orinocensis Margin of leaves conspicuously crenate ........................................... S. lasseri Margin of leaves entire, occasionally obscurely crenulate ....................... 4 Mature fruits 10–14 mm diameter, depressed globose; leaves chartaceous to coriaceous, the upper surface usually shiny ................................ S. atra Mature fruits 4–7(–9) mm diameter, globose; leaves membranaceous, the upper surface mostly dull ....................................................... S. micrantha

Stylogyne atra Mez in Engl., Pflanzenr. IV. 236(Heft 9): 273. 1902. —Ardisia nigricans Miq. in Mart., Fl. Bras. 10: 290. 1856. Stylogyne spruceana Mez in Engl., Pflanzenr. IV. 236(Heft 9): 274. 1902. Shrub or small tree 3–6 m tall. Lowland to lower montane evergreen forests, often on sandy soils, 100–800 m; Amazonas (Río Casiquiare, Río Mawarinuma, Sierra de la Neblina). Brazil (Amazonas). ◆Fig. 639. Stylogyne lasseri (Lundell) Pipoly, Ernstia 53: 4, fig. 1. 1989. —Ardisia lasseri Lundell, Wrightia 4: 59. 1968. Shrub or small tree 1.5–2 m tall. Montane gallery forests 700–900 m; Bolívar (Gran Sabana). Endemic. Stylogyne micrantha (H.B.K.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 276. 1902. —Ardisia micrantha H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 246. 1818 [1819]. —Bumelia micrantha (H.B.K.) Willd., herb. n. 4606 ap. Roem. & Schult., Syst. 4: 802. 1819. Stylogyne venezuelana Mez in Engl.,

Pflanzenr. IV. 236(Heft 9): 273. 1902. Cybianthus foliosus Rusby, Descr. S. Amer. Pl. 79. 1920. Stylogyne latifolia A.C. Sm., Lloydia 2: 203. 1939. Shrub or tree 2–10 m tall. Evergreen lowland to lower montane forests, near sea level to 500 m; Delta Amacuro (Caño Atoiba north of Caño Araguao, Paloma south of Tucupita, Río Cuyubini, Río Grande, Río Orocoima, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, El Dorado, El Palmar, Puerto Ordaz, Río Caura, Río Colorado east of Arekuna, Río Curutu, Río Hacha, Río Nichare, Río Paragua, Sierra de Maigualida). Anzoátegui, Apure, Barinas, Distrito Federal, Mérida, Miranda, Portuguesa, Trujillo, Sucre, Zulia; Colombia, Guayana, Suriname, French Guiana. ◆Fig. 638. Stylogyne orinocensis (H.B.K.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 270. 1902. —Ardisia orinocensis H.B.K., Nov. Gen. Sp. 3: (quarto ed.) 244. 1818 [1819]. —Tinus orinocensis (H.B.K.) Kuntze, Revis. Gen. Pl. 2: 974. 1891. Ardisia hostmannii Miq. in Mart., Fl.

774

M YRSINACEAE

Bras. 10: 288. 1856. —Tinus hostmannii (Miq.) Kuntze, Revis. Gen. Pl. 2: 974. 1891. Ardisia surinamensis Miq. in Mart., Fl. Bras. 10: 288. 1856. —Tinus surinamensis (Miq.) Kuntze, Revis. Gen. Pl. 2: 975. 1891. —Stylogyne surinamensis (Miq.) Mez in Engl., Pflanzenr. IV. 236(Heft 9): 271. 1902. Stylogyne amazonica Mez in Engl., Pflanzenr. IV. 236(Heft 9): 275. 1902. Stylogyne kappleri Mez in Engl., Pflanzenr. IV. 236(Heft 9): 275. 1902. —Ardisia nigricans Miq. in Mart., Fl. Bras. 10: 289. 1856, e.p., quoad cit. Kappler 1633. Stylogyne micans Mez in Engl., Pflanzenr.

IV. 236(Heft 9): 272. 1902. Shrub or tree 1.5–14 m tall. Seasonally flooded forests near black-water rivers, riparian forests, 100–900 m; Bolívar (Amaruay-tepui, near El Paují, Río Caura, Río Nichare, Río Paragua, San Pedro de las Bocas, Sierra de Maigualida), Amazonas (widespread). Colombia, Guyana, Suriname, French Guiana, Brazil. Stylogyne viridis (Lundell) Ricketson & Pipoly, Sida 18: 1171, fig. 2. 1999. —Parathesis viridis Lundell, Phytologia 56: 26. 1984. Small tree 2.5–4 m tall. Seasonally flooded forests, ca. 100 m; Amazonas (San Carlos de Río Negro). Endemic. ◆Fig. 640.

Fig. 638. Stylogyne micrantha

Stylogyne 775

Fig. 639. Stylogyne atra

Fig. 640. Stylogyne viridis

Appendix List of new names published in this volume

Aciotis viscida (Benth.) A. Freire-Fierro, comb. nov. ............................................. 276 Cladocolea intermedia (Rizzini) Kuijt, comb. nov. ................................................... 38

776

Index Entries in Roman type = accepted names of taxa and vernacular names Entries in italics = synonyms Page numbers in bold = illustrations

—A— Abarema, 586 acreana, 587 adenophora, 587 barbouriana, 588 var. arenaria, 588 claviflora, 680 ferruginea, 588 floribunda, 588 jupunba, 588 var. jupunba, 588 var. trapezifolia, 588 laeta, 589 leucophylla, 589 var. leucophylla, 589 levelii, 589 longipedunculata, 589, 590 microcalyx, 589 var. microcalyx, 589 trapezifolia, 588 villifera, 589 Abarkasa-dek, 665 Abelmoschus, 188 moschatus, 188, 188 Abira, 622, 626 Abira-yek, 622, 626 Abuta, 557 candicans, 568 grandifolia, 558, 559 grisebachii, 558 guyanensis, 558 imene, 558, 560

limaciifolia, 568 obovata, 558, 561 pahnii, 558, 562 rufescens, 559, 561 splendida, 559 velutina, 559 verruculosa, 575 wilson-brownei, 559 Abutilon crispum, 194 patens, 219 spicatum, 189 Acacia, 590 alemquerensis, 591 arenosa, 648 articulata, 591, 592 farnesiana, 591, 593 flexuosa, 595 formosa, 677 glomerosa, 594, 594 guachapele, 671 guianensis, 674 jupunba, 588 laxa, 602 leucocephala, 642 lutea, 595 macracantha, 595, 593 macroloba, 665 multiflora, 596 paniculata, 595 paniculiflora, 651 podadenia, 595 pulcherrima, 674 subdimidiata, 596

777

tamarindifolia, 595, 592 viridiflora, 666 Acaciella, 590 Acamosebi, 536 Acanthella, 268 montana, 270 plicata, 269 pulchra, 269, 269 sprucei, 270, 269 Acanthinophyllum, 699 ilicifolia, 701 strepitans, 701 Acanthospaera, 723 Acerola, 159 Aciotis, 270 acuminifolia, 271, 274 acutiflora, 272 var. parvifolia, 272 aequatorialis, 271 amazonica, 271 var. radicans, 271 annua, 271, 275 anomala, 272 aristata, 272 asplundii, 272 caulialata, 272 circaeifolia var. major, 276 dichotoma, 271 var. anomala, 271 var. longifolia, 271 discolor, 273 dysophylla, 271

778

INDEX

[Aciotis] fragilis, 273 var. lancifolia, 276 herbacea, 273 indecora, 272, 275 var. glabrescens, 272, 276 var. macrophylla, 272 var. sagotiana, 272 laxa, 272 var. kappleriana, 272 var. robusta, 272 levyana, 272 longifolia var. glabra, 276 martinicensis, 273 ornata, 273, 275 pellucida, 273 polystachya, 273, 275 purpurascens, 273, 274 var. alata, 272 var. alata, 276 var. longifolia, 276 var. martinensis, 272 var. pellucida, 273 rostellata, 272 rubricaulis, 272 sieberi, 273 sileniflora, 273 sp. A, 276 sphaeranthera, 271 trinitensis, 276 uliginosa, 273 viscida, 276 viscosa, 272 Acisanthera, 276 adscendens, 279 beccabunga, 277 bivalvis, 277, 279 boissieriana, 278 brevifolia, 277 crassipes, 278, 280 erecta, 368 gracilis, 278 hedyotidea, 278, 280 lasiophylla, 368 limnobios, 278, 279 nana, 278 pellucida, 278 quadrata, 279 recurva, 279 salzmannii, 278 trivalvis, 277

uniflora, 279, 280 Adelbertia, 384 Adelobotrys, 281 adscendens, 282, 284 barbata, 282, 286 ciliata, 282, 285 duidae, 282, 283 fruticosa, 282, 284 guianensis, 282 laxiflora, 490 linearifolia, 283 monticola, 283, 285 subsp. occidentalis, 283 permixta, 283 rotundifolia, 283 saxosa, 286 spruceana, 286 stenophylla, 286 Æphí, 696 Æræ æphi, 696 Affonsea, 615 Aijiadu, 719 Aik-yek, 744 Akodek, 739 Akwi æ’mi, 728 Albizia, 595 barinensis, 596, 597 corymbosa, 614 glabripetala, 596 guachapele, 671 inopinata, 673 longepedata, 671 subdimidiata, 596 Alcoceratothrix,106 rugosa, 124 stipulacea, 124 Algodón, 193, 194 Algodoncillo, 196 Almidón, 696 Alowata lua, 628 Alpinia latifolia, 226 Alstroemeria, 2 amazonica, 3, 3 edulis, 5 Althaea racemosa, 207 Amaryllis eguestris, 11 elegans, 11 longiflora, 11 punicea, 11 solandriflora, 11

Amatlania, 749 Ammannia ramosior, 79 Amor seco, 69 Anadenanthera, 597 peregrina, 598 var. peregrina, 598, 599 Anelasma domingense, 571 Anomospermum, 562 hirsutum, 575 japurense, 564 schomburgkii, 575 steyermarkii, 563, 563 Anthericum coarctatum, 8 Antonia, 23 ovata, 23, 24 var. excelsa, 23 var. pilosa, 23 pilosa, 23 pubescens, 23 Apatitia, 287 blakeoides, 287 Apodina, 48 cucullaris, 50 Arabakasa-yek, 665 Arana de gato negro, 652 Ardisia, 749, 760 acuminata, 749 acuminifolia, 750 amanuensis, 749 cookii, 749 guianensis, 749, 749 hostmannii, 773 lasseri, 773 longicaudata, 749 micrantha, 773 nigricans, 773 orinocensis, 773 pseudoicacorea, 760 quelchii, 762 stenobotrys, 751 subg. Graphardisia, 749 surinamensis, 774 Arepillo hoja fina, 680 Arespín, 666 Arthrosamanea, 595 corymbosa, 614 gonggrijpii, 614 panurensis, 615 Arthrostemma angusturenese, 493

INDEX

caulialatum, 272 glomerata, 493 nummularioides, 312 sect. Monocentrum, 469 sect. Monochaetum, 469 villosum, 313 Aru-maka, 226 Aruro, 661 Ashuhua, 745 Aspuela-guara, 567 Atauashima, 744 Atopocarpus, 128 Aura-tá-na-ca-tá, 82 Auriculardisia, 749 Auyamilla, 202 Awadu-di-venadu-ilu, 578 Awenadu, 624 Axinanthera, 287 macrophylla, 290 Ayahuasca, 93 Ayoco, 745 Ayudek, 744 Ayuko, 746 Ayuku, 744, 745 Azucarito, 741 —B— Balizia, 599 pedicellaris, 599, 600 Banisteria argentea, 95 caapi, 93 cristata, 94 fagifolia, 151 heterophylla, 179 leona, 144 leptocarpa, 95 lucida, 94 macradena, 144 martiniana, 95 metallicolor, 95 mossii, 146 multiflora, 144 muricata, 95 orbicularis, 94 orinocensis, 146 ovata, 177 pubera, 178 reticulata, 144 schomburgkiana, 95 sinemariensis, 167 sinuata, 179

splendens, 179 Banisteriopsis, 92 argentea, 95 caapi, 93 cabrerana, 137 cristata, 94 elegans var. pulcherrima, 97 inebrians, 93 krukoffii, 94 leptocarpa, 95 lucida, 94 lyrata, 94 maguirei, 84, 95 martiniana, 95 var. laevis, 95 var. martiniana, 95, 96 var. subenervia, 95 metallicolor, 95 muricata, 95 pulcherrima, 97 schomburgkiana, 95 wurdackii, 97 Barbasco lagartilla, 567 Baruta de lagartillo, 567 Basegua banero, 745 Bastardia sect. Gayoides, 194 Behebehedu, 665 Bejuco de cruz, 31 Bejuco de estribo, 31 Bejuco de gallineta, 251 Bejuco de iguana, 155 Bejuco de mono, 31 Bejuco de ratón, 94 Bejuco de sapo, 155 Bejuco jala pa’trás, 595 Bejuco pantalla, 250 Bejuco pene de diablo, 260 Bejuco rabo de guacamayo, 256 Bejuco rojo, 552 Bellucia, 287 arborescens, 363 aricuaizensium, 290 brasiliensis, 287 circumscissa, 287 grossularioides, 287, 288 hostmannii, 287 huberi, 288, 289 mespiloides, 363 pentamera, 290 quinquenervia, 287

779

Bertolonia, 290 sp. A, 290, 291 venezuelensis, 290, 291 Besleria bivalvis, 551 Biaslia, 261 Bihibihidu, 665 Bijau, 225 Billo, 701 Biscochuelo, 544 Biscochuelo amarillo, 545 Biscochuelo negro, 545, 546 Blakea, 292 calycanthus, 292 caudata, 292 quinquenervia, 287 rosea, 292, 292 Blepharandra, 97 angustifolia, 98, 99 cretacea, 100 fimbriata, 98, 99 heteropetala, 98 hypoleuca, 98, 99 ptariana, 100 Bogenhardia, 194 crispa, 194 Bomarea, 4 edulis, 5, 4 Bonyunia, 24 aquatica, 25, 25 cinchonoides, 25 minor, 25, 25 superba, 25, 26 Bora, 579 Borismene, 564 japurensis, 564, 564 Boyo boyo, 340 Boyoyo blanco, 272 Boyuyo, 271, 282 Boyuyo morado, 273, 424 Boyuyo palo de vieja, 413, 431 Boyuyo palo de viejo, 410 Boyuyo rebalsero, 403 Brachyandra, 492 Brachypterys, 176 borealis, 177 ovata, 177 Briquetia, 188 spicata, 189, 189 Brosimopsis, 695 lactescens, 696 Brosimum, 695

780

INDEX

[Brosimum] alicastrum, 695 subsp. bolivarense, 695, 697 bolivarense, 695 guianense, 696, 697 lactescens, 696, 697 lanciferum, 696 latifolium, 695 longifolium, 698 melanopotamicum, 696 mello-barretoi, 723 myristicoides, 696 ovatifolium, 698 palmarum, 696 paraense, 696 potabile, 696 rotundatum, 696 rubescens, 696 utile, 696 subsp. longifolium, 698 subsp. ovatifolium, 698 Brusquillo, 674 Bucarito, 79 Bumelia, 773 micrantha, 773 Bunchosia, 100 argentea, 101, 102 armeniaca, 101 decussiflora, 84, 101 glandulifera, 101 mollis, 103 petraea, 103 rhombifolia, 103 schomburgkiana, 103 squarrosa, 184 Burdachia, 103 atractoides, 104 prismatocarpa, 104, 105 var. sphaerocarpa, 104 sphaerocarpa, 84, 104, 105 var. glandifera, 104 williamsii, 104 Byrsonima, 106 aerugo, 114, 128 angustifolia, 98 baccae, 115 barkleyana, 123 basiliana, 115 bolivarana, 115 bracteolaris, 117 bronweniana, 115, 126

carmeniana, 118 carraoana, 115, 126 chalcophylla, 116 var. carraoana, 115 christianeae, 116 chrysophylla, 116 coccolobifolia, 116, 126 concinna, 117 coniophylla, 84, 117 coriacea var. spicata, 123 cowanii, 118, 127 crassifolia, 118 cretacea, 100, 121 crispa, 118 cuprea, 119 dubia, 119 duidana, 119, 126 fernandezii, 119 ferruginea, 118 fluminensis, 120 frondosa, 120 huberi, 120 inundata, 120 japurensis, 120 subsp. fluminensis, 120 subsp. silvatica, 120 kariniana, 120 laevis, 121 laurifolia, 118 leucophlebia, 121 linguifera, 121 luetzelburgii, 121 macrostachya, 122 maguirei, 122 nitidissima, 122 pachypoda, 122 poeppigiana, 123 propinqua, 123 punctulata, 123 reticulata, 144 rugosa, 124 schomburgkiana, 123 sessilifolia, 116 sp. A, 125 spicata, 123 steyermarkii, 124, 127 stipulacea, 124 tillettii, 124 uvulifera, 120 verbascifolia, 125 var. denudata, 125 wurdackii, 125

—C— Caapi, 93 Caballeria, 768 ferruginea, 770 pellucida, 772 Cabaya-cuaja, 260 Cabeza de mono, 82 Cabimbo, 536 Cacaguaru, 739 Cacani-caritsapi, 661 Cachete de vieja, 36 Cachicama, 607 Cadillo, 189 Café de venado, 736 Cafecillo, 701 Cahuabari, 623 Cajimán, 696, 701 Calathea, 221 acuminata, 224, 227 albicans, 226 allouia var. violacea, 226 altissima, 224, 226 barbillana, 226 cannoides, 225, 227 casupito, 225, 228 cataractarum, 225 cyclophora, 225 densa, 225 discolor, 226 duidae, 225 elliptica, 225 erecta, 225 fragilis, 225 grandis, 225 hirsuta, 230 inocephala, 225 klugii, 226 lasseriana, 226 latifolia, 226 laxa, 241 leucophaea, 235 liesneri, 226 lutea, 226, 229 magnifica, 226 micans, 226, 229 var amabilis, 226 mishuyacu, 229 neblinensis, 229 ovata, 229 panamensis, 229 pardina, 230

INDEX

polyphylla, 235 trinitensis, 225 variegata, 229 villosa, 228, 230 var. pardina, 230 violacea, 226 vittata, 225 zingiberina, 230 Calliandra, 600 affinis, 601 caripensis, 602 coriacea, 601, 603 cruegeri, 601, 604 flavida, 678 formosa, 677 glomerulata, 601 var. glomerulata, 601, 605 guildingii, 602 latifolia, 681 laxa, 602 var. laxa, 602 var. stipulacea, 602, 605 var. urimana, 602 marginata, 677 pakaraimensis, 602, 606 pilosifolia, 607 portoricensis, 678 resupina, 602 rigida, 602, 603 riparia, 602 stipulacea, 602 surinamensis, 602, 604 tenuiflora, 602 tergemina, 602 trijugata, 602 trinervia, 607 var. pilosifolia, 607 var. trinervia, 603, 607 tsugoides, 606, 607 vaupesiana, 607 var. oligandra, 604, 607 Calophysa tococoidea, 303 Calyptrella, 321 Camaticaro, 745 Camaticaro rojo, 745 Cambur, 732 Cañafistolillo, 674 Caoba, 544 Capau-rit-warei-yek, 746 Capuy-yen-cu-me-peu, 558 Caraashiji, 696

Caramacate rebalsero, 546 Carapa, 529, 531 guianensis, 530, 531 Carapillo, 535 Cara-re-yek, 287 Caro, 661, 674 Caro blanco, 613, 661 Caro montañero, 661 Carpeto, 745 Carutillo, 235 Caryomene, 564 olivascens, 565, 565 Casabe de murciélago, 661 Casabe murciélago, 661 Cascarito, 474 Cascarón, 661 Castilla, 699 elastica, 699 subsp. elastica, 698, 699 Castilloa, 699 Casupo, 229, 230, 241, 242 Casupo amarillo, 230 Catamajaca rebalsera, 725 Catarajaca, 536 Cathormion corymbosum, 614 Caucho, 699 Cayena, Malva, 196 Cedrela, 531 brownii, 533 fissilis, 533 odorata, 532, 533 Cedrelinga, 607 cateniformis, 607, 608 Cedril, 544 Cedrillo, 744, 746 Cedrito, 544 Cedro, 533 Cedro blanco, 746 Cedro colorado, 533 Cedro rojo, 533 Ceguera, 736 Centronia, 293 crassiramis, 384 neblinae, 293, 294 Cereza, 159 Chaenophora ferruginea, 414 Chaenopleura ferruginea, 414 hypoleuca, 414 longifolia, 416 Chaetogastra

781

callichaeta, 493 geitneriana, 506 glomerata, 493 lasiophylla, 368 rhynchanthera, 470 sect. Adesmogastra, 294 Chaetolepis, 294 anisandra, 294, 295 citrina, 294 phelpsiae, 294 subsp. chimantensis, 295, 295 subsp. phelpsiae, 295 Chalepophyllum pungens, 270 Chaparro, 116, 118 Chaparro manteco, 118, 123, 124 Chara, 695 Charo, 696, 725, 728 Charo blancho, 696 Charo colorado, 696 Charo macho, 696, 719, 728 Charo peludo, 719, 725 Chigüire, 594 Chinak, 558 Chincharrón, 674 Chitonia bubalina, 406 Chloretis, 11 Chloroleucon, 609 mangense, 610 var tetrazyx, 609, 610 Chlorophora, 719 tinctoria, 720 Chondrodendron candicans, 568 limaciifolium, 568 Cienfuegosia, 190 affinis, 190, 192 heterophylla, 190, 192 phlomidifolia, 192 Ciruela, 101 Ciruela de fraile, 101, 103 Ciruela montañera, 103 Cissampelos, 565 andromorpha, 566, 567 ovalifolia, 566, 567 pareira, 566, 567 ramiflora, 567 tropaeolifolia, 567 Cladocolea, 38 apiculata, 38

782

INDEX

[Cladocolea] intermedia, 38 micrantha, 38, 39 roraimensis, 39 Clarisia, 699 biflora, 701 ilicifolia, 700, 701 racemosa, 700, 701 spruceana, 701 strepitans, 701 Clavelino, 661, 674 Clavellín, 665 Clavellina, 661, 665 Clavellino, 665, 666, 680 Clidemia, 295 acurensis, 299 alternifolia, 299, 304 andersonii, 299 aphanantha, 299, 305 aristigera, 352 attenuata, 299 bernardii, 299 bracteata, 405 bullosa, 299, 304 buntingii, 299, 306 campestris, 406 var. pauciflora, 404 candolleana, 299 capillipes, 299, 307 capitata, 299, 306 capitellata, 300 var. capitellata, 300 var. dependens, 300 var. levelii, 300 ceramicarpa, 408 chaetodon, 352 coccinea, 408 conglomerata, 300, 308 coriacea, 303 debilis, 300 dentata, 300 dependens, 300 desmantha, 424 divaricata, 353 duidae, 300 epibaterium, 300, 306 heptamera, 300, 309 heterobasis, 301 heteroneura, 300 hirta, 301 var. elegans, 301, 308 var. hirta, 301 var. tiliaefolia, 301

involucrata, 301, 310 japurensis, 301 var. heterobasis, 301, 310 var. japurensis, 301 juruensis, 301 linearis, 301, 307 lutescens var. lindeniana, 355 maculata, 414 manoacensis, 299 marahuacensis, 302 martii, 407 miconioides, 414 micrantha, 302 microthyrsa, 302 minutiflora, 302, 310 morichensis, 302 neblinae, 302 novemnervia, 302, 305 var. depauperata, 299 octona, 302 subsp. guayanensis, 302 subsp. octona, 302 piperifolia, 302 pustulata, 302, 305 pycnaster, 303 subsp. pycnaster, 303, 304 subsp. robusta, 303 radulaefolia, 424 rariflora, 299 rhamnifolia var. macrodon, 353 rubra, 303 secunda, 357 sericea, 303 sessiliflora, 303 siapensis, 303 silvicola, 303 simpsonii, 303 stellipilis, 303 strigillosa, 303 tepuiensis, 303, 309 tiliaefolia, 301 tococoidea, 303, 307 trinitensis, 311 ulei, 311 urceolata, 311 violacea, 408 virgata, 414 Clidemiastrum, 350 Clonodia, 128

complicata, 128, 129 racemosa var. orinocensis, 129 Cobija, 674 Cocculus domingensis, 571 imene, 558 japurensis, 564 pahni, 558 pauper, 572 tamoides, 572 toxicoferus, 569 Codo rebalsero, 744 Cola de guaca, 256 Coleostachys hypoleuca, 98 vestita, 132 Coletia, 261 Collania, 4 Coloradito, 738 Colorado, 545 Cometure, 474 Cometure tierra firmero, 528 Comolia, 311 affinis, 315 angustifolia, 315 anomala, 279 coriacea, 312, 313 hirtella, 313 hoehnei, 498 kuhlmannii, 315 leptophylla, 312, 314 lythrarioides, 315 microphylla, 312, 313 montana, 312, 314 neglecta, 315 nummularioides, 312, 314 prostrata, 312 purpurea, 313 serpyllacea, 313, 314 smithii, 313 surinamensis, 315 vernicosa, 313, 314 veronicaefolia, 313 villosa, 313, 313 Comoliopsis, 315 neblinae, 315, 316 Comomyrsine, 751 Compsoneura, 735 costaricensis, 736 debilis, 736 sprucei, 736, 736

INDEX

Conejilla, 215 Conomorpha, 751 ampla, 758 apiculata, 758 candolleana, 763 crotonoides, 759 curvivenia, 761 depressa, 762 duidae, 759 fulvopulverulenta, 759 glaucorubens, 763 gracilis, 763 grandiflora, 763 guyanensis, 760 heterantha, 760 latifolia, 763 laxiflora, 763 var. latifolia, 763 lepidota, 761 f. acutata, 761 leprosa, 759 lithophyta, 759 macrophylla, 758 madeirensis, 763 magnoliifolia, 759 peruviana var. β guyanensis, 760 ptariensis, 759 punctata, 762 reticulata, 762 rigida, 759 riparia, 759 roraimae, 762 sect. Aconomorpha, 757 sect. Conomorphida, 757 sect. Euconomorpha, 757 sessilis, 762 spicata, 763 steyermarkiana, 763 subg. Euconomorpha, 757 utiarityi, 758 Conostegia superba, 316, 317 Copedesma nitens, 425 Coral, 69 Coralito, 79 Correlliana, 751 Coryphadenia, 527 Cozoiba, 745 Cremanium trinitatis, 419 Crenea, 60

maritima, 60, 61 surinamensis, 60 Crinum, 5 erubescens, 5, 6 Cruceta, 31, 36 Cruceta real, 32 Ctenanthe, 231 compressa, 230, 231 Ctenardisia, 750 stenobotrys, 750, 751 Cuajo, 738, 739, 744, 745, 746, 747 Cuajo blanco, 745 Cuajo de tierra firme, 738 Cuajo grande, 739 Cuajo hoja grande, 741 Cuajo montañero, 738 Cuajo morichalero, 746 Cuajo negro, 738, 745 Cuajo pepa amarilla, 745 Cuajo pequeño, 745 Cuajo sabanero, 739 Cuajo tierra firme, 739 Cudami, 627 Cudo, 745 Cují, 595 Cují cabro, 591 Cumala blanca, 741 Cumarawa-yek, 36 Cuphea, 60 anisoclada, 63, 73 annulata, 64, 71 antisyphilitica, 64 f. gracillima, 64 f. subhirsuta, 64 var. acutifolia, 64, 72 var. antisyphilitica, 65, 72 balsamona, 65 bolivariensis, 65 cardonae, 65 carthagenensis, 65 cataractarum, 65, 73 curiosa, 66 var. curiosa, 66, 75 var. oresbia, 66, 74 dactylophora, 66 distichophylla, 67 elliptica, 67 galeato-calcarata, 67 gracilis, 68, 76 var. major, 64 var. media, 68

783

maigualidensis, 68, 74 melvilla, 69, 71 micrantha, 69 multicaulis, 68 odonellii, 69, 76 pauciflora, 68 pleiantha, 69, 75 repens, 70 rhodocalyx, 70 var. setosa, 65 rigidula, 70, 75 speciosa, 69 tenuissima, 77 Curanbo-yek, 627 Curare, 31, 32, 36 Curare de pava, 36 Curarea, 567 candicans, 568, 568 toxicofera, 569 Curariua grande, 613 Curari-yek, 36 Curculigo, 5 scorzonerifolia, 6, 6 Cybianthopsis, 743 Cybianthus, 751 agostinianus, 758 amplus, 758, 764 apiculatus, 758, 765 breweri, 758 brownii, 763 cardonae, 758 comatus, 762 crotonoides, 759, 764 deltatus, 759, 764 duidae, 759 egensis, 763 foliosus, 773 fulvopulverulentus, 759 subsp. fulvopulverulentus, 759, 765 subsp. magnoliifolius, 759 grandifolius, 760 guyanensis, 760 subsp. guyanensis, 760 subsp. multipunctatus, 760 subsp. pseudoicacoreus, 760 holstii, 760 huberi, 760 julianii, 761

784

INDEX

[Cybianthus] lepidotus, 761, 766 liesneri, 761 lineatus, 761, 764 lithophytus, 759 longifolius, 761 macrophyllus, 758 magnoliifolius, 759 maguirei, 761 multicostatus, 761 multipunctatus, 760 nitidus, 762 penduliflorus, 761 piresii, 761 plowmanii, 761 potiaei, 761 prieurei, 762 pseudoicacoreus, 760 ptariensis, 762, 768 punctatus, 762, 766 quelchii, 762, 768 resinosus, 762 reticulatus, 762 roraimae, 762, 766 sect. Cybianthoides, 757 sect. Eucybianthus, 757 sect. Weigeltia, 758 sipapoensis, 762 spathulifolius, 763 spicatus, 763, 767 steyermarkianus, 763, 767 subg. Conomorpha, 757 subg. Cybianthus, 757 subg. Grammadenia, 757 subg. Laxiflorus, 758 subg. Weigeltia, 758 subspicatus, 762 surinamensis, 763, 767 sylvaticus, 760 venezuelanus, 763, 765 viridiflorus, 762 wurdackii, 763 —D— Dajaka, 546 Dan-anatu, 709 Dato, 739 Dau-joro, 199 Davya, 281 adscendens, 282 ciliata, 282

crassiramis, 384 guianensis, 282 sclerophylla, 385 Dayacabi, 698 Day’e’mon, 661 Dejaka, 545 Derello, 475, 474, 476 Derello hoja grande, 474 Dereye, 475 Dereyo, 474 Desmoscelis, 317 mollis, 317 villosa, 317, 318 Deyi, 661 Diacidia, 129 cordata, 131, 133 duckeana, 131 ferruginea, 131, 133 galphimioides, 131 glaucifolia, 131 hypoleuca, 132 kunhardtii, 132, 134 parvifolia, 131 rufa, 132 steyermarkii, 132, 133 stipularis, 132 vestita, 132 Dianella dubia, 8 Diaru, 744, 746 Dicella, 134 amazonica, 134 julianii, 134, 135 Dicrananthera hedyotidea, 278 salzmannii, 278 Dieru, 744 Diolena longidens, 379 repens, 379 Diplochita bubalina, 406 parviflora, 423 serrulata, 427 tomentosa, 430 Diplopterys, 137 cabrerana, 136, 137 includens, 172 involuta, 172 var. ovata, 170 sect. Mezia, 169 Disciphania, 569 ernstii, 569

var. ernstii, 569, 569 Dividive, 661 Dodecas, 60 maritimus, 60 surinamensis, 60 Dorstenia, 701 brasiliensis, 702, 702 sabanensis, 702 tubicina, 702 Ducuajo, 707 Dugandiodendron, 81 chimantense, 81, 82 ptaritepuianum, 81, 82 Dukaja, 709 Dukuajo, 708 —E— Eccremis, 7 coarctata, 7, 8 Echeandia, 8 bolivarensis, 8, 9 Embodium, 5 Entada, 610 myriadenia, 651 polyphylla, 610, 611 polystachya, 611 Entadopsis, 610 polyphylla, 610 polystachya, 611 Enterolobium, 612 barinense, 612, 613 schomburgkii, 613 Erenpi, 696 Erika-bai-yek, 745 Ernestia, 319 cataractae, 319, 320 cordifolia, 319 lata, 321 maguirei, 319, 320 pullei, 321 tenella, 321 var. sprucei, 320, 321 var. tenella, 321 Ero, 250 Escoba, 211, 215 Escoba negra, 215 Escobilla, 211 Espina de pescado, 681 Eugenia nigra, 475 Eurychaenia punctata, 423

INDEX

Excentradenia, 137 adenophora, 138, 138 Exoecaria ilicifolia, 701 Eyokoa, 729 —F— Farnesia, 590 Farringtonia fasciculata, 500 Feuilleea, 615 cayennensis, 620 edulis, 620 fagifolia, 622 heterophylla, 621 ingoides, 622 laurina, 622 lindeniana, 626 marginata, 623 myriantha, 628 pezizifera, 625 pilosula, 625 sapindoides, 626 sciadon, 628 setifera, 625 splendens, 626 umbellifera, 627 umbratica, 628 Ficus, 702 albert-smithii, 706, 711 amazonica, 706, 712 americana, 706, 713 angustifolia, 706 arukensis, 710 broadwayi, 706 caballina, 707, 714 catappifolia, 707 corpulenta, 708 crocata, 707 dendrocida, 707 donnell-smithii, 707, 711 ernstiana, 710 erratica, 708 erythrosticta, 710 eximia, 707 expansa, 707 fagifolia, 710 foveata, 707 foveolata, 707 gardneriana, 709 gemina, 710

glabrata, 708 glandulosa, 707 gleasonii, 708 gomelleira, 707, 715 guanarensis, 707 guianensis, 707 hebetifolia, 708 hydrophila, 709 insipida, 708 subsp. insipida, 708, 713 subsp. scabra, 708 involuta, 709 jacquiniifolia, 706 kanukuensis, 710 krukovii, 708, 717 longistipula, 708 maguirei, 709 malacocarpa, 708, 713 maroana, 708 mathewsii, 708, 715 matiziana, 709 maxima, 709 mendelsonii, 706 mensalis, 708 mollicula, 709, 714 nymphaeifolia, 709, 716 obtusifolia, 709 orinocensis, 710 pakkensis, 709 palmicida, 710 paludica, 709 panurensis, 709 paraensis, 709, 714 parkeri, 709 pascuorum, 709 perforata, 706 pertusa, 710, 716 piresiana, 710 plicato-ostiolata, 710 radula, 709 savannarum, 706 scabrida, 708 schumacheri, 710, 712 sphenophylla, 710 tamatamae, 707 tepuiensis, 710, 712 tigrensis, 709 trigona, 710 turbinata, 707 urbaniana, 709 yoponensis, 710 Flor de Jamaica, 197

785

Flor de mosquito amarillo, 473 Folianthera guianensis, 674 Fothergilla mirabilis, 419 Fruto de Paloma, 725 Fugosia campestris, 192 guianensis, 192 phlomidifolia, 192 retusa, 192 Fuiyu, 696 Furarium, 43 disjectifolium, 44 —G— Gaiadendron, 39 lanceolatum, 40 poasense, 40 punctatum, 39, 40 var. puracense, 40 puracensis, 40 revolutum, 40 tagua, 40 var. reductum, 40 var. revolutum, 40 Galactodendrum utile, 696 Galphimia longifolia, 157 Garrapata, 271 Gaya, 192 gaudichaudiana, 191, 192 Gayoides, 194 crispum, 194 Geniostoma, 28 Glandonia, 139 williamsii, 139, 139 Glinus, 687 radiatus, 687, 688 Glossocentrum collinum, 419 Goeppertia, 221 Gossypium, 192 barbadense, 193, 193 hirsutum, 194 latifolium, 194 peruvianum, 193 punctatum, 194 religiosum, 194 vitifolium, 193

786

INDEX

Graffenrieda, 321 candelabrum, 352 caryophyllea, 324, 337 cinnoides, 324 curta, 409 fantastica, 324 floribunda, 326 fruticosa, 324, 334 hitchcockii, 324, 332 var. parvifolia, 325 intermedia, 325, 335 jauana, 325, 335 kralii, 325 lanceolata, 325, 331 miconioides, 326 obliqua, 326, 330 ovalifolia, 329 patens, 326 pedunculata, 326, 330 polymera, 326 subsp. neblinensis, 327 subsp. polymera, 327, 334 reticulata, 327, 330 rotundifolia, 327, 336 rufa, 327 rupestris, 328, 338 sessilifolia, 328 subsp. cardonae, 328, 333 subsp. occidentalis, 328 subsp. sessilifolia, 328 subsp. A, 328 sipapoana, 328 stenopetala, 325 steyermarkii, 329, 331 tricalcarata, 329, 333 versicolor, 329, 331 weddellii, 329, 333 Grammadenia, 751, 757 lineata, 761 ptariensis, 762 subsp. auyantepuiensis, 762 subg. Cybianthopsis, 751 subg. Eugrammadenia, 757 Graphardisia, 749 Grischowia, 469 Grislea, 77 compacta, 79 secunda, 79 Guacamaya, 256

Guacha, 671 Guachinakua, 745 Guaimaro, 695 Guaímero, 696 Guama, 620, 622, 623, 625, 626, 627, 628 Guama comestible, 620 Guama de araguato, 623 Guama pompeya, 625 Guama yaragua, 622 Guamillo, 622 Guamita, 625, 627 Guamita yaragua, 623 Guamito, 588, 619 Guamito montañero, 619 Guamo, 619, 620, 622, 623, 624, 625, 626, 627, 628 Guamo amarillo, 625 Guamo balbino, 623 Guamo bejuco, 620 Guamo blanco, 622, 626 Guamo cambur, 626 Guamo camburito, 622 Guamo caraota, 622 Guamo chivo, 622, 628 Guamo cinta, 622 Guamo colorado, 619 Guamo de rano, 680 Guamo liso, 620, 622, 626 Guamo monero, 621 Guamo montañero, 589 Guamo morrocoyero, 626 Guamo negro, 621, 622, 627 Guamo pata de morrocoy, 622, 626 Guamo peludo, 620 Guamo rabo de mono, 620, 625 Guamo rastrojero, 626 Guamo rebalsero, 622 Guamo terciopelo, 625 Guamo verde, 626 Guanabillo, 746 Guanacaste, 696 Guaranaramuji, 613 Guaratarillo, 475, 476 Guarataro, 474, 475 Guarea, 533 adenocarpa, 545 delgadoi, 545 glabra, 535, 537 gomma, 535

grandifolia, 535, 538 guidonia, 536, 537 kunthiana, 536 macrophylla, 536 subsp. pachycarpa, 536 subsp. pendulispica, 536 pendulispica, 536 puberula, 536 pubescens, 536 subsp. pubescens, 536 subsp. pubiflora, 539, 540 pubiflora, 540 silvatica, 540, 539 sprucei, 540 trichilioides var. pachycarpa, 536 trunciflora, 540 Guarodek, 741 Guaura, 607 Guavillo, 475 Guayaba de danto, 744 Guayaba de danto rebalsera, 340 Guayaba paujicera, 474 Guayaba Pesjua, 475 Guayabito rojo, 159 Guayabo de danta, 287 Gueguewudu, 739 Guiejudo, 739 Güigüire sabanero, 594 Guigujuro, 740 Guildingia, 470 psidioides, 475 —H— Haplodesmium, 294 Helianthostylis steyermarkii, 717, 718 Helicostylis, 718 asperifolia, 719 bolivarense, 695 elegans, 718 lancifolia, 718 scabra, 718 tomentosa, 719, 719 Henriettea, 338 brunnescens, 363 granulata, 340, 341 horridula, 340 maroniensis, 340, 342

INDEX

martiusii, 340, 344 mucronata, 340, 343 multiflora, 340 oblongifolia, 340 orinocensis, 340 patrisiana, 340, 344 ramiflora, 340 spruceana, 345, 346 stellaris, 341, 346 succosa, 345, 346 var. guianensis, 340 surinamensis, 340 trinervia, 340 Henriettella, 346 caudata, 347 duckeana, 347 heteroneura, 347 longistyla, 347 micrantha, 347 ovata, 347, 349 prancei, 347, 349 ramiflora, 340 steyermarkii, 348, 348 Herissantia, 194 crispa, 194, 194 Heteroneuron, 362 Heteropterys, 140 acutifolia, 146 africana, 144 alata, 143 argentea, 95 atabapensis, 143 ayacuchensis, 143 beecheyana var. alata, 143 carinata, 143 castanea, 164 cristata, 143 cuatrecasasii, 144 dichromocalyx, 144 helicina, 146 huberi, 144 lasseri, 145 leona, 144 macradena, 144 macrostachya, 145 maguirei, 145 megaptera, 84, 145 molesta, 145 mossii, 146 multiflora, 144 neblinensis, 145 nervosa, 146

oblongifolia, 146, 148 oleifolia, 183 orinocensis, 146, 148 quetepensis, 146 reticulata, 144 siderosa, 147 stannea, 167 steyermarkii, 84, 147, 147 suberosa, 146 Heterotrichum novemnervium, 302 strigosum, 429 Hibiscus, 195 abelmoschus, 188 affinis, 192 bifurcatus, 196 bracteosus, 196 cancellatus, 202 cubensis, 199 dimidiatus, 196, 197 fraternus, 197 furcellatus, 196 ingratus, 199 lambertianus, 199 moschatus, 188 pernambucensis, 196, 198 peruvianus, 196 radiatus, 196, 197 sabdariffa, 197 salviifolius, 199 sororius, 197, 198 striatus, 199 subsp. lambertianus, 199 trilobus, 199 subsp. ingratus, 199 Higuerón, 707, 708 Higuerote, 707, 709 Higuillo, 706, 710 Higuito, 707, 708 Hippeastrum, 8 eguestre, 11 elegans, 10, 11 puniceum, 10, 11 purpureum, 11 solandriflorum, 11 Hiraea, 149 adenophora, 138 affinis, 150 apaporiensis, 150 bifurcata, 153 celiana, 151 chrysophylla, 151

787

complicata, 128 demerarensis, 151 divaricata, 165 fagifolia, 151 faginea, 151, 154 fimbriata, 152 fulgens, 151 var. demerarensis, 151 gracilis, 182 laurifolia, 167 macrodisca, 166 neblinensis, 152 nitida, 128 oblongifolia, 165 oleifolia, 183 ovatifolia, 166 platytriphylla, 153 poeppigiana, 166 schizoptera, 167 sect. Mascagnia, 160 sepium, 167 steyermarkii, 152 swartziana, 151 tepuiensis, 153, 154 ternifolia, 153 Hoja de mono, 31, 36, 558 Hoja de venado, 607 Hoja grande, 475 Hormocalyx, 512 hirsutus, 517 Huama, 627 Huatacurán, 558 Hueso de pescado, 600, 661, 670 Hueso de pescado blanco, 596 Hueso de pescado rebalsero, 614 Huevo de chiguire, 575 Hydrochorea, 613 acreana, 587 corymbosa, 614, 614 gonggrijpii, 614 marginata, 615 var. panurensis, 615 var. scheryi, 615 Hydrocleys, 16 nymphoides, 17, 17 Hylaeanthe, 231 unilateralis, 231, 232 Hymenocallis, 11 guianensis, 12 var. undulata, 12

788

INDEX

[Hymenocallis] moritziana, 12 petiolata, 12 undulata, 12 tubiflora, 12, 12 Hyperbaena, 571 cuatrecasasii, 564 domingensis, 570, 571 graciliflora, 571 Hypobrichia spruceana, 79 var. tenuifolia, 79 Hypoxis scorzonerifolia, 6 —I— Ibirkabá, 745 Icacorea, 749 acuminata, 749 acuminifolia, 750 amanuensis, 749 cookii, 749 guianensis, 749 longicaudata, 749 Indiecito, 79 Inga, 615 adiantifolia, 644 affinis, 625 alba, 619, 629 amazonica, 623 var. bracteifera, 623 var. lomatophylla, 623 var. membranacea, 623 antioquensis, 626 auristellae, 619 bijuga, 619 bourgonii, 619, 630 brachyptera, 623 bracteosa, 623 calocephala, 623 camuriensis, 626 capitata, 620, 631 caracasana, 626 carachensis, 619 cataractae, 680 cauliflora, 681 cayennensis, 620 coriacea, 626 cyclocarpa, 623 discolor, 644 disticha var. negrensis, 624

dysantha, 620 edulis, 620, 632 fagifolia, 622 fastuosa, 620, 633 floribunda, 626 forfex, 668 gladiata, 627 glomerata, 680 gracilifolia, 621 grandifolia, 626 guaremalensis, 621 heterophylla, 621, 634 huberi, 621 inaequalis, 681 inflata, 621, 634 inga, 620 ingoides, 622, 635 laeta, 589 lateriflora, 622 latifolia, 681 laurina, 622, 633 lawrenceana, 628 leiocalycina, 622 lindeniana, 626 lomatophylla, 623 macradenia, 627 macrophylla, 623, 625, 636 marginata, 623 melinonis, 623 micradenia, 624, 631 microphylla, 668 multiflora, 622 multijuga, 624 subsp. multijuga, 624 myriantha, 628 neblinensis, 624, 630 negrensis, 624 niopo, 598 nitida, 625 nobilis, 624 subsp. nobilis, 624, 637 paraensis, 625 pedicellaris, 600 pendula, 661 pezizifera, 625 pilosiuscula, 625 pilosula, 625, 638 protracta, 621 pubescens, 668 pubiramea, 644 racemiflora, 627 ramiflora, 681

riparia, 628 rosea, 668 rubiginosa, 625, 638 salicifolia, 626 sapindoides, 626, 639 sciandion, 627 sertulifera, 626 subsp. sertulifera, 626 setifera, 625 speciosa var. membranacea, 623 splendens, 626, 640 spuria, 628 stenoptera, 627 stipularis, 627, 641 tenuiflora, 627 thibaudiana, 627 subsp. thibaudiana, 627, 629 ulei, 627, 639 umbellata, 621 umbellifera, 627, 637 umbratica, 628 unguis-cati, 668 venosa, 620 vera, 620, 628 subsp. affinis, 628, 640 subsp. vera, 628 Ingaria, 615 Iryanthera, 737 congestiflora, 738 elongata, 739 hostmannii, 738 juruensis, 738 laevis, 739 lancifolia, 739 leptoclada, 738 longiflora, 739 macrophylla, 739 obovata, 739 paradoxa, 739 paraensis, 739, 740 porcata, 739 sessilis, 739 tricornis, 740 Ischnosiphon, 233 arouma, 234, 236 cannoideus, 234 enigmaticus, 235 gracilis, 235 subsp. gracilis, 235

INDEX

var. scabra, 235 hirsutus, 235 lasseriana, 235 leucophaeus, 235 subsp. leucophaeus, 235 longiflorus, 235 subsp. angustifolius, 235 obliquus, 235 plurispicatus, 241 polyphyllus, 235 puberulus, 235 var. scaber, 235 var. verruculosous, 236 secundus, 242 surumuensis, 236 ursinus, 236 verruculosous, 236 Isidrogalvia, 13 duidae, 13, 14 guianensis, 13 schomburgkiana, 13, 14 Iwakrete-piji, 546 —J— Jaguayak, 620 Jaico, 747 Jidiuey, 258 Jidiway, 260 Jidiwoi, 260 Jödöwoi, 258 Jonanae, 625 Josefina, 674 Juacha, 696 Juasudu, 540 Jubelina, 154 bracteosa, 155 grisebachiana, 155, 155 magnifica, 84, 156 Jucunda tomentosa, 430 Jupunba, 586 trapezifolia, 588 —K— Kairaimundek, 600 Kamadak, 622 Kanayek, 692 Kanwae, 707, 709 Karayek, 622

Kasujö, 226 Kerosene, 544 Klaprothia, 22 mentzelioides, 21, 22 Klugiodendron, 586 laetum, 589 Komik, 624 Konono, 226 Kuaho, 744 Kuarik-yek, 619 Küdadei, 273 Kuduvi, 738 Kuepiyek, 412 Kumaduwa, 36 Kunuri, 741 —L— Lala-ak, 346 Lancetillo, 745 Lanessania, 728 turbinata, 729 Lasiandra bipenicillata, 505 Lasiostoma, 28 bredemeyeri, 31 curare, 32 Lawsonia, 77 alba, 77 inermis, 77 speciosa, 77 Leandra, 350 aristigera, 352, 358 candelabrum, 352, 359 chaetodon, 352, 358 chimantensis, 356 clidemioides, 353 cuspidata, 411 divaricata, 353, 359 edentula, 354 fendleri, 355 francavillana, 354, 360 glandulifera, 354, 360 gorzulae, 354 hylophila, 353 lindeniana, 355, 359 linearis, 301 longisepala, 355 maguirei, 355, 361 neblinensis, 355 phelpsiae, 355, 361 polyadena, 356, 361 procumbens, 356, 359

789

purpurea, 356 rhamnifolia var. macrodon, 353 rufescens, 357 var. asperiuscula, 357 var. glandulosa, 357 sanguinea, 357 subsp. sanguinea, 357, 360 subsp. tepuiensis, 357 secunda, 357 solenifera, 362 stellipilis, 303 steyermarkii, 362 Lechero, 707 Leiostegia, 311 vernicosa, 313 Lengua de tigre, 718 Lengua de vaca, 241 Lengua de vaca hoja fina, 242 Leptoglottis leptocarpa, 653 Leucaena, 641 glabrata, 642 leucocephala, 642 subsp. glabrata, 642, 642 ulei, 661 Liliaceae, 1 Limnocharis, 18 flava var. minor, 18 laforestii, 18, 18 Limnocharitaceae, 16 Lissocarpa, 19 benthamii, 19, 20 stenocarpa, 19 Lissocarpaceae, 19 Li-wadi-wain-mue, 565 Loasaceae, 22 Loevigia, 469 sericea, 470 Loganiaceae, 22 Lombricera, 26 Lophanthera, 156 longifolia, 84, 156, 157 Lophopterys, 157 euryptera, 157, 158 inpana, 158 Lopimia dasypetala, 202 Loranthaceae, 37

790

INDEX

Loranthus acinarius, 49 acutifolius, 59 aduncus, 45 alveolatus, 41 amplexicaulis, 41 biternatus, 49 cinctus, 49 conduplicatus, 45 cucullaris, 50 cupulifer, 50 eugenioides, 59 falcifrons, 50 florulentus, 41 formosus, 55 lanceolatum, 40 magdalenae, 45 nitens, 44 occidentalis, 41 orinocensis, 45 paniculatus, 45 podopterus, 44 pterygopus, 44 punctatus, 40 puracensis, 40 pyrifolius, 45 retroflexus, 45 rhynchanthus, 50 robustus, 55 ruficaulis, 41 speciosus, 55 stelis, 45 syringifolius, 56 tagua, 40 Loreya, 362 acutifolia, 363 arborescens, 363 brunnescens, 363 collatata, 363 huberi, 288 maguirei, 363 mespiloides, 363, 364 minor, 363 mucronata, 340 ovata, 363, 365 quadrifolia, 363 spruceana, 363, 365 strigosa, 363 Loxania, 38 Lysiloma guachapele, 671 Lythraceae, 59 Lythrum

carthagenense, 65 —M— Macairea, 366 albiflora, 375 aspera, 375 axilliflora, 367, 369 calvescens, 368 cardonae, 367, 369 chimantensis, 367, 370 duidae, 367 lanata, 367, 370 subsp. eglandulosa, 368 lasiophylla, 368, 371 linearis, 368 maguirei, 375 maroana, 368, 370 multinervia, 368, 369 neblinae, 368, 372 pachyphylla, 368, 372 parvifolia, 368, 372 rigida, 368, 373 rufescens, 368, 371 schultesii, 375 spruceana, 368, 374 stylosa, 368, 374 thyrsiflora, 373, 375 Macairicao, 739 Macho, 475 Maclura, 719 tinctoria, 720 subsp. tinctoria, 720, 720 Macrocentrum, 375 angustifolium, 376, 377 anychioides, 376, 377 brevipedicellatum, 376, 378 chimantense, 376 cristatum, 376 var. parviflorum, 376 droseroides, 376, 378 glandulosum, 496 huberi, 379 longidens, 379 maguirei, 377, 379 minus, 377, 379 neblinae, 378, 379 pusillum, 496 repens, 377, 379 rubescens, 378, 379 steyermarkii, 378, 379

Macrosamanea, 643 amplissima, 644 aquatica, 644 basijuga, 680 consanguinea, 644 discolor, 644 pedicellaris, 600 pubiramea, 644 var. lindsaeifolia, 644 simabifolia, 644, 645 spruceana, 644 Magnolia chimantensis, 82 roraimae, 82 ptaritepuiana, 82 Magnoliaceae, 80 Maieta, 379 glandulifera, 516 guianensis, 380, 381 juruensis, 301 neblinensis, 380, 380 poeppigii, 380, 381 Maipá, 620 Majagua, 696, 736, 738 Majomo, 696 Makumik, 661 Malache chiapensis, 207 scabra, 207 Malachra, 199 alceifolia, 199, 200 fasciata, 199 heptaphylla, 200 kegeliana, 199 palmata, 200, 201 radiata, 200, 201 sect. Peltaea, 207 trinervis, 209 Mallophyton, 382 chimantense, 382, 382 Malpighia, 158 argentea, 101 armeniaca, 101 bannisterioides, 177 crassifolia, 118 elegans, 174 emarginata, 159 glabra, 159, 160 glandulifera, 101 punicifolia, 159 reticulata, 144 spicata, 123 ternifolia, 153

INDEX

verbascifolia, 125 volubilis, 167 Malpighiaceae, 82 Malva mansa, 211 Malvaceae, 186 Mancalina, 36 Manewa, 36 Manganaroa, 590 alemquerensis, 591 articulata, 591 Manglilla, 768 ferruginea, 770 Manichi, 696, 722 Mankowa, 36 Manteco, 116, 123, 124 Manteco de agua, 118, 124 Manteco merey, 116 Maquira, 720 calophylla, 721, 722 coriacea, 721, 721 costaricana, 722 guianensis, 721 subsp. costaricana, 722 subsp. guianensis, 722 sclerophylla, 722 Maquira guianensis, 721 Maracanthus, 42 Maranta, 237 amabilis, 226 amplifolia, 238 arouma, 234 arundinacea, 238, 239 cachibou, 226 casupito, 225 compressa, 231 divaricata, 238 gibba, 238 gracilis, 235 humilis, 238, 239 jacquinii, 247 linearis, 238, 240 lutea, 226, 247 micans, 226 obliqua, 235 protracta, 240 ruiziana, 240 rupicola, 240 spicata, 242 tonckat, 247 Marantaceae, 219 Marcetia, 383 andicola, 383 cordigera

var. andicola, 383 juniperina, 383 taxifolia, 383, 383 Marcgravia, 249 coriacea, 250, 252 eichleriana, 251 maguirei, 251 parviflora, 251 var. macrophylla, 251 var. sprucei, 252 patellulifera, 251 pedunculosa, 251 punctifolia, 251 purpurea, 251 roraimae, 250 sororopaniana, 251 sprucei, 252 steyermarkii, 252 Marcgraviaceae, 248 Marcgraviastrum, 253 mixtum, 253, 254 pendulum, 254 Marepillo, 589 Marima, 696, 698, 701, 721 Maripo, 589 Marmaroxylon, 678 claviflorum, 680 collinum, 681 ramiflorum, 686 Marshallfieldia, 281 Maruetije, 739 Maruetiji, 739 Marupa, 588 Masaguaro, 671, 674 Masarico, 738, 744, 745 Mascagnia benthamiana, 167 bracteosa, 155 castanea, 164 complicata, 128 cordifolia var. peruviana, 166 cynanchifolia, 164 dissimilis, 164 divaricata, 165 eggersiana, 165, 168 glandulifera, 165 hondensis, 167 lehmanniana, 129 liesneri, 165 macrodisca, 166, 168 materdei, 166 multiglandulosa

791

var. surinamensis, 169 nervosa, 166 nitida, 128 ovatifolia, 166 poeppigiana, 166 schizoptera, 167 schunkei, 166 sepium, 167, 168 sericans, 167 sinemariensis, 84, 167 stannea, 167 subsericea, 166 surinamensis, 169 tenuifolia var. eggersiana, 165 volubilis, 167 Masjrata, 709 Maskedek, 719 Matapalo, 696, 706, 707, 708, 709, 710 Matekesashe, 250 Mayaca, 261 fluviatilis, 261, 262 longipes, 261, 262 sellowiana, 261, 262 Mayacaceae, 260 Mayjama, 536 Meisneria cordifolia, 499 microlicioides, 499 Melastoma acinodendron, 401 adscendens, 282 alata, 402 albicans, 402 aplostachya, 403 aquatica, 483 arborescens, 363 biglomerata, 404 bivalve, 277 capitellata, 300 chrysophylla, 408 ciliata, 409 dichotoma, 494 dodecandra, 410 elata, 411 elegans, 301 fulva, 408 grandiflora, 494 grossularioides, 287 heteroneura, 301 hirta, 301 holosericea, 413

792

INDEX

[Melastoma] ibaguense, 414 impetiolaris, 415 laevigata, 415 longifolia, 416 maieta, 380 micrantha, 484 minutiflora, 419 nervosa, 420 octona, 302 physophora, 516 prasina, 422 punctata, 423 purpurascens, 273 purpurea, 273 racemosa, 424 ramiflora, 340 rubiginosa, 425 rubra, 303 rufescens, 425 scorpioides, 431 splendens, 428 strigillosa, 303 succosa, 346 theaezans, 430 tococo, 516 tomentosa, 430 trinervia, 431 trivalvis, 277 villosa, 317 Melastomataceae, 263 Melia, 540 azedarach, 540, 541 Meliaceae, 528 Meliandra, 527 Melvilla speciosa, 69 Mendoncia, 550 bivalvis, 551, 553 cardonae, 552, 553 coriacea, 552 hirsuta, 551 hoffmannseggiana, 552, 553 neblinensis, 552 obovata, 552 phalacra, 552 pubescens, 552 schomburgkiana, 552 sprucei, 552 steyermarkii, 554 williamsii, 554 Mendonciaceae, 550

Mendozia hirsuta, 551 pubescens, 552 Menispermaceae, 554 Menyanthaceae, 578 Menyanthes indica, 579 Merecucuri, 744 Meriania, 384 broccha, 384 crassiramis, 384, 386 duidae, 282 ferruginea, 282 neblinensis, 385 ornata, 385 paraensis, 385 sclerophylla, 385, 386 urceolata, 385, 385 Meseyek, 588 Mezclo, 544 Mezia, 169 curranii, 170 huberi, 170, 171 includens, 172 rufa, 172 Miconia, 387 abbreviata, 401 abysmophila, 401 acinodendron, 401, 433 acutifolia, 401, 434 affinis, 401, 433 aguitensis, 402, 432 alata, 402, 435 albicans, 402, 436 alborufescens, 402, 437 alternans, 403, 436 amacurensis, 403 ampla, 403, 438 amplexans, 430 anceps, 431 aplostachya, 388, 403, 434 argyrophylla, 404, 437 var. attenuata, 404 arirambae, 402 aristata, 404 bifrons, 416 biglandulosa, 404 biglomerata, 404, 434 borjensis, 405, 439 brachybotrya, 405 bracteata, 405, 440 brevipes, 405, 441 bubalina, 406, 441

buchtienii, 407 buntingii, 406 cachimbensis, 402 cacumina, 406 campestris, 406, 441 carassana, 407, 440 caryophyllea, 515 caudiculata, 431 cautis, 407, 442 centrandra, 407 centrodesma, 407, 439 ceramicarpa, 408, 435 var. violacea, 408 chrysophylla, 408, 443 ciliata, 409, 444 compacta, 407 congesta, 422 crassinervia, 409 cristulata, 422 cruegeriana, 414 curta, 409 subsp. curta, 410, 442 subsp. ptariensis, 410 decurrens, 410, 443 demerarensis, 405 dioica, 410, 445 dispar, 410, 446 disparilis, 416 dodecandra, 410, 442 egensis, 411 elaeoides, 411, 447 elata, 411, 447 erioneura, 516 erythrophylla, 516 erythropila, 408 eugenioides, 411, 443 fallax, 412, 444 fendleriana, 421 fragilis, 412 fragrans, 388, 412, 448 fulva, 408 var. angustifolia, 408 var. aubletiana, 408 glossocentra, 419 gratissima, 412 grossidentata, 413, 448 guaiquinimae, 413, 444 subsp. angustifolia, 413 holosericea, 413, 449 huberi, 413 hypargyrea, 428 hypoleuca, 414 ibaguensis, 414

INDEX

iluensis, 414, 445 impetiginosa, 425 impetiolaris, 415 var. spruceana, 415, 449 inaequalifolia, 415 involucrata, 403 iodopila, 408 kavanayensis, 415 laevigata, 415 lambertiana, 416 lasseri, 415, 450 lateriflora, 416 subsp. lateriflora, 416, 450 subsp. monticellensis, 416 lepidota, 416, 445 leschenaultiana, 408 longifolia, 416 var. aubletiana, 408 longispicata, 417, 451 longistyla, 404 lourteigiana, 417 macrophylla var. serrulata, 427 macrostachya, 403 macrothyrsa, 417, 452 macrotis, 417, 453 marginata, 418, 452 mariae, 418 maroana, 418, 451 matthaei, 418 megaphylla, 403 micrantha, 432 microcarpa, 401 minutiflora, 419, 454 mirabilis, 388, 419, 454 mituana, 419 mucronulata, 422 multinervulosa, 430 myriantha, 419, 455 neblinensis, 420 nervosa, 420, 456 nitens, 518 obovalis, 428 pachypoda, 429 pallida, 411 palmetorum, 428 panicularis, 411 parviflora, 423 perobscura, 420, 468 perturbata, 420 phaeophylla, 421, 454

pileata, 421 pilgeriana, 421 planinervia, 401 plebeia, 405 plukenetii, 421 plumosa, 426 poeppigii, 422 polita, 422, 456 prasina, 422, 458 pseudoaplostachya, 423, 458 pseudocapsularis, 423, 458 pseudorubiginosa, 417 pterophora, 402 pteropoda, 422 pubipetala, 423 punctata, 423, 457 purpurascens, 273 pyrifolia, 423 racemosa, 424 var. ciliata, 409 radulaefolia, 388, 424, 457 revoluta, 422 rhytidophylla, 424, 459 roraimensis, 425, 459 rubiginosa, 425, 459 var. kuhlmannli, 405 var. platyura, 425 rufescens, 425, 460 var. grandifolia, 425 ruficalyx, 425 rugosa, 426, 461 rupestris, 426 rupticalyx, 426 schomburgkii, 416 scorpioides, 431 scrobiculata, 424 semota, 407 septuplinervis, 420 serialis, 427 serrulata, 427, 462 setimarginata, 518 silicicola, 427, 463 solmsii, 427, 464 splendens, 388, 428, 465 spondylantha var. leschenaultiana, 408 sprucei, 428, 465 stenostachya, 428, 463 stephananthera, 428

793

steyermarkii, 429, 460 strigosa, 429 subciliata, 519 subtriloba, 407 superba, 388, 429, 466 surinamensis, 422 sylvestris, 407 tessmannii, 416 tetraspermoides, 429 theaezans, 430, 467 tillettii, 430, 462 tinifolia, 430, 455 var. roraimensis, 430 tomentella, 427 tomentosa, 388, 430 traillii, 431 trinervia, 388, 431, 467 trinitatis, 419 truncata, 431, 455 undata, 431 virgulata, 409 wittii, 432 yatuensis, 432 Microlicia, 468 benthamiana, 468, 469 bivalvis, 277 brevifolia, 277 bryanthoides, 468 guanayana, 468 limnobios, 278 recurva, 279 steyermarkii, 468 Microphysa, 512 rotundifolia, 518 Mijarro, 545 Mimosa, 645 adhaerens, 649 alba, 619 arenosa, 648 var. arenosa, 649 asperata, 652 bauhiniaefolia, 649 bipinnata, 611 bourgonii, 619 brachycarpoides, 649 brevispica, 650 camporum, 649 casta, 649 cataractae, 651 cauliflora, 681 colombiana, 649 coriacea, 626 corymbosa, 614

794

INDEX

[Mimosa] debilis, 649 var. panamensis, 649 var. debilis, 650 dormiens, 650 fagifolia, 622 farnesiana, 591 fasciculata var. ernestiana, 648 fastuosa, 620 flavescens, 649 flaviseta, 649 flexuosa, 595 guanchezii, 650 guianensis, 674 hirsutissima, 650, 656 hispidula, 653 hostmannii, 649 huberi, 650 humilis, 650 inga, 620, 628 ingoides, 622 intermedia, 650 invisa, 650 subsp. spiciflora var. spiciflora, 650 jiramenensis, 656 latifolia, 681 laurina, 622 leucocephala, 642 lucida, 625 lutea, 595 macracantha, 595 macrophylla, 623 mangensis, 610 martensis, 649 micracantha, 654 micrantha var. plurijuga, 654 microcephala, 651 subsp. cataractae, 651 var. lumaria, 651 subsp. microcephala, 651 var. communis, 651 var. microcephala, 651 multijuga, 624 myriadenia, 651 natans, 659 nitida, 625 notata, 649 orinocoënsis, 652

orthocarpa, 652, 657 palpitans, 655 panamensis, 649 parae, 621 pellita, 652 var. pellita, 652, 657 peregrina, 598 pigra, 652 pilosula, 625 piscatorum, 652 plena, 659 polydactyla, 652 polystachia, 611 portoricensis, 678 prostrata, 659 pudica, 653 var. tetrandra, 653, 657 var. unijuga, 653 pulcherrima, 674 pusilla, 649 quadrivalvis, 653 var. leptocarpa, 653, 656 rosea, 668 rubiginosa, 625 rufescens, 653 var. amnis-nigri, 654 var. rufescens, 654 sagotiana, 655 saman, 673 santanderensis, 655 sapindoides, 626 scabrella, 654 schomburgkii, 654 schrankioides, 654 var. sagotiana, 655 var. schrankioides, 655 sensitiva, 655 somnians, 655 subsp. somnians, 655 var. deminuta, 655 var. somnians, 655 somniculosa, 655 spiciflora, 650 splendens, 626 spuria, 628 surumuënsis, 655 tamarindifolia, 595 tarda, 655 tergemina, 602 tobagensis, 655 tomentosa, 650 trapezifolia, 588 umbellifera, 627

unguis-cati, 668 unijuga, 653 velloziana, 656 var. jiramenensis, 656 viva, 656 xanthocentra, 656 subsp. xanthocentra var. xanthocentra, 656 xantholasia, 648 Mimosaceae, 580 tribe Ingeae genus D Miocarpus paludosus, 278 Misionero negro, 695, 696 Molenillo, 739 Mollinedia, 689 glabricaulis, 690 killipii, 690 laurina, 690 neblinensis, 690 ovata, 690, 690 ptariensis, 690 repanda, 691 roraimensis, 691 yaviensis, 691 Molluginaceae, 686 Mollugo, 688 radiata, 688 verticillata, 688, 688 Môlôkoto kahi, 474 Monimiaceae, 689 Monochaetum, 469 bonplandii, 470, 470 sericeum, 470 Monostiche, 221 Monotaceae, 691 Monotagma, 240 duidae, 242 laxum, 241 ovatum, 241 plurispicatum, 241, 243 rhodanthum, 241 secundum, 241, 244 spicatum, 242, 243 surinamense, 242 tomentosum, 242 vaginatum, 242, 244 yapacanensis, 242, 242 Mora, 720 Moraceae, 693 Moringa, 731 oleifera, 730, 731

INDEX

Moringaceae, 729 Morongia, 645 Morus tinctoria, 720 subsp. tinctoria, 720 Moschoxylum cipo, 544 pleeanum, 545 surinamense, 546 Mouriri, 470 acutiflora, 473, 477 var. oligantha, 474 angulicosta, 474 angustifolia, 474 brevipes, 474, 476 cauliflora, 474 collocarpa, 474 densifoliata, 474 ficoides, 474, 477 grandiflora, 474, 478 guianensis, 474 huberi, 474 latihila, 475 longifolia, 475 myrtifolia, 475 myrtilloides, 475 subsp. orinocensis, 475 nervosa, 475 nigra, 475 pauciflora, 475 plasschaerti, 475 polyantha, 474 princess, 474 rhizophorifolia, 475, 479 sagotiana, 476 sideroxylon, 476 subumbellata, 476 ulei, 474 uncitheca, 476, 478 vernicosa, 476 weddellii, 474 Moy sebe, 260 Moya, 622 Mucumi, 567 Mure, 607 Musa, 731 ×paradisiaca, 732, 732 sapientum, 732 Musaceae, 731 Mutumutu, 706, 707, 709, 710 Myriaspora, 479 decipiens, 480

egensis, 480, 480 surinamensis, 480 Myrica, 734 rotundata, 733, 734 Myricaceae, 733 Myristica calophylla, 744 cuspidata, 744 debilis, 736 elongata, 744 fatua, 746 hostmannii, 738 laurifolia, 736 longicuspis, 744 macrophylla, 739 membranacea, 744 mexicana, 736 mocoa, 746 panamensis, 746 paradoxa, 739 pavonis, 745 platysperma, 741 punctata, 744 rugulosa, 745 sebifera, 745 var. cordifolia, 746 var. curvinervia, 746 var. longifolia, 746 sect. Compsoneura, 735 sect. Iryanthera, 737 sect. Sychnoneura, 742 sect. Virola, 742 sprucei, 736 surinamensis, 746 theiodora, 746 uaupensis, 744 venosa, 747 var. pavonis, 745 var. poeppigii, 747 Myristicaceae, 734 Myrmidone, 512 lanceolata, 517 macrosperma, 517 Myrosma, 245 cannifolia, 245, 245 unilateralis, 231 Myrsinaceae, 748 Myrsine, 768 coriacea, 770 subsp. coriacea, 770 subsp. reticulata, 770 ferruginea, 770 guianensis, 770, 771

795

macrocarpa, 770 maguireana, 770 minima, 770, 771 myricoides, 770 nitida, 771, 772 ovalifolia, 770 pellucida, 772 perpauciflora, 772 picturata, 772 resinosa, 772 roraimensis, 772 spicata, 763 Myrtus longifolia, 475 —N— Narda, 28 Naucleopsis, 723 mello-barretoi, 723 oblongifolia, 723, 723 Neblinanthera, 481 cumbrensis, 481, 482 Nepsera, 482 aquatica, 482, 483 Neptunia, 658 natans, 659 oleracea, 658, 659 plena, 659 prostrata, 659 surinamensis, 659 Newtonia sect. Neonewtonia, 669 suaveolens, 670 Nietneria, 15 corymbosa, 14, 15 paniculata, 14, 15 Niopa peregrina, 598 No’samo adü, 225 Nombró-poh, 271 Nopo, 739 Norantea, 254 guianensis subsp. guianensis, 256 subsp. japurensis, 255, 256 var. gracilis, 256 japurensis, 256 mixta, 254 paraensis, 256 peduncularis, 254 pendula, 254

796

INDEX

[Norantea] subsect. Marcgraviastrum, 253 tepuiensis, 257 Notanthera lanceolata, 40 Noterophila beccabunga, 277 brevifolia, 277 limnobios, 278 Notocentrum, 384 Noyera, 723 Nymphoides, 579 humboldtiana, 579 indica, 579, 579 —O— Octopleura ciliata, 416 micrantha, 484 Odontandra acuminata, 544 Odontocandra karstenii, 544 Odontocarya, 571 hederifolia var. canescens, 572 mallosperma, 572, 573 paupera, 572 sp. A, 574 tamoides, 572 var. canescens, 572 var. tamoides, 574 Oerstedianthus, 749 Ogcodeia, 723 oblongifolia, 723 Oí Pharatæ, 708 Olisbea, 470 rhizophorifolia, 475 Olmedia calophylla, 721 caurensis, 726 laevis, 725 maxima, 721 obliqua, 721 polycephala, 719 tomentosa, 719 virgata, 728 Olmedioperebea, 720 calophylla, 721 sclerophylla, 722 Olmediophaena, 720

maxima, 721 obliqua, 721 Onoctonia calcarata, 278 crassipes, 278 pauciflora, 278 Opisthocentra, 483 clidemioides, 483, 484 Oreja de tigre, 567 Orthomene, 574 hirsuta, 575 schomburgkii, 574, 575 verruculosa, 575 Oryctanthus, 40 alveolatus, 41 var. amplexicaulis, 41 var. kuijtii, 41 amplexicaulis, 41 botryostachys, 41 florulentus, 41, 42 laceratus, 41 occidentalis, 41 phthirusoides, 42 ruficaulis, 41 Oryctina, 42 myrsinites, 43, 43 Osbeckia aubletiana, 494 glomerata, 493 sect. Chaetolepis, 294 sect. Pterolepis, 492 striphnocalyx, 512 Ossaea, 484 ciliata, 416 disparilis, 416 mavacana, 484, 485 micrantha, 484, 485 trichopoda, 353 trinitensis, 311 Osteophloeum, 740 platyspermum, 741, 741 sulcatum, 741 Oxymeris, 350 aristigera, 352 asperiuscula, 357 chaetodon, 352 cuspidata, 411 glandulifera, 354 lindeniana, 355 rufescens, 357 secunda, 357

—P— Pachygone domingensis, 571 graciliflora, 571 Pachyloma, 486 huberioides, 486, 487 nanum, 488 pusillum, 487, 488 scandens, 486 setosum, 487, 488 Pachymeria, 384 Pakaraimaea, 691 dipterocarpacea, 692 subsp. nitida, 692, 692 Pal de brasil, 728 Palala calophylla, 744 cuspidata, 744 debilis, 736 elongata, 744 macrophylla, 739 membranacea, 744 mexicana, 736 mocoa, 746 pavonis, 745 platysperma, 741 punctata, 744 rugulosa, 745 sprucei, 736 surinamensis, 746 theiodora, 746 uaupensis, 744 venosa, 747 Palmolmedia, 723 Palo aro de atarraya, 736 Palo blanco, 666 Palo carbón, 747 Palo de agua, 423 Palo de Brasil, 696, 725, 729 Palo de carbón, 19 Palo de escoba, 363 Palo de gallineta, 745 Palo de lameya, 19 Palo de loro, 32 Palo de morrocoy, 718 Palo molenillo, 747 Palo vieja, 708 Pancratium guianense, 12 petiolatum, 12 tubiflorum, 12 undulatum, 12

INDEX

Paraclarisia, 726 Paranovillo, 649 Parathesis viridis, 774 Pareira-brava, 567 Parkia, 659 auriculata, 661 barnebyana, 660 discolor, 661, 662 igneiflora, 661 nitida, 661 oppositifolia, 661 panurensis, 661 pectinata, 661 pendula, 661, 663 truncata, 661 ulei, 661 ulei var. surinamensis, 661 velutina, 664 Passovia, 43 guyanensis, 44 micrantha, 38 myrsinites, 43 orinocensis, 45 pyrifolia, 45 suaveolens, 45 Pata de morrocoy, 622 Pata de Morrocoy, 719 Pate, 625 Pavonia, 201 angustifolia, 202 cancellata, 202, 203 castaneifolia, 202, 204 chiapensis, 207 dasypetala, 202, 205 fruticosa, 202 guanacastensis, 202 imatacensis, 203 malacophylla, 203, 206 paludicola, 205, 207, racemosa, 207 scabra, 207 sessiliflora, 209 sidifolia, 203, 207 speciosa, 209 spicata, 207 surumuensis, 209 ulbrichiana, 196 Peckia, 751, 757 guyanensis, 760 longifolia, 761

penduliflora, 761 prieureii, 762 purpurea, 763 surinamensis, 763 Pega-pego, 661 Pehria, 77 compacta, 78, 79 Peine de morocoto, 256 Peltaea, 207 aliculata, 209 sessiliflora, 209 speciosa, 208, 209 surumuensis, 208, 209 trinervis, 208, 209 Pentaclethra, 665 filamentosa, 665 macroloba, 664, 665 Peonía, 536 Perebea, 723 costaricana, 722 elegans, 718 guianensis, 725 subsp. guianensis, 724, 725 laurifolia, 722 Petaloma, 470 mouriri, 474 myrtilloides, 475 Phainantha, 488 laxiflora, 490 maguirei, 489, 490 myrteoloides, 489, 490 steyermarkii, 489, 490 Pharmacosycea, 702 Phõi æ’mi, 728 Phõi pharatæ, 708 Phõi tæphi, 728 Phrygilanthus, 39 acutifolius, 59 coriaceus, 59 eugenioides lateovatus, 59 vulgaris, 59 lanceolatum, 40 megathermicus, 47 punctatus, 40 subgen. Tripodanthus, 58 Phrynium altissima, 224 densum, 225 ellipticum, 225 micans, 226 ovatum, 229

797

pusillum, 226 Phthirusa, 43 adunca, 45 var. magdalenae, 45 var. orinocensis, 45 var. rigidifolia, 45 anastyla, 45 bernardiana, 38 bisexualis, 44 calloso-albida, 44 castillana, 44 cothurnata, 45 disjectifolia, 44, 46 gracilis, 56 guyanensis, 44, 47 hippocrateoides, 56 huberi, 45 intermedia, 38 maculata, 42 maritima, 45 megathermica, 45 micrantha, 38 var. bolivariensis, 38 myrsinites, 43 var. savannarum, 43 nitens, 44, 47 orinocensis, 45 paniculata, 45 percrassa, 45 perdivergens, 45 perforata, 44 podoptera, 44 ptariana, 56 pyramidalis, 45 pyrifolia, 45 retroflexa, 45 roraimense, 39 rubromicans, 45 savannarum, 43 stelis, 45, 46 stenophylla, 45, 46 Phyllodes inocelphalum, 226 platyphyllum, 226 Phyllopus, 338 martiusii, 340 Piassám, 745 Pica pica, 567 Pilón, 622 Pio-yek, 272 Piptadenia, 665 biuncifera, 666 catenaeformis, 607

798

INDEX

[Piptadenia] guianensis, 674 imatacae, 666 leucoxylon, 666, 667 moniliformis, 666 niopo, 598 peregrina, 598 polystachya, 674 rubescens, 666 sect. Pityrocarpa, 665 speciosa, 667 suaveolens, 670 tocantina, 674 viridiflora, 666 Piratinera, 695 guianensis, 696 Pirboi-yek, 260 Pir-boi-yek, 258 Pirboyek, 258 Piruói, 260 Pithecellobium, 667 acreana, 587 adenophorum, 587 adiantifolium, 644 amplissimum, 644 aquaticum, 644 arenarium, 588 arenicola, 644 basijugum, 680 bijugatum, 615 catenaeformis, 607 cauliflorum, 681 claviflorum, 680 collinum, 681 consanguineum, 644 corymbosum, 614 dinizii, 686 discolor, 644 divaricatum, 680 ferrugineum, 588 floribundum, 588 forfex, 668 glabripetalum, 596 glomeratum, 680 gonggrijpii, 614 guachapele, 671 guaricense, 668 inaequale, 681 inopinatum, 673 jupunba, 588 laetum, 589 larensis, 669 lasiopus, 681 latifolium, 681

leucophyllum, 589 levelii, 589 lindseifolium, 644 longepedatum, 671 longipedunculatum, 589 marginatum, 615 microcalyx, 589 microchlamys, 669 microphyllum, 669 multiflorum, 596 nuriense, 673 panurense, 615 parviflorum, 668 pedicellare, 600 pilosulum, 681 pubescens, 668 pulchellum, 669 racemosum, 686 ramiflorum, 681, 686 roseum, 668, 668 saman, 673 samanigua, 671 schomburgkii, 613 sect. Abaremotemo, 586 sect. Caulanthon, 678 sect. Chloroleucon, 609 sect. Samanea, 671 ser. Carnosae, 671 ser. Coriaceae, 643 ser. Corymbosa, 613 sect. Unguis-cati, 667 simabifolium, 644 spruceanum, 644 subaquaticum, 644 trapezifolium, 588 unguis-cati, 668 unifoliolatum, 686 villiferum, 590 Pityrocarpa, 665 viridiflora, 666 Plananillita, 235 Plátano, 732 Platycentrum, 350 clidemioides, 353 Pleroma bipenicillata, 505 Pogonorhynchus amplexans, 430 Pomarrosa, 287, 419 Poponax, 590 canescens, 595 farnesiana, 591 flexuosa, 595 lutea, 595

macracantha, 595 Poroi, 741 Poté, 624 Poteranthera, 490 calcarata, 278 crassipes, 278 duidae, 500 foliosa, 500 minor, 500 pauciflora, 278 pusilla, 490, 490 Potojimujii, 729 Prooiji, 741 Psathyranthus, 48 Pseudabutilon, 210 spicatum, 189 umbellatum, 209, 210 Pseudolmedia, 725 coriacea, 721 laevigata, 725, 725 laevis, 725, 726 multinervis, 725 scabra, 718 Pseudopiptadenia, 669 suaveolens, 669, 670 Pseudosamanea, 671 guachapele, 670, 671 Pseudosorocea, 726 sprucei, 728 uaupensis, 728 Psittacanthus, 48 acinarius, 49, 51 biternatus, 49, 51 brachynema, 49, 52 calyculatus, 55 var. wurdackii, 55 chrismarii, 50 cinctus, 49, 51 var. guainiae, 49 clusiifolius, 49, 53 var. pseudojulianus, 49 collum-cygni, 50, 54 corynocephalus, 50 cucullaris, 50, 52 cupulifer, 50 falcifrons, 50 intermedius, 55 julianus, 50, 54 lasianthus, 50 montis-neblinae, 50 peronopetalus, 50, 53 rhynchanthus, 50 subsp. wurdackii, 55 robustus, 54, 55

INDEX

semiarticulatus, 55 warmingii, 49 Pterandra, 172 flavescens, 173, 174 guianensis, 174 sericea, 174 Pterogastra, 491 divaricata, 491, 492 glabra, 492 major, 492 minor, 491, 492 Pterolepis, 492 glomerata, 493 lasiophylla, 368 pumila, 493 var. procera, 493 striphnocalyx, 512 trichotoma, 493, 493 Ptilanthus, 321 —R— Rabo de arara, 256 Rabo de guaca, 256 Rapanea, 768, 770 ambigua, 770 coriacea, 770 ferruginea, 770 guianensis, 770 minima, 772 myricoides, 770 nitida, 772 oblonga, 770 ovalifolia, 770 pellucida, 772 perforata, 772 resinosa, 772 reticulata, 770 roraimensis, 772 Raspa, 651 Redutea, 190 heterophylla, 192 Resedá, 77 Reventillo, 744 Rhexia acisanthera, 279 angusturensis, 493 aquatica, 483 bivalvis, 277 bonplandii, 470 canescens, 470 divaricata, 492 glomerata, 493 gracilis, 506

incana, 470 indecora, 272 juniperina, 383 leptophylla, 312 longifolia, 506 nummularioides, 312 polystachya, 273 pumila, 493 recurva, 279 rotundifolia, 327 sileniflora, 273 taxifolia, 383 tenella, 321 trichotoma, 493 trivalvis, 277 uniflora, 279 villosa, 313 Rhynchanthera, 494 ambigua, 494 cardonae, 494 dichotoma, 494 grandiflora, 494, 495 modesta, 495 orinocensis, 494 serrulata, 495 Roemeria, 768 Roezlia, 469 Rosa Jamaica, 197 Rotala, 79 apetala, 79 mexicana, 79 var. apetala, 79 var. spruceana, 79 ramosior, 79, 80 Rouhamon, 28 bredemeyeri, 31 guianensis, 32 pedunculatum, 31 Rutæ, 729 —S— Sacau-yek, 421 Saccolena, 495 Sacuma, 741 Sada, 728 Sagraea capillipes, 299 minutiflora, 302 neurocarpa, 486 Sahagunia, 699 Sajurua, 696 Sakau, 430 Sakuma, 739

799

Salpinga, 495 cristata, 376 glandulosa, 496, 496 maguirei, 496, 497 parviflora, 376 pusilla, 496, 497 secunda, 496, 497 Salvadora surinamensis, 763 Samán, 589, 673 Samanea, 671 adiantifolia, 644 corymbosa, 614 inopinata, 673 pedicellaris, 600 saman, 672, 673 samanigua, 671 simabifolia, 644 Samanigua, 671 Samara, 768 coriacea, 770 Samyda guidonia, 536 Sandemania, 498 glandulosa, 498 hoehnei, 498, 498 Sangrino blanco, 745 Sangrito, 738 Sangrito blanco, 745 Saranthe urceolata, 238 Sarcopera, 256 flammifera, 257 tepuiensis, 257 subsp. coccinea, 258 subsp. tepuiensis, 257, 258 Sarmentaria, 281 decora, 282 Sarsa hueca, 196 Schrankia, 645 leptocarpa, 653, 645 Schwerinia, 384 Sciadotenia, 575 cayennensis, 576, 577 sprucei, 576, 577 Scleroxylum, 768 Sebophora, 742 Semeruco, 159 Senegalia, 590 glomerosa, 594 paniculata, 595 podadenia, 595 tamarindifolia, 595

800

INDEX

Serianthes inopinata, 673 Sesejudi, 709 Shada, 696 Shara, 728 Sharama, 728 Shiigui, 708 Shina-teu, 568 Shopa, 719 Shú-she, 496 Sida, 210 acuta, 211, 212 aggregata, 211, 213 angustissima, 211, 214 carpinifolia, 211 ciliaris, 211, 214 contracta, 219 cordifolia, 211, 212 crispa, 194 excelsior, 219 fruticosa, 202 glomerata, 215 hernandioides, 219 jamaicensis, 215 linifolia, 214, 215 malacophylla, 203 micrantha, 216 periplocifolia, 219 radiata, 200 rhombifolia, 215 savannarum, 211 serrata, 213, 215 setifera, 211 setosa, 215 surinamensis, 215 umbellata, 210 viarum, 215 Sidastrum, 215 micranthum, 216, 216 Sijanama, 546 Simure, 19, 707 Sipapoa, 129 cordata, 131 ferruginea, 131 glaucifolia, 131 hypoleuca, 132 kunhardtii, 132 rufa, 132 steyermarkii, 132 stipularis, 132 vestita, 134 Siphanthera, 499 capitata, 502 cordifolia, 499, 501

var. glomerata, 499 cowanii, 500 dawsonii, 500 duidae, 500, 501 fasciculata, 500, 501 foliosa, 500, 501 hostmannii, 501, 502 jenmanii, 499 microlicioides, 499 paraensis, 502 tatei, 502 Siphantheropsis, 366 Sirichá, 273 Skoliopterys, 128 lehmanniana, 129 Sorocea, 726 amazonica, 728 guayanensis, 728 hirtella, 728 muriculata, 727 subsp. muriculata, 728 subsp. uaupensis, 727, 728 opima, 728 pubivena, 728 subsp. hirtella, 727, 728 subsp. oligotricha, 728 sprucei, 728 subsp. sprucei, 728 uaupensis, 728 Souroubea, 258 dasystachya, 258, 259 guianensis, 259 subsp. cylindrica, 259, 260 var. cylindrica, 260 var. tomentella, 260 Spachea, 174 elegans, 174, 175 Spennera, 270 acuminifolia, 271 acutiflora, 272 alata, 272, 276 annua, 271 anomala, 271 aquatica, 483 caulialata, 272 chaetodon, 352 dichotoma, 271 var. lanceolata, 271 dysophylla, 271 fragilis, 273 grandifolia, 273 indecora, 272

kappleriana, 272 lancifolia, 276 laxa, 272 martinicensis, 273 ornata, 273 pellucida, 273 polystachya, 273 rostellata, 272 rubricaulis, 272 sieberi, 273 sileniflora, 273 sphaeranthera, 271 tetraptera, 271 uliginosa, 273 viscida, 276 viscosa, 272 Sphaerine, 4 Sphaerogyne, 512 Spigelia, 26 anthelmia, 26, 27 filipes, 28 gracilis, 27, 28 humilis, 27, 28 multispica, 28 nervosa, 26 polystachya, 28 Staphidiastrum aphananthum, 299 attenuatum, 299 coriaceum, 303 Staphidium conglomeratum, 300 sessiliflorum, 303 Stenocalyx involutus, 172 Stigmaphyllon, 176 adenodon, 177 var. adenodon, 177 bannisterioides, 177 brachiatum, 179 convolvulifolium, 177 fulgens, 179 grenadense, 177 heterophyllum, 179 hypoleucum, 179 martianum, 179 monancistrum, 179 ovatum, 177 puberum, 178 sinuatum, 84, 178, 179 splendens, 179 Stratiotes nymphoides, 17 Stromanthe, 246

INDEX

jacquinii, 247 lutea, 247 tonckat, 246, 247 Struthanthus, 55 chimantensis, 56 cupulifer, 56 dichotrianthus, 56, 57 var. dichotrianthus, 56 var. lasserianus, 56 dissimilis, 56 eichlerianus, 56 gracilis, 56 var. mucronatus, 56 granulatus, 56 micranthus, 38 mucronatus, 56 nitens, 44 pseudolepidotus, 44 pterygopus, 44 syringifolius, 56, 57 terniflorus, 56 translucens, 56 vulgaris, 56 yavitensis, 56 Strychnos, 28 blackii, 31 bovetiana, 32 brachiata, 31 bredemeyeri, 31, 34 cogens, 31 curare, 32 darienensis, 31 davidsei, 31 diaboli, 31 erichsonii, 32 fendleri, 32, 34 guianensis, 32, 35 jobertiana, 32 lanceolata, 32 mattogrossensis, 32 mitscherlichii, 32 var. mitscherlichii, 33, 36 panamensis, 33 panurensis, 33, 35 peckii, 33, 36 pedunculatum, 31 rondeletioides, 33 ruizii, 31 schomburgkiana, 31 smilacina, 33 smithiana, 33 syntoxica, 33 toxifera, 33

trichostyla, 32 trinitensis, 31 urbanii, 32 Stryphnodendron, 673 angustum, 674 floribundum, 674 guianense, 674 f. floribundum, 674 subsp. guianense, 674 inaequale, 674 levelii, 674 melinonis, 674 microstachyum, 674, 675 polystachyum, 674, 676 pulcherrimum, 674, 675 purpureum, 674 Stylogyne, 772 amazonica, 774 atra, 773, 775 kappleri, 774 lasseri, 773 latifolia, 773 micans, 774 micrantha, 773, 774 orinocensis, 773 spruceana, 773 surinamensis, 774 venezuelana, 773 viridis, 774, 775 Suiguichasto, 741 Suipo, 546 Surenus brownii, 533 Surja, 719 Syena, 261 —T— Tabaca, 671 Tabacilla, 739 Taguapira, 668 Taguaria, 39 puracense, 40 Takuramo, 696 Tani, 745 Tanyeechemen, 661 Tapata, 739 Tararujuru, 741 Tártago, 739 Tateanthus, 503 duidae, 502, 503 Telitoxicum, 577 inopinatum, 578, 578 Tëpo po mën, 250

801

Tereyo, 475 Tetrameris, 311 villosa, 313 Tetrapodenia, 103 glandifera, 104 Tetrapterys, 179 aristeguietae, 181 benthamiana, 167 boliviensis, 184 crebriflora, 183 crispa, 181 discolor, 181 fimbripetala, 182, 185 glaberrima, 183 gracilis, 182 huachamacariensis, 182 includens, 172 julianii, 134 maranhamensis, 182 mucronata, 183 subsp. crebriflora, 183 var. crebriflora, 183 oleifolia, 183, 185 phylladenophora, 184 poeppigiana, 166 pusilla, 84, 183, 184 rhodopteron, 184 silvatica, 183 squarrosa, 184 styloptera, 184, 185 Thalia, 248 geniculata, 247, 248 latifolia, 226 trichocalyx, 248 unilateralis, 231 Thymocarpus, 221 cannoides, 225 Tiamo güire, 666 Tibouchina, 503 adscendens, 279 aspera, 505 var. aspera, 505, 507 var. asperrima, 505 bipenicillata, 505, 508 brachyanthera, 505 catharinae, 505, 509 cryptadena, 505, 508 dissitiflora, 505 duidae, 505, 509 fraterna, 506 subsp. fraterna, 506, 510 subsp. paruana, 506 geitneriana, 506

802

INDEX

[Tibouchina] gracilis, 506, 511 huberi, 506 impressa, 506 kunhardtii, 506, 510 lasiophylla, 368 llanorum, 506, 511 longifolia, 506 multinervia, 368 pseudomollis, 506 sipapoana, 507, 512 spruceana, 507, 512 steyermarkii, 509, 512 striphnocalyx, 512 yavitensis, 512 Tinus guianensis, 749 hostmannii, 774 orinocensis, 773 surinamensis, 774 Tirita, 225, 241 Tirita cania grulla, 235 Tirite, 235 Tirripi, 747 Tococa, 512 acuminata, 516 aristata, 515, 520 aubletii, 516 bolivarensis, 515 subsp. bolivarensis, 515, 520 subsp. occidentalis, 515 bullifera, 515 cardiophylla, 516 caryophyllea, 515 castrata, 516 ciliata, 515, 521 cinnamomea, 515 cordata, 515 coriaceae, 516 coronata, 515, 521 didymophysca, 516 discolor, 516 egensis, 516 erioneura, 516 erythrophylla, 516, 521 erythrostachya, 516 ferruginea, 516 glandulosa, 516 guianensis, 516, 522 hirta, 517 juruensis, 516 lancifolia, 517, 523

lasiostyla, 519 liesneri, 517 longisepala, 516 loretensis, 516 macrophysca, 517, 523 macrosperma, 517, 519 maieta, 380 montana, 518 nitens, 518, 520 obovata, 518 subsp. neblinensis, 518 subsp. obovata, 518, 525 occidentalis, 516 oligantha, 518, 524 pachystachya, 518, 524 pauciflora, 518 planifolia, 519 roreimi, 516 rotundifolia, 518, 522 spruceana, 515 subciliata, 519, 524 subnuda, 516 tepuiensis, 519 subsp. tepuiensis, 519, 525 subsp. glabrata, 519 traillii, 516 ulei, 515 wedellii, 518 Tofieldia duidae, 13 schomburgkiana, 13 Topobea, 526 ferruginea, 526, 526 Tortolito blanco, 536 Traga venado, 564, 575 Tragwena, 564 Trebol sabanero, 215 Tricentrum, 311 leptophyllum, 312 Trichilia, 542 acuminata, 544 cipo, 544 elegans, 544 subsp. elegans, 544 gamopetala, 544 grandis, 546 inaequilatera, 544, 547 lanceolata, 546 lepidota, 544 subsp. leucastera, 544, 547

mazanensis, 544 micrantha, 544 montana var. fendleriana, 545 moritzii, 546 pallida, 545, 548 pleeana, 545, 546 pubescens, 536 quadrijuga, 545 subsp. quadrijuga, 545, 549 rubra, 545, 549 schomburgkii, 545 subsp. javariensis, 546 subsp. schomburgkii, 546, 548 septentrionalis, 546 subsimplex, 545 surinamensis, 546 tetrapetala, 546 Trimeranthus, 294 Triopterys discolor, 181 Tripodanthus, 58 acutifolius, 58, 59 eugenioides, 59 Trompillo, 536, 744 Trompo de lamorado, 363 Truncaria, 512 caryophyllea, 515 Trymatococcus, 728 amazonicus, 729, 729 paraensis, 729 turbinatus, 729 Tschudya, 350 asperiuscula, 357 ibaguensis, 414 rufescens, 357 strigillosa, 353 Tuædu Pharatæ, 708 Tulasnea foliosa, 500 Tumba potro, 627 Tusilla, 702 —U— Uaira tombepe, 664 Ududu, 613 Uéi, 746 Uranthera dicranophora, 279

INDEX

Urena, 217 lobata, 217, 218 var. sinuata, 217 moenchii, 200 palmata, 200 sinuata, 217 Urodesmium, 486 huberioides, 486 Urostigma, 702 amazonicum, 706 angustifolium, 706 crocatum, 707 erythrostictum, 710 gardnerianum, 709 geminum, 710 involutum, 709 leucostictum, 709 mathewsii, 708 paraense, 709 schumacheri, 710 Uruichá, 32 Usibo akuanetete, 256 —V— Vachellia, 590 farnesiana, 591 Vainefá, 674 Valdesia rosea, 292 Valerioanthus, 749 Villarsia humboldtiana, 579 Virola, 742 boliviensis, 746 calophylla, 744 cuspidata, 744 var. membranacea, 744 duckei, 744 elliptica, 745 elongata, 744 var. longicuspis, 744 var. punctata, 744 elongata var. subcordata, 744 glaziovii, 746 incolor, 744 melinonii, 745 michelii, 745 minutiflora, 745 mocoa, 746 mycetis, 746

panamensis, 746 parvifolia, 745 pavonis, 745 peruviana var. tomentosa, 746 rufula, 746 rugulosa, 745 sebifera, 745 sp. A, 747 sp. B, 747 steyermarkii, 746, 747 surinamensis, 746 theiodora, 746 venezuelensis, 746 venosa var. poeppigii, 740 venosa, 740 warburgii, 746 Vomitivo, 546 Votomita, 527 orinocensis, 527, 528 ventuarensis, 528 —W— Wadasa, 740, 746 Wadimashu, 696 Wajuko, 709 Wajunuma, 628 Walleria laxiflora, 763 Wanawadudu, 709 Warada-kuma-tare, 661 Wãrãke duthæ, 696 Warusey-yek, 746 We yó, 728 Weigeltia, 751, 758 grandifolius, 760 longifolia, 761 potiaei, 761 quelchii, 762 surinamensis, 763 sylvatica, 760 Wichaurea, 4 Wishoguatemosi, 696 Wissadula, 217 amplissima, 219 contracta, 219 excelsior, 219 hernandioides, 219 patens, 219 periplocifolia, 218, 219 spicata, 189

803

zeylanica, 219 Wi-wi-hijú-deu, 739 —X— Xeracina, 281 —Y— Yagé, 93, 137 Yagua rosorina, 739 Yaque, 591 Yaragua, 622, 627 Yara-yara, 690 Yigüire, 670 Yoeru, 719 Yopo, 598, 745 Yopo-Ayuco, 744 Yunckeria, 750 Yuquito, 696 —Z— Zapoteca, 676 formosa, 677 formosa subsp. formosa, 677, 677 portoricensis, 678 subsp. flavida, 678, 678 Zarcillo, 661 Zarzaparilla, 649 Zunilia, 749 Zygia, 678 basijuga, 680, 682 cataractae, 680, 683, 684 claviflora, 680, 682 collina, 681, 683 divaricata, 680 glomerata, 680 inaequalis, 681, 684 latifolia, 681 var. communis, 681 var. lasiopus, 681, 685 var. latifolia, 686 palustris, 686 potaroënsis, 686 racemosa, 686 ramiflora, 686 unguis-cati, 668 unifoliolata, 685, 686